il CATBK N+ APRIL 29, 2005 * VOLUME 56 * NUMBERS 1-17 7 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES G Dallas Hanna (1887-1970) Diatomist, Geologist, and Invertebrate Paleontologist CALIFORNIA ACADEMYorSCIENCES FOUNDED 1853 SAN FRANCISCO, CALIFORNIA Copyright © 2005 by the California Academy of Sciences All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage or retrieval system, without permission in writing from the publisher. SCIENTIFIC PUBLICATIONS Alan E. Leviton, Ph.D., Editor Gary C. Williams, Ph.D., Associate Editor Michele L. Aldrich, Ph.D., Consulting Editor COVER IMAGE G Dallas Hanna, Diatomist and Invertebrate Paleontologist (1887-1970); Curator of Geology, California Academy of Sciences, 1919-1970 In the early 1920s, G Dallas Hanna established the Academy’s diatom collection, reintroducing a tradi- tion for the study of fossil and Recent marine and freshwater microorganisms begun with the founding of the California Academy of Sciences in the early 1850s. Hanna’s interest in diatoms began in 1911 during the period when, as an employee of the U.S. Bureau of Fisheries, he was assigned to carry out a fur seal census in the Pribilof Islands. Thus, Hanna, when he came to the Academy in 1919, was to carry on the tradition begun by Academy co-founder John Boardman Trask and Academy member Henry G. Hanks, a tradition he reenergized when he established the first laboratory for micropaleontology on the Pacific Coast. Today, the California Academy of Sciences houses one of the World’s premier diatom research col- lections, under the capable curatorship of Dr. J. Patrick Kociolek, and is a beehive of research activity dealing with Recent and fossil, freshwater and marine diatoms emphasizing systematics and environ- mental studies. ISSN 0068-547X The Proceedings of the California rbeademy of Sciences accepts manuscripts for publication in the Natural Sciences that stress systematics and evolutionary biology and paleontology, biogeography, and biodiversity, also relat- ed areas in the Earth Sciences, such as biostratigraphy, regional and global tectonics as they relate to biogeography, and paleoclimatology, and topics in astrobiology, anthropology, as well as the history of science as they relate to insti- tutions of natural history, to individuals, and to activities, such as expeditions and explorations, in the natural sciences. Priority consideration will be given to manuscripts submitted by Academy staff and others who are affiliated with the research programs of the institution and/or manuscripts that include reference in part or in whole to Academy research collections or to Academy-sponsored expeditions. Others who do not meet the above criteria may submit manuscripts for consideration for publication on a space-available basis. Manuscripts accepted for publication are subject to page charges; charges may be waived on a case-by-case basis. Nore: All manuscripts submitted for publication in any of the Academy's scientific publication series (Proceedings, Occasional Papers, Memoirs) are subject to peer review. Peer review includes both internal and exter- nal review, internal review by at least one Academy scientist whose interests parallel those of the submission, and external review, ordinarily by two individuals who are recognized experts in the field. Published by the California Academy of Sciences 875 Howard Street, San Francisco, California 94103 U.S.A. Printed in the United States of America by Allen Press Inc., Lawrence, Kansas 66044 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1, pp. 1-22, 6 figs., 1 table (Appendix) April 29, 2005 Redescriptions of the Nudibranch Genera Akiodoris Bergh, 1879 and Armodoris Minichev, 1972 (Suborder Doridacea), with a New Species of Akiodoris and a New Family Akiodorididae Sandra V. Millen! and Alexander Martynov2 ! Department of Zoology, University of British Columbia, 6270 University Blvd., Vancouver, B.C., Canada V6T 124, Email: millen@ zoology.ubc.ca; 2 Zoological Museum, Moscow State University, Bolshaya Nikitskaya Str. 6, Moscow, 103009, Russia, Email: martynov @ zmmu.msu.ru The type species of the genus Akiodoris, A. lutescens, Bergh, 1880 is redescribed from specimens found in the Sea of Okhotsk, and from the Commander Islands, Russia. A second species in the genus, Akiodoris salacia Millen sp.nov., is described from the Pacific waters off British Columbia, Canada. The type species of the genus Armodoris, A. antarctica Minichev, 1972 from the Davis Sea, is redescribed from the type and two additional specimens. The relationship of these genera with other gen- era of phanerobranch dorids is explored and a new family Akiodorididae is formed for the suctorian genera Akiodoris, Armodoris, Echinocorambe, Doridunculus, and Prodoridunculus. Key worbDs: Opisthobranchia, Akiodoris, Armodoris, dorid phylogeny, Akiodorididae. During the second half of 19th and early 20th centuries representatives of phanerobranch dorids of the genera Akiodoris, Prodoridunculus and Doridunculus, all with a remarkably similar radula, were discovered (G.O.Sars 1878; Bergh 1880; Odhner 1907; Thiele 1912). Bergh (1880) arranged the genus Akiodoris within the Suctoria, but Doridunculus was placed between the gen- era Goniodoris and Ancula. Following this arrangement, Fisher (1883-1887) placed the genus Doridunculus within the genus Goniodoris and the genus Akiodoris (as Aciodoris) within the genus Acanthodoris, in subgeneric ranks. Bergh (1883) created, for the first time, a phylogenetic scheme for the phanerobranch dorids where both genera Akiodoris and Doridunculus appeared at the base of Goniodorididae sensu latu as descendants of cryptobranch dorids. Later authors usually grouped the genera Akiodoris, Prodoridunculus and Doridunculus together into the base of the family Onchidorididae, probably considering them the most archaic forms (e.g., Thiele 1931; Odhner in Franc 1968), but Hgiseter (1986) arranged Doridunculus near the genus Lophodoris in the family Goniodorididae. In the second part of last century Minichev (1972) added the genus Armodoris as closely allied to Akiodoris. Millen (1987:19), in an abstract referring to a cladistic analysis of the family Onchidorididae, isolated the four genera mentioned above as a separate clade. Martynov (1997) suggested for these genera a new family, Akiodoridae, but he did not give a formal description for this taxon. The main goal of the present study is to revise the two principal genera, Akiodoris and Armodoris, and to explore their relationship with the families Goniodorididae and Onchidorididae. The genus Akiodoris was erected by Bergh (1879 a) for a phanerobranch nudibranch which has tripinnate gills, a sessile buccal pump, smooth lip cuticle, two cuspidate lateral teeth per side, D PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. | numerous rectangular marginal teeth, a penis armed with hooks and a vagina containing cilia coy- ered villi. The type species, Akiodoris lutescens Bergh, 1880 was found in the Alaskan Aleutians, and this species has not been reported since. One of the authors, Alexander Martynoyv, found a large number of specimens in several museums and research institutions in Russia, all from the Commander Islands and from the Sea of Okhotsk, mainly in the Kuril Islands. Bergh’s (1880) description was based on two specimens, one of which he considered a variety, although why he separated it is unclear. He never illustrated the external features and described them only briefly, although he described the anatomy in detail and included illustrations of some of the organs. A redescription of this species with emphasis on the morphology and additional illustrations of some of the anatomy is included in this paper. Martens (1879) emended the name to Aciodoris, replac- ing the Greek Akio (= sharp) with the Latin Acio. Because doris is a Greek name, we follow the recommendation of the Code that the components of a compound name should agree (Appendix D, II, 11) and revert to the spelling Akiodoris. Aciodoris is an unjustified emendation under the Code (art.32.5 and 33.2.3) and therefore a junor objective synonym of Akiodoris. The other author, Sandra Millen, found 6 specimens of a small white dorid which possesses the internal characters of Akiodoris, in the waters near Vancouver, British Columbia, Canada. These animals differ in a number of external features from Akiodoris lutescens and represent a new species belonging to the genus Akiodoris. This species is described. and the relationship of the genus to other members of the family is discussed. The genus Armodoris was created by Minichev (1972) for A. antarctica Minichev, 1972 from Tokarev Island in the Davis Sea. It has pear-shaped or conical, spiculose tubercles and a radula shape similar to that of Akiodoris, but with more teeth. There are up to six inner lateral teeth and up to eight outer lateral teeth per side. The specimen is 16 mm long and sexually mature; it is pos- sibly a mature specimen of Prodoridunculus gaussianus as only very small specimens of Prodoridunculus have been found. Without having a size range of specimens to examine, we shall retain them as separate taxa. Minichey (1972) separated Armodoris from Akiodoris because in Armodoris, the gonad surrounds the stomach, he observed a single seminal bursa, the spermathe- ca, on a long duct, and the vagina did not have a villous lining. In our observations of Akiodoris lutescens, we found that the stomach is anteriorly buried and posteriorly exposed so that the gonad- covered digestive glands surround it. The spermatheca of Akiodoris is short stalked, but has a tubu- lar shape similar to that of Armodoris antarctica. Akiodoris salacia sp. nov. lacks villi on the vagi- na. Thus, the taxonomical position of the genus Armodoris needs clarification. During a study of the collection of the Zoological Institute RAS (St. Petersburg), two addition specimens of Armodoris antarctica were found along with the dissected holotype of A. antarctica, which are used here for a redescription of this taxon. Subclass Opisthobranchia Order Nudibranchia Suborder Doridacea Superfamily Anadoridoidea Family Akiodorididae Millen and Martynov, fam. nov. TYPE GENUS: Akiodoris Bergh, 1879 Akiodorididae, nom. nudum: Martynoy, 1997:233. DIAGNOsIS. — The notum is spiculose, covered by rounded or elongate tubercles. Posterior part of the notum is round or transformed into two lobes. Branchial pocket is absent, gills are MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS — 3 arranged in semicircle with the exception of aberrant Echinocorambe where they are reduced to one simple leaf. Head has small four-corned oral veil. Anus dorsal, except for Echinocorambe where it is ventral. Buccal pump is a diverticulum of the dorsal surface of the buccal bulb. Peripheral muscle is absent and there are small, separate muscles at the anterior part of buccal pump. Lip disk thin and smooth. Radular formula is 2-14.2-6. 0-1. 2-6. 2-14. Central tooth is usu- ally present, ranging from a small plate to wide arch-shaped structure, sometimes with a central cusp. The first and following up to six lateral teeth have an irregular rectangular base and strong cusp directed downward and several strong denticles on one or both sides. Remaining laterals are in varying degrees of reduction, with a rectangular shape. Receptaculum seminis vaginal or insert- ed on the uterine duct. Stomach is entirely or partially free from the digestive gland. Prostate tubu- lar or enlarged. Penis with or without spines. The new family includes the following five genera: Akiodoris Bergh, 1879, Armodoris Minichey, 1972, Doridunculus G.O. Sars, 1878, Echinocorambe Valdés and Bouchet, 1998, and Prodoridunculus Thiele, 1912. Genus Akiodoris Bergh, 1879 TyPE SPECIES Akiodoris lutescens Bergh, 1880, by monotypy DIAGNOsIS.— The notum is spiculose, covered by rounded tubercles. Posterior part of the notum is round and fully covers the foot and tail. Gills are in a semicircle. Head has a small four- corned oral veil. Anus dorsal. Buccal pump is slightly prominent anteriorly. Radular formula is 8- 13.2.1.2.13-8. The central tooth is wide and arch-shaped. The first and second laterals have only 1—2 blunt denticles on both sides. Remaining laterals are in various degrees of reduction. Cerebral and pleural ganglia are fully fused. Stomach is very large and fully free from the digestive gland. Gonad does not cover most of the digestive gland and stomach. Prostate long, tubular. Vagina may have villi. Ejaculatory duct has simple or complex spines. The genus contains only the type species and the new species described in this paper. Redescription of Akiodoris lutescens Bergh, 1880 (Figs. 1-2) Akiodoris lutescens Bergh, 1879a:354—355 (nomen nudum). Akiodoris lutescens Bergh, 1879b:4-8 (plates only). Akiodoris lutescens Bergh, 1880:55—58. MATERIAL EXAMINED?.— HOLoryPeE: The type specimen cannot be found in the University of Copenhagen Museum (Kathe Jensen, pers. commun.). OTHER MATERIAL: | specimen, 18 September 1949, Shikotan Island, Kuril Islands, Russia, 13.5 m depth, collected by E.F. Gurjanova. 1 specimen, 10 September, 1966, 44°41’N, 148°57’E, 780 m., Segsbee trawl, R/V “Vityaz,” St. 5640. Collected by V.M. Koltun. 1 specimen, 31 July, 1971, Vasilieva Pt., Paramushir Is, Kuril Islands, Russia, 20 m depth, large stones, collected by P.G. Kraynyuk. 2 specimens, 5 August, 1970, Gilyak Pt., Paramushir Is, Kuril Islands, Russia, 20 m. depth, rock, collected by A.M. Murakhveri. 1 specimen, 10 August, 1970, Anciferova Is., Kuril Islands, Russia, 15 m depth, large stones, col- lected by A.M. Murakhveri. 1 specimen, 20 September, 1971, Gromky Pt., Iturup Is., Kuril Islands, Russia, collected by V.I. Lukin. 1 specimen, 20 September 1971, Gromky Pt., Iturup Is., Kuril Islands, Russia, 15 m depth, collected by V.I. Lukin. 1 specimen, 22 September 1971, Shutka Inlet, Iturup Is., Kuril Islands, Russia, 20 m depth, rock, collected by V.I. Lukin. 1 specimen, 14 July, ° Except as indicated, no specimen numbers are given inasmuch as most are in field station collections. 4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 1972, Korabel’naya Inlet, | km to sw of Korabel’ny Pt., Medny Is., Commander Islands, Russia, 20 m depth, collected by V.I. Lukin. 2 specimens, 5 August, 1972, Kozyrevskogo Pt., Paramushir Is., Kuril Islands, Russia, 15 m depth, stones, collected by V.N. Romanov. | specimen (juvenile), 5 September 1973, Peregrebnogo Pt., Beringa Is., Kuril Islands, Russia, 20-22 m depth, collected by B.I. Sirenko. 2 specimens, 17 September, 1973, Cherny Pt., Medny Is., Commander Islands, Russia, 10 m depth, rock, collected by B.I. Sirenko. 1 specimen, 31 July, 1978 Iona Is., Okhotsk Sea, Russia, 80 m depth, big and small stones, collected by V.A. Pavlyuchkov. 4 specimens, 11 July, 1985, Lopatka Pt., Kamchatka Peninsula, Russia, 17 m depth, rock and stones, collected by V.I. Shalukhanoy. | specimen, 10 July, 1985, Vladimira Pt., Atlasova Is., Kuril Islands, Russia, 25 m depth, rock. 1 specimen, ZMMU Lc-25738. 14 July, 1985, Sakhalinsky Bay, 54°57.5.N, 141°01.0.E, 95 m depth, R/V “G. Popov,” dredge 5, collected by Yu. I. Kantor. 1 specimen, 4 August, 1985, Okhotsk Sea, 55°26’8”N, 145°55’3”W, 160 m depth, stones, collected by A.V. Smirnov. | specimen, 20 Aug. 1986, Gladkovsky Pt., Medny Is., Commander Islands, Russia, 10-12 m depth, rock, collected by V.V. Oshurkov. | specimen, 25 August, 1986 Monati Pt., Beringa Is., Commander Islands, Russia, 25 m depth, rock, collected by V.V. Oshurkov. 1 specimen, 22 July 1991, Podutesnaya Inlet, Beringa Is., Commander Islands, Russia, 20 m depth, rock, collected by V.I. Shalukhanov. 2 specimens, 14 August 1991, Buyan Pt., Beringa Is., Commander Islands, Russia, 10 m depth, rock, collected by V.I. Shalukhanov. | specimen, 23 July 1992, Sivuchy Kamen’ Kekur Islet, Medny Is., Commander Islands, Russia, 20 m depth, rock, collected by K.E. Sanamyan. Specimens in Kamchatka’s Institute of Ecology and Environment, Petropavlovsk- Kamchatsky, Russia, the Institute of Marine Biology, Vladivostok, Russia, the Zoological Institute, Saint Petersburg, and in the Zoological Museum, Moscow State University, Russia. 1 specimen, 11 July, 1985, Lopatka Pt., Kamchatka Peninsula, Russia, 17 m depth, rock and stones, collected by V.I. Shalukhanov. EXTERNAL MORPHOLOGY.— The body is elongate-oval and reaches a maximum preserved length of 40 mm (Fig. 1A). The dimensions of one specimen are 40 x 25 x 13 mm. The mantle edge is 4-5 mm, barely covering the sides and tail in preserved animals, although the tail probably extends beyond the mantle in live animals. The notum is densely covered with differentially sized tubercles. The small tubercles, up to 0.4 mm in width, are conical or spindle shaped, the large tuber- cles, up to 1.4 mm in width, are rounded or cylindrical with round tips (Fig. 1B). Spicules lie in the notum radially around the tubercles, most of the spicules in the tubercles have been dissolved by formalin but traces remain, radiating loosely from the centre outward. The rhinophores have 10-15 lamellae and numerous spicules. The rhinophores contract into openings which do not have raised sheaths but are edged all around with 11—12 large tubercles. There are 10—17 tri- or quadripinnate gills which contract into depressions arranged in a broad horseshoe, which is broken posteriorly by a few tubercles. The gill size decreases towards the posterior, but sometimes a few small tripinnate gills are present anteriorly between the larger gills. Within the gill circlet are many large and some small tubercles, about 80 in total. The anal opening is not raised and is slightly left of centre towards the posterior end of the circlet. The color in life, according to Dall, is light yellow (Bergh 1880). Preserved specimens are dirty white or pinkish-tan. The head is veliform, not high, and slightly narrower than the foot. The mouth opening is round and the rounded tentacles are thicker on the anterior edge (Fig. 1C). The foot is broad and bilabiate anteriorly, posteriorly it extends into a small leaf-like tail, which is papillate on the dor- sal surface. ANATOMY.— The oral tube is short and muscular. The anterior half of the buccal bulb has a poorly developed, sessile, rounded buccal pump (sucking crop) with no median muscular band MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS — 5 FiGureE 1. Akiodoris lutescens, drawn with a camera lucida. A. Whole animal, dorsal view. B. Tubercles, one showing traces of spicules. C. Head and anterior of foot. D. Radula, one half row and rachidian tooth. A. adult, B. juvenile. E. Buccal bulb, A. lateral view B. dorsal view. F. Reproductive system. Key: al, albumen gland; am, ampulla; be, bursa copulatrix; hd, hermaphroditic duct: id, insemination duct; mu, mucous gland; p, penis; pr, prostate; rs, receptaculum seminis; v, vagina; vd, muscular vas deferens. (Figure 1E). The anterior part of the buccal pump has longitudinal muscular fibres, followed by a band of transverse fibres which extends down the sides of the buccal bulb. Posteriorly there are three longitudinal bands of muscles on each side of the esophagus. Ventrally there is a small pos- teriorly projecting radular sac. The round lip disk is muscular with a thick outer layer, smooth and yellowish in color. This outer layer is fleshy with only a thin cuticle coating. The radula has 52-69 rows of reddish brown teeth with the formula (8-13.2.1.2.13-8) (Figs. 1D, 2A—B). The central tooth 6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 FiGure 2. Akiodoris lutescens. Specimen from Lopatka Pt., tip of Kamchatka Peninsula, Russia, 11/7/85. A. SEM micro- graph of radula, scale bar = 400 um. B. SEM micrograph of central region of radula. Scale bar = 200 um. C. SEM micro- graph of entire penis. Scale bar = | mm. D. SEM micrograph of one posterior penial spine. Scale bar = 40 um. is wide and flat in the shape of a raised arch. Sometimes a small, posteriorly directed, chitinised denticle can be seen. The two lateral teeth per side can be seen in Fig. 2A—B. The first lateral teeth have a rectangular base and a short, wide, recurved cusp with one inner and one or two outer den- ticles. The second lateral teeth have a rectangular blade with a large quadrangular cusp on the inside and 5—6 outer denticles. The marginal teeth have rectangular bases and a small projection on the inner posterior corner. They diminish in size towards the outside, the last two being very small and narrow. The remainder of the digestive tract is as described by Bergh (1880), except that the ribbon shaped salivary glands are shorter, ending at the digestive gland. The large blood glands are pinkish, not white, in color. The central nervous system is as described by Bergh (1880). The reproductive system is shown in Figure 1F. Bergh (1880) described this system well although he only illustrated portions of it. The vagina is lined by villi, 0.6—0.8 mm long, with a dense axis and non glandular, ciliated epithelium. The penis in Bergh’s specimen was retracted, the glans only partly everted. A fully everted penis is shown in Figure 2C and a closeup view of a pos- terior penial spine in Figure 2D. The reproductive openings are located on the right side about one third of the body’s length from the anterior mantle edge. The surface around the genital openings is covered with numerous soft villi approximately 0.3 mm in length. NATURAL History.— The known range is expanded from its original localities of Atka Island and Kyska Island in the Aleutians, westward and northward to the Commander Islands, southward along the Kamchatka peninsula and down the chain of Kuril Islands to Shikotan near Hokkaido and into the Sea of Okhotsk to Iona Island near its centre and Sakhalinskiy Bay in the west. It has been MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS — 7 collected from low water line to depths of 780 m, usually between 10 and 25 m in the months of July, August and September. The only juvenile was collected in September. All specimens have been collected from rocky areas. Akiodoris salacia Millen, sp. nov. (Figs. 3-4) Akiodoris sp. 1 Behrens, 1991:51, #68 EtTyMoLoGy.— Named for the Roman sea-goddess Salacia, wife of Neptune. MATERIAL.— HovoryPe: California Academy of Sciences, CASIZ 110807, 14 September 1985, Tyee Point, Copper Cove, British Columbia, Canada (49°22’8”N, 123°16’5”W), 20 m depth, on hydroids, rocky substrate, collected by S. Millen. PARATYpPES: Royal British Columbia Provincial Museum, RBCPM 005-000013-001, 1 specimen, collected with the holotype. Zoologi- cal Museum of Moscow University, ZMMU, LC-25737, | specimen, collected with the holotype. EXTERNAL MORPHOLOGY.— Preserved specimens of this small dorid range in length from 4 to 6.5 mm. The largest animal measures 6.5 x 3 x 2.5 mm (I X w x h). The body shape (Fig. 3A) is elongate-oval, wider in front than behind, with a slightly trailing tail. The mantle margin is fairly wide, covering the high sides and head but not the tail. Posteriorly, above the tail, the marginal edge is indented. The notum bears elongate, slightly inflated, spear shaped tubercles with pointed tips (Fig. 4A). There is little variation in tubercle size, which ranges from 0.08 to 0.4 mm in height and 0.06 to 0.18 mm in width. The numerous tubercles are spaced evenly over the notal surface, with a few small ones in the mid-dorsal area and some medium sized ones near the mantle edge. Spicules are found in the lower “4 of each tubercle, and their tips protrude slightly. The pro- truding spicules are rod-like, slimmer than those forming the central shaft inside each tubercle. At the bases of the tubercles, spicules extend in a radial, star-like pattern which can be seen through the notum. Within the notum there are scattered spicules forming a mesh-like network which forms a radial pattern towards the mantle margin. This pattern is only visible on the ventral surface of the mantle margin. There are scattered quadrate spicules stiffening the sides of the body. In the foot, spicules form a mesh pattern on the sole, and the dorsal surface of the foot flange and the tail are heavily spiculated. Numerous sharp pointed spicules protrude in these areas, espe- cially on the tail. The slightly raised margins of the rhinophores each bear 4—5 tubercles, the largest being the innermost anterior one. There are no tubercles within the branchial arch. The rhinophores are sturdy and end in a rectangular, blunt tip. The stalk is short and most of the clavus bears wide, thick lamellae supported by spicules. The 8—10 lamellae are attached along a vertical, anterior line which appears inset due to the sudden indentation of the wider lamellae. The lamellae slope ventrally and become narrower towards the posterior where they meet forming a chevron. The 4-6 contractile gills are bipinnate and incompletely tripinnate, non-retractable, separated on the outside, but joined at their bases towards the inside. They are arranged in a semi-circle which is open posteriorly. The gills are largest anteriorly and directed towards the posterior. The gills are surrounded by tubercles, but none are within the small semicircle. The anus 1s at the posterior end of the gill arch. Living specimens (Fig. 3A) are white or faintly yellow in color. There are sometimes a few opaque white specks in the dorsum along the sides and in front of the rhinophores. In immature animals the central area appears pinkish due to the underlying color of the reddish-brown digestive gland. In more mature animals, the digestive gland is covered by creamy-yellow gonads giving the central area a peach hue. Ventrally the digestive gland shows only faintly through the foot. The 8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 FiGureE 3. Akiodoris salacia sp. nov. Specimen from Tyee Pt., near Vancouver, British Columbia, Canada, 14/9/85. A. Photograph of a live specimen of A. salacia, 4.6 mm in length. B. SEM micrograph of the radula. Scale bar = 20 tum. C. SEM micrograph of the lateral teeth. Scale bar = 10 um. D. SEM micrograph of the penis. Scale bar = 40 Um. anterior edge of the foot and the head are sometimes pale yellow. The rhinophores are pale yellow and the gills are pale yellow or white. An unusual feature of this species is a series of opaque white, round, granular glands along the overhanging posterior border of the mantle. There are 8—10 of these glands, the smallest being located on the midline. The head (Fig. 4B) is composed of two triangular lobes which end in flap-like tentacles sepa- rated from the rest of the head by a dorsal groove. The foot is narrow and elongate, wider and trun- cate anteriorly. The anterior edge is bilabiate. The triangular tail protrudes up to 0.5 mm. It is thick but does not have a dorsal keel. Noteworthy are its many, posteriorly projecting spicules. ANATOMY.— The oral tube is a muscular ring with short labial glands surrounding it. The buc- cal bulb (Fig. 4C) has a dorsal, rounded, sucking crop which is sessile, with no median muscular band and weak peripheral muscles. The radular sac is short and protrudes from a wide, muscular odontophore. The round lip disk is thinly cuticularized and smooth. The radula has 35—39 tooth rows. The radula formula is 3-5.2.1.2.5-3 (Figs. 3B—C, 4D). There is no well-developed rachidian tooth but the central area has a thickened, raised plate which begins at the fourth or fifth tooth row. The plate is 3—6 um long. The innermost lateral tooth has a rectan- gular base and a large, recurved, pointed cusp. It has one inner and two or three outer, well devel- oped denticles. These laterals range in width from 12.5—24 um and 13-19 um in length. The outer lateral teeth have a rectangular base with a large, recurved hook on the inside and three or four well developed outer denticles. The outer laterals are 18—19 um wide and 12—18 um long. Each mar- ginal tooth is a flat, rectangular plate with slightly rounded outer corners and a small spine on the inner posterior corner. The marginal teeth diminish in width towards the outside. The marginal MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS 9 Ficure 4. Akiodoris salacia, drawn with a camera lucida. A. Tubercles, one showing internal spicules. B. Head and ante- rior of foot. C. Buccal bulb. A. dorsal view, B lateral view. D. Radula. One half row including rachidian plate. E. Reproductive system. Key: al, albumen gland; am, ampulla; be, bursa copulatrix; hd, hermaphroditic duct; id, insemination duct; mu, mucous gland; p, penis; pr, prostate; rs, receptaculum seminis; v, vagina; vd, muscular vas deferens. 10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 teeth range in width from 6-19 um and in length from 6-10 um. The salivary glands are small, flattened, elongate-oval masses lying under the buccal bulb and attaching to it at the anterior end of the esophagus (Fig. 4C). The esophagus is a narrow tube which becomes slightly wider and softer before entering the digestive glands. The digestive glands appear as one elongate mass, wider in front than behind and truncated on the anterior right side. The stom- ach is sac-shaped, almost entirely separated from the digestive glands. It has a small, round cae- cum on the right border of the anterior part of the stomach, tucked in the loop of the intestine. The intestine is broad and runs to the right of the midline curving inward to the anus. The anal opening is not raised. The pericardial sac contains a posterior, thin-walled, triangular auricle and a smaller, more muscular, diamond shaped ventricle. The flattened, compact blood glands are located just posteri- or to the central nervous system. The fused cerebro-pleurals are oval and joined by a short commissure. The olfactory nerves have bulbous bases with small attachment points. The large, black eyes have very short stalks. The oval pedal ganglia, smaller than the cerebro-pleurals, are located directly below them and are con- nected to them by a short commissure. They are connected together by a commissure which is slightly longer than the cerebral one. The small oval, paired buccal ganglia adjoin each other. The reproductive system is illustrated in Figure 4E. It lies on the right side with the loop of the vas deferens extending from the region of the mouth to a short distance in front of the gills. The ovotestis consists of oval, creamy-yellow lobules lying on the dorsal and lateral surfaces of the digestive glands. The branched gonoducts are broad bands which unite to form one central pre- ampullar duct. This duct widens slightly into a tubular ampulla. There is a short post-ampullar duct which bifurcates into an oviduct and a long vas deferens. A short distance from its origin, the vas deferens becomes prostatic, loops posteriorly towards the ovotestis, encircles the bursa copulatrix and continues anteriorly. It then becomes non-prostatic, loops back on itself and bends downward. The long, tubular, non-prostatic portion enlarges slightly to form a penal bulb containing the uncoiled vas deferens. The penis joins a common atrium with the vagina. It is a cylindrical papil- la. Inside, the distal portion of the vas deferens is weakly chitinised and has small spines, 5-8 um in length. The penis (Fig. 3D) is eversible, but the spines were not everted on any of the specimens. The vagina is long and cylindrical, wider near its opening posterior to the penis in the com- mon atrium, and gradually narrowing. It loops back on itself and then gives off a short duct to the large, round bursa copulatrix. The insemination duct, after a short distance, is joined by a short duct from the tubular receptaculum seminis. The insemination duct is long and winding. It terminates at the oviduct near its junction with the post ampullary duct and entrance to the female gland mass. The female gland mass has an oval, yellow albumen gland. The hermaphroditic ampulla runs down its anterior surface.Surrounding it, anteriorly, posteriorly, and on the inner side is a white mucous gland. The separate oviductal opening is ventral to the common genital atrium. The two reproductive openings are located high on the right side about '/ of the body’s length from the ante- rior mantle edge. NATURAL History.— The specimens described in this paper were found in September at a depth of 20 m. on a rocky bottom swept by moderate currents. They were all found on an old hydroid colony intermingled with the ctenostome bryozoan Farrella elongata, which is presumed to be their food. The larger specimens had partially mature reproductive systems and were copu- lating, but no spawn was present or laid in the laboratory during the week subsequent to their cap- ture. REMARKS.— The newly described species is placed in the genus Akiodoris Bergh, 1879 because it has a nearly identical reproductive system, bi- and tripinnate gills, smooth lip disk, ses- MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS 11 sile pharyngeal bulb, stomach which is free from the digestive glands, digestive caecum, and armed penis. However, in the presence of a slight posterior mantle reduction and gills arranged in a small, non-tuberculose semi-circle with a posterior anus, A. salacia is more similar to the genus Doridunculus. We considered this similarity as superficial due to the small size of A. salacia and relative frequency of appearance of a pair of lobe-like structures in the posterior part of the notum in phanerobranch dorids, apparently caused by partially delayed ontogenetic processes in notum formation. There are a number of differences between Akiodoris lutescens and A. salacia. Externally, the tubercles differ in that A. salacia has long, similar sized, lanceolate tubercles with projecting spicules, whereas A./utescens has smaller, conical and rounded tubercles of varying sizes. In A. salacia, the gill opening is in a small semi-circle with no enclosed tubercles and the anus is at the posterior edge. In A./utescens, the gill pocket forms a large horseshoe enclosing many tubercles and the anus. The posterior mantle margin is slightly indented in the new species, but this does not appear to be the case in A. /utescens. Both species have a flat leaf like tail which is dorsally spic- ulated. The distinctive mantle glands of A. salacia can only be seen in live specimens, so their pres- ence cannot be used as a distinguishing feature. Only preserved Akidoris lutescens have been observed. Internally, the radula has fewer (4-5) marginal teeth than a similar sized (6 mm) juvenile A. lutescens, which had 7—9 marginal teeth (Fig.1D). The thickened cuticular central tooth observed in specimens of A. salacia, is not well developed, but it may develop in older specimens. Only in larger A. lutescens does the central cuticle form a chitinised spine on the rachidian tooth. The central nervous system of the two species varies in that the eyes are large and sessile in A. sala- cia and smaller and on long stalks in A. /utescens and the unusual lateral position of the pedal gan- glia found in A. lutescens was not found in A. salacia. Genus Armodoris Minichevy, 1972 TYPE SPECIES Armodoris antarctica Minichey, 1972, by monotypy DIAGNOsIs.— The notum is spiculose, with rounded tubercles. Posterior part of the notum is round and fully covers the tail. Gills are in a semicircle. Head has a small four-corned oral veil. Anus dorsal. Buccal pump is oval and prominent. Radular formula is (4-8.4-6.1.4-6.4-8). The cen- tral tooth is a square plate of moderate size with a slightly prominent central cusp. The lateral teeth have a rectangular base and one long cusp to the inside of center. Denticles are found on the first 6 teeth, the outer 4-8 have one reduced cusp or are reduced to rectangular bases. Cerebral and pleu- ral ganglia are fully separated. Only a small dorsal part of the stomach is free from the digestive gland. Gonad covers most of the digestive gland and stomach. Prostate massive. Interior of vagina is simple. Ejaculatory duct has simple spines. The genus contains only the type species. Redescription of Armodoris antarctica Minichev, 1972 (Figs. 5-6) Armodoris antarctica Minichev, 1972:366—368, fig 4; Cattaneo-Vietti et. al., 2000:175. TYPE MATERIAL.— HOobortyPe: ZIN N 14 (dissected, most internal organs are removed and are 4 ZIN — The Zoological Institute, Russian Academy of Sciences, St. Petersburg; the numbers refer to systematic cat- alogue, without inventory numbers in this case. 5 There are no exact data in the first description of Armodoris antarctica, nor on the label that accompanies the holo- type. except for Minichev’s personal label “19-2”. The present date, depth and substrata is reconstructed by comparison with neighboring labels “19-1” and “19-3”, which both have complete or semi-complete data. 1, PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 not traceable, radula slide not found), around January 19, 19665, 11 Soviet Antarctic Expedition, Davis Sea, Tokarev Id., Between Samples 45 and 55, Depth about 30—32 m, rock, collected by A. F. Pushkin. OTHER MATERIAL: 2 specimens (ZIN N 2%, N 34), Zoological Institute, Russian Academy of Sciences, February 1, 1969, 13 Soviet Antarctic Expedition South Shetland Ids., King-George Id., La Guardia Nacional Bay, transect I, St. XXI, sample 60, 36 m depth, stones, col- lected by A. F. Pushkin. EXTERNAL MORPHOLOGY.— Preserved specimens range in length from 13 to 16 mm. The holotype measures 16 x 9 x 5 mm (1 X w x h) (Fig. 5 A-C). The specimen, ZIN N 3, is 13 x 7.4 (1 x w). The body shape is elongate-oval, slightly wider in front than behind. The mantle margin is amply wide, fully covering the sides and tail. Posteriorly, the marginal edge is without any traces of an indent. The notum bears tubercles of different sizes, scattered small tubercles between the larger ones. In the middle of notum, longitudinally, there are some rather large, pear-shaped tuber- cles. They become more elongate or cylindrical toward to the notal margins. Tubercles sizes range from 0.07 to | mm in height and 0.06—0.7 mm in diameter. Spicules within a tubercle are rod-like, varying in size, nearly straight or slightly crooked. The spicules sometimes slightly extruded through upper part of the tubercle. The margins of the rhinophore pockets are very low and bear 6 blunt tubercles (right side of the specimen ZIN N 3) and 4 in the holotype. The rhinophores have about 10 lamellae. Six (ZIN N 3) to nine (holotype) contractile, but non-retractile, gills are unipinnate and incom- pletely bipinnate (Fig. 5D). They are arranged in a semi-circle, which is posteriorly completed by a large tubercle. The gills are surrounded by elongate tubercles. The holotype and ZIN N 2 speci- men have three tubercles inside the gill semicircle, whereas example ZIN N 3 has none. The anus is in front of the posterior tubercle in the 13-mm length specimen. Preserved specimens are whitish-pinkish or yellowish. The head is a small semi-oval oral veil with two short blunt lobes extending posterio-laterally (Fig. 6C) The foot is rather wide, anteriorly truncated without any traces of post oral lobes. Posteriorly, the foot narrows slightly to form a flat, blunt tail (Fig. 6C). ANATOMY.— The oral tube is short and muscular. Almost the entire upper part of the buccal bulb has a well-developed, oval buccal pump (Fig. 6E). The buccal pump has no peripheral mus- cle (median muscular band). The anterior part of the buccal pump has longitudinal muscular fibers, followed by a band of transverse fibers, which extends down the sides of the buccal bulb. The round lip disk is smooth, covered by a colourless cuticle. Radular formula is 36—52 (4-8.4-6.1.4- 6.4-8) (Fig. 6D). The central tooth is a thin square plate of moderate size with a slightly prominent central cusp. The inner laterals are denticulate, and they gradually transform to the simple small plates of the outermost laterals. The first and at least 4-5 following laterals have a rectangular base and short or moderately long cusp (cusp of 2"4 and 3" laterals, in general, stronger than first ones) directed downward, usually straight or slightly curved. The first three or four inner laterals have 1—3 denticles on both sides of the cusp, the following 3-5 laterals have a serration (or it is reduced) on the outer side of the cusp. The remaining two to four laterals have a greatly reduced cusp, devoid of denticles, and the outermost tooth is just a simple plate. The elongate-oval salivary glands are small and attach to the buccal bulb at the anterior end of the esophagus (Fig. 6E). The latter is a rather wide, flattened tube. The stomach is sac-shaped, only a small dorsal portion is separate from the digestive gland (Fig. 6B). There is no caecum on the stomach. The pericardial sac contains a posterior, thin walled narrow-triangular auricle and smaller, oval shaped ventricle (Fig. 6B). The blood glands are slightly visible and located posterior to the cen- tral nervous system. The cerebral and pleural ganglia (Fig. 6B, 6F) are separated and the later are smaller. The MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIJODORIS AND ARMODORIS 13 FIGURE 5. Armodoris antarctica, holotype. A. dorsal view. B. Ventral view. C. lateral view. D. Close up of the gills and tubercles. large, black eyes have very short stalks. The pedal ganglia, similar in size to the cerebral, lie below them and are connected to them by short commissures. The rhinophoral ganglia are roundish. The buccal ganglia are roundish-oval. Gastro-esophageal ganglia were not found. In the cerebrals, three pairs of cerebral nerves were detected, in the pleurals, four, and in the pedals, three. The ovotestis is characterised by large female follicles (Fig. 6B). The ampulla is voluminous 14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 (9, eT a NY i Kv) werceprc /<* : ——- MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIJODORIS AND ARMODORIS 15 and kidney-shaped (Fig. 6G). There is a short post-ampullar duct, which bifurcates into an oviduct and a long vas deferens. The pre-prostatic duct is relatively long and wide. The prostatic part of vas deferens is a very massive tangle of a few thick, flattened loops. The prostate doesn’t encircle the bursa copulatrix. The vas deferens narrows rapidly to a non-prostatic portion, loops back on itself, and bends downward. The deferent duct is a thin, muscular, not very long, gradually enlarging duct, which widens to an elongate penial bulb. The penial bulb joins a common atrium with the vagina. The ejaculatory duct of vas deferens has inside tiny, densely placed, spines. A partially everted penis is very narrow. The vagina is rather short and wide, similar in diameter its whole length, bending in the middle and entering the large, irregularly-triangular, bursa copulatrix. Villi inside the vagina are absent. The receptaculum seminis is oval and has a long, muscular, gradually widen- ing duct, which joins the middle of the insemination duct. In the holotype, the receptaculum sem- inis is more elongate. The insemination duct is long, leaving the vagina near the middle and enter- ing the female gland mass near the oviduct. REMARKS.— The genus Armodoris has at least four to six well developed inside lateral teeth and no clear border between inner and outer laterals, whereas the genera Akiodoris, Prodoridunculus and Doridunculus have two well-differentiated inner laterals and rest of the later- als demonstrate reduction. Echinocorambe has three inner laterals, markedly different from the outer teeth. The radulas of A. lutescens and A. salacia are similar in the shape of the two first lat- erals and in the reduction of most of the outer laterals. This is a parallel case with the genus Adalaria, where the first lateral is more or less stable, but outer laterals have tended to reduce. The stomachs of Akiodoris and Doridunculus are massive and mostly free from the digestive gland, whereas Armodoris has a rather small free part of the stomach. Armodoris has large female folli- cles, which may be due to direct embryonic development, but the stomach is not fully surrounded by the gonad, as erroneously reported by Minichev (1972). The ampulla of Armodoris is large, more like that of Doridunculus echinulatus. Both Akiodoris species have similar long, but tubular prostates, whereas the prostate of Armodoris is massive. The receptaculum seminis is very narrow and tube-shaped in both Akiodoris and Doridunculus, but in the King-George’s examples of Armodoris it is oval and in the holotype it is narrow-oval. The villi of the vaginal duct are not pres- ent in Armodoris. The buccal pump in Armodoris antarctica is more developed, although it has the same structure as found in Akiodoris, Doridunculus and Echinocorambe. Due to above differences between Armodoris and the other four genera with similar radular teeth, it is suggested that we maintain the genus Armodoris. Compared to the King George Island examples, the dissected holotype has well preserved gills and oral veil. Minichev (1972) incorrectly interpreted the reproductive system and partially the digestive system of A. antarctica. He found only one receptaculum (“spermatheca” in his terms) and although he noted that “a “spermatocyst’ (= bursa copulatrix) is not evident in the present mate- rial”, nevertheless, he considered this case to be a “unique feature” (Minichev 1972:366). The holo- type originated from the Davis Sea, and the two additional specimens originated from another cor- ner of Antarctic, the South Shetland Islands. An additional specimen of A. antarctica is the “Italian FiGuRE 6. Armodoris antarctica drawn with a camera lucida. A. Dorsal view, scale bar = 1 mm. B. Dissected view, scale bar = | mm. Key: bb, buccal bulb; be, bursa copulatrix; cns, central nervous system; fgm, female gland mass; g, gonad; h, half row including rachidian tooth. E. Buccal bulb, Dorsal and lateral views. Scale bar = 0.5 mm. F. Central nervous sys- tem. Scale bar = 0.5 mm. Key: C 1-3, cerebral nerves; Cg, cerebral ganglia; P 1-3, pedal nerves; PC, pedal connective; PPC, pleuro-pedal connective; Pg, pedal ganglia; Pl 1-4, pleural nerves; Plg, pleural ganglia; R, rhinophoral nerve; Rhg, rhinophore ganglia: St, Statocyst; VL, visceral loop. G. Reproductive system. Scale bar= 1 mm. Key: am, ampulla; bc, bursa copulatrix: fgm, female gland mass; id, insemination duct; ps, penis sheath; pr, prostate; rs, receptaculum seminis; v, vagi- na: vd, muscular vas deferens. 16 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 sample” from the stomach of a fish, Trematomus bernacchii, caught in the Ross Sea, Terra Nova Bay (Cattaneo-Vietti 2000). By the courtesy of Stefano Schiaparelli, we have studied a SEM image of the radula of the Italian specimen of A. antarctica. The radulas of all four specimens of A. antarctica are similar. The rarity of Armodoris antarctica is caused by its inhabiting only shallow waters on rocky substrates, and it can only be found by diving collections, which is certainly rare in the Antarctic, especially around 30 m (both holotype and two additional samples were collected slightly below 30 m on rocky substrates and in remote localities). PHYLOGENETIC ANALYSIS Dorid nudibranchs are currently divided into two groups, those with gills and rhinophores which retract into a closable pocket, the Eudoridoidea (=Cryptobranchia), and those whose gills and rhinophores have no pocket or an open pocket, the Anadoridoidea (= Phanerobranchia). The phanerobrachs dorids were in turn subdivided by Bergh (1892) into two groups, those with a suc- torial buccal apparatus (suctorians) and those without (non-suctorians). These are informal groups, but the three families that Bergh placed into the suctorians, Corambidae, Goniodorididae, and Onchidorididae, are commonly arranged together. There have been numerous rearrangements with- in these three families over the years, but currently the family Onchidorididae is believed to con- tain the genera Acanthodoris, Adalaria, Akiodoris, Arctadalaria, Armodoris, Calycidoris, Diaphorodoris, Doridunculus, Onchidoris and Prodoridunculus, although Wagele and Willan (2000) consider the genera Akiodoris and Armodoris to belong to the Goniodorididae following Bergh (1883), who placed them at the base of the Goniodorididae. Four of these genera form a distinct clade based primarily on their radular characteristics (Millen 1987). Most genera in the family Onchidoridae have on each side a single, large, strongly hooked lateral with a triangular base and narrow, hooked, marginal teeth. The rachidian, where present, has an elongate, rectangular shape. In contrast, a clade of four genera, Akiodoris, Armodoris, Doridunculus, and Prodoridunculus, have two or three (sometimes up to 6), wide, rec- tangular lateral teeth per side. The first lateral has internal and external denticles, the second and third, when present, have external denticles. The rachidian tooth, if present, is arch-shaped. The marginal teeth are wide and rectangular with a small hook. These four genera also have a relative- ly simple buccal pump. Recent cladograms using either molecular or morphological data, which have included a vari- ety of cryptobranch and phanerobranch dorids, have suggested that the phanerobranch dorids may, in fact, be polyphyletic (Wollscheid and Wiagele 1999; Wagele and Willan 2000; Wollscheid- Lengeling et al. 2001; Vallés et al. 2001; Valdes 2002). These studies have not shown suctorian dorids to be polyphyletic, nor have they been able to use the full range of genera in the family Onchidoridae. The genera used were those with hook-shaped teeth, Acanthodoris, Adalaria, Calycidoris, Diaphorodoris, and Onchidoris, which generally clustered together. To observe the characters which distinguish Akiodoris and Armodoris from other genera and to establish their closest relationships, and to which family they should be assigned, we produced a cladogram using all of the possible Onchidorididae genera. These were compared to sister taxa consisting of three genera from the Corambidae (Corambe, Loy and Echinocorambe) and one Goniodorididae (Goniodoris). One cryptobranch dorid, family Chromodorididae (Cadlina), was used as a sister taxa to the phanerobranchs. The outgroup and basal rooting was based on the basal Anthobranchiate dorid genus Bathydoris, using B. spiralis Valdés, 2002, which is closest to the cryptobranch and phanerobranch dorids (fide Valdés 2002). In total, 15 taxa and 29 characters were analysed using MacClade and PAUP 3.1.1. MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS_ 17 The 29 characters used in this analysis are listed below: 23 characters are binary and 6 are mul- tistate. Character states are indicated by numbers; 0 is the plesiomorphic condition and 1—2 are apo- morphic. Non-applicable and unknown characters are coded with a ?. Polarities are the result of outgroup comparison. Table | (see Appendix) shows the distribution of plesiomorphic and apomor- phic states. 1. Mantle rim: The mantle rim (projecting edge) 1s narrow, partially absent in Bathydoris and some phanerobranch dorids (0) whereas it covers the sides and foot in most dorids (1). 2. Posterior mantle rim: The posterior mantle rim, where distinct, is usually entire in Bathydoris and most dorids (0). It is bilobed in some phanerobranch dorids (1). 3. Tail: The tail is extended in Bathydoris and some phanerobranch dorids (Q). It is small and covered by the mantle in most dorids (1). 4. Ridge on tail: The tail is usually smooth in Bathydoris, cryptobranch dorids and most phanerobranch dorids (0). It has a mid-dorsal crest or ridge in some phanerobranch dorids (1). 5. Notal spicules: Some Bathydoris and most dorid nudibranchs have integumentary spicules in the notum (Q). These are absent in a few dorid species (1). 6. Notal sculpture: Sculpture in the form of deciduous villi is present in Bathydoris (0); most species of dorid nudibranchs have well attached tubercles (1). Only a few species are smooth (2). 7. Head: The head in Bathydoris and most phanerobrach dorids is veliform (0). Cryptobranch dorids have a round head and separate tentacles (1). 8. Rhinophore sheath: Bathydoris and some phanerobranch dorids have rhinophores which lack sheaths and are attached directly to the dorsum (0). Other phanerobranch dorids have non-con- tractile sheaths into which the rhinophores can retract (1). Cryptobranch dorids have contractile rhinophore sheaths which close over retracted rhinophores (2). 9. Rhinophores: Bathydoris and most other dorids have lamellate rhinophores (0). A few dorids have smooth rhinophores (1) which may have vertical envelopes (2). 10. Gill protection: Bathydoris and some phanerobranch dorids have gills which lack sheaths and are inserted directly on the notum (0). Some phanerobranch dorids have sheaths into which the gills can retract but which do not close over the gills (1). Cryptobranch dorids have contractile gill sheaths which close over the retracted gills (2). For taxa with no mantle gills, this character has been treated as non-applicable. 11. Gill number: Bathydoris and most dorids have more than five gills (0). A few species of phanerobranch dorids have less than five gills which is considered a reduction (1). 12. Gill location: Bathydoris and most dorids have the gills located on the mantle some dis- tance from the edge (0). Some phanerobranch dorids have the gills posterior at the mantle edge or below it (1). 13. Notal sculpture within the gill circlet: Bathydoris and cryptobranch dorids do not have papillae or tubercles within the area encircled by the gills (0). Some phanerobranch dorids have notal sculpture in this area (1). For taxa with no mantle gills, this character has been treated as non- applicable. 14. Anus location: Bathydoris and most dorids have the anus located on the mantle some dis- tance from the edge (0). Some phanerobranch dorids have the anus posterior at the mantle edge or below it (1). 15. Lip disk: Bathydoris has thick chitinous labial armature and other dorids may have vari- ous rods or papillae to strengthen the lip disk (0); a smooth, thin cuticle is considered apomorphic (1). 16. Buccal pump: A sucking pump on the pharynx is absent in Bathydoris, non-suctorian 18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 1 phanerobranch dorids and cryptobranch dorids (0). This pump distinguishes the suctorian phaner- obranch dorids (1). 17. Peripheral muscles: A median longitudinal band of muscles is not found on the phanynx of Bathydoris and most dorids (0). This muscle is found on some of the suctorian phanerobranch dorids (1). 18. Rachidian tooth: A rachidian tooth is present in Bathydoris and some dorids (0). Many dorids have lost this central tooth (1). 19. Number of teeth per half row: Primitive dorids and Bathydoris have many teeth, >10, per half row (0). Reduction has resulted in narrower radulas with fewer than 10 lateral teeth (1). 20. Number of inner laterals: Inner lateral teeth tend to be more highly developed than the outer lateral (marginal) teeth which are often reduced. In Bathydoris, and some dorids, only the first lateral tooth differs from the outer lateral teeth (0). In other dorids there are 2 or more teeth which can be differentiated from the outer laterals (1). 21. Shape of innermost lateral tooth: Bathydoris, some cryptobranch and some phanero- branch dorids have cuspidate innermost lateral teeth. The cusp is towards the center of a broad tooth and denticles may appear on one or both sides (0). Other, phanerobranch dorids have beak- like teeth, with a long cusp on the outer edge, and denticles, when present, only on the inner side (1). 22. Denticulation on innermost lateral tooth: Bathydoris has denticulations on both sides of the inner lateral teeth, as do some dorids (0). Other dorids, used here, may have denticulations only on the inner side (1) or they may be absent (2). 23. Marginal teeth or outer lateral teeth: In Bathydoris and many cryptobranch dorids, these are elongate, hamate teeth, with or without denticles on the outer side (0). In some phanerobranch dorids they are reduced to either an elongate-oval plate (1) or a wide, rectangular plate (2). 24. Ampulla: The hermaphroditic ampulla is narrow and undifferentiated in Bathydoris and many dorids (Q). It is wide and usually short in some dorids (1). 25. Penial spines: Spines on the eversible vas deferens of the penis are absent in Bathydoris and some dorids (0). Small spines are present in a number of dorids and their presence 1s consid- ered an apomorphy (1). 26. Seminal receptacle: This sac for sperm storage is absent in most Bathydoris species, but present in Bathydoris spiralis as a small sac with one duct (stalked) (0). In some dorids, it has two ducts (unstalked), which is considered derived (1). 27. Seminal receptacle insertion: This sac is inserted by its duct on the vagina in Bathydoris spiralis and many other dorids (0). In some dorids, it is inserted part way along the insemination (uterine) duct (1). 28. Oviduct: The oviduct carrying the eggs usually passes directly into the female gland mass (O). In a few dorids, it enters the seminal receptacle before entering the female gland mass, which is considered apomorphic (1). 29. Cerebro-pleural ganglia: These ganglia are separate in Bathydoris and a few dorids (0). They are fused in most dorids (1). PHYLOGENETIC RESULTS The data matrix results in one tree (Fig. 7), 64 steps long, with a consistency index of 0.562 and a retention index of 0.692. The Bremmer support index values (values to the right of the line in Fig. 7) show that most of the branches are poorly supported, with the exception of the clade con- taining those phanerobranch dorids with beak-shaped laterals (value 2) and the node containing Corambe and Loy (value 2+). MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS_ 19 Bathydoris Cadlina Goniodoris Diaphorodoris Calycidoris Acanthodoris Adataria Onchidoris Akiodoris Armodoris Doridunculus Echinocoerambe Prodoridunculus wo loo vo fe a _ —) = — ~~ a in WN is oe Mim _— + 2 3 18 5 18 19 24 A 26 ; 24 20 13 27 16 Figure 7. Strict consensus tree of a preliminary phylogeny. Apomorphies are indicated by character numbers included in the analysis. Characters which exhibit reversal are underlined. Bremer support values are 0 unless indicated by large numerals on the right side of the nodes. In the resulting tree, the cryptobranch dorid Cadlina separates as a sister group to the reman- ing suctorian phanerobranch dorids, which are supported by the synapomorphies of non-contrac- tile rhinophore sheaths (#8), the presence of papillae within the gill circlet (#13) and the presence of a buccal pump (#16). These suctorian phanerobranch dorids traditionally belong to three families, Corambidae, Goniodorididae and Onchidorididae, but some members of the three families clustered together into one clade. This clade is distinguished by the presence of a peripherial muscle in the buccal pump (#17), a reduced half row of the radula (#19), one inner lateral tooth (#22), beak-shaped inner laterals (#21), and elongate oval outer laterals (#23). The Corambidae nests within the Onchidorididae and the genus Goniodoris nests with other members of the Onchidorididae and next to Diaphorodoris. Already the Superfamily Onchidoridoidea Gray, 1827 has been used to combine family Corambidae and Onchidorididae by Valdés and Bouchet (1998). Our data suggest that the families Corambidae and Onchidorididae are closely related (see Martynov 1994) and should be combined into one family, as suggested by Martynov (1999); the family group name Onchidorididae Gray, 1827 has priority. Further study is needed to clarify the position of Goniodoris and other members of the Goniodorididae in relation to this family. 20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. | The second clade contains members of the ‘basal’ Onchidorididae, which have sometimes been placed into the Goniodorididae. In this clade are the genera studied in this paper, Akiodoris and Armodoris, together with the closely related Doridunculus and the poorly known Prodoridunculus. Another poorly known genus, Echinocorambe, initially was placed in the family Corambidae by Valdés and Bouchet (1998) inasmuch as it has similar paedomorphic features. It was also suggested by Martynov (2000) that it is close to Doridunculus because of its similar radu- la and buccal pump; it may, in fact, be a juvenile of the latter. The cladogram confirms that it belongs in the same clade as Doridunculus. These five genera are in a clade with similar radulas, buccal pumps, and reproductive systems. Synapomorphies which distinguish this clade are the presence of a smooth, thin, lip disk (#15), two or more inner lateral teeth (#20), and the possession of rectangular, reduced outer laterals (#23). The distant position of Goniodoris in relation to this clade suggests that these five genera should not be in the family Goniodorididae. This clade forms a sister group to the clade containing the Onchidorididae (including Corambidae) and Goniodoris. Thus, this clade is considered a distinct family, Akidorididae Millen and Martynov, fam. nov. ACKNOWLEDGMENTS We thank Dr. Tanya Korshunova (Moscow, IHNA and NF RAS) for drawings and photographs of Armodoris antarctica. We are also thankful to Dr. Alexi Chernyshev for figures used in plate 1 and additional sketches of Akiodoris lutescens. Ron Long, Simon Fraser University, kindly sup- plied the photograph of Akiodoris salacia. Stefano Schiaparelli provided an SEM image of the radula of A. antarctica. We also thank Sven Donaldson and John McNickol for diving assistance. This research was partly funded by the Department of Zoology, University of British Columbia to Sandra Millen. Partial funding to Alexander Martynov was from the Russian Federal Program “Study of Antarctic biota, Project N 16.” Funding for publication costs is from the National Science Foundation, PEET Program (DEB-9978155) to Dr. Terrence Gosliner. LITERATURE CITED BEHRENS, D.W. 1991. Pacific Coast Nudibranchs, A Guide to the Opisthobranchs. Alaska to Baja Califorina, 2nd ed. Sea Challengers: Monterey, California, USA. 107 pp. BERGH, R. 1879a. Gattungen nordischer Doriden. Archiv. fiir Naturgeschichte 45(1):340—369, pl. 19. BERGH, R. 1879b. On the nudibranchiate gasteropod mollusca of the north Pacific Ocean, with special refer- ence to those of Alaska. Part I. Proceedings of the Academy of Natural Sciences of Philadelphia 31:71-132, pls 1-8. BERGH, R. 1880. On the nudibranchiate gasteropod mollusca of the north Pacific Ocean, with special refer- ence to those of Alaska. Part Il. Proceedings of the Academy of Natural Sciences of Philadelphia 32:40—127, pls. 9-16. BERGH, R. 1883. Beitrage zu einer Monographie der Polyceraden, II. Verhandlungen der konglich-kaiserlich Zoologisch-botanischen Gesellschaft in Wein. Abhandlungen 33:135—180. BERGH, R. 1892. Malacologische Untersuchungen, 3. System der Nudibranchiaten Gasteropoden. Pages 995—1165 in Carl Semper, Reisen im Archipel der Philippinen, Zweiter Theil. Wissenschaftliche Resultate Band 2, Theil 3, Heft 18. CATTANEO-VIETTI, R., M. CHIANTORE, S. SCHIAPARELLI, AND G. ALVERTELLI. 2000. Shallow and deep water mollusc distribution at Terra Nova Bay (Ross Sea), Antarctica. Polar Biology 23:173-182. FISCHER, P. 1833-1837. Manuel de Conchyliologie et de Palentologie conchyliologique ou histoire naturelle des mollusques vivants et fossiles suivi d'un appendice sur les Brachiopodes. Librairie F. Savy, Paris, France. 1369 pp. FRANC, A. 1968. Sous-classe des opisthobranches. Pages 834-888 in P.P. Grasse, ed., Traite de Zoologie, vol. 5. Masson et Cie., Paris, France. MILLEN AND MARTYNOV: NUDIBRANCH GENERA AKIODORIS AND ARMODORIS 21 Hols#TER, T. 1986. An annotated check-list of marine molluscs of the Norwegian coast and adjacent waters. Sarsia 71:73-145. Martens, E. 1879. Emend. for Akiodoris Bergh, 1879. Zoological Record, Mollusca 16:61. Martynoy, A. 1994. Materials for the revision of the nudibranch family Corambidae (Gastropoda, Opisthobranchia). Part 2. Origin of the Corambidae. Zoologicheskiy Zhurnal 73(11): 36-43 (In Russian). 1995. Hydrobiological Journal 31(7):59-67 (In English). Martynoy, A. 1997. Opisthobranch molluscs of the coastal waters of Commander Islands with notes on their Fauna of the Far-Eastern Seas of Russia. Pages 230-241 in A. Rzhavsky, ed., Donnaya fauna i flora Komandorskikh ostrovoyv. {Benthic Fauna and Flora of the shelf of Commander Ids.] Dalnauka, Vladivostok, Russia. (In Russian) Martynoy, A. 1999. [Abstract] Buccal pumps, gills pockets and new understanding of suctorial phanero- branchial dorids. Pages 13-14 in Systematic, Phylogeny and Biology of Opisthobranch Molluscs. 24 International Workshop of Malacology, Menfi, Italy, June 10-14, 1999. Martynovy, A. 2000. On the taxonomic placement of the genus Echinocorambe Valdés et Bouchet, 1998 (Gastropoda: Nudibranchia). XIV (5) All-Russian Malacological Conference, St. Petersburg, pp. 50-52. (In Russian) MILLEN, S. 1987. [Abstract] The nudibranch family Onchidorididae: a cladistic analysis. Western Society of Malacol-ogists Annual Report 20:19. MINICHEV, Y. 1972. Opisthobranchiate molluscs of the Davis Sea. [ssledovaniya Fauny Moreyi 19:358—-382. ODHNER, N.H. 1907. Northern and Arctic invertebrates in the collection of the Swedish State Museum (Riksmuseum). III. Opisthobranchia and Pteropoda. Kungelige Svenska Vetenskaps Akademiens Handlingar 41:1—114. Sars, G.O. 1878. Bidrag til kundskaben om Norges Arktiske fauna. I. Mollusca Regionis Arcticae Norvegiae, Oversigt Over de I Norges Arktiske Region Forekommende. Bloddyr., xii + 466 pp., 34 pls. Universitets- program, Christiania, Norway. THIELE, J. 1912. Die antarktischen schnecken und muscheln. Pages 183-285, pls. 11-19 in Drygalski, ed., Deutsche Sudpolar-Expedition 1901-1903. Zoologie, 13(5, 2). THIELE, J. 1931. Handbuch der systematishen Weichtierkunde. Bd. 1. Verlag von Fischer, Jena, Germany. 778 Pp- VaLpEs, A., AND P. BOUCHET. 1998. A blind abyssal Corambidae (Mollusca, Nudibranchia) from the Nor- wegian Sea, with a reevaluation of the systematics of the family. Sarsia 83:15—20. Vatpes, A. 2002. A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia). Zoological Journal of the Linnean Society 136(4):535—636. VALLES, Y., M. MEDINA, AND T. GOSLINER. 2001. [Abstract] Phanerobranch dorids: clade or grade? Page 368 in Abstracts of the World Congress of Malacology 2001. Vienna, Austria. WAGELE, H., AND R. WILLAN. 2000. Phylogeny of the Nudibranchia. Zoological Journal of the Linnean Society 130:83-181. WOLLSCHEID, E., AND H. WAGELE. 1999. Initial results on the molecular phylogeny of the Nudibranchia (Gastropoda, Opisthobranchia) based on 18A rDNA data. Molecular Phylogeny and Evolution 13: 215-226. WOOLSCHEID-LENGELING, E., J. BOORE, W. BROWN, AND H. WAGELE. 2001. The phylogeny of Nudibranchia (Opisthobranchia, Gastropoda, Mollusca) reconstructed by three molecular markers. Organisms Diversity and Evolution 1(4):241—256. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. De, Appendix TABLE |. Data matrix showing the distribution of character states (see text for details). PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. | So N ro) Acanthodoris Corambe Diaphorodoris Doridunculus Onchidoris Prodoridunculus Echinocorambe Goniodoris Calycidoris Bathydoris Adalaria Akiodoris Armodoris Cadlina PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 2, pp. 23-30, 2 figs. April 29, 2005 First Hawaiian Record of the Grenadier Lepidorhynchus denticulatus (Macrouridae: Gadiformes: Teleostei) Tomio Iwamoto Department of Ichthyology, California Academy of Sciences, 875 Howard St., San Francisco, CA 94103 USA; Email: tiwamoto:calacademy.org On August 16, 2003, while night lighting for squid off the northwest coast of Oahu, Mr. Richard Cadaoas caught on hook and line an 18 to 20-inch (about 46-51 cm) squid. In its stomach was a small fish about 20 cm long in excellent condition, having suffered little deterioration from its stay in the squid’s stomach. The specimen was taken to Mr. Arnold Suzumoto, Collections Manager for the ichthyological collections of the Bernice P. Bishop Museum, Honolulu. The fish could be readily identified as a grenadier, owing to its long tapered body that ends in a point and lacks a caudal fin, the relative form and positions of the fins, the spinulated body scales, and the general form of the head. The snout in the specimen was narrow and rounded, however, unlike the pointed and often stoutly reinforced snout of most other members of the family (excluding the bathygadids), and none of the ridges on the head were strengthened by stout spiny scutelike scales. The fish did not fit the description of any previously known species from the Hawaiian Islands and was subsequently sent to me, as one of the primary workers on this group of deepsea fishes, for identification. Even before I had taken it out of the jar, I recognized the fish as a Lepidorhynchus denticula- tus, a Species originally described by Sir John Richardson in 1846 from a dried specimen that “was thrown up on the beach of South Australia, and has lost part of its tail” (Richardson 1846:54). Despite the desiccated state of the type specimen, the original description and illustration more than adequately characterized this distinctive grenadier, which is the sole representative of the genus. The species was previously known only from the coasts of south and southeastern Australia and off the coasts of New Zealand in depths ranging from less than 100 m to more than 1000 m. It is one of the most common grenadier in these waters and forms a significant part of the by-catch of trawlers fishing at upper-slope depths. So far as known, the species has not been recorded north of latitude 34°S off New Zealand and about 30°S off southeastern Australia. Its distribution is gen- erally confined to subtropical waters. Its occurrence off Hawaii is thus most astounding and prompts the recording of this extraordinary specimen. METHODS Methods for making counts and measurements follow Iwamoto and Sazonov (1988). Institutional abbreviations follow Leviton et al. (1985). ep 24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 2 DESCRIPTION Lepidorhynchus denticulatus Richardson, 1846 Figs. 1-2 Macrourus vel Lepidorhynchus denticulatus Richardson, 1846:1—2, pl. 32, fig. 1-3 (holotype: BMNH 1845.11.31.6; type locality: coast of South Australia). Coryphaenoides denticulatus: Gunther, 1862:396. Macrurus (Optonurus) denticulatus: Giinther, 1887:147. Optonurus denticulatus: Gilbert and Hubbs, 1916:144. Lepidorhynchus denticulatus: Phillipps, 1927:58. MATERIAL EXAMINED.— BPBM 39286 (200 mm total length, tail complete; sex indetermi- nate); Hawaii, Oahu, northwest side about 2 miles (3.2 km) offshore; collected by Richard Cadaoas, 16 August 2004 from stomach of 18—20-inch (46-51 cm) squid caught by hook and line at night. CAS 220920 (6 spec., 240+-456 mm total length); Australia; New South Wales, off Bermagui; 36°46’-41’S, 150°21’E; 285-310 fm [521-567 m]; collected by Ken Graham on F/V “Shelley,” by trawl; 15 February 2000. COUNTS AND MEASUREMENTS.— (Hawaiian specimen first, followed by range of Australian specimens in parentheses.) First dorsal-fin rays II,11 (10-12); pectoral-fin rays 117 and 116 (115-118); pelvic fin rays 9 (9); branchiostegal rays 6; gill rakers first arch (outer/inner) 0+11 /3+13 (0+10—14 / 3-44+13-15 = 17-18), second arch 3+14 / 2+13 (2—3+12-15 = 14-17 / 2-3+12-14 = 14-17); scale rows below origin of first dorsal fin about 8 (about 7-8), below mid-base of first dor- sal fin about 5 (5.5), below origin of second dorsal fin about 6 (5—6.5); pyloric caeca 16 (15-25). Head length 31.5 mm (41.7—70.3 mm); the following in mm for Hawaiian specimen followed in parentheses by percent of head length for Hawaiian specimen and then Australian specimens: snout length 7.6 (22%, 20-24%): preoral length 5.1 (16%, 9—-13%); internasal width 6.8 (22%, 16-21%); interorbital width 8.8 (28%, 20-25%); orbit diameter 12.6 (40%, 35-39%); suborbital width 2.8 (9%, 8-8%):; postorbital length 12.6 (40%, 41-45%); distance orbit to angle of preopercle 11.9 (38%, 39-42%); length upper jaw 15.3 (49%, 47-50%); length barbel 3.1 (10%, 9—-15%); length Ye — ¥ . % a 4 t ¢ rf ¢ x : ’ ia a , FIGURE 1. Lepidorhynchus denticulatus, BPBM 39286 (31.5 mm head length, 200 mm total length), from off Oahu, Hawaii. IWAMOTO: GRENADIER LEPIDORHYNCHUS DENTICULATUS IN HAWAII DES outer gill slit 8.6 (27%, 22-27%); preanal length 47.5 (151%, 158-177%); distance outer pelvic fin to anal-fin origin 17.7 (56%, 53-69%); distance isthmus to anal-fin origin 32 (102%, 108-139%); greatest body depth 24.5 (78%, 79-90%): body depth at anal-fin origin 20.0 (63%, 64-80%); inter- space between first and second dorsal fins 23.5 (75%, 74-95%); height first dorsal fin 28.5 (90%, 78+—92); length base of first dorsal fin 11.1 (35%, 34-42%); length pectoral fin 23 (73%, 72-82%); length outer pelvic-fin ray 20.5 (65%, 57-62%). General description of Hawaiian specimen, with condition in larger Australian specimens in parentheses, if different. Body long and slender, laterally compressed, tapering posteriorly from abdomen into a long tail that ends in a fine point. Width of body across pectoral-fin bases less than half body depth below origin of first dorsal fin; greatest depth about equal to distance from anteri- or margin of orbit to end of opercle. Head about 6.2 into total length, compressed, its greatest width slightly greater than orbit diameter. Snout rounded and narrow, its tip slightly behind vertical of tip of jaws (slightly forward in larger Australian specimens). Orbits huge, more than postorbital length (slightly less than postorbital), about 2.4 into head length; its dorsal margin entering dorsal profile of head. Mouth large, essentially terminal, unrestricted by lip folds at angle; posterior margin of premaxillary approximately below middle of orbit (below posterior margin of pupil). Suborbital region narrow, shelf and ridge well defined, although this may be partly caused by slightly desic- cated state of specimen. Preopercle broadly rounded, its ridges well defined, covering most of interopercle (except anteroventrally and distal tip) and anterior part of subopercle, which ends ven- trally in a short tab. Chin barbel short, fine, its length about equal to diameter of posterior nostril. Gular membrane narrowly con- nected at isthmus far forward under middle of orbit (posterior half of orbit), overlapping and forming a deep fold over bran- chiostegal membrane. Anus immediately in front of anal-fin origin, separated at most by one or two scale rows. Swimbladder large, elongated, with eight or nine broad, flat retia, each con- nected to a globular to elongated bean-shaped gas gland (Fig. 2). Teeth on premaxilla in mod- erately wide band extending entire length of jaw opening; teeth small and conical or spike- like, except outer series of much larger, widely spaced canines in a single row. Dentary teeth small anteriorly and in one or two irregular series, becoming in- creasingly larger posteriorly and Ficure 2. Lepidorhynchus denticulatus, CAS 202920, 41.7 mm _ head ina single file. Larger teeth end length, 240+ mm total length. Saggital otolith, (a) mesial and (b) dorsal views; (c) swimbladder, showing retia and gas glands; (d) mesial view of lower jaw showing comblike teeth (arrow). abruptly and followed at posteri- 26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 2 or angle of jaws by a series of seven (four or five) comb-like teeth, slightly reclined, closely set, and somewhat overlapping at bases. These comblike teeth unlike those seen in any other grenadier. (In much larger Australian specimens, these teeth fewer and mostly embedded in gum tissue.) As with all other macrouroid fishes, no teeth on vomer and palatines. First dorsal fin high, triangular, with two spinous rays and 11 segmented rays, anteriormost segmented rays appear to be longest. First spinous ray short and splintlike, tightly coalesced with proximal part of long second spinous ray. Distally on second spine a few small thornlike denticles on leading edge (absent in larger Australian specimens). Interspace between first and second dor- sal fins wide, much greater than length base of first dorsal fin. Rays of second dorsal fin low and fine to end of tail, although gradually increasing in height posteriorly. Anal fin well developed throughout, its origin slightly behind vertical through posterior edge of first dorsal fin. Upper edge of pectoral fin about at mid-lateral line; fin well developed, relatively long, its length about equal to postrostral length of head, its tip extending to above 10 (4% or 5th) anal ray. Pelvic fins well developed; distal tip of outermost ray filamentous and hairlike, just reaching anal-fin origin. Luminescent tissue on ventral surfaces of body extensive and consisting of extremely fine transverse parallel black lines, the ventral striae. Luminescent tissue extends from isthmus over abdomen up to and slightly behind pectoral-fin base (as well as lateral surface of fin base), narrow- ing posteriorly to above anus, gradually tapering above anal fin to approximately 57'* (53-57) anal- fin ray. Tissue readily observed in preserved specimens as black areas underlain by extremely fine parallel black lines; in fresh specimens, these areas of striae usually overlain by a silvery reflection and striae much less apparent. (Ventral striae are also found in several unrelated groups such as the cardinalfishes of the genus Siphamia [Apogonidae], and the berycoids Aulotrachichthys spp. and Paratrachichthys spp. |Trachichthyidae]). A small, flat, black, triangular light organ situated with- in median line of abdominal wall closely adherent to base of rectum. Light organ externally man- ifested as a round white or translucent body encircled by a narrow black ring continuous with anal surround. Body scales of moderate size throughout, somewhat deciduous, and fully covered with fine, conical, needle-like spinules arranged in more or less parallel rows. (In larger Australian speci- mens, 30 or more rows of spinules on scales below interspace between dorsal fins.) Body scales almost entirely missing in Hawaiian specimen, but a few remain on nape and top of head. (Scales cover most of head, although lower surface of snout naked. Tiny scales incompletely cover subor- bital region.) Mandibular rami fully scaled. Gular and branchiostegal membranes naked. (Australian specimens have lowermost branchiostegal rays scaled, and in one specimen, exposed anteroventral margin of interopercle densely covered with tiny oval scales, each having a few short, upright spinules.) None of scales on head enlarged or thickened. Color in alcohol of denuded Hawaiian specimen overall flesh colored with prominent black areas corresponding to luminescent tissue. Gill membranes, lower parts of head, and opercle pos- terior to preopercular ridge blackish. Fins clear, although first dorsal fin somewhat light dusky. (In fresh specimens, species silvery over most lateral surfaces of head and body, with dorsum gray and area Over luminescent tissue black. Pelvic fins blackish; pectoral fins light dusky.) Comparisons The small size of the Hawaiian specimen made comparison of the morphometry of head and body structures uncertain. Several proportions of head parts in BPBM 39286 were greater than in the larger Australian specimens: snout length, preoral length, internasal width, interorbital width, orbit diameter, and suborbital width. The postorbital and orbit-to-preopercle measurements were slightly lower, but those were probably related to the relatively larger orbit diameter, which typi- IWAMOTO: GRENADIER LEPIDORHYNCHUS DENTICULATUS IN HAWAII pol cally shows allometric growth in grenadiers. That the interorbital and suborbital widths were not narrower is somewhat contrary to what one would expect. The slightly lower values for the preanal length, the distance of isthmus to anal-fin origin, and the greatest body depth are not uncommon in smaller individuals of grenadier species. The weakly and sparsely denticulated distal portion of the spinous dorsal ray is also seen in the smallest Australian specimen and depicted in the original illus- tration of the holotype. In the larger Australian specimens, these denticulations are lost. All other characters appear exactly the same in Hawaiian and Australian specimens, and despite the small differences in some head proportions, there is little reason to doubt the conspecificity of the two populations at this time. The possibility of there being two species cannot be completely discount- ed, however, and a more-definitive conclusion can only be made after additional specimens from Hawaii become available. DISCUSSION The unusual Hawaiian record of this species leads to speculation as to how this particular indi- vidual found its way so far across the Pacific from its normal grounds some 3700 nautical miles and more than 54 degrees of latitude distant. Many species of grenadiers have broad distributions, but those can generally be separated into three categories: (1) abyssal or lower-slope species, such as Coryphaenoides armatus (Hector, 1875) and Cetonurus spp.; (2) bathypelagic species, such as Cynomacrurus piriei Dollo, 1900 and Odontomacrurus murrayi Norman, 1939; and (3) species having a long-lived pelagic juvenile stage, the prime example being Malacocephalus laevis Lowe, 1843. Lepidorhynchus denticulatus does not appear to fit into any of these categories (but its early life history is not known), so its presence in the Hawaiian Islands is a mystery. Were it to be found in intervening areas of the western Pacific, its presence in Hawaiian waters would not be so sur- prising. Yet, despite relatively extensive collecting at appropriate depths throughout the central and North Pacific, especially by Japanese vessels, L. denticulatus has yet to be recorded north of New Zealand and southeastern Australia. Extensive trawling on the Lord Howe Rise and Norfolk Ridge north of New Zealand during the cooperative (New Zealand and Australia) NORFANZ cruise of the R/V Tangaroa in 2003 captured the species only once, and that was west of Three Kings Islands off the northern tip of New Zealand’s North Island at 34°S. The extensive French survey cruises to New Caledonia and adjacent areas failed to produce any specimen of the species. Most species of grenadiers are localized in their distributions, usually to basins, island groups, oceanic seamounts and ridges, or along continental margins that are bounded by differing oceano- graphic or geologic conditions. So far as known, besides Lepidorhynchus denticulatus, only five species of grenadiers from Hawaii are also known from Australia or New Zealand: Malaco- cephalus laevis, Kuronezumia bubonis (Iwamoto, 1974), Hymenocephalus aterrimus Gilbert, 1905, Nezumia propinqua (Gilbert and Cramer, 1897), and Trachonurus sentipellis (Gilbert and Cramer, 1897). The last three are found in Australian waters but not off the coasts of New Zealand’s main islands. Malacocephalus laevis and K. bubonis are widespread throughout most of the Pacific, Atlantic and Indian oceans, and N. propinqua is widespread in the Pacific and Indian oceans, although there is some question as to whether that name is currently being applied to more than one species. Trachonurus sentipellis is recorded from Hawaii, the New Caledonian region, and Australia, but there is again some doubt that the Hawaiian populations are the same species as the austral populations. If the identifications are correct, this species comes closest in its distribution to that of Lepidorhynchus. It has been theorized (Marshall 1965, 1973) that most grenadiers have pelagic eggs that hatch near the bottom; the larval stage is very short before metamorphosis into a pelagic juvenile stage (sometimes called postlarva or prejuvenile) takes place. The young probably spend relatively little 28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 2 time in the midwaters before descending to the bottom to live a benthopelagic life. A short pelag- ic early life would ensure that the settled young are over appropriate living depths and are not waft- ed too far from the narrow continental slopes on which adult populations live. The scarcity of lar- val grenadiers in collections, despite the group’s abundance throughout the world’s oceans, adds support to this idea of a short pelagic stage lived close to the bottom. Perhaps there are some species that can tolerate an extended pelagic larval or juvenile stage when wafted far beyond their normal range. The concept of “expatriate” individuals is well known for some fishes, and this may come into play with some unusual range extensions in grenadiers. Whether any of these factors may account for the extraordinary occurrence of L. dénticulatus in Hawaii 1s yet to be learned, as the early life history of the species is not known, nor is it known that L. denticulatus is actually established in the Hawaiian Islands. It seems unlikely, however, that the captured specimen repre- sents a single stray individual in these waters. There are few examples of other demersal shelf and upper-slope fishes of southern Australia and New Zealand that have a distribution also in the Hawaiian Islands. Mr. Suzumoto informs me (in litt., Nov. 2004) that the Bishop Museum has “‘a specimen of Cheilidonichthys kumu” caught off Hawaii Island in 1926 and never seen here since.” That Bishop Museum sea robin (family Triglidae) was recorded by Pietschmann (1930) and considered of questionable validity by Springer (1982:103). It was said (Gomon et al. 1994:496) to occur off South Africa, southern Australia, and New Zealand, and also in the North Pacific off Japan, Korea, and China; however, recently published books on the Japanese fish fauna (e.g., Masuda et al. 1984; Nakabo 2002) list C. spinosus and C. ischyrus but not C. kumu. Martin F. Gomon (NMV) has informed me (in Iitt., 12 Dec. 2004) that Peter Last (of CSIRO) “has differentiated several species of Cheilidonichthys in Australia, probably none of them also in South Africa, and the Japanese species are probably something else again.” Gomon adds that “there are other Hawaiian cognates, like the Bodianus oxycephalus-like species. . .with 3 cognates in Japan, southern Pacific and SW Australia, respec- tively.” The morwong Cheilodactylus vittatus was thought to have a north-south disjunct distribu- tion: the Hawaiian Islands in the northern hemisphere, the Lord Howe Island and New Caledonia in the southern hemisphere (Springer 1982:33). Burridge (2004), however, has reported that the New Caledonia and Norfolk Island Cheilodactylus represents a different species, which he described as C. francisci. The mirror dory, Zenopsis nebulosus, is a wide-ranging shelf to upper- slope species (62—550 m) found off Australia, New Zealand and Chile in the southern hemisphere, and also in the North Pacific off Japan, Korea, Hawaii and California (Gomon et al. 1994:420-421). Of the three deep-dwelling (100-400 m) Emmelichthyidae in Hawaii, one species also occurs off Japan, Australia and the Philippines (Springer 1982:41). In light of these few exam- ples, it is apparent that many fish species formerly thought to have a broadly disjunct distribution in Hawaii and the southwest Pacific are, in fact, separable into more than one closely related species. Other examples are likely to surface after closer study using tools and knowledge not for- merly available. That possibility for Lepidorhynchus cannot at this time be discounted. Springer (1982:117), in his valuable study on Pacific Plate biogeography, found 121 families of shorefishes that are nonmarginally represented on the Pacific Plate. Of these, 15 families (12%) are found only in the Hawaiian Islands, and 10 of the 15 occur there only in deep water. These “Hawaiian exceptions,” as so designated by Springer, “denote Indo-Pacific taxa that occur nonmar- ginally on the Pacific Plate only at the Hawaiian Islands. These taxa usually occur also in Japan and/or the Ruykyu [sic; misspelling for Ryukyu] Islands, and many are antitropically distributed in the Indo-West Pacific; some are restricted to north of the tropics and some are moderately deep dwelling.” (Springer 1982:135.). Lepidorhynchus should be added as an Hawaiian exception, but the underlying cause of its apparent disjunct distribution is unknown. IWAMOTO: GRENADIER LEPIDORHYNCHUS DENTICULATUS IN HAWAII 29 Central to understanding the vicariant event that produced the current distribution of Lepidorhynchus, one should have a firm grasp of its closest relatives. “If one knows the cladisti- cally determined closest relatives (sister groups) of endemic taxa, and the distributions of those rel- atives, one can narrow the geographic and temporal search for the pertinent vicariant events.” (Springer 1982:6). Unfortunately, the sister-group relationships of Lepidorhynchus within the Macrouridae are uncertain and a thorough cladistic analysis of macrourid genera has yet to be made. Iwamoto and Sazonovy (1988:39), in their tentative phylogeny of macrourines with six bran- chiostegal rays, included the genera Cynomacrurus and Odontomacrurus in a clade, with Lepidorhynchus as the clade’s questionable sister group. Odontomacrurus murrayi perhaps comes closest in overall morphology to Lepidorhynchus, having a compressed head, large, terminal jaws beset with canines, and a small light organ. That bathypelagic species, however, lacks ventral stri- ae, a developed swimbladder, and a chin barbel, and it has many reductions and differences in its morphology and organ systems that reflect its bathypelagic existence and thereby distance it from L. denticulatus. The broad, almost worldwide distribution of O. murrayi in tropical to subtropical waters shows a noteworthy hiatus — so far as known, there are no records of its presence on the Pacific Plate except marginally. Of the grenadiers with seven branchiostegal rays, the luminescence on the body, the rounded snout with little protrusion of the rostrum, and the scale spinulation are most suggestive of Hymenocephalus, but the six branchiostegal rays, the eight or nine retia and gas glands, the absence of a light organ lens on the chest, the very different dentition, and the narrowly compressed head in Lepidohynchus are among several characters that separate it from members of Hymenocephalus. The head shape, especially the narrow snout with little protrusion of the median nasal ridge, is somewhat like that of Ventrifossa atherodon (Gilbert and Cramer, 1897) and the scale morphology is quite similar, but members of Ventrifossa lack ventral striae, they have seven branchiostegal rays and two retia and gas glands, and there is a small dermal window of the light organ between the pelvic-fin bases that is absent in Lepidorhynchus. Steindachneria argentea Goode and Bean, 1886, in the monotypic family Steindachneriidae, bears some resemblance to L. denticulatus in its ven- tral striae, large terminal mouth beset with canine-like teeth, and similar retia and gas glands in the swimbladder (see Marshall 1966, fig. 3C), but differs radically in having second dorsal rays well developed and higher than most anal-fin rays, an elevated lobe in the otherwise low anal fin, vomerine teeth, no chin barbel, anus located between pelvic-fin bases and distantly separated from urogenital opening, which is immediately before the anal fin. It is apparent that a more thorough comparative study of the morphology of Lepidorhynchus denticulatus, perhaps combined with molecular investigations, will be needed to clarify the phylogenetic position and discover the sis- ter group of this perplexing species. ACKNOWLEDGMENTS I thank Arnold Suzumoto (BPBM) for sending me the specimen of Lepidorhynchus and for providing additional information; Ken Graham (New South Wales Fisheries, Australia) for provid- ing comparative material from Australia; Jon Fong and Mysi Hoang (CAS) assisted in the imaging of the specimen. Drs. Martin Gomon (NMV) and Clive Roberts (NMNZ) critically reviewed the manuscript and provided valuable insights and information, as well as editorial corrections. LITERATURE CITED BURRIDGE, CHRISTOPHER P. 2004. Cheilodactylus (Goniistius) francisi, a new species of morwong (Perciformes: Cirrhitoidea) from the southwest Pacific. Records of the Australian Museum 56:23 1-234. 30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 2 GILBERT, CHARLES H., AND CARL L. Hupss. 1916. Report on the Japanese macrouroid fishes collected by the United States fisheries steamer “Albatross” in 1906, with a synopsis of the genera. Proceedings of the United States National Museum 51(2149):135—214, pls. 8-11. GoMoNn, MArtIN F., JOHN C.M. GLOVER, AND RupIE H. KulTer. 1994. The Fishes of Australia’s South Coast. State Print, Adelaide, Australia. 992 pp. GUNTHER, ALBERT. 1862. Catalogue of the Acanthopterygii, Pharyngognathi and Anacanthini in the collection of the British Museum. Catalogue of the Fishes of the British Museum, vol. 4:1—xxi, 1-534. Trustees of the British Museum, London, UK. GUNTHER, ALBERT. 1887. Report on the deep-sea fishes collected by H.M.S. Challenger during the years 1873-76. Report on the Scientific Results of the Voyage of H.M.S. “Challenger” During the Years 1873-76, vol. 22 (part 57):1—268, pls. 1-66. London, UK. IwAMoTO, TOMIO, AND YURI I. SAZONOV. 1988. A review of the southeastern Pacific Coryphaenoides (sensu lato) (Pisces, Gadiformes, Macrouridae). Proceedings of the California Academy of Sciences, ser. 4, 45(3):35-82. LEVITON, ALAN E., ROBERT H. GIBBS, JR., ELIZABETH HEAL, AND CHARLES E. DAwSoNn. 1985. Standards in her- petology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in her- petology and ichthyology. Copeia 1985(3):802—832. MARSHALL, NORMAN B. 1965. Systematic and biological studies of the macrourid fishes (Anacanthini- Teleostii). Deep-Sea Research and Oceanographic Abstracts 12(3):299-322. MARSHALL, NORMAN B. 1966. The relationships of the anacanthine fishes, Macruronus, Lyconus, and Steindachneria. Copeia 1966(2):275—280. MARSHALL, NORMAN B. 1973. Family Macrouridae. Pages 496-665 in Daniel M. Cohen, ed., Fishes of the Western North Atlantic, Part 6. Sears Foundation for Marine Research Memoir 1. Masuba, HAJIME, KUNIO AMAOKA, CHUICHI AARGA, TERUYA UYENO, AND TETSUO YOSHINO, EDS. 1984. The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, Japan. 437 pp.; 370 pls. NAKABO, TETSUJI, ED. 2002. Fishes of Japan with Pictorial Keys to the Species, English edition. Tokai University Press, Tokyo. Vol. I: pp. i-xx, 1-866; Vol. I: pp. i-vii, 867-1749. PHILLIppSs, W.J. 1927. Bibliography of New Zealand fishes. New Zealand Marine Department, Fisheries Bulletin (1):1-68. PIETSCHMANN, VIKTOR. 1930. Remarks on Pacific fishes. Bernice P. Bishop Museum, Bulletin 73:\—24, pls. 1+. RICHARDSON, JOHN. 1846. Fishes. In: The Zoology of the Voyage of H.M.S. Erebus and Terror, under the com- mand of Capt. Sir James Clark Ross during the years 1839 to 1843. Vol. 2. viii + 739 pp., 40 pls. London, UK. SPRINGER, VICTOR G. 1982. Pacific Plate biogeography, with special reference to shorefishes. Smithsonian Contributions to Zoology No.67. 182 pp. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 3, pp. 31-41, 2 figs. April 29, 2005 The Phylogenetic Position of the Recently Rediscovered Philippine Forest Turtle (Bataguridae: Heosemys leytensis) Arvin C. Diesmos!:2, James F. Parham>-4, Bryan L. Stuart5-6, Rafe M. Brown’ ! Department of Biological Sciences, National University of Singapore, Block S3 14 Science Drive 4, Singapore 117543. E-mail: kaloula@i-manila.com.ph; *Herpetology Section, Zoology Division, National Museum of the Philippines, Padre Burgos Avenue, Ermita 1000, Manila, Philippines; 3Evolutionary Genomics Department, Joint Genome Institute, 2800 Mitchell Drive, Walnut Creek, CA 94598, USA. Email: parham®@ socrates.berkeley.edu; (Museum of Paleontology, University of California, Berkeley, CA 94720, USA; >The Field Museum, Department of Zoology, Division of Amphibians and Reptiles, 1400 S. Lake Shore Drive, Chicago, IL 60605-2496, USA. Email: bstuart@ fmnh.org; “Department of Biological Sciences (M/C 066), University of Mlinois at Chicago, 845 W. Taylor, Chicago, IL 60607-7060, USA; ‘Division of Herpetology, Natural History Museum, University of Kansas, Lawrence, KS 66045-2454, USA. Email: rafe@mail.utexas.edu The Philippine forest turtle (Heosemys leytensis) is one of the least-known Asian tur- tles. Until recently it was only known from a single specimen, the neotype (CAS 60930). The rediscovery of this enigmatic turtle in the wild provides access to fresh genetic material, allowing us to rigorously test its phylogenetic position among Asian pond and wood turtles of the clade Bataguridae for the first time. We sequenced 1174 bp of mitochondrial DNA (cytb) and 1076 bp of nuclear DNA (R35 intron) from a newly acquired specimen of H. leytensis (PNM 8488) and compared our sequences to a previously published genetic survey of batagurids. Our main goal is to determine if H. leytensis is closely related to other species of Heosemys (H. spinosa [type species], H. grandis, H. depressa, and H. annandalii [comb. nov.]). Parsimony and likelihood analyses of mitochondrial and nuclear DNA data strongly support H. leytensis as the sister taxon to the Southeast Asian species Siebenrockiella crassicollis (commonly known as the ‘black marsh turtle’ or ‘smiling terrapin’). The close rela- tionship between H. leytensis and S. crassicollis is a novel hypothesis, although both share an easily recognizable diagnostic feature: vertebrals 2-4 are mushroom- shaped or ‘ginkgo leaf’-shaped. In order to eliminate two potentially monotypic gen- era, the genus Siebenrockiella is phylogenetically defined to accommodate the close relationship between S. crassicolis and H. leytensis whereas a new name, Panyaenemys, is defined for the distinct H. leytensis stem lineage. The rediscovery of wild populations of the Philippine forest turtle (Heosemys leytensis Taylor 1920; Fig. 1B-E) in the Palawan island group of the western Philippines (Fig. 1A; Diesmos et al. 2004a), resolved a decades-long search for this species. Prior to this, our knowledge of H. leyten- sis was restricted to the original published description (the type series was lost in the destruction of the Bureau of Science, predecessor to the National Museum of the Philippines, during World War II; Brown and Alcala 1978; Buskirk 1989), one museum specimen (CAS 60930, the neotype, Fig. 1B,C) reported by Buskirk (1989), and one specimen from Palawan (UF/FSM 67515) report- ed by Timmerman and Auth (1988). Thus, for over forty years, no museum specimens of H. leyten- 3] 32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 3 sis were known and so it remained one of the least-known Asian turtles. The rediscovery of H. leytensis in the wild (Diesmos et al. 2004a) provides access to fresh genetic material, allowing us to rigorously test its phylogenetic position within the ‘Asian pond and wood turtles’ (Bataguridae Gray, 1870 = “Geoemydidae Theobald, 1868’; Joyce et al. 2004) for the first time. Batagurids are a species-rich group (60+ spp.) of highly threatened (van Dijk et al. 2000), but poorly studied non-marine turtles from Asia. A taxonomic reassessment of H. leytensis is nec- essary because the content and diagnosis of batagurid genera have changed dramatically since Taylor (1920) described the Philippine forest turtle as Heosemys leytensis (e.g., Ernst and Barbour 1989; Iverson 1992). The twelve characters that Taylor (1920) used to diagnose Heosemys are now known to be either plesiomorphic for batagurids or else appear in other genera as homoplasy (Joyce and Bell 2004). One of these characters, the derived absence of a temporal arch in the skull, has played a particularly important role in maintaining H. Jeytensis in the genus Heosemys. Besides being noted by Taylor (1920), this character was later used by McDowell (1964) to diagnose Heosemys (including H. leytensis) and was also described in the H. leytensis neotype (Buskirk 1989). This character is also known in Hieremys annandalii (Boulenger, 1903) as well as some species of testudinoid box turtles from China and North America (Cuora Gray, 1855 and Terrapene Merrem, 1820; Zangerl 1948; McDowell 1964; Joyce and Bell 2004). Because H. leytensis lacks the characteristic hinged plastron of a box turtle, its referral to Heosemys seems logical. Although H. leytensis shares the absence of the temporal arch with other Heosemys, it is mor- phologically divergent from other Heosemys in other respects. For example, H. leytensis usually has mushroom-shaped or ‘ginkgo leaf’-shaped vertebral scales 2-4 (Fig. 1B, E), a feature that is otherwise restricted to the “smiling terrapin’ of southeast Asia, Siebenrockiella crassicollis (Gray 1831) (Fig. 1F—G). Additionally, the projecting gular region (anterior region of the plastron) of H. leytensis clearly distinguishes it from all other batagurid lineages, including other species referred to Heosemys. In this character, it superficially resembles some species of terrestrial tortoises (Testudinidae Gray, 1825), the sister-taxon to batagurids (Spinks et al. 2004). Given the extensive homoplasy within batagurid morphology (Joyce and Bell 2004) and the overall distinctiveness of H. leytensis, it is difficult to confidently place H. leytensis into any exist- ing genus on morphological grounds alone. Moreover, weak nodes generated from cladistic analy- ses of morphological variation in Bataguridae (Hirayama 1985; McCord et al. 1995) are strongly rejected by molecular studies (McCord et al. 2000; Spinks et al. 2004). For this reason, we tested the phylogenetic position of H. /eyrensis with cladistic analyses of mitochondrial and nuclear DNA sequences. Our primary goal was to determine whether H. leytensis should continue to be recognized as a member of the genus Heosemys. But in order to avoid confusion, we must establish an explicit concept of Heosemys and what species names this informal clade name should include. Spinks et al. (2004) reported DNA sequences from three species of Heosemys including one sample attrib- uted to the type species Heosemys spinosa (Gray, 1831). According to their study, Heosemys form a well-supported clade with Hieremys annandalii, and so Spinks et al. (2004) raised the possibili- ty that H. annandalii might be better placed within an expanded Heosemys. However, Spinks et al. (2004) refrained from making a taxonomic change pending future evidence for the possible place- ment of Hieremys annandalii relative to Heosemys species. We think this reticence is unwarranted inasmuch as Hieremys Smith, 1916 is a monotypic genus and H. annandalii forms a well-support- ed clade with other Heosemys. The Spinks et al. (2004) suggestion to expand the older genus name (Heosemys) to the well-supported node creates a much more stable taxonomy. Under this scheme, the widely-used genus name will remain stable regardless of the ultimate resolution of the position within the Heosemys basal polytomy. We refer Hieremys annandalii to the genus Heosemys, but DIESMOS ET AL.: PHYLOGENETIC POSITION OF HEOSEMYS LEYTENSIS 38 FIGURE 1. A: Map showing the known distribution of Siebenrockiella crassicollis (dots) based on Iverson (1992) and Stuart and Platt (2004). The black rectangle includes the known and hypothesized range of Heosemys leytensis based on Diesmos et al. (2004b): B-C: Neotype specimen of Heosemys leytensis (CAS 60930), dorsal [note the missing “ginkgo leaf’- shaped vertebral 2] (B) and ventral view (C); D: Dorsal view of H. leytensis head showing diagnostic markings; E: Dorsal view of H. leytensis possibly from Dumaran Island (see Diesmos et al., 2004a); F: Dorsal view of S. crassicollis (FMNH 224070): G: View of S. crassicollis head showing the diagnostic markings. refrain from defining the genus Heosemys phylogenetically until we discuss our results, especially regarding the placement of H. leytensis. In summary, our concept of Heosemys is the clade that includes the type species Heosemys spinosa, the aforementioned Heosemys [orig. Cyclemys] annandalii (Boulenger, 1903) comb. nov., Heosemys grandis (Gray, 1860), and Heosemys depres- sa (Anderson, 1875). INSTITUTIONAL ABBREVIATIONS.— CAS, The California Academy of Sciences, San Francisco, 34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 3 California, USA; FMNH, The Field Museum of Natural History, Chicago, Illinois, USA; PNM, National Museum of the Philippines, Manila, Philippines; UF, Florida Museum of Natural History, University of Florida, Gainesville, Florida. MATERIALS AND METHODS Ethanol-preserved muscle was taken from a museum specimen of Heosemys leytensis (PNM 8488, see Material Examined). Total genomic DNA was extracted from the muscle using PureGene Animal Tissue DNA Isolation Protocol (Gentra Systems, Inc.). Primers for amplifying and sequencing parts of cytb and tRNA-Thr (collectively ‘cytb’) were taken from Spinks et al. (2004) and R35 primers were taken from Fujita et al. (2004). PCR products were electrophoresed in a 1% low melt agarose TALE gel stained with ethidium bromide and visualized under ultraviolet light. The bands containing DNA were excised and agarose was digested from bands using GELase (Epicentre Technologies). PCR products were sequenced in both directions by direct double-strand cycle sequencing using Big Dye version 3.1 chemistry (Perkin Elmer). Cycle-sequencing products were precipitated with ethanol, 3 M sodium acetate, and 125 mM EDTA, and sequenced with a 3730 DNA Analyzer (ABI). Sequences were edited and protein-coding regions were translated into amino acids with Sequencher version 4.1 (Genecodes). We compared new cytb and R35 sequences with the sequences reported by Spinks et al. (2004). Sequences were aligned by eye into the matrix used by Spinks et al. (2004) and deposited into TreeBASE (www.treebase.org, accession number $1002). Our sequences (cytb, 1174 bp; R35, 1076 and 1078 bp) were longer than those used in the Spinks et al. (2004) alignment; the extra base pairs were excluded from the analysis. For the parsimony analysis of combined data sets, we included 72 of the 79 taxa sequenced by Spinks et al. (2004), excluding seven putative hybrids. Some of these hybrids were described as new species from pet trade samples leading to taxonom- ic confusion that is not yet fully resolved (Parham et al. 2001; Wink et al. 2001; Spinks et al. 2004). We avoid these issues by removing the suspect taxa from consideration. We feel that we are justi- fied in doing this because an initial phylogenetic analysis (not shown) demonstrated that H. leyten- sis is not closely related to any of these taxa and the inclusion or exclusion of these sequences does not affect the phylogenetic position of H. leytensis. Besides H. leytensis, the only addition to the Spinks et al. (2004) data set was the sequence of one vouchered Siebenrockiella crassicollis sam- ple from Cambodia (see below). In addition to the aforementioned combined analysis, we analyzed the mitochondrial and nuclear data sets separately in order to compare the potential congruence or conflict in the phylo- genetic signal of our markers. We also performed a maximum likelihood and Bayesian analyses on a restricted data set that includes 28 of the 79 species from Spinks et al. (2004). These 28 sequences represent all the non-hybrid taxa from the Spinks et al. (2004) study that have both cytb and R35 sequenced. The restricted data set is still appropriate for placing H. leytensis because it contains every major lineage (1.e., genus) of batagurid. Parsimony analyses were performed using the 1000 random addition-sequence replicates of the heuristic search algorithm in PAUP*4.0b10 (Swofford 2002). Support for the parsimony analy- ses was determined using bootstrap and decay indices. For the combined analysis, we used 1000 bootstrap replicates with 10 random addition-sequence replicates each. Decay indices were calcu- lated using a PAUP command file generated by MacClade 4.0 (Maddison and Maddison 2000). For the likelihood analyses, we used hierarchical likelihood-ratio tests with ModelTest (Posada and Crandall 1998) to determine which model was appropriate. For the standard likelihood analysis the model GTR+I+G was selected, with proportion of invariable sites 0.3836, gamma distribution DIESMOS ET AL.: PHYLOGENETIC POSITION OF HEOSEMYS LEYTENSIS 35 shape parameter 0.4762, and base frequencies as A=0.3198, C=0.3039, G=0.1336, and T=0.2427. Maximum likelihood analyses were performed with 100 random addition replicates with stepwise addition of taxa using the heuristic search algorithm and TBR branch swapping. For the Bayesian analysis, we separated the cytb and R35 data into their own partitions and estimated the best mod- els for each (GTR+I+G and HKY respectively). Using these parameters, we performed three mixed-model analyses of five million generations and four chains each using MrBayes v3.0b4 (Huelsenbeck and Ronquist 2001). We plotted the log likelihood scores against generation to deter- mine when the analysis achieved stationarity for each analysis. In each case, the seven thousand trees recovered prior to stationarity were discarded as “burn in’ and a 50% majority rule consensus was used to determine the posterior probabilities for each clade. RESULTS In our initial analyses, the sequence from our sample of H. leytensis is consistently placed as the sister taxon to S. crassicollis with high statistical support. None of the samples from the Spinks et al. (2004) study have vouchered specimens, so we confirmed our initial results by sequencing a vouchered specimen of S. crassicollis from Cambodia (see Material Examined). All tree statistics were calculated with the additional sample included. All analyses (parsimony, likelihood, Bayesian) place H. leytensis and S. crassicollis as sister taxa (Fig. 2). In the parsimony and Bayesian analyses, H. leytensis and S. crassicollis are united with high statistical support (100% parsimony bootstrap, 23 decay index, 100% Bayesian posterior probability). Parsimony analyses of the separate cytb and R35 data sets (not shown) show that the H. leytensis + S. crassicollis clade is independently supported by our mitochondrial and nuclear markers. Meanwhile, the other species of the genus Heosemys form a well-supported clade that is separated from H. leytensis + S. crassicollis clade by several branches (Fig. 2). DISCUSSION Our study demonstrates that the continued placement of H. leytensis in the genus Heosemys 1s no longer tenable. Inasmuch as the other species of Heosemys form a well-supported clade, which is readily-diagnosable with molecular and morphological data, we phylogenetically define the genus Heosemys as the crown clade arising from the last common ancestor of Heosemys [orig. Emys| spinosa (Gray, 1831), Heosemys |orig. Geoemyda] grandis (Gray, 1860), Heosemys [orig. Geoemyda| depressa (Anderson, 1875), and Heosemys [orig. Cyclemys| annandalii (Boulenger, 1903). The hypothesis that H. leytensis and S. crassicollis are sister taxa within Bataguridae is a novel result. Both H. leytensis and S. crassicollis share two diagnostic characters that may have been inherited from a common ancestor: 1) Vertebrals 2-4 are ‘ginkgo leaf’-shaped. Orlitia borneensis Gray 1873 has a superficially similar condition, but it is restricted to the first vertebral; 2) Light spots or lines in the posterior dorsal region of the head. In S. crassicollis (Fig. 1G), this character occurs as two circles or spots that may fade with age (Lim and Das 1999) whereas in H. leytensis (1C) it is a straight line that is faint or absent in 35% of individuals (Diesmos et al. 2004a, b). A similar, but more dramatic, condition of posterior head marking occurs in parallel in the ‘eyed tur- tles’ of the genus Sacalia Gray 1870. Additional morphological comparisons of S. crassicollis and H. leytensis, including skeletal material, may reveal more potential synapomorphies. Meanwhile, the character that has long been used to place H. leytensis in the genus Heosemys, the absence of a complete temporal arch, is more 36 __ <50 Bataguridae PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Emydidae Rhinoclemmys rubida Rhinoclemmys areolata 7O\ “Ament aemean CLOCIC Hardella Ca Melanochelys trijuga 3 S11 Volume 56, No. 3 Testudinidae Rhinoclemmys pulcherrima Rhinoclemmys annulata Rhinoclemmys punctularia Rhinoclemmys diademata Rhinoclemmys melanosterna Rhinoclemmys funerea Malayvemys subtrijuga 9 Orlitia borneensis mys hamiltonii Morenia ocellata thurjii llagur borneoensis Kachuga dhongoka Batagur baska Pangshura tecta Pangshura smithii Pangshura tentoria Pangshura tentoria 50 changes | Geoemyda spengleri 10 Sacalia bealei 94 100— Melanochelys trijuga <50, 3 Geoemyda japonica Sacalia quadriocellata Heosemys annandalii Heosemys gran Heosemys depressa | 1] Leucocephalon yuwonoi hee qi) Notochelys platynota <50|_ 14 Cyclemys atripons —p 8 | | 89 | tole 8| 98 | | ale bs Cyclemys tcheponensis Cyclemys dentata Mauremys leprosa Mauremys reevesii Mauremys reevesii | Mauremys japonica —— Mauremys sinensis Mauremys rivulata Mauremys caspica Mauremys annamensis Mauremys mutica Mauremys mutica Cuora amboinensis Cuora amboinensis |= Cuora amboinensis * - Cuora amboinensis Cuora picturata Cuora mccordi Cuora flavomarginata Cuora flavomarginata Cuora mouhotii =| Cuora zhoui Cuora pani ‘— Cuora trifasciata — Cuora aurocapitata * 3 "Heosemys" leytensis pei: 100 67-— Siebenrockiella crassicollis 100— Sitebenrockiella crassicollis -—— Heosemys spinosa (type species) i DIESMOS ET AL.: PHYLOGENETIC POSITION OF HEOSEMYS LEYTENSIS 3)// homoplastic than previously thought. If this character is optimized onto the phylogeny (Fig. 2), it predicts that the temporal arch was lost at least three independent times within Bataguridae: once in the H. Jeyrensis stem-lineage, once in the in the Heosemys stem lineage, and at least once with- in Cuora. A polytomy at the base of Cuora (e.g., Spinks et al., 2004; Parham et al., 2004) and a poor understanding of the morphological variation in extent of the temporal arch (see Joyce and Bell 2004) preclude a definitive assessment of the polarity of this character within Cuora. From a biogeographic perspective, the S. crassicollis + H. leytensis clade seems plausible. Heosemys leytensis is known only from the Palawan Pleistocene Aggregate Island Complex (or Palawan PAIC; Brown and Diesmos 2001). Although the Palawan region contains a significant level of endemicity in fauna and flora not found in the Sundaic region (see Widmann 1998, McGuire and Alcala 2000, Brown and Guttman 2002, Evans et al. 2004), in general the fauna of the Palawan PAIC has a much closer affinity to the Islands of the Sunda Shelf (e.g., Kalimantan/‘Borneo’, Madura, Sumatra) than do the other Philippine islands (Heaney 1986; Brown and Diesmos 2001). Therefore, the close relationship of H. leytensis to a Sunda Shelf species (S. crassicollis) is not surprising. The combined distribution of the S. crassicollis+H. leytensis clade (Fig. 1A) is very similar to that of Cyclemys dentata (Gray 1831) and Cuora amboinensis (Daudin 1802), the two other turtle species known from Palawan PAIC (Iverson 1992). Their shared distributions may indi- cate that these three lineages may have a shared biogeographical history as well, the details of which could be compared through additional DNA sequencing of S. crassicollis, Cu. amboinensis and Cy. dentata from throughout their range. Given the strong phylogenetic signal placing it as the sister taxon to S. crassicollis, we feel jus- tified in referring H. leytensis to a phylogenetically defined Siebenrockiella Lindholm 1929. By using the genus name Siebenrockiella to highlight this close relationship, we eliminate two mono- typic genera (Siebenrockiella and a hypothetical new genus for H. leytensis) from a literature that is lamentably crowded with monotypic genera (Spinks et al. 2004). Polytypic genera are function- ally superior to monotypic genera because they maximize the information content in each widely- used name (Parham and Feldman 2002; Feldman and Parham 2004). With this is mind, we phylo- genetically define Siebenrockiella as the crown clade arising from the last common ancestor of Siebenrockiella |orig. Emys| crassicollis (Gray 1831) and Siebenrockiella [orig. Heosemys| leyten- sis (Taylor 1920) (comb. nov.). In addition to developing a conservative nomenclature for the widely-used genus name, we also want to name the stem-lineage that includes the morphologically and molecularly divergent S. leytensis. For example, compared to S. crassicollis, S. leytensis has a proportionately larger head and correspondent nuchal emargination, is larger in overall body size, and has an extremely pro- truding gular region of the plastron (Diesmos et al. 2004b). The morphological distinctiveness of the two known Siebenrockiella species is demonstrated by the fact that no author has ever suggest- ed a close relationship between them until now. Moreover, the genetic distinctiveness of the S. leytensis lineage is shown by a high sequence divergence from S. crassicollis (~13% cytb) that is greater than that shown by other batagurid congeners (5.0-10.7%, see Spinks et al. 2004). In order FIGURE 2 (left). Parsimony phylogram of one of the four most parsimonious trees (4387 steps) recovered by the par- simony analysis of the combined cytb and R35 data set for batagurid turtles. For taxonomy of suprageneric clades see Joyce et al. (2004). The four equally parsimonious results differ in the placement of species within the genus Mauremys or between suprageneric genera not related to Heosemys or Siebenrockiella. Nodes that are not supported in all of the four equally parsimonious trees are indicated by ‘-’ on either side of the stem. The numbers at each of the other nodes represent support values. The top numbers are decay indices and the bottom numbers are parsimony bootstrap percentages. Support values for species clades within genera are not shown but see Spinks et al. (2004). The batagurid lineages that lose a com- plete temporal arch are shown by ‘*’. 38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 3 to provide a name for this distinct lineage that is divorced from the vagaries of species concepts, we name a new subgenus, Panyaenemys, for S. leytensis. SYSTEMATICS Panyaenemys, new clade name, new subgenus TYPE SPECIES: Siebenrockiella leytensis (Taylor, 1920) comb. nov. We designate a type species for this phylogenetically defined clade name so that it is valid under the rules of the International Code of Zoological Nomenclature (1999). ETYMOLOGY.— Panyaen-, from the word ‘panya-en’ which means ‘enchanted’ in the lan- guage of the Pala’ wan (one of seven ethnic groups that inhabit the Palawan islands). A resident of Palawan interviewed by ACD during forest surveys in November, 2003 referred to a S. leytensis as a ‘panya-en’, because it is apparently a favorite pet of forest spirits; -emys, turtle. PHYLOGENETIC DEFINITION.— We define Panyaenemys as the most inclusive clade containing Siebenrockiella |orig. Heosemys| leytensis (Taylor 1920) but not Siebenrockiella (orig. Emys| cras- sicollis (Gray 1831). DIFFERENTIAL DIAGNOSIS.— The sole known member of the subgenus Panyaenemys can be distinguished from all other turtles by the combination of the following four characters: 1) Strongly projecting epiplastra; 2) “Ginkgo leaf’-shaped vertebral scales 2-4; 3) No temporal arch in the skull; 4) Light lines on the back of the head. MATERIAL EXAMINED HEOSEMYS LEYTENSIS.— PHILIPPINES: CAS 60930, the neotype specimen discussed in detail by Buskirk (1989) and Diesmos et al. (2004a); PHILIPPINES: PNM 8488, GenBank accession numbers = AY954911 (cytb) and AY954914 (R35), a dead specimen that was donated to the National Museum of the Philippines (PNM) by a wildlife collector in 2003 and examined by Diesmos et al. (2004b). The specimen was apparently bought from a wildlife trader from Palawan. An additional 47 live specimens of H. leytensis in captivity were examined by Diesmos et al. (2004b) comprised of 34 turtles on Palawan being held by a private collector and 13 government-confiscated turtles housed at the Widlife Rescue Center of the Protected Areas and Wildlife Bureau, Quezon City, Philippines. SIEBENROCKIELLA CRASSICOLLIS.— CAMBODIA: FMNH 259055 (see Stuart and Platt 2004), GenBank accession numbers = AY954912 (cytb) and AY954913 (R35), Koh Kong Province, Sre Ambel District, Prek Kroch River (tributary of Sre Ambel River), 11°06’20”N 103°39’35”E, <10 m elevation, in a flooded paddy at edge of mangrove and Melaleuca forest, captured by a fisherman in bamboo fish trap set at that location, 27 August 2000, B.L. Stuart and S.G. Platt; UNCERTAIN/ MALAYSIA?: FMNH 224070, Perak, Batu Gajah, purchased in a pet shop, 19 November 1975, E.O. Moll. ACKNOWLEDGMENTS We thank the Protected Areas and Wildlife Bureau of the Department of Environment and Natural Resources of the Government of the Philippines for issuing a Memorandum of Agreement to conduct biological research in the Philippines and for facilitating collecting and export permits for this and related studies. We are indebted to Rogelio Sison (PNM) for providing access and a tissue sample of the S. /eytensis specimen at PNM. Field surveys of S. leytensis (between 2001 and 2003) were made possible through funding and technical support provided by the BP Conservation DIESMOS ET AL.: PHYLOGENETIC POSITION OF HEOSEMYS LEYTENSIS 39 Programme, Royal Melbourne Zoological Gardens, North of England Zoological Society, Conservation International Philippines, Turtle Conservation Fund, Fauna & Flora International, and the National Museum of the Philippines. ACD and RMB are indebted to colleagues at “Herp Watch Palawan 2001” especially Mae Leonida-Diesmos, Genevieve Gee-Das, Carlos Infante, Nonito Antoque, Jude Dimalibot, Jason Diesmos, and Jeffrey Diesmos. We also thank Peter Widmann, Indira Widmann, Siegfred Diaz, Deborah Villafuerte, Rolito Dumalag, Snapper Poche (Philippine Cockatoo Conservation Project) and Sabine Schoppe (Western Philippines University) for their various assistance in the field. Financial support to JEP was from a National Science Foundation Postdoctoral Fellowship. Financial support to BLS was provided by The John D. and Catherine T. MacArthur Foundation (with Harold Voris and Robert Inger). The opportunity for BLS to collect the sample of Siebenrockiella crassicollis 11 Cambodia was made possible by the Wildlife Conservation Society/Ministry of Agriculture, Forestry and Fisheries/Ministry of Environment Collaborative Program. JFP thanks Daniel G. Mulcahy (Utah State) and Ted Papenfuss (University of California, Berkeley) for technical assistance regarding the phylogenetic analyses when he was “Bayesed and confused’ and also Jens Vindum (CAS) for facilitating his examinations of the H. leytensis neotype. Sarah Drasner photographed the Field Museum carapace of S. crassicollis. Sequencing was con- ducted in The Field Museum’s Pritzker Laboratory for Molecular Systematics and Evolution oper- ated with support from the Pritzker Foundation. This work is LBNL-57255 and was performed under the auspices of the U.S. Department of Energy, Office of Biological and Environmental Research. This is University of California Museum of Paleontology Contribution # 1878. LITERATURE CITED ANDERSON, J. 1875. Description of some new Asiatic mammals and Chelonia. Annals and Magazine of Natural History, ser 4, 16:282—285. BOULENGER, G.A. 1903. Report on the batrachians and reptiles. Pages 131-176 in N. Annandale and H.C. Robinson, eds., Fasciculii Malayaenseis, Anthropological and Zoological Results of an Expedition to Perak and the Siamese Malay States. 1901-1902, Undertaken by Nelson Annandale and Herbert C. Robinson. Vol 1, Zoology. University Press, Liverpool, UK. Brown, R. M., AND A.C. Diesmos. (2001) 2002. Application of lineage-based species concepts to oceanic island frog populations: the effects of differing taxonomic philosophies on the estimation of Philippine bio- diveristy. The Silliman Journal 42:133—162. Brown, R.M., AND S.I. GUTTMAN. 2002. Phylogenetic systematic of the Rana signata complex of Philippine and Bornean stream frogs; reconsideration of Huxley’s modification of Wallace’s Line at the Oriental- Australian faunal zone interface. Biological Journal of the Linnean Society 76:393—-461. Brown, W.C., AND A.C. ALCALA. 1978. Philippine Lizards of the Family Gekkonidae. Silliman University Press, Dumaguete City, Philippines. 146 pp. BuskirK, J.R. 1989. A third specimen and neotype of Heosemys leytensis (Chelonia: Emydidae). Copeia 1989:224-227. DAUDIN, F.M. 1802. Histoire Naturelle, Générale et Particuliére des Reptiles, vol.2. Dufart, Paris, France. 342 pp- Diesmos, A.C., G.V.A. GEE, M.L. DiesmMos, R.M. Brown, P.J. WIDMANN, AND J.C. DIMALIBoT. 2004a. Rediscovery of the Philippine forest turtle Heosemys leytensis (Chelonia: Bataguridae), from Palawan Island Philippines. Asiatic Herpetological Research 10:22-27. Diesmos, A.C., R.V. Sison, M. DG. PEDREGOSA, AND M.J.C. CEnIzA. 2004b. The Conservation Status of Heosemys leytensis Taylor, 1920. Technical report submitted to National Museum of the Philippines, Fauna & Flora International, Royal Melbourne Zoological Gardens, North of England Zoological Society, Conservation International Philippines, and Turtle Conservation Fund. Manila, Philippines. 31 pp. 40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 3 ERNST, C.H., AND R.W. BARBOUR. 1989. Turtles of the World. Smithsonian Institution Press, Washington, DC, USA. 290 pp. Evans, B.J., R.M. Brown, J.A. McGuire, J. SUPRIATNA, N. ANDAYANI, A. DiEsmMos, D. ISKANDAR, D.J. MELNICK, AND D.C. CANNATELLA. 2003. Phylogenetics of fanged frogs: testing biogeographical hypothe- ses at the interface of the Asian and Australian faunal zones. Systematic Biology 52:794-819. FELDMAN, C.R., AND J.F. PARHAM. 2004. Molecular systematics of Old World stripe-necked turtles (Testudines: Mauremys). Asiatic Herpetological Research 10:28—37. Fusita, M.K., T.N. ENGsTROM, D.E. STARKEY, AND H.B. SHAFFER. 2004. Turtle phylogeny: insights from a novel nuclear intron. Molecular Phylogenetics and Evolution 31:1031—1040. Gray, J.E. 1825. A synopsis of the genera of reptiles and amphibia, with a description of some new species. Annals of Philosophy 10:193—217. Gray, J.E. 1831. Synopsis Reptilium, Part. 1, Cataphracta Crocodiles, Enaliosaurians. Treuttel, Wurtz, London, UK. 85 pp. Gray, J.E. 1855. Catalogue of Shield Reptiles British Museum, Pt. 1, Testudinata (Tortoises). Taylor and Francis, London, UK. 79 pp. Gray, J.E. 1860. Description of a new species of Emys lately living in the gardens of the Zoological Society. Proceedings of the Zoological Society of London 1860:232—233. Gray, J.E. 1870. Supplement to the Catalogue Collection of the British Museum, Pt. 1, Testudinata. Taylor and Francis, London, UK. 120 pp. Gray, J.E. 1873. On a new freshwater tortoise from Borneo (Orlitia borneensis). Annals and Magazine of Natural History, ser. 4, 11:156—157. HEANEY, L.R. 1986. Biogeography of the mammals of Southeast Asia: Estimates of colonization, extinction, and speciation. Biological Journal of the Linnean Society 28:127-—165. HIRAYAMA, R. 1985. Cladistic analysis of batagurine turtles. Studia Palaeocheloniologica 1:140-157. HUELSENBECK, J.P, AND F. RoNQuist. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17:754—755. INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE, 4TH ED. 1999. International Trust for Zoological Nomenclature, London, UK. 306 pp. IVERSON, J.B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. Privately pub- lished, Richmond, Indiana, USA. 363 pp. Joyce, W.G., AND C. BELL. 2004. A review of the comparative morphology of extant testudinoid turtles (Reptilia: Testudines). Asiatic Herpetological Research 10:53—109. Joyce, W.G., J.F. PARHAM, AND J.A. GAUTHIER. 2004. Developing a protocol for the conversion of rank-based taxon names to phylogenetically defined clade names, as exemplified by turtles. Journal of Paleontology 78(5):989-1013. LINDHOLM, W.A. 1929. Revidiertes Verzeichnis der Gattungen der rezenten Schildkroten nebst Notizen zur Nomenklatur einiger Arten. Zoologischer Anzeiger 81:275—272. MApDISON, D.R., AND W.P. MApDDISON. 2000. MacClade 4.0. Sinauer Associates, Inc. Sunderland, Massachusetts, USA. McCorp, W.P., J.B. IVERSON, AND BOEApI. 1995. A new batagurid turtle from northern Sulawesi, Indonesia. Chelonian Conservation and Biology 1(4):311-316. McCorp, W.P., J.B. IVERSON, P.Q. SPINKS, AND H.B. SHAFFER. 2000. A new genus of geoemydid turtle from Asia. Hamadryad 25:20-24. McDoweE LL, S.B. 1964. Partition of the genus Clemmys and related problems in the taxonomy of the aquatic Testudinidae. Proceedings of the Zoological Society of London 143:239-279. McGuire, J.A., AND A.C. ALCALA. 2000. A taxonomic revision of the flying lizards (Iguania: Agamidae: Draco) of the Philippine Islands, with a description of a new species. Herpetological Mononographs (14):81-138. MERREM, B. 1820. Versuch eines Systems der Amphibien. Johann Christian Krieger, Marburg, Germany. 191 Pp- PARHAM, J.F., AND C.R. FELDMAN. 2002. Generic revisions of emydine turtles. Turtle and Tortoise Newsletter 6:28-30. DIESMOS ET AL.: PHYLOGENETIC POSITION OF HEOSEMYS LEYTENSIS 4] PARHAM, J.F., W.B. SIMISON, K.H. KOZAK, C.R. FELDMAN, AND H. Sut. 2001. New Chinese turtles: endangered or invalid? Animal Conservation 4:357-367. Posaba, D., AND K.A. CRANDALL. 1998. ModelTest: Testing the model of DNA substitution. Bioinformatics 14:817-818. SmitH, M.A. 1916. A list of the crocodiles, tortoises, and turtles and lizards at present known to inhabit Siam. Journal of the Natural History Society of Siam 2:48-57. SPINKS, P.Q., H.B. SHAFFER, W.P. MCCorD, AND J.B. IVERSON. 2004. Phylogenetic hypotheses for the turtle family Geoemydidae. Molecular Phylogenetics and Evolution 32:164—-182. STUART, B.L., AND S.G. PLattr. 2004. Recent Records of Turtles and Tortoises from Laos, Cambodia, and Vietnam Asiatic Herpetological Research 10:129-150. SWOFFORD, D.L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts, USA. TayLor, E.H. 1920. Philippine turtles. Philippine Journal of Science 16:111—144. THEOBALD, W. 1868. Catalogue of reptiles in the museum of the Asiatic Society of Bengal. Journal of the Asiatic Society 1868:8-88. TIMMERMANN, W.W., AND D.L. AuTH. 1988. Geographic distribution, Heosemys leytensis. Herpetological Review 19(1):21. WIDMANN, P. 1998. A Guide to the Ecosystems of Palawan Philippines. VisCA-GTZ and Times Edition, Singapore. 118 pp. Wink, M., D. GUIKING, AND U. Fritz. 2001. Molecular evidence for hybrid origins of Mauremys iversoni Pritchard and McCord, 1991, and Mauremys pritchardi McCord, 1997 (Reptilia: Testudines: Bataguridae). Zoologische Abhandlungen, Staatliches Museum fiir Tierkunde Dresden 52:4\1—49. VAN Duk, P.P., B.L. STUART, AND A.G.J. RHODIN, EDS. 2000. Asian Turtle Trade: Proceedings of a Workshop on Conservation and Trade of Freshwater Turtles and Tortoises in Asia. Chelonian Research Monographs, No. 2. Chelonian Research Foundation, Lunenburg, Massachusetts, USA. 164 pp. ZANGERL, R. 1948. The methods of comparative anatomy and its contribution to the study of evolution. Evolution 2:351—374. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 4, pp. 42-52, 4 figs., 1 table [Appendix] April 20, 2005 A New Species of Rhacophorus (Anura: Rhacophoridae) from Myanmar (Burma) Jeffery A. Wilkinson!3, Thin Thin2, Kyi Soe Lwin?, and Awan Khwi Shein2 'Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103, Email: jwilkinson@ calacademy.org; 2Nature and Wildlife Conservation Division, Forest Department, Ministry of Forestry, Bayintnaung Road, West Gyogone, Insein, Yangon, Myanmar; 3H. T. Harvey & Associates, 3150 Almaden Expressway, Suite 215, San Jose, CA 95118 A new species of the genus Rhacophorus is described from Myanmar. The new species is most similar to R. bipunctatus but differs in the male having a larger body size, a bright green dorsal coloration, yellow in the outer portion of the iris, fainter crossbands on the limbs, a more extensive dermal fringe along the arm, more exten- sive projection on the heel, more extensive webbing on the hand, and typically two large equal-sized black spots, one in the axillary region and one on the middle of the flank. Ten species of Rhacophorus are thought to occur in Myanmar (R. appendiculatus (Giinther), R. bipunctatus Ahl, R. bisacculus Taylor, R. dennysi Blanford, R. feae Boulenger, R. maximus Giinther, R. reinwardtii (Schlegel), R. taronensis Smith, R. turpes Smith, and R. verrucosus Boulenger). Here, we report on an additional species of Rhacophorus collected during expeditions in 2001 and 2002 to Rakhine State in western Myanmar near the Bay of Bengal and to Kachin State in northeastern Myanmar near the Chinese border (Fig. 4). METHODS AND MATERIALS Specimens were collected by hand, euthanized, tissue samples removed, then fixed in 10% buffered formalin before preserving in 70% ethanol. Latitude and longitude were recorded with a Garmin 12 GPS, datum WGS84. Specimens are housed in the Department of Herpetology, California Academy of Sciences (CAS) and the Division of Amphibians and Reptiles, National Museum of Natural History, Smithsonian Institution (USNM). The preserved specimens were examined, measured, and compared with available specimens (see material examined; museum acronyms follow Leviton et al. [1985]) and published descrip- tions of currently recognized (Frost 2004) species of Rhacophorus and Polypedates from Myanmar and neighboring countries (Boulenger 1920; Smith 1924; Smith 1940; Bourret 1942; Liu and Hu 1961; Taylor 1962; Inger 1966; Berry 1975; Dring 1983; Inger et al. 1985; Sarkar and Sanyal 1985; Inger and Dutta 1986; Kiew 1987; Daniel and Sekar 1989; Yang et al. 1991; Brown and Alcala 1994; Manthey and Grossman 1997; Inger and Stuebing 1997; Iskandar 1998; Chan-ard et al. 1999; Fei 1999; Inger et al. 1999; Das 2000; Ohler et al. 2000; Vasudevan and Dutta 2000; Orlov et al. 2001; Ziegler and Kohler 2001; Chanda 2002; Harvey et al. 2002; Malkmus et al. 2002). Measurements were taken using dial calipers to the nearest 0.1 mm as follows: snout-vent length (SVL, from tip of snout to vent); head length (HL, from tip of snout to hind border of angle of jaw); 42 WILKINSON ET AL.: NEW RHACOPHORUS FROM MYANMAR 43 head width (HW, width of head at its widest point); internarial distance (IND, distance between nares); interorbital distance (IOD, minimum distance between upper eyelids); snout length (SL, from anterior border of eye to tip of snout); distance from nostril to eye (DNE, from nostril to ante- rior border of eye); forelimb length (FLL, from elbow to tip of third finger); hand length (HAL, from base of outer palmer tubercle to tip of third finger); thigh length (THL, from vent to knee); tibia length (TIL, from knee to foot); foot length (FL, from proximal end of metatarsal tubercle to tip of fourth toe); width of disk of third finger (3FDW, greatest horizontal width); and width of disk of fourth toe (4TDW, greatest horizontal width). SPECIES DESCRIPTION Rhacophorus htunwini Wilkinson, Thin Thin, Kyi Soe Lwin, and Awan Khwi Shein, sp. nov. Figs. 1-3. Htun Win’s treefrog DIAGNosis.— Rhacophorus htunwini can be distinguished from all other species of Rhacophorus and Polypedates by the following combination of characters: intermediate body size in the male (SVL 37.8-50.4 mm); extensive yellow webbing between fingers; thick dermal fringe on forearm and foot, dermal projection (calcar) on heel; squared-off supracloacal fold, snout point- ed: bright green dorsal color (slate blue in alcohol); yellow in the outer portion of the iris; very faint crossbands on the limbs; reddish-orange foot webbing; two large black spots on each side of body, one in the axillary region and one at the middle of the flank. Ho.LotyPe.— CAS 229893 (Fig. 1), an adult male, collected from Nagmung Township, Au Yin Ga Camp (27°17'36.9’N, 97°51’45.3”E), Putao District, Kachin State, Myanmar, elevation approximately 878 m, collected on 2 May 2002 by Htun Win, Young Ngai Thi Na, Ram Sar, and Hpe Ram. DESCRIPTION OF HOLOTYPE.— An adult male with slender habitus and head slightly longer than wide, 36% of SVL; snout pointed in dorsal view, gently slopes in lateral view to nostrils, then becomes slightly spatulate extending beyond mandible (Figs. la, c); nostrils closer to eye than tip of snout and anteriorly protuberant; canthus distinct, rounded, and inwardly curved; medial rostral areas between eyes and nostrils and between nostrils and tip of snout slightly concave; lores con- cave to nostril; eye directed anterolaterally with horizontal pupil. Tympanum distinct and circular; weak supratympanic fold, curving ventrally from dorso-posterior edge of tympanum to posterior edge of axilla. Vomerine processes with 8/6 rounded teeth respectively and approximately equal to transverse plane, separated medially by a space equal to two times their width, and laterally in contact with anteromedial edge of choanae; choanae small, ovoid to a medial point, and wholly visible at edge of lingual shelves of maxillae when viewed ventrally; tongue deeply bifurcates posteriorly; paired vocal slits oval and lateral to tongue. Dorsal body surface smooth; ventral abdominal surface and region below thighs areolate; pec- toral and gular regions much less areolate. Vent protrudes posteriorly, squared supracloacal fold medially notched. Arms short and slender; hand 73% as long as foot; when adpressed, relative length of fingers is3>4>2> 1: tips of fingers rounded; digital pads on hands and feet well developed and oval, with circummarginal grooves; distal phalanges bifurcate (as seen from dorsal aspect of digital tips). Hands extensively webbed, webbing formula for digits is I2-2111-1111-11V following Myers and Duellman (1982); narrow dermal fringe extends along lateral margin of fourth finger to base of hand. Subarticular tubercles between penultimate and adjoining proximal phalange round and well developed; proximal subarticular tubercles on finger 3 smallest; right hand with one, three, and two 44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 4 Ficure 1. (A) Dorsal and (B) ventral views of the body, (C) lateral view of the head, and ventral views of the (D) left hand and (E) left foot of the holotype of Rhacophorus htunwini sp. nov. (CAS 229893). small supernumerary tubercles in a row between proximal tubercle and base of hand on fingers two, three, and four, respectively; left hand with less obvious supernumerary tubercles; thenar tubercle low, extends medially at base of first finger, palmar tubercle absent. Thick dermal flange extends from lateral base of fourth finger to elbow, at widest approximately 18% of width of fore- arm. Hind limbs moderately long and slender; when adpressed to body, tibiotarsal articulation reaches beyond anterior edge of eye: webbing on foot reaches to base of pads on all toes; when adpressed, relative length of toes is 4>5 =3>2> 1: thick dermal fringe from base of pad of toe 5 extends along lateral edge of tarsus to heel where it develops into broad flange with laterally pro- jecting calcar on lateral edge of heel. A single subarticular tubercle on toes 1 and 2, two subartic- ular tubercles on toes 3, 4, and 5; proximal tubercle on toe 5 approximately same size as distal tubercle; supernumerary tubercles and an outer metatarsal tubercle absent; inner metatarsal tuber- cle flat, oval, and pointed medially. WILKINSON ET AL.: NEW RHACOPHORUS FROM MYANMAR 45 Coloration in preservative (Fig. 1). Dorsal color of body slate blue, extending laterally and ventrally '/4 of flanks and limbs, from elbow to halfway up middorsal aspect of hindarm, and along dorsal aspect of forearm, lateral fringe, and lateral half of fourth finger to base of pad, and from vent to knee along middorsal aspect of thigh, entire dorsal aspect of tibia, and lateral half of foot, lateral fringe, and fifth toe to just short of base of pad. Lateral margins of tarsal and supracloacal fringes light cream; first to third fingers and webbing, medial side of forearm, and all but middor- sal distal half of hindarm cream yellow; similarly, all but middorsal aspect of thigh, lateral and medial sides of tibia, medial half of foot, and first through fourth toes cream yellow. Webbing between toes two through five with streaks of orange, giving an orange appearance, but cream yel- low between toes one and two; small, elongated patch of slate blue on lateral side of second pha- lange of fourth toe and within webbing at same position between fourth and fifth toes; venter and flanks cream yellow. Posterolateral fringe of lower jaw and midventral aspect of fifth toe with some slate blue pigmentation; cream yellow margin on upper lip. Two large black oval spots on sides in axillary and mid-flank; axillary spot smaller, 8.7 mm horizontal diameter, mid-flank spot larger, 10.8 mm horizontal diameter. Color in life based on a color transparency (Fig. 2a). Dorsum bright green with sparsely scat- tered black and white pin-sized spots. Dorsal surface of fingers one through three, toes one through four, all digital pads, webbing on hand, lateral margin of fringe from foot to heel, ventrum, and sides yellow to yellowish orange. Two large jet black spots, with clusters of light blue spots on the dorsal margins, on axillary and mid-flank. Faint crossbands present on dorsal aspect of thigh and tibia. Pupil surrounded by light grayish brown horizontally rectangular iris, with yellow above and below, more so above. Yellow color extends posterior onto interior of orbit. Black thin line sur- rounds eye at margin with eyelid. VARIATION.— The holotype, paratypes, and referred specimens of R. htunwini are male, sex- ual dimorphism could not be determined. The paratypes and referred specimens are similar to the holotype except for the following. CAS 222065 and 222136 have smaller calcars than holotype. CAS 222065 is much smaller (SVL 37.8 mm), and specimen CAS 222136 has one large black spot in axillary region, without posterior spot on flank. Supratympanic fold of CAS 221351 strong on both sides of head and covers dorsoposterior edge of tympanum, but does so only on left side of CAS 222065, in all other specimens supratympanic fold similar to holotype. The dorsum of CAS 221351 is lighter in coloration than other specimens in alcohol, although color pattern is same, however darker bluish green in life. CAS 221351 has scattered black pin-size spotting on head, dor- sum, and dorsal aspect of thighs. Snout of this specimen rounded instead of pointed as in all other specimens, and tips of pads more squared off than other specimens. ETYMOLOGY.— The name /itunwini is given in honor of the late U Htun Win, who devoted the last eight years of his life to the pursuit of knowledge of the diversity and natural history of the rep- tiles and amphibians within his country. As team leader of the Myanmar Herpetological Survey Team, he first recognized this frog as potentially new to science. COMPARISONS.— Because R. htunwini is a medium size tree frog that possesses expanded discs on the fingers and toes, an intercalary element between the penultimate and terminal pha- langes, a narrow bony metasternum, a flange on the distal end of the third metacarpal, Y-shaped terminal phalanges, and extensive webbing between the fingers and toes, it has been placed with- in Rhacophorus (Wilkinson and Drewes 2000). Rhacophorus is a relatively large genus of approx- imately 60 species from Asia (Frost 2004), and members of this genus closely resemble the approx- imately 28 species of the genus Polypedates (Liem 1970). Because of this close resemblance, sev- eral species have been moved back and forth between the two genera, or Polypedates has not been 46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES recognized as a genus separate from Rhacophorus (Dubios 1986; Fei 1999). Recently, Wil- kinson et al. (2002) provided molecular evidence to separate the genera Polypedates and Rhacophorus, and to move two species in Polypedates (P. den- nyst and P. prasinatus) back into Rhacophorus. We believe that generic level reversals are still required in order to ensure the correct taxonomic placement of many species within these two genera and therefore have includ- ed members of both genera in the comparisons below. Following the taxonomic designations in Frost (2004), R. htunwini can be distinguished from other species of the follow- ing Rhacophorus and Poly- pedates from Bangladesh, Cam- bodia, China, India, Indonesia, Laos, Malaysia, Myanmar, Thai- land, and Vietnam as follows: from members of Polypedates in the P. leucomystax species group (P. colleti, P. cruciger, P. eques, P. leucomystax, P. macrotis, P. maculatus, P. mutus, P. otilo- phus, and P. zed), P. insularis, P. megacephalus, P. naso, P. pseu- docruciger, members of Rha- cophorus in the subgenus Rha- cophorus in the R. appendicula- tus species group (R. appendicu- Volume 56, No. 4 A FiGuRE 2. Photos in life of (A) a male Rhacophorus htunwini sp. nov. (CAS 229893), (B) a female R. bipunctatus (CAS 229902), and (C) a male R. bipunctatus (CAS 224676). latus, R. bisacculus, R. verrucopus), R. baliogaster, R. baluensis, R. barisani, R. bimaculatus, R. calcaneus, R. catamitus, R. cyanopunctatus, R. exechopygus, R. margaritifer, R. modestus, R. nam- daphaensis, R. orlovi, R. poecilonotus, R. translineatus, R. tuberculatus, and R. verrucosus by a bright green dorsal color; from members of Polypedates in the P. chenfui species group (P. chen- fui, P. hungfuensis, and P. yaoshanensis), the P. dugritei species group (P. dugritei, and P. omei- montis), P. dorsoviridis, P. duboisi, P. nigropunctatus, P. pingbianensis, P. puerensis, P. zhaojuen- sis, members of Rhacophours in the subgenus Rhacophorus and the R. dennysii species group (R. dennysi and R. feae), the R. pardalis species group (R. annamensis, R. notator, R. pardalis, and R. robinsonii), R. achantharrhena, R. angulirostris, R. taronensis, and R. variabilis by the presence of a sharp dermal calcar at the heel; from members of Rhacophorus in the subgenus Rhacophorus and WILKINSON ET AL.: NEW RHACOPHORUS FROM MYANMAR 47 FiGuRE 3. Dorsal view of representative specimens of Rhacophorus htunwini sp. nov. (top row), representative speci- mens of female R. bipunctatus (middle row), and representative specimens of male R. bipunctatus (bottom row). the R. malabaricus species group (R. calcadensis and R. malabaricus), R. lateralis, R. pseudoma- labaricus, and R. turpes by axillary spots. Rhacophorus htunwini closely resembles members of the R. reinwardtii species group in the subgenus Rhacophorus (R. bipunctatus, R. dulitensis, R. maximus, R. nigropalmatus, R. promi- nanus, R. reinwardtii) and R. hoangliensis, but can be distinguished from all but R. bipunctatus and R. reinwardtii by the presence of axillary spots. It can be distinguished from R. reinwardtii (SVL male 68 mm) by its smaller size and yellow hand webbing (black hand webbing in R. reinwardtii). Rhacophorus htunwini most closely resembles R. bipunctatus, which also has axillary spots, dermal calcars at the heels, a pointed snout, and sometimes a green dorsal color (Fig. 2b). However, it can be distinguished from R. bipunctatus by size (Table 1, Fig. 3); the males of R. htunwini are larger (average SVL =45.7 mm) than the males of R. bipunctatus (average SVL = 34.9 mm). Rhacophorus htunwini also has a bright green dorsal color in life that becomes slate blue when pre- served, whereas some members of R. bipunctatus have an olive green dorsal color in life that becomes light bluish gray or brown when preserved and others are orange to tan in life with a dark- er brown pattern (blotching or an X mark) on the dorsum, which become brown when preserved (Figs. 2-3). The eye of R. htunwini contains yellow at the upper and lower portion of the iris, which is absent in R. bipunctatus (Fig. 2). Rhacophorus bipunctatus has distinct crossbands on the fore and hindlimbs (Figs. 2b-c), whereas R. htunwini has faint crossbands that disappear in alcohol (Fig. 2a). Rhacophorus htunwini has a more extensive dermal fringe on the forearm, a more exten- sive dermal calcar at the heel, and more extensive webbing on the hand than the male of R. bipunc- tatus. In all but one specimen, R. htunwini has two large black equal size spots on the sides, one in 48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 4 the axillary region and one on the flank, whereas, the major- ity of male specimens of R. bipunctatus examined had only one spot in the axillary region, and in specimens that had two spots the posterior spot was much smaller than the ante- rior spot. DISTRIBUTION AND NATURAL HISTORY.— At present Rhacophorus htunwini is known from Nagmung and Machanbaw Townships, Putao District, Kachin State, and from much further south in Rahkine State in the southwest- ern foothills of Rakhine Yoma, Gwa Township, and Kyauktaw Township, Sittawe District (Fig. 4). This distribu- tional pattern indicates that this species may be restricted to the Indo-Burman Mountain Range that arcs from southwest- ern Myanmar along the border with India, and the Eastern Himalayas in northern Myanmar. The absence of specimens from the Chin Hills of western Myanmar is probably due solely to a lack of surveys in the region. The type specimens including the holotype (CAS 229913, USNM 561869) were found approximately 2 m off the ground in bamboo. Referred specimens were found in undisturbed habitat near a spring (CAS 222136) or seasonal (CAS 221351) and permanent (CAS 222065) streams. Other species of Polypedates and Rhacophorus found in the vicinity of the type locality were P. leucomystax, R. bipunctatus, and R. dennysi. kilometers bese Ss FiGure 4. Distribution of Rhacophorus htunwini sp. nov. in Myanmar with type locality indicated by a star (at tip of arrow). MATERIAL EXAMINED Rhacophorus htunwini (paratypes): CAS 229913, USNM 561869 adult males collected at the same local- ity and date as the holotype. Rhacophorus htunwini (referred specimens): CAS 221351, an adult male, collected between Ahtan Ga and Au Rin Ga (27°15’27.2’N, 97°50’32.4”E), Ma Chan Baw Township, Putao District, Kachin State, Myanmar, collected on 4 September 2001 by Htun Win and Ran Shaung; CAS 222136, an adult male, collect- ed from Yea Pu Camp (17°56’02.6’N, 94°38’02.9”E), Gwa Township, Rakhine State, Myanmar, collected on 8 June 2001 by Hla Tun, Kyi Soe Lwin, and Awan Khwi Shein; CAS 222065, an adult male, collected from Pin Lone Camp, Pe Chaung, near Saba Sate Village (21°00°54.9”N, 92°52’06.6”E), Kyaut Taw Township, Sittawe District, Rakhine State, Myanmar, collected on 4 July 2001 by Htun Win, Kyi Soe Lwin, and Awan Khwi Shein. Polypedates chenfui: FMNH 232963, 232964 (China). Polypedates colletti: FANH 234773, 235631 (Malaysia). Polypedates cruciger: CAS 85280 (Sri Lanka). Polypedates dugritei: CAS 64273 (China). Polypedates eques: CAS 85281, 85282 (Sri Lanka). Polypedates leucomystax: FMNH 239159 (Malaysia); FMNH 254649 (Lao PDR); CAS-SU 15163 (India); CAS 14943 (China); CAS 94573 (Bangladesh); CAS 103624 (Indonesia); CAS 105003 (Vietnam); CAS 105972 (Malaysia); CAS 111336 (Cambodia); CAS 172691 (Thailand); CAS 221962, 224461 (Myanmar). Polypedates macrotis: FMANH 239107, 239119 (Malaysia): CAS 60630, 60631, 60684, 60804, 62138 (Philippines); CAS 62581 (Malaysia); CAS 64074 (Philippines); CAS 64077 (Indonesia); CAS 64089-64092 (Philippines); CAS 105974, 105975 (Malaysia). Polypedates maculates: CAS 16922-16924 (Sri Lanka); CAS 94571, 94572, 104152, 125365—125370 (India). WILKINSON ET AL.: NEW RHACOPHORUS FROM MYANMAR 49 Polypedates megacephalus: ROM (field numbers) 18038, 18045 (Vietnam). Polypedates otilophus: FANH 230836, 239147 (Malaysia). Rhacophorus angulirostris: FANH 235035 (Malaysia). Rhacophorus annamensis: FMNH 253933, 253940 (Vietnam); ROM 29889, 29890, 29891, 29892, 29897, 29901, 29904 (Vietnam). Rhacophorus appendiculatus: CAS 60169-60174, 62261, 64078-64086 (Philippines). Rhacophorus bimaculatus: CAS 61840, 133178—133180, 133251 (Philippines). Rhacophorus bipunctatus: FMNH 253122, 253124 (Vietnam); NMNS 3220 (China); CAS 224676, 228808, 229887, 229889, 229890, 229898, 229899, 229901—229907, 229910 (Myanmar). Rhacophorus calcaneus: FANH 256456, 257933 (Lao PDR); ROM 29849, 29850, 29854, 29855, 29875, 29877, 29879, 29880 (Vietnam). Rhacophorus dennysi: FMNH 256449, 256450 (Lao PDR); ROM 29839, 29840, 29841, 29842, 29843, 29846, 30245 (Vietnam); CAS 64224 (China); CAS 221535, 224496, 224659 (Myanmar). Rhacophorus dulitensis: FMNH 235741 (Malaysia). Rhacophorus feae: FMNH 257910 (Lao PDR); CAS-SU 6387, 6388 (Vietnam). Rhacophorus gaunt: FMNH 235044, 239238 (Malaysia). Rhacophorus maximus: CAS 221516, 221517 (Myanmar). Rhacophorus microtympanum: CAS 85283 (Sri Lanka). Rhacophorus nigropalmatus: FMNH 230901, 230902 (Malaysia). Rhacophorus pardalis: FANH 235750 (Malaysia); FMNH 259530 (Philippines); CAS 60472-60476, 61386, 128725, 129267—126270 (Philippines). Rhacophorus reinwardtit: FANH 235034 (Malaysia), FMNH 255305 (Lao PDR); NMNS 3213 (China). Rhacophorus rufipes: FANH 231377 (Malaysia). Rhacophorus taronensis: BMNH 1947.2.8.17 (Myanmar). Rhacophorus turpes: BMNH 1947.2.8.69, 1947.2.8.70 (Myanmar). Rhacophorus verrucosus: CAS 224441, 224442, 224469, 224737, 224754, (Myanmar). ACKNOWLEDGMENTS We thank U Shwe Kyaw, Director General, Forest Department, Ministry of Forestry, and U Khin Maung Zaw, Director, Nature and Wildlife Conservation Division, Forest Department, Ministry of Forestry for their continued support of the Myanmar Herpetological Project. We also thank Barry Clarke, Mark Wilkinson, and Nick Arnold of the British Museum, Alan Resetar and Harold Voris of the Field Museum of Natural History, Wen-Hao Chou of the National Museum of Natural Science, Taiwan, and Robert Murphy and Ross MacCulloch of the Royal Ontario Museum for loaning specimens. Fieldwork was supported by National Science Foundation Grant (DEB- 9971861) to the late Joseph B. Slowinski (Alan E. Leviton, current PI) and George Zug. Michelle Koo provided Figure 1. Dong Lin provided the photographs for Figures 2 and 4, and Hla Tun pro- vided the photographs for Figure 3. Jens Vindum and Guinevere Wogan critically read and provid- ed valuable comments on the manuscript. LITERATURE CITED Berry, P.Y. 1975. The Amphibian Fauna of Peninsular Malaysia. Tropical Press, Kuala Lumpur, Malaysia 130 pp. BOULENGER, G.A. 1920. III. Reptiles and batrachians collected in Korinchi, West Sumatra, by Messrs. H.C. Robinson and C. Boden Kloss. Journal of the Federated Malay States Museums 8:285—306. Bourret, R. 1942. Les Batraciens de |’ Indochine. Memoires de L’ Institut Oceanographique de L’ Indochine. Gouvernement General de L’ Indochine. Hanoi, Indochina (Vietnam). 547 pp. Brown, W.C., AND A.C. ALCALA. 1994. Philippine frogs of the family Rhacophoridae. Proceedings of the California Academy of Sciences 48:185—220. 50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 4 CHANDA S.K. 2002. Hand Book of Indian Amphibians. Zoological Survey of India, Kolkata, India. 335 pp. CHAN-ARD, T., W. GROSSMANN, A. GUMPRECHT, AND K.-D. SCHULZ. 1999. Amphibians and Reptiles of Peninsular Malaysia and Thailand, An illustrated Checklist. Bushmaster Publications, Wuerselen. 240 pp. DANIEL, J.C., AND A.G. SEKAR. 1989. Field guide to the amphibians of western India. Part 4. Journal of the Bombay Natural History Society 86:194—200. Das, I. 2000. Nomenclatural history and rediscovery of Rhacophorus lateralis Boulenger, 1883 (Amphibia: Rhacophordae). Current Herpetology 19:35—40. DRING, J. 1983[1984]. Some new frogs from Sarawak. Amphibia-Reptilia 4:103—112. Dusois, A. 1986. Miscellanea taxinomica batrachologica (I). Alytes 5:17—95. Fer, L. 1999. Atlas of Amphibians of China. Publishing House for Scientific and Technological Literature, Hunan, China. 432 pp. Frost, D.R. 2004. Amphibian Species of the World: an Online Reference. Version 3.0 (22 August, 2004). Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/index.html. American Museum of Natural History, New York, USA. Harvey, M.B., A.J. PEMBERTON, AND E.N. SmitH. 2002. New and poorly known parachuting frogs (Rhacophoridae: Rhacophorus) from Sumatra and Java. Herpetological Monographs 16:46—92. INGER, R.F. 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana: Zoology 52:1-402. INGER, R.F., AND S.K. Dutta. 1986. An overview of the amphibian fauna of India. Journal of the Bombay Natural History Society 83(Suppl.):135—146. INGER, R.F., N.L. ORLOV, AND I.S. DAREVSKY. 1999. Frogs of Vietnam: a report on new collections. Fieldiana: Zoology 92:1-46. INGER, R.F., H.B. SHAFFER, M. KOSHY, AND R. BAKDE. 1985 [1984]. A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, south India. Journal of the Bombay Natural History Society 81:551-570. INGER, R.F., AND R.B. STUEBING. 1997. A Field Guide to the Frogs of Borneo. Natural History Publications, Kota Kinabalu. 205 pp. ISKANDAR, D.T. 1998. Amphibians of Java and Bali. Research and Development Centre for Biology, Jakarta, Indonesia. 117 pp. Krew, B.H. 1987. An annotated checklist of the herpetofauna of Ulu Endau, Johore, Malaysia. Malayan Nature Journal 41:413-423. LEVITON, A.E., R.H. GipsBs, JR., E. HEAL, AND C.E. DAwson. 1985. Standards in herpetology and ichthyolo- gy: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832. Liem, S.S. 1970. The morphology, systematics, and evolution of the Old World treefrogs (Rhacophoridae and Hyperoltidae). Fieldiana Zoology 57:1—145. Liu, C.C., AND S.Q. Hu. 1961. Tailless Amphibians of China. Science Press, Peking, China. 364 pp. Matckmus, R., U. MANTHEY, G. VOGEL, P. HOFFMANN, AND J. KosucH. 2002. Amphibians and Reptiles of Mount Kinabalu. A.R. G. Gantner Verlag Kommanditgesellschaft, Ruggel, 424 pp. MANTHEY, U., AND W. GROSSMANN. 1997. Amphibien and Reptilien Siidostasiens. Natur und Tier Verlag, Miinster, Germany. 511 pp. Myers, C.W., AND W.E. DUELLMAN. 1982. A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama. American Museum Novitates (2752):1—32. OHLER, A., O. MARQUIS, S. SWAN, AND S. GROSJEAN. 2000. Amphibian biodiversity of Hoang Lien Nature Reserve (Lao Cai Province, northern Vietnam) with description of two new species. Herpetozoa 13:71-87. OrLov, N.L., A. LATHROP, R.W. MurPHy, AND H.T. Cuc. 2001. Frogs of the family Rhacophoridae (Anura: Amphibia) in the northern Hoang Lien Mountains. Russian Journal of Herpetology 8:17-44. SARKAR, A.K., AND A.K. SANYAL. 1985. Amphibia. Records of the Zoological Survey of India 81:285-—295. SMITH, M.A. 1924. New tree-frogs from Indo-China and the Malay Peninsula. Proceedings of the Zoological Society, London 1924:225-234. SmitH, M.A. 1940. The amphibians and reptiles obtained by Mr. Ronald Kaulback in upper Burma. Records of the Indian Museum 42:465-486. WILKINSON ET AL.: NEW RHACOPHORUS FROM MYANMAR 51 TayLor, E.H. 1962. The amphibian fauna of Thailand. University of Kansas Science Bulletin 43:265—599. VASUDEVAN, K., AND S.K. Dutta. 2000. A new species of Rhacophorus (Anura: Rhacophoridae) from the Western Ghats, India. Hamadryad 25:21-28. WILKINSON, J.A., AND R.C. DREwes. 2000. Character assessment, genus level boundaries, and phylogenetic analyses of the family Rhacophoridae: a review and present day status. Contemporary Herpetology 2000:2. WILKINSON, J.A., R.C. DREWES, AND O.C. Tatum. 2002. A molecular phylogenetic analysis of the family Rhacophoridae with an emphasis on the Asian and African genera. Molecular Phylogenetics and Evolution 24:265-273. YANG, D.T., S. Li, W. Lu, AND S. Lu. 1991. Amphibian Fauna of Yunnan, China. Forestry Publishing House, Beijing. 259 pp. ZIEGLER, T., AND J. KOHLER. 2001. Rhacophorus orlovi sp. n., a new tree frog from Vietnam (Amphibia: Anura: Rhacophoridae). Sauria 23:37-46. SZ PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 4 Appendix TABLE |. Measurements of the type series and referred specimens of Rhacophorus htunwini and comparative material of R. bipunctatus. Mean (in mm) followed by range (in parenthesis) and ratio of SVL (below). See text for abbreviations. Rhacophorus htunwini Male N=6 Rhacophorus bipunctatus Rhacophorus bipunctatus Female N=3 Male N= 16 SVL HL HW IND IOD SN DNE BEE HL THL AUG IRIE 3FDW 4TDW aS (SURV 9) 16.4 (13.5-17.7) 39.9 16.5 (14.9-17.5) 36 4.4 (4.3-4.9) 9.6 5.2 (4.8-5.9) 11.4 7.3 (6.1-8.1) 16 3.7 (3.1-4.2) 8 22.0 (18.6-25.1) 48.1] 13.8 (11.2-15.5) 30.3 22.5 (17.6-26.1) 49.3 23.0 (19.4-25.1) 50.3 5.4 1.9 (1.6-2.2) 4.2 Copyright © 2005 by the California Academy of Sciences 55.9 (51.2-60.2) 18.3 (17.2-19.8) S25; 17.9 (16.9-19.3) S251 4.5 (4.4-4.7) 8.1 6.6 (5.4-7.5) 11.8 8.0 (7.7-8.4) 14.2 3.6 (3.2-4.0) 6.4 27.9 (26.8-29.2) 49.8 17.6 (16.5-19.0) 31.4 27.0 (26.7-27.4) 48.2 26.2 (25.0-27.9) 46.9 25.9 (24.8-27.9) 46.3 3.2 (3.1-3.4) 5.8 2.4 (2.0-2.7) 4.3 San Francisco, California, U.S.A. 34.9 (32.0-37.7) 12.5 (11.5-14.0) 35.9 12.6 (11.4-14.1) 36 3.4 (2.7-4.2) 9.8 3.8 (2.9-4.5) 10.9 5.4 (4.6-6.0) 15.6 2.6 (2.1-2.9) 7.4 17.9 (16.1-19.7) 51.2 11.2 (10.0-12.5) 32 17.6 (14.3-19.7) 50.5 17.5 (15.6-18.7) 50.3 15.6 (13.9-16.9) 44.7 1.9 (1.5-2.4) 5.4 1.5 (1.2-1.9) 4.2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 5, pp. 53-65, 10 figs. April 29, 2005 New Taxa of Octocorals (Anthozoa: Octocorallia) from the Northeastern Pacific Ocean Gary C. Williams Department of Invertebrate Zoology and Geology, California Academy of Sciences, S875 Howard Street, San Francisco, California 94103, U.S.A, Email: gwilliams @ calacademy.org The octocorallian fauna of the Alaskan region is poorly known. Recently, several research projects have been initiated to explore the faunal elements of the continen- tal shelf in the Gulf of Alaska. A wealth of material, including octocorals (mostly gor- gonians with some pennatulaceans), has been collected by trawl nets during field research conducted by agencies such as the National Marine Fisheries Service (NMFS) of the National Oceanic and Atmospheric Administration (NOAA). A new genus of gorgonian has recently been described from the Aleutian Islands as Alaskagorgia Sanchez and Cairns, 2004. The present paper reports the discovery of an additional plexaurid gorgonian from the same region, and provides a description of it as a new genus and species. Additionally, an unusual new species of sea pen, genus Cavernularia (Family Veretillidae), is described from the central Aleutian island chain. The geographically nearest veretillid taxa outside of the type locality occur in the northwestern and eastern Pacific — Japan and Panama. According to Wing and Barnard (2004), approximately 64 species of octocorals are presently known to occur in Alaskan waters. The authors also provide a key to the known octocoral taxa and color figures for twenty species. Other kinds of coral also included are scleractinians, stylasterine hydrocorals, and antipatharians. The gorgonian family Plexauridae Gray, 1859 has relatively recently had a plethora of genera allocated to it along with the amalgamation of several groups at the ranks of family and subfami- ly. Included here are the Muriceidae, Paramuriceidae, Plexaurinae, and Stenogorgiinae (= Paramuriceinae) (Bayer 1981). The tropical western Atlantic genera were treated by Bayer (1961). Recent works have compiled previously described taxa and have added new generic names mak- ing a total of approximately 38 genera that are presently considered to comprise the family. These works include Grasshoff (1977) for the northern Atlantic Ocean and Mediterranean Sea, Grasshoff and Barbigant (2001) and Frabricius and Alderslade (2001) for the tropical western Pacific, and Sanchez and Cairns (2004) for Alaskan waters. The present paper describes an additional new genus and species from the Alaskan Aleutian Islands, making a total of 39 worldwide genera in the family Plexauridae. The new taxon does not closely resemble other plexaurid taxa. In addition, a new species of the veretillid pennatulacean genus Cavernularia is described, also from the Aleutian Archipelago. The only other known taxa of vertetillid pennatulaceans from the northwestern and eastern Pacific Ocean include several species from Japan, Panama, and the Galapagos Islands (L6pez-Gonzalez, Gili, and Williams 2000; Kiikenthal 1915; Hickson 1921; and Williams 1989, 1995). 53 54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 5 METHODS Material was collected on board a research vessel (the F/V “Vesteraalen” or the M/V “Sea Storm’’) using trawling gear, and was preserved in 75% ethanol. Sclerites were isolated from sur- rounding tissues by the use of sodium hypochlorite (household bleach). Micrographs were made using a Nikon Coolpix 990 digital camera, sometimes in combination with a Nikon SMZ-10 dis- secting microscope or an Olympus CH-2 compound microscope. Original drawings of sclerites were made with an Olympus CH-2 compound microscope and an Olympus drawing tube. Scanning electron micrographs were taken using a LEO 1400 series scanning electron microscope. Digital images for the making of plates were manipulated using Adobe Photoshop software. Abbreviations used in the text are CAS (California Academy of Sciences, San Francisco), NMFS (National Marine Fisheries Service), and NOAA (National Oceanic and Atmospheric Administration — of the United States Department of Commerce). SYSTEMATIC ACCOUNT Order Alcyonacea Lamouroux, 1816 Family Plexauridae Gray, 1859 REMARKS.— Most of the taxa allocated to this family have relatively large sclerites, over 0.3 mm long (up to a maximum length of approximately 5 mm), with tubercular sculpture rarely arranged in whorls. In addition, both Bayer (1981:924) and Fabricius and Alderslade (2001:59) use the following axial characteristics to describe the family. The plexaurid axis often appears woody and fibrous, often brown or darker. The central 5 ; core is wide, hollow, and cross-chambered. The surrounding axial cortex is soft, usually with hollow spaces called loculi that are sometimes filled with a fibrous substance or non-spicular calcite. In contrast, the family Gorgoniidae usually has small scerites, less than 0.3 mm long, with tubercles arranged in whorls. The axis is often dark brown or black. It has a nar- row core and the cortex is very dense with few or no locules. The new taxon described here is somewhat enigmatic in that it has small scerites, less than or equal to 0.15 mm in length, which is charac- teristic of the Gorgoniidae. Additionally, the tubercles of the spindles and radiates are some- times arranged in whorls, but often they are not. Regardless of the problematic nature of these features, the new taxon is here placed in the family Plexauridae because the axis has a FiGURE 1. Non-type specimen (CAS 164584) of woody or fibrous appearance and wide and hol- low central core, whereas the surrounding axial cortex is soft with easily observable amounts of non-spicular calcite (Fig. 1). Cryogorgia koolsae gen. and sp. nov. Transverse section of a branch near the apical end showing the axis and thin coenenchyme; scale bar = 3.8 mm; ca — calcitic material, em — coenenchyme, ex — axial cortex, he — hollow core. WILLIAMS: NEW OCTOCORALS FROM NORTHEASTERN PACIFIC OCEAN 55 Genus Cryogorgia Williams, gen. nov. TYPE SPECIES: Cryogorgia koolsae Williams, sp. nov., by monotypy DrAGNosis.— Branching relatively sparse, mostly lateral but rarely dichotomous, planar, anas- tomosis absent. Terminal branches elongate, somewhat clavate. Azooxanthellate. Axis proteina- ceous, flexible with wide hollow core, brown to dark brown, cylindrical near holdfast, slightly compressed in higher branches. Polyps monomorphic, retractile into polyp mounds or completely into coenenchyme, calyces absent. Coenenchymal sclerites leaf clubs, coarsely-tuberculated spin- dles, radiates, modified and often clavate radiates (some approaching wart clubs), and crosses. Sclerites colorless. Wet-preserved coenenchyme tan-white. ETyMOLOGyY.— The new generic name is derived from the Greek, Kryos (cold or chilly) and the commonly used suffix for a gorgonian octocoral, -gorgia; in reference to the cold nature of the type locality of this gorgonian. Cryogorgia koolsae Williams, sp. nov. Figs. 1-8 MATERIAL EXAMINED.— HOo_ortyPe: CAS 151364, Northeastern Pacific Ocean, United States, Alaska, Aleutian Islands, 52.25°N 171.70°W, 406 m depth, Haul 79, 11 June 2000, collected by E.J. Kools aboard F/V “Vesteraalen,’ NMFS Aleutian Survey 2000, one whole colony wet preserved in 75% ethanol. PARATYPE: CAS 150650, Northeastern Pacific Ocean, United States, Alaska, Aleutian Islands, 52.07° N 177.25° W, 86 m depth, Haul 146, 26 June 2000, collected by E.J. Kools aboard F/V “Vesteraalen,’ NMFS Aleutian Survery 2000, one whole colony wet preserved in 75% ethanol. OTHER MATERIAL: CAS 164584, Northeastern Pacific Ocean, United States, Alaska, Aleutian Islands, 51.75°N 175.67°W, 83 m depth, Haul 101, 7 July 2002, collected by R.J. Van Syoc aboard M/V “Sea Storm,” three whole colonies wet preserved in 95% ethanol. CAS 168893, Northeastern Pacific Ocean, United States, Alaska, Aleutian Islands, 52.31°N 173.65°W, 402 m depth, Haul 144, collected by R.J. Van Syoc aboard M/V “Sea Storm,” one whole colony wet preserved in ethanol. DESCRIPTION OF HOLOTYPE.— Growth form and size (Figs. 2—3): The holotype measures 370 mm long by 150 mm wide. The basal trunk is 115 mm in length before the first side branch arises. It averages approximately 10 wide and 8 mm deep, as it is slightly compressed. The branching is irregular (lateral) and sparse. The terminal branches are 30-100 mm in length. Polyps (Figs. 3A—E): The polyps of the holotype exhibit various states of retractility. Some are preserved totally retracted and flush with the surrounding coenenchyme — especially those along the main stem and along the thicker branches, whereas others are preserved in various stages of exsertion. The length of the exserted polyps is usually less than 2 mm. They vary in shape from mound-like and hemispherical to cone-shaped or somewhat cylindrical (Fig. 3E). These mounds or cones are thickly set with coenenchymal sclerites (Fig. 5). In all cases, the anthocodiae are at least partly retracted or hidden. Examination of dissected polyps has not shown the anthocodiae to con- tain sclerites. Sclerites (Figs. 2C, 4-7): Coenenchymal sclerites of the polyp-bearing branchs, branch tips, and polyp mounds are of several different forms: foliates or leaf clubs (0.07—0.12 mm in length); seven- and eight-radiates (0.07—0.10 mm long); clavate forms (0.08—0.15 mm in length); crosses (0.10—0.11 mm long); and modified radiates, cylinders, and spindles (0.07—0.15 mm in length). Anthocodial sclerites are apparently absent. Color (Figs. 2—3): The axis is dark brown to black. The coenenchyme and retracted polyps are grayish white to tan-white. The sclerites are colorless. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 5 FIGURE 2. A-B. Holotype of Cryogorgia koolsae gen. and sp. noy.; scale bars = 65 mm. C. Three leaf club sclerites from the surface coenenchyme of a terminal branch; scale bar = 0.05 mm. WILLIAMS: NEW OCTOCORALS FROM NORTHEASTERN PACIFIC OCEAN oy/ FIGURE 3. Holotype of Cryogorgia koolsae gen. and sp. nov.; details of branches and polyps. A—B. Details of branch- ing patterns. CD. Details of a single branch and polyps. E. Detail of a single polyp. Scale bars: A = 25 mm, B = 25 mm, C=5 mm, D=2 mm, E=1 mm. 58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 5 Ficure 4. Cryogorgia koolsae gen. and sp. nov. Coenenchymal sclerites from a polyp-bearing branch; scale bar = 0.10 mm. A. Holotype (CAS 151364). B. Paratype (CAS 150650). C. Non-type specimen (CAS 164584). D. Non-type specimen (CAS 168893). Scale bar = 0.10 mm. WILLIAMS: NEW OCTOCORALS FROM NORTHEASTERN PACIFIC OCEAN 59 FiGuRE 5 (above) Cryogorgia koolsae gen. and sp. noy. Coenenchymal sclerites from a mound formed by a retracted polyp: scale bar = 0.10 mm. FiGurRE 6 (below). Holotype of Cryogorgia koolsae gen. and sp. nov. Coenenchymal sclerites from the base of the holdfast; scale bar = 0.10 mm. 60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 5 FIGURE 7. Holotype of Cryogorgla koolsae gen. and sp. noy. Scanning electron micrographs of leaf clubs from the coenenchyme of a terminal branch. A. 0.08 mm. B. 0.11 mm. C. 0.08 mm. D. 0.08 mm. E. 0.09 mm. F. 0.08 mm. G. 0.07 mm. H. 0.09 mm. I. 0.08 mm. J. 0.10 mm. K. 0.08 mm. L. 0.06 mm. M. 0.10 mm. N. 0.12 mm. O. 0.10 mm. WILLIAMS: NEW OCTOCORALS FROM NORTHEASTERN PACIFIC OCEAN 61 ETyMOLOGyY.— This species is named for its discoverer, Elizabeth J. Kools, (Department of Invertebrate Zoology and Geology, California Academy of Sciences, San Francisco). DISTRIBUTION.— This new taxon is presently known from four localities in the central Aleutian Archipelago of Alaska, northern Pacific; 83-406 m depth (Fig. 8). RUSSIA Beaufort Sea Bering Sea CANADA * CAS 150650 C CAS 168893 av CAS 164584 al See CAS 151364 x Gulf of Alaska FiGuRE 8. Map of the Aleutian Island Chain showing collecting stations (with collection numbers) for Cryogorglia koolsae gen. and sp. nov. (); arrow shows type locality. Order Pennatulacea Verrill, 1865 Family Veretillidae Herklots, 1858 Genus Cavernularia Valenciennes in Milne-Edwards and Haime, 1850 Cavernularia vansyoci Williams, sp. nov. Figs. 9-10 MATERIAL EXAMINED.— HOLOTYPE: CAS 168894, Northeastern Pacific Ocean, United States, Alaska, Aleutian Islands, 52.0663° to 52.07043°N; 175.30231° to 175.2824°W, 86-93 m depth, Haul Number 66, 13 June 2002, collected by Robert Van Syoc on board M/V “Sea Storm,’” NMFS Aleutian Survey 2002, one whole colony cut longitudinally into two halves and wet preserved in 75% ethanol. DESCRIPTION OF HOLOTYPE.— Growth form and size (Figs. 9A-B, 10A): The colony is clavate, 33 mm in total length. Polyps (Figs. 9A—D, 10A): The polyps of the holotype are all preserved completely retracted. They are flush with the surface of the rachis and nowhere do they extend past the surface. Calyces are absent. The polyps and polyp walls contain minute sclerites, similar to the sclerites of the surperficial coenenchyme and peduncular interior. As in the coenchymal tissues, these sclerites can only accurately be detected at microscope magnifications at 400x or higher. Internal Anatomy (Figs. 9C—D): The specimen was cut longitudinally into two halves to reveal internal structures. Gastric cavities are approximately | mm wide and 4-5 mm long. Numerous ova can be concentrated in interior-most parts of some of the cavities. In the center of the colony, sev- eral vertical canals can be seen. A calcified axis is absent altogether. Sclerites (Figs. 9E, 1OB—-C). The sclerite complement is composed entirely of very small ovals 62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 5 FIGURE 9. Cavernularia vansyoci sp. nov. A—B. External views of both halves of the longitudinally sectioned holotype. C—D. Internal views of both halves of the longitudinally sectioned holotype. E. Photomicrographs of three sclerites from the superficial coenenchyme taken from the region of juncture between the rachis and peduncle of the holotype; the scle- ] } rites are actually colorless, color shown here is due to properties of the light microscope at high power. Scale bar for A-D = 16 mm: scale bar for E = 0.005 mm. WILLIAMS: NEW OCTOCORALS FROM NORTHEASTERN PACIFIC OCEAN 63 a. ~. uty Aleutian Archipelago Gulf of Alaska 153° Ficure 10. Cavernularia vansyoci, sp. nov. A. Whole wet-preserved holotype, 33 mm in length. B. Coenenchymal scle- rites from the surface of the junction between the rachis and the peduncle. C. Sclerites from the interior of the peduncle. Scale bar for B and C = 0.01 mm. D. Map showing locality of the holotype (Mf = collecting station). 64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 5 (0.003—0.007 mm in length). The shape of these sclerites is only accurately discernable at micro- scopic magnifications of 400x or more. They vary in shape from rounded-rectilinear to oval, more- or-less round, or elliptical. Some sclerites are irregular in shape. These minute sclerites are rela- tively dense is all parts of the colony examined, including the coenenchyme of the surface of the rachis and peduncle, as well as the interior of the colony, but seem to be less densely distributed in the anthocodiae and polyp walls. Color (Figs. 9A—D). The color of the preserved holotype is grayish white throughout. The scle- rites are colorless. ETYMOLOGY.— This species is named for its discoverer, Robert Van Syoc, Department of Invertebrate Zoology and Geology, California Academy of Sciences, San Francisco. DISTRIBUTION.— The new species is known only from the type locality in the Aleutian Islands of Alaska; 86—93 m depth (Fig. LOD). DISCUSSION The most recent compendium of Alaskan corals is that of Wing and Barnard (2004), which includes a listing of twelve species of plexaurid octocorals and twelve species of pennatulaceans. Of these, three plexaurids and three pennatulaceans are figured with color photographs. The taxa figured are identified as belonging to five genera — Muriceides, Swiftia, Protoptilum, Ptilosarcus, and Halipteris. The present paper describes a new genus and species of plexaurid gorgonian and a new species of the pennatulacean genus Cavernularia — these are newly recorded taxa to the Alaskan octocoral fauna. ACKNOWLEDGMENTS I gratefully appreciate the contributions and services of Elizabeth Kools and Robert Van Syoc (Department of Invertebrate Zoology and Geology, California Academy of Sciences), and the National Marine Fisheries Service (NMFS) of the National Oceanic and Atmospheric Administration (NOAA), for making this project possible. LITERATURE CITED BAYER, F.M. 1961. The Shallow-water Octocorallia of the West Indian Region. A Manual for Marine Biologists. Marinus Nijhoff, The Hague, Netherlands. 373 pp. BAYER, F.M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: Anthozoa), with diagnoses of new taxa. Proceedings of the Biological Society of Washington 94(3):902-947. FABRICIUS, K., AND P. ALDERSLADE. 2001. Soft Corals and Sea Fans — a Comprehensive Guide to the Tropical Shallow-water Genera of the Central-West Pacific, the Indian Ocean and the Red Sea. Australian Institute of Marine Science, Townsville, Queensland, Australia. 264 pp. GrassHorF, M. 1977. Die Gorgonarien des ostlichen Nordatlantik und des Mittelmeeres Ill. Die Familie Paramuriceidae (Cnidaria, Anthozoa). “Meteor” Forschungsinstitut-Ergebnisse D 27(5—76):5—76. GRASSHOFF, M., AND G. BARGIBANT. 2001. Coral Reef Gorgonians of New Caledonia. IRD Editions, Institut de Recherche Pour Le Développment, Collection Fauna et Flore tropicales 38. Paris, France. 335 pp. Hickson, S.J. 1921. On some Alcyonaria in the Cambridge Museum. Proceedings of the Cambridge Philosophical Society 20:366—373. KUKENTHAL, W. 1915. Pennatularia. Das Tierreich 43:1—132. LOPEZ-GONZALEZ, P.J., J.-M. GILI, AND GC.WILLIAMS. 2000. On some veretillid pennatulaceas from the east- ern Atlantic and western Pacific Oceans (Anthozoa: Octocorallia), with a review of the genus Cavernularia, and descriptions of new taxa. Journal of Zoology, London 250:201-216. WILLIAMS: NEW OCTOCORALS FROM NORTHEASTERN PACIFIC OCEAN 65 SANCHEZ, J.A., AND S. CAIRNS. 2004. An unusual new gorgonian coral (Anthozoa: Octocorallia) from the Aleutian Islands, Alaska. Zoologische Mededelingen, Leiden 78(15):265—274. WituiaMs, GC. 1989. The pennatulacean genus Cavernularia Valenciennes (Octocorallia: Veretillidae). Zoological Journal of the Linnean Society 95:285-—310. Wituiams, G.C. 1995. Living genera of sea pens (Coelenterata: Octocorallia: Pennatulacea): illustrated key and synopses. Zoological Journal of the Linnean Society 113:93-140. WinG, B.L., AND D.R. BARNARD. 2004. A Field Guide to Alaskan Corals. NOAA Technical Memorandum NMEFS-AFSC-146. U.S. Department of Commerce, Washington, DC, USA. 67 pp. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 6, pp. 66-75, 9 figs., 1 table April 29, 2005 A New Species of Marionia (Gastropoda: Nudibranchia) from the Caroline Islands Victor G. Smith and Terrence M. Gosliner Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco CA 94103, U.S.A., Email: vsmith@calacademy.org, tgosliner@ calacademy.org A new species of tritoniid nudibranch is described from the western Caroline Islands. Marionia bathycarolinensis is known only from its type location of Palau. This animal exceeds 10 cm in length, is reddish in color and is the second record from the Indo-Pacific region of a tritoniid with more than 20 pairs of branchial plumes. Internally, it is distinguished from other Indo-Pacific tritoniids by jaws with masti- catory borders composed of multiple rows of rodlets, a broad radula with elongate and distinctively shaped rachidian teeth, and a proportionally long esophagus. A muscular band carries approximately 50 chitinous plates arranged so their distal edges face the lumen of the stomach. Analysis of stomach contents shows that the holotype was feeding on octocorals of the genus Paracis. This is the first record of a nudibranch feeding on this genus. Placement of the new species in the genus Marionia is discussed in the light of taxonomic problems brought about by the treat- ment of the group by Odhner (1963). The current state of knowledge of the tritoniid nudibranchs from the Indo-Pacific region leaves workers with a difficult set of taxonomic and nomenclatural problems to overcome as they attempt to sort out the relationships within this diverse family. Descriptions from the early 1800's through the mid 1900’s lack consistency and are based primarily on external morphology, with only scat- tered references to important anatomical details. Subsequently, many species names have been syn- onomized, and other species have not been recorded since their original description. In some cases this may be due to the relative rarity of the taxa, in others the descriptions are insufficient to con- vince modern workers of their validity. Initial taxonomy was established by and expanded upon by Odhner (1936, 1963) but other workers still find difficulties with the most recent system (Willan 1988). Odhner proposed three plate bearing genera: Marionia, Marioniopsis and Paratritonia. His division was based primarily on characters of the digestive gland (liver), jaws and radula. Marionia was defined as having a digestive gland in two masses leaving the stomach uncovered, a jaw with 3 to 6 rows of fine denticles and a radula possessing tricuspid central teeth and differentiated first lateral teeth. Marioniopsis was defined by a digestive gland in a single mass covering the upper and left side of the stomach, jaws with a single row of strong denticles and a radula as in Marionia. In his 1963 key, Odhner defined Paratritonia as having a radula with unicuspidate central teeth and undifferentiated first laterals. This does not agree with the original description (Baba 1949), which stated that the first lateral teeth differ from the outer laterals. Baba also describes the jaw as hay- ing up to 10 rows of fine denticles, and a digestive gland in a single mass. Odhner’s classification scheme relies heavily on the morphology of the digestive gland, which he felt was of utmost importance in dividing the genera. This presents problems on more than one 66 SMITH AND GOSLINER: A NEW SPECIES OF MARIONIA (NUDIBRANCHIA) 67 level. Firstly, the progression of the nudibranch digestive gland from a compact single mass to a divided state and then to a diffuse distribution is on a continuum, is not composed of discrete steps, and appears in multiple lineages. Secondly, detailed dissections of numerous tritoniids by the sen- ior author have not always yielded results in agreement with published descriptions (e.g., Odhner 1963). In addition, there have been descriptions of new animals (e.g., Willan 1988) that do not fit well into any of the current genera, including the present study. Accordingly, the authors feel that it is most prudent for all new species of plate-bearing tritoniids to be placed in the genus Marionia, which precedes the more controversial genera Paratritonia and Marioniopsis. Nudibranchs of the family Tritoniidae are thus far known to feed exclusively on soft corals, sea pens, and gorgonians, but exact feeding patterns are unknown for many species (McDonald and Nybakken 1999). The gut contents of Marionia bathycarolinensis sp. nov. contained skeletal ele- ments of an undescribed octocoral of the genus Paracis Ktikenthal, 1919; Family Plexauridae Gray, 1859. This is the first record of a nudibranch feeding on this genus of octocoral. METHODS Dissection was performed by a ventral incision through the length of the sole of the foot and around the genital and anal openings, allowing the removal of the entire visceral mass in one piece. The jaws and radula were freed from the buccal mass by partially dissolving tissues in 10% KOH solution. They were then rinsed in deionized water, air dried, mounted and coated for electron microscopy. Scanning electron micrographs (SEMs) were produced with a Hitachi S-520 or Leo 1450VP scanning electron microscope. Digestive contents were treated with undiluted household bleach to dissolve the tissue from the octocoral skeletal elements. After multiple rinses in water and 75% EtOH, sclerites and axial samples were mounted, dried and coated for electron microscopy. Slides of sclerites were also prepared for optical microscopy. SPECIES DESCRIPTION Suborder Dendronotacea Odhner, 1934 Family Tritoniidae Lamarck, 1809 Genus Marionia Vayssiére, 1877 TYPE SPECIES: Marionia berghi Vayssiere, 1877 Marionia bathycarolinensis Smith and Gosliner, sp. nov. (Figs. 1-9) TYPE MATERIAL.— All material examined was collected by the Coral Reef Research Foundation (CRRF) and has been deposited at California Academy of Sciences Department of Invertebrate Zoology (CASIZ). HoLorype: CASIZ 156081, one specimen collected at 191.4 m depth, on rock, Mutremdiu 3, Palau, Caroline Islands, 07°16.27’N, 134°34.37’E, 18 March, 2001, L.J. Bell aboard the Deep Worker Submarine, dissected. PARATYPE: CASIZ 156082, buccal mass only, remainder of body not sent, Mutremdiu 3, Palau, Caroline Islands, 07°16.27'N, 134°34.37E, 25 March, 2001, P.L. Colin aboard the Deep Worker Submarine, 222.5 m depth, on rock, dissect- ed. EXTERNAL ANATOMY.— The size of the living paratype specimen was approximately 118 mm in length. Only the buccal mass of this specimen was sent. The color of the living animal (Fig. la) is brick red, with scattered patches of lighter red and greenish-red on the notum. There are also small white flecks, patches, and short linear white markings on the body. The sheaths and shafts of 68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 6 the rhinophores are brick red, with the apical portions a con- trasting dark greenish-red color. The preserved holotype speci- men is 155 mm in length, and 65 mm wide by 40 mm high in its largest section. The size of this animal alive was not recorded. The notum is a dull red color, with patches of a greenish tinge. The sides are of a more even red color. The foot, rhinophores and branchial plumes all have a greenish tinge. The body is elon- gate, subquadrilateral in cross section, with the largest section at about *4 of the way to the pos- terior end of the foot. The taper- ing posterior of the notum over- hangs the posterior end of the foot by about 20 mm. The broad oral veil is slightly incised at midpoint and extends about 5 mm beyond the front of the body. There are 12 velar papillae on each side of the veil. The rela- tively short velar papillae are arranged in multifid groups, with blunt rounded apices. The grooved oral tentacles usually associated with the tritoniids are present but are not distinctly FiGuRE |. Marionia bathycarolinensis sp. nov., digital photographs. (a). demarcated, instead being incor- photo of living animal provided by CRRF. (b). Jaws of paratype, 35 mm in porated as the outer margins of _ length. the veil. The body is finely gran- ular all over, except for the smooth sole of the foot, with some development of low rounded tuber- cles on the notum and sides. The broad foot is rounded anteriorly. From the narrowly overhanging margin of the notum are produced the rhinophores and branchial plumes. In the holotype, the rhinophoral shafts are completely retracted into widely spaced, 6 mm-long sheaths. The photo- graph of the living animal (Fig. la) shows a typical tritoniid arrangement, with a central clavus sur- rounded by a series of pinnate projections. The branchial plumes are relatively short, the largest being slightly greater in length than the extended rhinophores in the photograph. The anterior 8 or so plumes are discrete and easy to count, whereas the more posterior plumes tend to run together, making an exact count difficult. There are approximately 22 plumes per side, with the largest in the middle third of the body. The branchial plumes are divided into three or four main branches, which then subdivide one or two more times. The gonopore is located on the right side, at about midline below and between the 4" and 5" branchial plume. It has the three-valve appearance of some tri- SMITH AND GOSLINER: A NEW SPECIES OF MARIONIA (NUDIBRANCHIA) 69 toniids, and the tip of the penial papilla was visible. A floret of distended tissue surrounds the large opening of the anus, which is situated close to the notal margin below the 10 and 11 branchial plume on the right. The nephroproct opens between the 8" and 9‘ plume, just anterior to the anus. DIGESTIVE SYSTEM.— The jaws are yellowish brown in color, with a darker, thickened red- dish brown masticatory margin slightly overhanging the length of the base (Fig. 1b). The mastica- tory margin appears smooth to the naked eye, but scanning electron microscopy reveals the pres- ence of 25 or more rows of jaw rodlets visible in approximately the distal 5 mm of the half jaw examined (Figs. 2-3). The jaw rodlets exhibit a complex ultrastructure, with multiple vertical ele- ments encased in an outer layer showing horizontal sculpture. The radula of the paratype is large compared with other tritoniids, with a formula of 72 (142.1.1.142) at its broadest point (Figs. 4a- d, 5a—b). The base of the rachidian is roughly rectangular, and is wider than it is long. The upper margin of the base (as oriented in the figures) is incised in the shape of a ‘V’. The central cusp aris- es as a sturdy wedge-shaped process oriented nearly 90° from the plane of the base. The top of the wedge flattens and forms a long, emarginate, blade-shaped triangular denticle. Behind the base of this denticle the top of the wedge forms two projecting ridges that follow the ‘V’ shape of the mar- gin. The rows of teeth are arranged very closely together, so that the projecting wedge of the cen- tral cusp fits neatly into the ‘V’ shaped margin and ridges of the next tooth, with the apex of the flat blade extending over the top. At the base of each wedge-shaped ridge is a small, flattened, emarginate denticle, considered here as accessory. The two outer cusps of the rachidian are short- er than the median, and are formed from the thickened outer margins of the base. Their surface and that of the accessory denticles bears a pattern of folds or ridges resembling the veins on a leaf. The $36743 15KY FIGURE 2. SEMs of jaw of paratype. (a). Proximal portion FiGurE 3. SEMs of jaw of paratype. (a). View of rodlets of masticatory border showing eruption of jaw rodlets. (b). near distal end of masticatory border. (b). Detail of rodlets Distal portion of masticatory border, showing part of over- from same area. (c). View of rodlets at distal end of mastica- hang broken off in preparation. (c). Higher magnification tory border. (d). Detail of rodlets seen in Figure 3c. view of proximal masticatory border. (d). Close up of rodlets in proximal masticatory border. 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 6 first lateral teeth are differentiated from the remaining laterals, being more heavily constructed with a short, hooked, chisel-shaped denticle fitting closely with the edges of the outer cusps of the rachidian. The remaining laterals are straight to slightly curved, and relatively broad and stout (Figs. 5 ab). A pair of floccose salivary glands extended backwards from the proximal portion of the buccal mass, with attachments to the buccal mass on either side of the esophagus. The pre- served salivary glands exhibit a greenish coloration. The esophagus extends about 8 cm from the median posterior buccal mass as a long straight tube, widening into a distended sac-like portion before turning and entering the stomach on the ventral side (Fig. 5c). A large bolus of gorgonian food material was present in the proximal esophagus, and will be described and discussed in a later section. The stomach 1s relatively small, 2 cm in diameter, with a muscular girdle visible in the proximal portion (Fig. 5c). Inside the stomach are approximately 50 stout, dark brown plates arranged so that the bases are attached to the muscular girdle and the plates point to the lumen of the stomach (Figs. 6a—b). The plates are roughly rectangular, with the exposed corners rounded, and most are about 7 mm long by 3mm high. A pair of larger, thicker and higher plates surrounds the typhlosole that continues through the stomach from the esophagus and into the intestine. There are also some thinner plates and some plates that are only 4 mm long interspersed in no discern- able pattern. The intestine (Fig. 5c) exits the stomach from the anterior left portion, curving up and around the anterior portion of the digestive gland towards the right, becoming wider for a portion before narrowing again to terminate at the anus. The compact digestive gland is divided into two lobes, each with a duct leading into the stomach. The large, spheroid posterior portion has an ante- rior hollow that fits around the posterior stomach, leaving the top exposed. The smaller anterior FIGURE 5. SEMs from paratype, drawing from holotype. fication view of radula, older portion. (b). Rachidian teeth, (a). Middle latteral teeth from older portion of radula. (b). first and inner lateral teeth at older portion. (c). Detailed Outer laterals from newer portion of radula. (c). Visceral view of rachidian at area shown in Figure 4b. Scale bar= 300 mass (from holotype). an = anus; dga = anterior digestive tm. (d). Rachidian teeth, first and inner lateral teeth at newer gland; dgp = posterior digestive gland: esd = distal esopha- portion. gus; esp = proximal esophagus; in = intestine; st = stomach. SMITH AND GOSLINER: A NEW SPECIES OF MARIONIA (NUDIBRANCHIA) 71 FiGuRE 6. SEMs from holotype. (a). Portion of band of stomach plates, showing typhlosole area. Scale = 1 mm. (b). Isolated plate. Scale = 400 um. (c). Portion of gut contents, showing overview of characteristic plates and 'woody' stem. Scale = 200 um. (d). Closer view of plates. Scale = 100 Lm. lobe abuts the right anterior portion of the stomach, and is loosely connected to the posterior diges- tive gland (Fig. 5c). REPRODUCTIVE SYSTEM.— The reproductive system is triaulic. The diffuse ovotestis covers much of the surface of the digestive glands. A narrow hermaphroditic duct connects to the narrow end of the ampulla (Fig. 7a). The ampulla is muscular and convoluted, its wide end encased with- in the compact female gland mass. The smaller albumen gland is easily discernable from the larg- er membrane and mucos glands. The proximal vas deferens emerges from the female gland mass as a thin, relatively straight tube for about 10 mm, becoming thickened and sinuous in its median portion. The distal portion then extends, thinning slightly before entering the base of the conical unarmed penis, lying near the gonopore. The muscular bursa copulatrix is an inflated oval 18 mm long. The vaginal duct is approximately equal to the bursa in length, widening at the vaginal atri- um. The oviduct exits the female gland mass and opens to the vaginal atrium. NERVOUS SYSTEM.— The ganglia of the central nervous system sit on the dorsal esophagus, just behind the buccal mass (Fig. 7c). A thin transparent membrane not apparent in dissections of other tritoniids covers the ganglia and large nerves. The paired cerebral and pleural ganglia are dis- tinct but somewhat fused, and the pairs are joined by a short connective. The pedal ganglia are on either side of the cerebropleurals, joined to them by short connectives, and to each other by the cir- cum-esophageal nerve ring, which also contains commisures joining the cerebropleurals. A pair of buccal ganglia is present on the ventral esophagus, just anterior to the nerve ring. These are joined by a short connective (Fig 7c). The central nervous system and buccal ganglia are distinctly asym- V2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 6 metrical and ‘lumpy’ in appear- ance, with irregular nodules and a distinct giant nerve cells scat- tered about. The darkly pigment- ed eyes are small, connected to the central nervous system by long nerves. Gut CONTENTS.— The con- tents of the digestive system con- tained both free sclerites and large pieces consisting of a “woody” appearing axis which is covered by distinctive large plates (Figs. 6c—d; 8a—d; 9a—b). The exposed sides of the plates have a pearly appearance, the bumpy surface giving the impression of sclerites immersed in a smooth nacreous coating. There is a distinct transition between the smooth exposed side and the rougher surface of the attached side. A crack in the membrane of the attached side shows a few sclerite-like objects imbedded in a matrix. Although it is not possible to know with any certainty the origin or exact vdp fgm : : 3 S FiGuRE 7. Drawings from holotype. (a). Reproductive system. Al = albu- identity of the octocoral from min gland; am = ampulla; be = bursa copulatrix; fem = female gland mass; hd which the free sclerites were =hermaphroditic duct; od = oviduct; pn = penis; va = vaginal atrium; vd = dis- derived, the larger pieces are tal portion of vas deferens; vdp = proximal portion of vas deferens. (b). Central nervous system. ceg = cerebral ganglia; peg = pedal ganglia; plg = consistent with the genus Paracis ae g 2 me Beis pleural ganglia. (c) Buccal ganglia. Scale bars as indicated. Kiikenthal, 1919, Family Plex- auridae Gray, 1859 and appear to belong to an undetermined species (G.C. Williams, pers. com- mun.). Further references to octocoral information and biology can be found in Smith and Gosliner (2003). DISTRIBUTION.— Marionia bathycarolinensis sp. nov. is known only from the type locality of Palau, in the Caroline Islands. ETYMOLOGY.— The first part of the specific name is derived from the Greek, bathys (deep); the latter part refers to the type locality in the western-most fringe of the Caroline Islands. DISCUSSION In two recent papers (Avila et al. 1999; Jensen 1994), the anatomical and morphological char- acters of plate-bearing tritoniids have been tabulated. Avila et al. compared characteristics of species of the genus Marioniopsis, and included eight other species along with their newly described species. Jensen compared anatomical characters of species of Marionia and Marioniopsis, listing 17 species plus her newly described species. The combined species discussed SMITH AND GOSLINER: A NEW SPECIES OF MARIONIA (NUDIBRANCHIA) Ws FiGureE 8. Detailed view of detached octocoral plate. (a). Close up view of exposed (front) side. Scale bar = 20 um (b). A single detached plate, viewed from the attachment (back) side. Scale bar = 100 ttm. Small arrow indicates area detailed in 8c, large arrow indicates area detailed in 8d. (c). Detailed view of area indicated by smaller arrow in 8b. Scale bar = 100 um. (d). Detailed view of area indicated by larger arrow in 8b. Scale bar = 20 um. FiGure 9. A Portion of the 'woody' axis of the octocoral. (a). Scale bar = 1 mm. (b). Scale bar = 100 um. in both papers are summarized in Table 1, along with Paratritonia lutea Baba, 1949, and Marionia bathycarolinensis sp. nov. There are two external characters that differentiate all other plate-bearing tritoniids from Marionia bathycarolinensis sp. nov.: the number of branchial plumes and the number of velar papillae. Marionia bathycarolinensis sp. noy. has 22 pairs of branchial plumes and 12 pairs of com- pound velar papillae. All other species listed have 16 or fewer branchial plumes, with the excep- 74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES TABLE |. Comparison of Indo-Pacific species of the genus Marionia Volume 56, No. 6 Species Branchial Velar papillae Jaws ‘h row of Stomach plumes radula teeth __ plates M. albotuberculata 9 pairs 5 pairs, some com- | row of teeth, trace 95 >100 pound of second M. babai 7 pairs 6 pairs, compound > 100 teeth, number Ds present of rows not stated M. blainvillea 10-12 pairs 7 pairs 2—4 rows of teeth 15-21 ca 40 M. chloanthes 9+ pairs 4 pairs, some com- — 5—6 rows of teeth 22 70 pound M. cucullata 12-16 pairs _ 7-11 pairs 6 rows of teeth 58 30-40 M. cyanobranchiata 9-13 pairs 4-7 pairs, simple __1 row of teeth, trace 15-50 100-120 of second M. dakini 13 pairs 6-7 pairs, compound 1 row of 45 teeth 135 present M. echinomuriceae 10-14 pairs 6-8 pairs 1 row of teeth, 65 28 indistinct M. fulvicola 7-9 pairs 3-4 pairs, most simple 4-5 rows,>100 38-42 22-32 teeth M. granularis 13-14 pairs 6 pairs unknown 50 present M. levis 9-10 pairs 3-5 pairs, compound 1-3 rows, 14-30 80-130 ca 150 teeth M. pambanensis 12 pairs 6 pairs unknown 43 present M. pellucida 13 pairs 6 pairs unknown 22 70 M. platyctena 100 pairs 5-7 pairs, simple 10 rows of teeth 71-103 30-35 M. putulasa 15 pairs 6 pairs unknown 112 25 M. rubra 10-12 pairs 6 pairs, compound | row of 100-120 50-55 present teeth M. tessellata 13 pairs 7 pairs 3 rows of teeth unknown present M. bathycarolinensis 22 pairs 12 pairs, compound 25-100 rows of 142 50 rodlets M. viridescens 10 pairs 7 pairs, some single row of teeth 90 25 compound M. olivacea 9-15 pairs 7 pairs, some 3-7 row of teeth 70-80 50-60 compound Paratritonia lutea 6—7 pairs 34 pairs 3—10 rows of teeth 110 Md) tion of M. platyctena, which has 100. However, this animal is recorded as having a jaw with 10 rows of teeth, a velum with 5—7 pairs of simple papillae, a radular half-row of up to 103 lateral teeth and a maximum of 35 stomach plates. This is in contrast with M. bathycarolinensis sp. nov., which has a jaw armed with rodlets, a velum with 12 pairs of compound papillae, a radula with a half-row of 143 lateral teeth, and 50 stomach plates. Only M. cucullata (which may be a synonym for M. blainvillea) has a recorded number of velar papillae approaching that of the new species. However, M. cucullata does not match our new species in any of the other characters discussed, differing in the number of branchial plumes, jaw morphology, radular formula and number of stom- ach plates. Internally, the principal apomorphy of Marionia bathycarolinensis sp. nov. is a jaw armed with SMITH AND GOSLINER: A NEW SPECIES OF MARIONIA (NUDIBRANCHIA) TS multiple rows of rodlets. This type of denticulation has not been recorded for any other species of tritoniid nudibranch. The basic form of the tritoniid radula is, with few exceptions, consistent throughout the gen- era, consisting of a tricuspid central tooth flanked by a pair of first lateral teeth differentiated from the remaining curved outer laterals. The species described here does not differ from the basic form, but is distinctive in the appearance of the central tooth. The median cusp is elongate, overlapping the next row when the radula is contracted. The leaf-like shape of this flattened central cusp has not been recorded for any other species of tritoniid. The remaining distinguishing characters are more general: large size and a relatively compact visceral mass with an elongate esophagus. Body lengths of over 10 cm are at the large end of the tritoniid size continuum, mostly occurring in temperate species without stomach plates. Both spec- imens described here are of this large size. Figure 5b illustrates the long straight esophagus and rel- atively compact visceral mass, which contrasts with published drawings of other tritoniids (Odhner 1936) and the personal observations of the present authors. ACKNOWLEDGEMENTS The authors would like to thank Pat Colin and Lori Bell of the Coral Reef Research Foundation (Palau) for providing the specimens and a photograph of the live animal. Dr. Gary Williams of the California Academy of Sciences provided expertise and advice on octocoral iden- tification, and edited an early draft of the manuscript. An anonymous reviewer made valuable com- ments and suggestions, which led to improvements of the final manuscript. This work was partial- ly supported by NSF PEET grant #0329054. LITERATURE CITED AviLa, C., D. KELMAN, Y. KASHMAN, AND Y. BENAYAHU. 1999. An association between a dendronotid nudi- branch (Mollusca, Opisthobranchia) and a soft coral (Octocorallia, Alcyonaria) from the Red Sea. Journal of Natural History 33:1433-1449. BaBa, K. 1949. Opisthobranchs of Sagami Bay Collected by His Magisty The Emperor of Japan. wanami Shoten, Tokyo, Japan. 194 pp., 50 pls. JENSEN, K.R. 1994. Sublittoral Notaspidea and Nudibranchia (Opisthobranchia) from Hong Kong, with a description of a new species. Pages 117—139 in B. Morton, ed., The Malacofauna of Hong Kong and Southern China III. Proceedings of the Third International Workshop on the Malacofauna of Hong Kong and Southern China, Hong Kong 13 April-1 May 1992. Hong Kong University Press, Hong Kong, China. McDONALD, G.R., AND J.W. NYBAKKEN. 1999. A worldwide review of the food of nudibranch mollusks I. The Suborder Dendronotacea. The Veliger 42(1):62—66. ODHNER, N. 1936. Nudibranchia Dendronotacea — A revision of the system. Mémoires du Musée Royal d Histoire Naturelle de Belgique, series 2, 3:1057—1128, pl. 1. OpHNER, N.H. 1963. On the taxonomy of the family Tritoniidae (Mollusca: Opisthobranchia). The Veliger 6:48-62. 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):155-278. WILLAN, R.C. 1988. The taxonomy of two host-specific, cryptic dendronotoid nudibranch species (Mollusca: Gastropoda) from Australia including a new species description. Zoological Journal of the Linnean Society 94:39-63. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 7, pp. 76-79, 1 fig. April 29, 2005 Perulibatrachus aquilonarius, a New Toadfish Species from India (Teleostei: Batrachoididae) David W. Greenfield! California Academy of Sciences, 875 Howard Street, San Francisco, California 94103; Email: greenfie @ hawaii.edu The fourth known species in the genus Perulibatrachus is described. It is the north- ernmost species in the genus, and the second known from the Indian Ocean. It has a wider head than the other described species and differs from others in counts: dor- sal-fin elements II]-17, anal-fin rays 13, pectoral-fin rays 21, vertebrae 26. It has a shallow, funnel-shaped pocket in the pectoral-fin axil. The genus Perulibatrachus is currently represented by three species. Two species are known only from the west coast of Africa: P. elminensis (Bleeker, 1863) from Ghana south to Walvis Bay, Nambia, and P. rossignoli (Roux, 1957) from Gabon south to Walvis Bay (Roux 1981; Hutchins 1986). A third species, P. kilburni Greenfield, 1996, is known only from Natal, southeastern South Africa. In 1941, A.W.C.T. Herre collected a toadfish specimen, purportedly in India, which he or someone else had identified as Chatrabus damaranus. The specimen had been deposited in the ichthyological collections of Stanford University and later transferred to the California Academy of Sciences along with the rest of Stanford’s collections. Recently, I found the specimen on the shelves at the Academy and on examination discovered that it does not belong to the genus Chatrabus but to the genus Perulibatrachus. The genus Chatrabus was described by Smith in 1949, whereas Perulibatrachus was not described by Roux and Whitley until 1972. Thus, at the time of its identification, Chatrabus would have been a logical choice in which to place this spec- imen. The most recent key to toadfish genera is that of Smith (1952). In that key, Perulibatrachus would fall in section BIH, which includes Chatrabus, Barchatus, Tharbacus, Batrachoides, and Halobatrachus. Hutchins (1986) considers Tharbacus to be a synonym of Chatrabus. Roux (1981) presents a key to separate Perulibatrachus from Halobatrachus, which has a distinct foramen (axil- lary pore) on the upper part of the pectoral-fin axil, whereas the other genera have only a funnel- shaped pocket (Perulibatrachus) or neither a foramen nor pocket (Chatrabus and Batrachoides). In species without the pocket, the skin stretches straight across the pectoral-fin axil, whereas in Perulibatrachus there is a clear indentation, which is shallow in P. rossignoli or relatively deep in both P. elminensis and P. kilburni. The specimen from India has three dorsal-fin spines and two subopercular spines, placing it in the subfamily Batrachoidinae. The body is mostly scaled, it lacks a foramen in the pectoral-fin axil, and it has a shallow funnel-shaped pocket in the pectoral-fin axil, placing it in the genus Perulibatrachus. | Research Associate, Department of Ichthyology, California Academy of Sciences and Emeritus Professor, University of Hawaii. Mailing address: Moss Landing Marine Laboratory, 8272 Moss Landing Road, Moss Landing, CA 95039. 76 GREENFIELD: NEW TOADFISH OF THE GENUS PERULIBRACHUS FROM INDIA Teh MATERIALS AND METHODS All counts and measurements follow Hubbs and Lagler (1964) except that the last two fin rays are not counted as one unless it is clear that they are joined at the base. Measurements were made to the nearest 0.1 mm using dial calipers. All measurements are expressed as percentage of stan- dard length (SL). Counts were made from a radiograph. Neither the holotype nor any additional material of P. rossignoli could be located at the Museum National d’ Histoire Naturelle, Paris, by M.L. Bauchout and J.E. Randall, so comparisons with that species are based on the literature. Institutional abbreviations are as listed in Leviton et al. (1985). SPECIES DESCRIPTION Perulibatrachus aquilonarius Greenfield, sp. nov. (Figs. 1A—B) MATERIAL EXAMINED.— HOLotyPE: CAS-SU 41322, 191.4 mm SL, India, Tamil Nadu State, Ennur Fisheries Station, Madras, January, 1941, A.W.T.C. Herre. ADDITIONAL MATERIAL EXAMINED: Perulibatrachus kilburni, RUSI 28203 (1; 56.6 mm SL). Perulibatrachus elminensis, MNHN 1967-909 (1; 204 mm SL), MNHN 1970-43 (1; 150 mm SL; cleared and stained). DIAGNOsISs.— A species of Peruli- batrachus with a wide head (43.6% SL), 17 dorsal-fin rays, 13 anal-fin rays, 21 pectoral-fin rays, 26 vertebrae, and a shal- low, funnel-shaped pocket in the pectoral- fin axil. DESCRIPTION.— Dorsal-fin elements I-17; anal-fin rays 13; pectoral-fin rays 21; vertebrae 26; head length 45.0; head width 43.6; head depth 25.3; bony interor- bital width 9.5; orbit diameter 7.6; snout length 8.8; upper jaw length 25.9; mouth width at rictus 36.1; first predorsal-fin dis- tance 42.1; second predorsal-fin distance 61.6; preanal-fin distance 72.5; greatest body depth 25.2; caudal-peduncle depth 9.8; caudal-peduncle length 13.6; first dorsal-fin base length 11.4; second dorsal- fin base length 39.6; anal-fin base length ~ SO Par NS X 21.5; caudal-fin length 22.7, pectoral-fin a FICHE 1. Holotype of Perulibatrachus Rpailennite CAS-SU length DSO: pelvic-fin length 26.7; dis- 41322. A. lateral view; B. dorsal view. tance between pelvic-fin bases 15.3. Head moderately depressed and wide, eyes medium in size and not raised above head profile. Pectoral-fin axil with a shallow, fun- nel-shaped pouch in upper half, glandular tissue present in rest of axil and on sides under pectoral fins. Some glandular tissue present on inner surface of pectoral fins. Body with small scales extend- ing from middle of first dorsal fin back to caudal-fin base. Scales extending forward on ventral sur- face to pelvic fins. Two lateral lines: the upper one originating above upper opercular spine, run- ning posteriorly in a straight line to below second dorsal-fin origin where it curves up to run along 78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 7 dorsal-fin base to its end, with about 41 pores present, each with two short skin flaps; the second lateral line, with about 17 pores, runs along anal-fin base. Vomer and palatine each with single row of slightly curved conical teeth. Dentary laterally with single row of conical teeth, double row near symphysis. Premaxilla with single row of conical teeth posteriorly, double row anteriorly. Subopercle with two spines. Anterior nostrils tubular; nostril on left side with a simple major ten- tacle; nostril on right side with three terminal tentacles; a cluster of multifid cirri surrounds each tubular nostril. Cirrus at distal end of maxilla with many tips. Ventral margin of dentary with numerous cirri. Dorsal surface of neurocranium completely covered with muscles, no exposed bone. Color in alcohol: After 64 years in preservative, general background color is cream, overlaid with rusty brown pigment pattern. Anterior portion of head from posterior margin of eyes forward rusty brown. A cream band running from posterior margins of eyes posteriorly to opercular spine forms a distinct band across the head. The area posterior to this and extending back to first dorsal- fin origin overlaid with distinct, small, rust-colored spots. Sides of body, second dorsal fin, anal fin, pectoral, and caudal fins with scattered, irregular rust-colored spots. Ventral surface of head cream, area from pelvic fins posteriorly to anal-fin origin rust. ETYMOLOGY.— The specific epithet is an adjective from the Latin, aquilonarius, meaning northern, referring the fact that this species has the northernmost distribution of any member of the genus Perulibatrachus. COMPARISONS.— Perulibatrachus aquilonarius differs from all other species in the genus by having a wider head: 43.6 versus 29.7 in P. kilburni, 29.1—37.9 in P elminensis, and 37.8-40.9 in P. rossignoli. \t differs from P. rossignoli by having 17 versus 19 second dorsal-fin rays, 21 versus 23 pectoral-fin rays, and 26 versus 29 vertebrae. It differs from P. elminensis by having 13 versus 14-17 anal-fin rays, 21 versus 26 pectoral-fin rays, and 26 versus 27—28 vertebrae. It differs from P. kilburni by having 17 versus 18 dorsal-fin rays, 13 versus 14 anal-fin rays, and 21 versus 19 pec- toral-fin rays. It has a shallow, funnel-shaped axillary pocket, whereas it is deep in P. elminensis and P. kilburni. ACKNOWLEDGMENTS C. Roux and M.L. Bauchot were very helpful in locating and providing specimens of P. elmi- nensis. M.L. Bauchot and J.E. Randall also searched for specimens of P. rossignoli. | would also like to thank Guy Duhamel and Patrice Pruvost for their hospitality while visiting the MNHN. Special thanks to the staff at CAS for lending material and providing assistance: D. Catania, W.N. Eschmeyer, J. Fong, M. Hoang, and T. Iwamoto. LITERATURE CITED BLEEKER, P. 1863. Mémoire sur les poissons de la céte de Guinée. Natuurkundige Verhandelingen van de Hollandsche Maatschappij der Wetenschappen te Haarlem, ser. 2, 18:1—136, 28 pls. GREENFIELD, D.W. 1996. Perulibatrachus kilburni, a new toadfish from East Africa. Copeia 1996:901—904. Huss, C.L., AND K.F. LAGLER. 1964. Fishes of the Great Lakes Region. University of Michigan Press, Ann Arbor, Michigan, USA. 213 pp. HuTcuins, J.B. 1986. Family No. 100: Batrachoididae. Pages 358-361 in M.M. Smith and P.C. Heemstra, eds., Smith’s Sea Fishes. Macmillan South Africa, Johannesburg, South Africa. 1047 pp. Leviton, A.E., R.H. Gipss, JR., E. HEAL, AND C.E. DAwson. 1985. Standards in herpetology and ichthyology. Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832. GREENFIELD: NEW TOADFISH OF THE GENUS PERULIBRACHUS FROM INDIA 72 Roux, C. 1957. Poissons marins. Pages 139-253 in J. Collignon, M. Rossignol, and C. Roux. Mollusques, Crustacés, Poissons marins des cétes d’A.E.F. en collection au Centre d’Océanographie de I’ Institut d’E- tudes Centrafricaines de Point-Noir. Office de la Recherche Scientifique et Technique Outre-Mer, Paris, France. 369 pp. Roux, C. 1981. Batrchoididae. Vol. 1. Jn W. Fisher, G. Bianchi, and W.B. Scott, eds., FAO species identifica- tion sheets for fishery purposes. Eastern Central Atlantic; fishing areas 34, 47 (in part). Vols. 1-7. Canada Funds-in-Trust. Ottawa, Department of Fisheries and Oceans Canada, by arrangement with the Food and Agriculture Organization of the United Nations, Ottawa, Ontario, Canada. (No page numbers.) Roux, C., AND G. WHITLEY. 1972. Perulibatrachus, nouveau nom de genre de Poissons Téléostéens de la famille des Batrachoididae, en remplacement de Parabatrachus Roux, 1970. Bulletin du Muséum National d Histoire Naturelle Zoologie 6(1971):349-350. SmiTH, J.L.B. 1949. The Sea Fishes of Southern Africa. Central News Agency, Ltd., Cape Town, South Africa. 550 pp. SmitH, J.L.B. 1952. The fishes of the family Batrachoididae from South and East Africa. Annals and Magazine of Natural History, ser. 12, 5(52):313-339. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 8, pp. 80-85, 3 figs. April 29, 2005 Brotula flaviviridis, a New Species of Brotula from Fiji (Teleostei: Ophidiidae: Brotulinae) David W. Greenfield! California Academy of Sciences, 875 Howard Street, San Francisco, Callifornia 94103 The third known species of Brotula from the Indo-Pacific, B. flaviviridis sp. nov., is described from Fiji. Brotula townsendi is an antiequitorial species, known from the Hawaiian Islands, Johnston Island, and the Marshall Islands in the northern hemi- sphere, and Chesterfield Islands, New Caledonia, Loyalty Islands, Vanuatu, and Tonga in the southern hemisphere. Brotula multibarbata is widespread in the Indo- Pacific. Brotula flaviviridis differs from both in coloration, being yellow-green. It is similar to B. townsendi in having an eye that is narrower than the fleshy interorbital width, whereas B. multibarbata has an eye that is wider. It differs from B. townsendi by having a smaller eye and narrower interorbital width. While conducting a survey of the marine fishes of Fiji, two specimens of a yellow-green bro- tula were collected using rotenone at the barrier reef on the north side of Kanacea Island in the Northern Lau Group, Fiji. The fresh coloration of these specimens was distinctly different from the other two species in the genus Brotula known from the Indo-Pacific. The new species is placed in the genus Brotula because: the anterior nostril is well above the upper lip, near the eye; the supramaxilla is present; the dorsal-fin rays are equal to or longer than opposing anal-fin rays; and six barbels are present on the snout and six on the chin. Only five species in the genus Brotula are currently considered to be valid. Brotula barbata (Bloch in Bloch and Schneider, 1801) occurs in the tropical and subtropical Atlantic Ocean. Brotula clarkae Hubbs (1944) occurs in the tropical eastern Pacific from the Gulf of California to Peru, and B. ordwayi Hildebrand and Barton (1949) occurs at Peru and the Galapagos Islands. In the Indo- Pacific Ocean, B. townsendi Fowler (1900) is known from the Hawaiian Islands, Johnston Island, Marshall Islands, Chesterfield Islands, New Caledonia, Loyalty Islands, Vanuatu, and Tonga, and B. multibarbata Temminck and Schlegel (1846) is widespread. Hubbs (1944) listed ten other described species as synonyms of B. multibarbata, and these were also listed by Nielsen, et al. (1999). MATERIALS AND METHODS Information on the holotype is presented first, followed by that of the paratype in parentheses. The paratype died with its mouth open wide, making measurements less accurate than those for the holotype. All measurements are presented as percentage of standard length (SL), and some as per- centages of other body parts. Information on Brotula clarkae and B. ordwayi is from Allen and ! Research Associate, Department of Ichthyology, California Academy of Sciences and Emeritus Professor, University of Hawaii. Email: greenfie@hawaii.edu. Mailing address: Moss Landing Marine Laboratory, 8272 Moss Landing Road, Moss Landing, CA 95039. GREENFIELD: A NEW BROTULA FROM FIJI 81 Robertson (1994) and Hubbs (1944). Pectoral-fin ray counts were obtained by slitting the skin on the inside of the right pectoral fin. Other fin-ray counts and vertebral counts were obtained from radiographs. Methods of counting and measuring follow Nielsen et al. (1999), and the format of the description follows Cohen and Nielsen (1982). SPECIES DESCRIPTION Brotula flaviviridis Greenfield, sp. nov. (Figs. 1-3) MATERIAL EXAMINED.— HOLOTYPE: CAS 221531, 125.0 SL, Fiji, Northern Lau Group, north side of Kanacea Island at copra planta- tion, 17°14.890’S, 179°08.475’W, isolated piece of barrier reef, sand and dead coral, 7—9.5 ~ m, 5 January 2003, field number G03-15, FIGURE 1. Drawing of holotype (CAS 221531) of Brotula rotenone, collected by D.W. Greenfield, K.R. “avvirdis. Longenecker, and R.C. Langston. PARATYPE: CAS 221532, 151.0 SL, Fiji, Northern Lau Group, north side of Kanacea Island at copra plantation, 17°14.890’S, 179°08,475’W, over- hang on barrier reef with sand at base, 12-14 bi : = m, 4 January 2003, field number GO3-I1, FIGURE 2. Fresh color of holotype (CAS 2 rotenone, collected by D.W. Greenfield, K-R. — Brotula flaviviridis. Photo D.W. Greenfield. Longenecker, R.C. Langston, and Bio Koroi Mataitini. ADDITIONAL MATERIAL EXAMINED: Brotula townsendi: Hawaiian Islands, Oahu-CAS 221533. Johnston Island-BPBM 33975 (2), BPBM 29609 (1), BPBM 34043 (1). Chesterfield Islands-BPBM 33794 (5). New Caledonia-BPBM 22550 (1). Brotula multibarbata: Hawaiian Islands, Oahu-CAS 78987 (5), CAS 78901 (2), CAS 210003 (4). Fiji-CAS 218868 (4), CAS 219394 (5), CAS 219404 (5), CAS 219405 (4). DiAGNosis.— A species of Brotula that is yellow-green in coloration, with an eye that is nar- rower than the interorbital width, 98-103 dorsal-fin rays, 77—79 anal-fin rays, 24 pectoral-fin rays, 43-44 vertebrae, 2—3 developed gill rakers, a fleshy interorbital width of less than 4.5 percent SL, and an orbit diameter 3.5 percent SL or less. DESCRIPTION.— Dorsal-fin rays 97 (103). Anal-fin rays 77 (79). Pectoral-fin rays 24 (24). Vertebrae 12+32 =44 (11+32 =43). Total developed gill rakers 3 (2 one side, 3 other). Longitudinal scale series about 133. Predorsal length 26.6 (25.8). Preanal length 48.8 (48.3). Preventral length 14.2 (21.0). Body depth at vent 19.2 (21.5). Head length 22.3 (23.5). Eye diam- eter 3.2 (3.5). Snout length 4.6 (6.0). Fleshy-interorbital width 4.2 (3.8). Upper-jaw length 12.0 (13.8). Greatest maxillary width 9.7 (12.4). Pectoral-fin length 14.2 (13.8). Pectoral-fin peduncle width 6.6 (6.3). Ventral-fin length 20.3 (17.8). Body deepest just before end of pectoral fin, tapering to tail, which is not greatly attenuate. Body scales relatively large for the genus, about 133 rows along the side of the body. Head cov- ered with scales. Scales on belly forward to ventral fins. Scales present on pectoral-fin base and also out onto the dorsal and anal fins. Head compressed. Snout bluntly rounded, longer than orbit diameter. Upper lip terminal, extending slightly beyond lower jaw. Posterior nostril with raised rim, adjacent to anterior margin of eye. Anterior nostril immediately anterior to posterior nostril, with a raised rim and barbel on posterior margin. Two barbels at front of snout, one on each side of mid- » 21531) of 82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 8 dle of snout. Another barbel on each side of snout, about at level of ventral margin of eye, for a total of six barbels on snout. Maxilla notably expanded posteriorly, reaching about one pupil diam- eter past rear margin of orbit. A sheath of tissue on upper third of maxilla, with a notch at end exposing top end of maxilla. Underside of lower jaw with six barbels, three on each side. Two pores at symphasis and one adjacent to posterior side of anteriormost barbel. Teeth small and granular, some with fine points, on dentary, premaxilla, palatine, and vomer. Vomerine tooth patch rounded anteriorly and V-shaped. First gill arch with four (five) short, stubby rakers on upper arm, a longer (developed) raker at angle, lower arm with two (one) longer and ten (14) short protuberances. Longest raker about one- half eye diameter. Longest gill filament a little longer. Branchial cavity and palate pale. Lateral line not obvious. Dorsal fin originating over end of pectoral-fin peduncle, at origin of pectoral-fin rays. Pectoral fin on a short, rounded, fleshy peduncle. Opercle with a strong spine hid- den under skin which continues as a prominent flap of skin dorsal to pectoral-fin base. Ventral fins inserted well behind symphysis of cleithra, about under opercular spine. Lining of peritoneal cav- ity pale, with scattered, small, black spots. Stomach and intestine pale. Color in alcohol: Background color cream, overlaid with gray-brown pigment on center of scales, leaving the scales outlined. Nape, snout, and front of jaws with greater concentration of gray-brown. Barbels and posterior ends of maxilla and premaxilla white. Ventral surface of head gray. Pectoral fins clear. Pelvic fins white. Bases of dorsal and anal fins gray where scales extend out onto them. Distal margin of anterior two-thirds of dorsal fin clear. Posterior third gray with black margin. Anal fin clear distally on anterior two-thirds of fin, posterior third like posterior part of dorsal fin. Caudal fin edged in black. Color of fresh specimen: Background color yellow with a greenish tinge. Head yellow-green. Pupil of eye black, surrounded by yellow iris. Barbels on snout orange, barbels under chin white. Snout and tip of upper and lower jaws dusky. Anterior two-thirds of body primarily yellow-green, posterior third more dusky green. Belly yellow-white. Pectoral fin yellow. Anterior half of dorsal fin yellow with orange margin. Posterior half of dorsal fin reddish grading into black posteriorly as it merges with black caudal fin. Anal fin similar to posterior half of dorsal fin. Pelvic fins white. ETYMOLOGY.— The specific epithet is a compound adjective, combining the Latin flavus, meaning golden-yellow, plus viridis, meaning green, alluding to the yellow-green color of the species. CoOMPARISONS.— Brotula flaviviridis differs from all known valid Pacific Ocean species by its yellow-green coloration. It differs from the two eastern Pacific species, B. clarkae and B. ordwayi, by having fewer dorsal-fin rays, 97-103 versus 108-112 (B. clarkae) and 118-125 (B. ordwayi), and fewer anal-fin rays, 77-79 versus 78-89 (B. clarkae) and 86—94 (B. ordwayi). It also differs from B. clarkae in having 24 pectoral-fin rays verses 27—28. As pointed out by Hubbs (1944), “the distinction between the relatively large scales of the Indo-Pacific species and the small scales of the American ones, particularly of B. clarkae, was obvious at sight.” Brotula flaviviridis differs from the Indo-Pacific species B. multibarbata by having an eye that is narrower than the interor- bital width, verses one that is wider. It differs from the other Indo-Pacific species, B. townsendi, by having a smaller eye (3.5 % S.L. or less) and narrower interorbital width (3.8% S.L or less) ( Fig. 3). DiscussIONn.— Because one of the eleven species listed by Hubbs (1944) as a synonym of B. multibartata, B. townsendi, is valid, it was necessary to investigate the status of the other ten species because in many cases he worked only from the literature. Patrice Pruvost (MNHN) pro- vided me with a photograph of the dried holotype (A.8468) of Brotula burbonensis Kaup (1858). It has a large eye that is typical of B. multibarbata, not the smaller eye of B. flaviviridis. The types GREENFIELD: A NEW BROTULA FROM FII 83 of three species of Hubbs’ synonyms are at the = ad 6 . British Museum of Natural History: B. ensi- = (ae K | formis Giinther (1862) (stuffed syntypes) from eS = | Vanuatu, B. jayakari Giinther (1909) (BMNH S Wu =} ge g @ | 1888.12.29.193) from Oman, and three syn- = 4.5 ¢ | types of B. miilleri (corrected to B. muelleri) Whos i | Giinther (1909) (BMNH 1881.10.28.8 from 2 ape 4 | Ponape. 1868.8.1.7° from Tahiti, and ; ae f ie g 1876.5.19.45 from Tahiti). In his description of B. ensiformis, Giinther (1862) states “The ORBIT DIAMETER width of the interorbital space is less than the FiGurE 3. Interorbital diameter versus orbit diameter, horizontal diameter of the orbit,” which clearly both as percentage SL for Brotula flaviviridis (squares), and places itn B. multirate, Oe Cr (BMNH) has kindly examined the types of joands (circles). a ; these three species for me, and confirmed that B. ensiformis, and B. jayakari have the large eye typical of B. multibarbata. Brotula muelleri is rep- resented by three syntypes and is a mixed type series, containing both large and small eyed species. In the original description, Giinther stated that the width of the interorbital space is much smaller than the diameter of the eyes, which clearly places this species as a synonym of B. multibarbata as concluded by Hubbs (1944). To avoid further confusion, | am designating one of the syntypes, BM(NH)1876.5.19.45 from Tahiti, which has the large eye, as the lectotype of B. muelleri, and as such, it is a synonym of B. multibarbata. The other two syntypes, now paralectotypes of B. muel- leri, have the small eye, typical of B. townsendi and B. flaviviridis. Brotula palmietensis Smith (1935) (RUSI 299) from South Africa was listed as a synonym of B. multibarbata by Nielsen and Cohen (1986). Brotula formosae Jordan and Evermann (1902) (ZUMT 359) from Taiwan was listed as a synonym of B. multibarbata by Chen and Shao (1991). Brotula japonica Steindachner and Doderlein (1887) (NMW) from Japan was listed as a synonym of B. multibarbata by Lindberg and Krasyukova (1975) and also was not listed as a valid species in Nakabo (2002). Two of the species were described from the Hawaiian Islands, B. marginalis Jenkins (1901) and B. multicirrata Vaillant and Sauvage (1875). Hubbs (1944) examined the type of B. marginalis at the USNM (49694) and found it to be B. multibarbata with a damaged tail. Both Fowler (1900) and Hubbs (1944) considered B. multicirrata to be a synonym of B. multibarbata. It was described as having eight rather than six barbels on the snout. Many individuals of B. multibarbata have been collected at the Hawaiian Islands and none have had eight barbels. Hubbs (1944) speculated that the nasal flap had been counted as a barbel. Considering that only two species of Brotula are known from the well-collected Hawaiian Islands, and that B. townsendi is relatively rare, treating B. mul- ticirrata as a synonym of B. multibarbata is justified. Finally, B. ferruginosus (Tickell in Day, 1888) is not available (Eschmeyer 1998). Brotula flaviviridis appears to be most similar to B. townsendi, which was previously known only from the Hawaiian Islands and Johnston Island but now is known to be antiequatorial. Randall (in press) lists its distribution as “Known from the Hawaiian Islands, Johnston Island, Marshall Islands, Tonga, Loyalty Islands, and Vanuatu.” It is also known from the Chesterfield Islands and New Caledonia (this paper). Brotula flaviviridis shares the relatively small eyes with B. townsen- di, and has similar counts. Brotula townsendi is an orange brown with a yellowish pectoral fin in fresh coloration. Two lots at the Bishop Museum from the South Pacific were identified as B. townsendi: BPBM 22550 from New Caledonia and BPBM 33794 from Chesterfied Islands in the 84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 8 Coral Sea. Examination of these specimens confirmed that they appear to be conspecific with B. townsendi, clustering together in Figure 3. Brotula townsendi thus joins a number of other species from the Hawaiian Islands that demonstrate an antitropical or antiequatorial distribution (Randall, 1982). Brotula flaviviridis most likely was derived from these populations of B. townsendi to the southwest. ACKNOWLEDGMENTS I would like to thank K.R. Longenecker, R.C. Langston, and Bio Koroi Mataitini for assistance in collecting specimens. | also thank Captain B. Vasconcellos and the crew of the Moku Mokua Hine for assistance in our work. I am grateful to J. Seeto, GR. South, R.R. Thaman , and R.W. Tuxton of the University of the South Pacific, Fiji for facilitating our collections in Fiji. I also thank the Fijian Government and local village chiefs for permission to collect fishes. I thank Susan Monden for drawing the figure. Patrice Pruvost (MNHN) kindly photographed the holotype of B. burbonensis for me. Special thanks goes to Oliver Crimmen (BMNH) who examined the types of three species for me . W.N. Eschmeyer provided valuable guidance in nomenclatural matters. I also thank the staff at CAS for lending material and providing assistance: D. Catania, W.N. Eschmeyer, J. Fong, M. Hoang, and T. Iwamoto. This research was supported by National Science Foundation grants INT97-29666 and DEBO-1027545, and Sea Grant Project R/FM-6PD. LITERATURE CITED ALLEN, G.R. AND D.R. ROBERTSON. 1994. Fishes of the tropical Eastern Pacific. University of Hawaii Press, Honolulu, Hawaii, USA. 332 pp. BiocH, M.E., AND J.G SCHNEIDER. 1801. Systema ichthyologiae iconobus ex illustratum. Sanderiano Commissum, Berlin, Germany. 584 pp. CHEN, L.J., AND K.T. SHAO. 1991. Fishes of the families Ophidiidae and Bythitidae from Taiwan. Bulletin of the Institute of Zoology, Academia Sinica (Taipei) 30(1):9-18. COHEN, D.M., AND J.G. NIELSEN. 1982. Spottobrotula amaculata, a new ophidiid fish from the Philippines. Copeia 1982(3):479-S00. ESCHMEYER, W.N., ed. 1998. Catalog of Fishes. Vol. I. California Academy of Sciences, San Francisco, California, USA. 958 pp. Fow Ler, H.W. 1900. Contributions to the ichthyology of the tropical Pacific. Proceedings of the Academy of Natural Sciences of Philadelphia (1900):493—528. GUNTHER, A. 1862. Catalogue of the fishes in the British Museum, 4. The Trustees of the British Museum, London, England, UK. 534 pp. GUNTHER, A. 1909. Andrew Garrett’s Fische der Siidsee Journal des Museum Godeffroy 6:261-388. HILDEBRAND, S.F., AND O. BARTON. 1949. A collection of fishes from Talara, Peru. Smithsonian Miscellaneous Collections 111(10):1—36. Huss, C.L. 1944. Species of the circumtropical fish genus Brotula. Copeia 1944(3):162-178. JENKINS, O.P. 1901. Descriptions of fifteen new species of fishes from the Hawaiian Islands. Bulletin of the United States Fish Commission 19(1899):387-404. Jordan, D.S., and B.W. Evermann. 1902. Notes on a collection of fishes from the island of Formosa. Proceedings of the United States National Museum 25:315-368. Kaup, J.J. 1858. Ubersich der familie Gadidae. Archiv fiir Naturgeschichte 24:85-93. LINDBERG, G.U., AND Z.J. KRASYUKOVA. 1975. Fishes of the Sea of Japan and adjacent territories of the Okhotsk and Yellow Sea. Part 4. Teleostomi. XXIX. Perciformes. 2. Blennioidei. 13. Gobioidei (CXLYV. Fam. Anarhichthididae — CLXXV. Family Periophthalmidae). Akademiya Nauk, SSSR. 463 pp. NaKabo, T.(ed.). 2002. Fishes of Japan with Pictorial Keys to the Species, English edition. Vol. 1. Tokai University Press, Tokyo, Japan. 866 pp. GREENFIELD: A NEW BROTULA FROM FIJI 85 NIELSEN, J.G., AND D.M. COHEN. 1986. Family No. 96: Ophidiidae. Pages 344—350 in M.M. Smith and P.C. Heemstra, eds., Smith’s Sea Fishes. Macmillan South Africa, Johannesburg, South Africa. NIELSEN, J.G., D.M. COHEN, D.F. MARKLE, AND C.R. Rosins. 1999. Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated calagogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO species catalogue. FAO Fisheries Synopsis. 18(125). FAO, Rome, Italy. 178 pp. RANDALL, J.E. 1982. Examples of antitropical and antiequatorial distribution of Indo-West Pacific fishes. Pacific Science 35(3):197—209. RANDALL, J.E. (In Press.) Reef and shore fishes of the south Pacific: New Caledonia to Tahiti and the Pitcairn Islands. University of Hawaii Press, Honolulu, Hawaii, USA. 707 pp. SmitH, J.L.B. 1935. New and little known fishes from South Africa. Records of the Albany Museum 4:169-235. STEINDACHNER, F., AND L. DODERLEIN. 1887. Beitrage zur Kenntniss der Fische Japan’s (IV). Denkschriften der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Klasse 53: 257-296. TEMMINCK, C.J., AND H. SCHLEGEL. 1846. Pisces. Pages 173-269 in P. F. von Siebold, ed., Fauna Japonica, parts 10-14. Nagasaki, Japan. 324 pp. TICKELL, S.R. 1888. in F. Day. 1888. Fishes of India. Supplement. London, England, UK. pp. 779-816. VAILLANT, L., AND H.E. SAUVAGE. 1875. Note sur quelques espéces nouvelles de poisons des Iles Sandwich. Revue et Magasin de Zoologie Pure et Appliquée 38 (sér. 3, vol 3):278-287. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. 86 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, Short Communications Short Communications First RECORD OF THE EASTERN ATLANTIC SQUIRRELFISH SARGOCENTRON HASTATUM (HOLOCENTRIDAE) FROM THE WESTERN ATLANTIC OCEAN.— Each summer a ReefNet team con- ducts a trip to a different Caribbean destination to survey marine life using film and video to doc- ument their sightings. In 2003 the team surveyed St. Vincent Island, and in 2004, San Andres, Colombia. At both of these sites the team observed and photographed a squirrelfish that they never had seen before. The results of this most recent trip to San Andres were made available on the web (hittp://www.reefnet.ca) and sent to the third author by e-mail. Upon viewing the photographs of this squirrelfish, the third author recognized it as Sargocentron hastatum, a distinctive species previ- ously known only from the eastern Atlantic, where it has been recorded from Portugal southward to Angola, including the Cape Verde Islands. At St. Vincent the squirrelfish was first seen at a location called the Bat Cave, a cavern with a swim-through, located on the west side of the island north of Kingstown. The fish was observed at 9.1 meters (Figs. 1-3). At a second site about 90 meters south of the first location, two more spec- imens were observed swimming upside-down along the ceiling of a smaller cave. During dives along the southwest coast of San Andres, the team encountered cracks, caves, and caverns in the shallow water near shore, where craggy limestone drops steeply into the water. In two instances they sighted and attempted to photograph S. hastatum, and were successful in one case (Fig. 4). These fish also were at a depth of about 9.1 meters. All observed individuals were well back in a cave. The photographed fish was in the northeast corner of the cave that was about 9.1 meters wide, 1.2 meters high, and 6.1 meters deep. It was seen hiding from the dive lights among a few large rocks, was extremely shy, and would only reveal itself after a few minutes if the lights were turned off. Sargocentron hastatum (Cuvier 1829), is easily recognized by the distinct, alternating, red and white lines on the entire side of the body, a dorsal fin with distinct white spots on the basal half of the membranes, a very long third anal-fin spine reaching, when folded back, to the caudal-fin base, and a very long spine on the opercle (Greenfield 1981, in prep.). In earlier literature this species was referred to as Adioryx hastatus. The broad, deep-water region between the West African coast and the western tropical Atlantic is known as the Mid-Atlantic Barrier (Briggs 1974, 1995). Briggs (1974) calculated that for fish- es, this barrier was 91 percent efficient in preventing transatlantic movement, and that the predom- inant migratory movement across this barrier is from west to east. He goes on to say that “The west- ward colonization traffic appears to be restricted to certain dominant species that originated in the Indo-West Pacific and then gained access to the Atlantic by rounding the Cape of Good Hope. So far, there are no indications that species originating in the eastern Atlantic, and belonging to gen- era typical of that area, have been successful in becoming established on the western side.” Four holocentrid species are known from the eastern Atlantic Ocean, Corniger spinosus, Holocentrus adscensionis, Myripristis jacobus, and Sargocentron hastatum. All of these species, except S. has- tatum, are well known from the western Atlantic Ocean, and are thought to have crossed from there to the eastern Atlantic Ocean. Sargocentron hastatum, thus, is an exception. Although the genus Sargocentron has its greatest diversity in the Indo-Pacific Ocean (Randall 1998), S. hastatum is a species restricted to the eastern Atlantic and does not occur south of Angola. Luiz-Junior et al. (2004) reported on the occurrence of four eastern Atlantic fishes that have migrated from east to west, Acanthurus monroviae, Aulostomus strigosus, Epinephelus margina- WILK ET AL.: NEW RECORD FOR EASTERN ATLANTIC SQUIRRELFISH 87 tus, and Parablennius pilicornis, or 3.7% of the 106 reef-associated amphi-Atlantic fishes. The addition of S. hastatuwm raises this number to five, or 4.7%. ACKNOWLEDGMENTS.— We thank the following divers for assistance in field work: Any Wilk, Les Wilk, Scott Chandler, Patti Chandler, and Bill Tewes. We also thank Dive St. Vincent and Scuba San Andres for assistance in the surveys. LITERATURE CITED Briaccs, J.C. 1974. Marine Zoogeography. McGraw-Hill Book Co., New York, USA. 475 pp. Briaas, J.C. 1995. Global Biogeography. Elsevier, Amsterdam, The Netherlands. 472 pp. GREENFIELD, D.W. 1981. Holocentridae. Four pages (unnumbered) in W. Fischer, G. Bianchi, and W.B. Scott, eds., FAO Species Identification Sheets for Fishery Purposes. Eastern Central Atlantic, vol. 2. FAO and Department of Fish. and Oceans, Canada. GREENFIELD, D.W. (In prep.) Holocentridae. /n K. Carpenter, P. Oliver, M. Lamboeuf, and J. Martin, eds., The Living Marine Resources of the Eastern Central Atlantic. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, Italy. Luiz-JUNtor, O.J., S.R. FLOETER, J.L. GASPARINI, C.E.L. FERREIRA, AND P. Wirtz. 2004. The occurrence of Acanthurus monroviae (Perciformes: Acanthuridae) in the south-western Atlantic, with comments on other eastern Atlantic reef fishes occurring in Brazil. Journal of Fish Biology 65:1173-1179. RANDALL, J.E. 1998. Revision of the Indo-Pacific squirrelfishes (Beryciformes: Holocentridae: Holocentr- inae) of the genus Sargocentron, with descriptions of four new species. Indo-Pacific Fishes (27):1—105. — Kris Wilk!, Keri Wilk?, and David W. Greenfield3:4. 1.2 ReefNet Inc., 3610 Walnut Grove, Mississauga, Ontario L5L2W8, Canada; 3 California Academy of Sciences, 875 Howard Street, San Fran- cisco, CA 94103. FIGURES 1—3. Sargocentron hastatum, St. Vincent. Photo by Les Wilk. FiGURE 4. Sargocentron hastatum, San Andres. Photo by Keri Wilk. + Research Associate, Department of Ichthyology, California Academy of Sciences and Emeritus Professor, University of Hawaii. Mailing address: Moss Landing Marine Laboratory, 8272 Moss Landing Road, Moss Landing, CA95039. Email: <greenfie @ hawaii.edu>. Corresponding author. 88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, Short Communications ADDITIONS TO THE ACANTHACEAE OF ARIZONA.— Two species of Acanthaceae are reported from Arizona for the first time. Both Carlowrightia texana and Ruellia parryi have been collected recently in Chihuahuan desertscrub in the southeastern portion of the state (Cochise County). Illustrations of both species and identification keys for all species of Carlowrightia and Ruellia in Arizona are provided. Fifteen species of Acanthaceae are now recognized as native to the state. Carlowrightia texana Henr. & T.F. Daniel, Madrono 26:27. 1979. ARIZONA: Cochise Co.: San Pedro Riparian National Conservation Area, Upper San Pedro River floodplain near old Tombstone River gauge, ca. 2 mi N of Hwy. 82, 31°45.244’N, 110°12.87’'W, 6 Sep 2002, E. Makings 1299 (ASU). This represents the first collection of the species in Arizona, where it is infrequent on a rocky granite substrate at 1153 meters elevation in Chihuahuan desertscrub dominated by Acacia greg- gii, Larrea tridentata, Rhus microphylla, Prosopis velutina, Flourensia cernua, Talinum angustis- simum, and Acourtia wrightii. The locality in western Cochise County represents the westernmost known occurrence of the species. A detailed description C. texana was provided by Daniel (1983); illustrations of the species are available in the protologue (Henrickson and Daniel 1979) and in Powell (1988); its pollen was figured in Daniel (2004); and Figure 1 shows some of the major mor- phological attributes of this taxon. It can be distinguished from other species of Carlowrightia in Arizona by the following key: 1. Shrubs to 2 m tall; corolla purplish, lower-central lobe not keel-like or enclosing stamens; anthers yellow; leaves sessile to subsessile, lamina linear, 4-35 times longer than wide, mid- vein only evident; head of capsule subglobose to partially flattened......... C. linearifolia 1. Perennial herbs, usually less than | m tall; corolla white (often with colored markings or veins), lower-central lobe keel-like, at least partially enclosing stamens; anthers maroon (turning blackish); leaves generally petiolate, lamina lanceolate to ovate to cordate to elliptic, 1—3.7 (—5.3) times longer than wide, several orders of venation evident; head of capsule distinctly flattened laterally. 2. Corolla 8-18 mm long, upper lip with a yellow “eye” surrounded by maroon markings; bracteoles subulate to triangular, 0.3—1.3 mm wide; seed margin dentate; Sonoran desertscrub 23... 82s See Se et ee Seo ae oe eee C. arizonica 2. Corolla 5.5—7 mm long, upper lip lacking a yellow “eye,” veins of all lobes usually maroon; bracteoles foliaceous, usually petiolate, ovate to elliptic-lanceolate, 0.7—-5 mm wide; seed margin entire; Chihuahuan desertscruby. 2222-2. 22242425540 sen eee C. texana Carlowrightia texana occurs in Chihuahuan desertscrub, mesquite woodlands, and oak-juniper woodlands in the southern United States (Arizona, New Mexico, Texas) and northern Mexico (Chihuahua, Coahuila, Nuevo Le6én, San Luis Potosi, Sonora). Plants have been collected at eleva- tions from 25 to 1700 meters. Throughout its range, C. texana flowers from March until November. Arizona plants flowered in September (possibly in response to summer monsoonal rainfall). In Arizona the species is undoubtedly rare, but it likely occurs elsewhere among the isolated regions of Chihuahuan desertscrub in the southeastern portion of the state. The nearest known occurrence of C. texana to that reported here is about 75 km to the southeast in northeastern Sonora, Mexico, from which state the species was recently (Daniel 2004) reported for the first time. Ruellia parryi A. Gray, Syn. Fl. N. Amer. 2(1):326. 1878. ARIZONA: Cochise Co.: S-facing slopes of big limestone hill, N of Guadalupe Canyon Road, 31°21’01”N, 109°06’54’W, 1372 m, 8 Apr 2001, M. Chamberland 1900 (ARIZ). SS ee DANIEL ET AL: ADDITIONS TO THE ACANTHACEAE OF ARIZONA 89 FiGURE 1. Carlowrightia texana (a—c) and Ruellia parryi (d-f). a. Habit (Daniel 100). b. Flower from above (Daniel 108). c. Seed (McVaugh 8204). d. Habit with flower (Daniel 581). e Branched trichomes from leaf surface (Reina G. et al. 2003-957). £. Seed (Breedlove & Mahoney 71916). Ruellia parryi (Fig. 1) 1s reported here for the first time from Arizona, where it occurs in south- eastern Cochise County. Daniel (1984, 2004) provided descriptions of R. parryi; its pollen was fig- ured in Daniel (2004); and illustrations of the species can be found in Vines (1960), Wasshausen (1966), and Powell (1988). The species occurs in the southwestern United States (Arizona, New Mexico, western Texas) and northern Mexico (Chihuahua, Coahuila, Durango, Nuevo Leon, Sonora, Zacatecas). Plants occur on limestone in Chihuahuan desertscrub and juniper chaparral at elevations from 500 to 1950 meters. Throughout its geographic range flowering takes place from March through October. In Arizona, flowering plants were collected in April from a region of 90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, Short Communications Chihuahuan desertscrub (associated species: Fouquieria splendens, Atriplex canescens, Quercus pungens, Rhus microphylla, Tecoma stans, Salvia parryi). Ruellia parryi was also recently collect- ed in northeastern Sonora for the first time (Daniel 2004), about 45 km southwest of its occurrence in Arizona. The two species of Ruellia known from Arizona can be distinguished as follows: 1. Herbs from woody caudex; leaves 20-170 mm long, 10-90 mm wide, attenuate at base, surfaces lacking any branched or stellate trichomes; inflorescence of (1—) 3— many-flowered dichasia borne on peduncles up to 85 mm long from axils of leaves or distal bracts; calyx (9—) 11-25 mm long, anterior lobes not fused for half or more of their length; corolla lobes 10-12 mm long; capsule 12—22 mm long, glandular pubescent; seeds 3-4 m long......... R. nudiflora 1. Subshrubs; leaves 9-35 mm long, 3-15 mm wide, acute at base, surfaces with at least some tri- chomes branched or stellate; inflorescence of | (—3)-flowered dichasia borne on peduncles 0.5—6 mm long from leaf axils; calyx 6—11 mm long, anterior lobes fused for half or more of their length; corolla lobes 5-10 mm long; capsule 10-13 mm long, eglandular; seeds 2.5—3 MT TOMS aie Foe. seth ansdeee Fe are: & Sue oes sls Mimslonal allot eee tay ay ne ae R. parryi Fifteen species of Acanthaceae are now known to occur in Arizona: Anisacanthus thurberi, Carlowrightia arizonica, C. linearifolia, C. texana, Dicliptera resupinata, Dyschoriste decumbens, Elytraria imbricata, Henrya insularis, Justicia californica, J. candicans, J. longii, J. sonorae, Ruellia nudiflora, R. parryi, and Tetramerium nervosum. The distributions of all of these species in the state are either concentrated in or restricted to the southern counties. Among counties in the state, Cochise County in southeastern Arizona has the most species of Acanthaceae (12). This coun- ty also possesses nearly all of the Chihuahuan desertscrub in the state. The recent collections of both Carlowrightia texana and Ruellia parryi in southeastern Arizona and northeastern Sonora (Daniel 2004) suggest that additional botanical exploration of the isolated regions of Chihuahuan desertscrub in these states is warranted. Srenandrium barbatum is another species of Acanthaceae that might be expected to occur in southeastern Arizona and/or northeastern Sonora based on its Known occurrence in western portions of the Chihuahuan Desert in New Mexico and Chihuahua (Daniel 1985). LITERATURE CITED DANIEL, T.F. 1983. Carlowrightia (Acanthaceae). Flora Neotropica 34:1—116. DANIEL, T.F. 1985. A revision of Stenandrium (Acanthaceae) in Mexico and adjacent regions..Annals of the Missouri Botanical Garden 71:1028—1043. DANIEL, T.P. 2004. Acanthaceae of Sonora: taxonomy and phytogeography. Proceedings of the California Academy of Sciences 55:692-807. HENRICKSON, J., AND T.F. DANIEL. 1979. Three new species of Carlowrightia (Acanthaceae) from the Chihuahuan Desert Region. Madrono 26:26—36. POWELL, A.M. 1988. Trees & Shrubs of Trans-Pecos Texas, including Big Bend and Guadalupe Mountains National Parks. Big Bend Natural History Association, Big Bend National Park, Texas, USA. 536 pp. VINES, R.A. 1960. Trees, Shrubs, and Woody Vines of the Southwest. University of Texas Press, Austin, Texas, USA. 1104 pp. WASSHAUSEN, D.C. 1966. Acanthaceae J. St. Hil., Expos. Fam. 1:236. 1805, in C. Lundell, Flora of Texas 1(3): 223-282. Texas Research Foundation, Renner, Texas, USA. — Thomas F. Daniel!, Elizabeth Makings?, and Michael Chamberland?. ! Department of Botany, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103; Email: tdaniel@ calacade- my.org; 2 Tonto National Forest, 2324 E. McDowell, Phoenix, AZ 85006; 3 Cheekwood Botanical Garden, 1200 Forrest Park Drive, Nashville, TN 37205. MOSER ET AL.: FIRST RECORD OF PLACOBELLA PARASITICA IN CALIFORNIA 91 First REPORT OF AN EASTERN UNITED STATES SPECIES OF BLOOD-FEEDING LEECH, PLACOBDELLA PARASITICA (EUHIRUDINEA, GLOSSIPHONIIDAE), IN CALIFORNIA.— Placobdella par- asitica (Say, 1824) is an opportunistic blood-feeding leech on turtles. It has been reported from 22 species and subspecies of turtles (Moser 1995; Watermolen 1996) and 4 species of amphibians (Meyer and Moore 1954; Waffle 1963; Watermolen 1998). Any turtle species occurring in North America is considered a potential host. Placobdella parasitica is abundant and widely distributed throughout the north-central and eastern United States, and southern Canada to Alberta (Sawyer 1972; Klemm 1982, 1985). It is most abundant in the Great Lakes region, where it reaches great numbers (e.g., 768 on one host tur- tle) (Sawyer 1972; Klemm 1982, 1985; Brooks et al. 1990). Placobdella parasitica 1s less known in the western United States. MATERIALS AND METHODS.— Placobdella parasitica specimens (n=4) were collected on Trachemys scripta elegans (Red-Eared Slider) in Mission Valley, San Diego River (San Diego County, California) on 24 July, 2003. Additionally, two free-living specimens of P. parasitica were collected at Birch Lake, Camp Mather (Tuolumne County, California) on 4 May, 2004. Leech spec- imens were deposited in the Invertebrate Collections of the National Museum of Natural History, Smithsonian Institution (USNM 1026199) and the California Academy of Sciences (CASIZ 171366). RESULTS AND DISCUSSION.— This is the first record of P. parasitica from the state of California. To date, no published records of species of the genus Placobdella have been reported from California (Sawyer 1972; Klemm 1982, 1985). Holland (1991, 1994) mentioned that leeches attached to approximately 7—10% of the population of Actinemys marmorata (Western Pond Turtle) in northern California and central Oregon, but referred to them as presumably Placobdella. These specimens could possibly have been P. parasitica, P. ornata or P. papillifera. Except for two reports (Arizona and Nevada), P. parasitica has not been reported from west of the Rocky Mountains. The likely source of the geographic voucher of P. parasitica from Nevada (USNM 33987) was re-examined and has been re-identified as P. ornata. The voucher record of P. parasitica from Arizona is unknown. It is possible that P. parasitica may have been accidentally introduced to California with the introduction of exotic eastern and central United States turtles. Placobdella parasitica will likely be found in more localities in the western United States upon fur- ther collection. ACKNOWLEDGMENTS.— We thank the National Wildlife Health Center (USGS; Madison, Wisconsin) for collection of this species in Southern California, Dr. Dan Holland for permission to cite reports and Dr. T’Shaka Toure (USGS, Irvine, California). LITERATURE CITED Brooks, R.J., D.A. GALBRAITH, AND J.A. LAYFIELD. 1990. Occurrence of Placobdella parasitica (Hirudinea) on snapping turtles, Chelydra serpentina, in southeastern Ontario. Journal of Parasitology 76(2): 190-195. HOLLAND, D.C. 1991. A Synopsis of the Ecology and Status of the Western Pond Turtle (Clemmys marmora- ta) in 199]. United States Fish and Wildlife Service, National Ecology Research Center, San Simeon Field Station, San Simeon, California, USA. 141 pp. HOLLAND, D.C. 1994. The Western Pond Turtle: Habitat and History. United States Department of Energy, Bonneville Power Administration, Portland, Oregon, USA. DOE/BO-62137-1. 334 pp. Kiem, D.J. 1982. Leeches (Annelida: Hirudinea) of North America. EPA-600/3-82/025. United States Environment Protection Agency, Environmental and Support Laboratory, Cincinnati, Ohio. USA. 177pp. KLEMM, D.J. 1985. A Guide to the Freshwater Annelida (Polychaeta, Naidid and Tubificid Oligochaeta, and Hirudinea) of North America. Kendall/Hunt Publishing Company, Dubuque, Iowa, USA. 198 pp. 92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, Short Communications Meyer, M.C., AND J.P. Moore. 1954. Notes on Canadian leeches (Hirudinea) with the description of a new species. The Wasmann Journal of Biology 12(1):63—96. Moser, W.E. 1995. Placobdella parasitica (Rhynchobdellida: Glossiphoniidae) from the Eastern River coot- er (Chelonia: Emydidae) in Oklahoma. The Texas Journal of Science 47(1):71-74. SAWYER, R.T. 1972. North American Freshwater Leeches, Exclusive of the Piscicolidae, with a Key to All Species. Illinois Biological Monographs 45. 154 pp. WAFFLE, E.L. 1963. An Ecological Study of the Iowa Glossiphoniidae (Annelida: Hirudinea) with Emphasis on Feeding and Reproductive Habits. MS Thesis. University of lowa, lowa City, Iowa, USA. 54 pp. WATERMOLEN, D.J. 1996. Notes on the leech Desserobdella picta (Hirudinea: Glossiphoniidae). Journal of Freshwater Ecology 11(2):211-217. WATERMOLEN, D.J. 1998. A new parasite record from the tiger salamander (Ambystoma tigrinum). Bulletin of the Chicago Herpetological Society 33(12):262. — William E. Moser!, Brian D. Quelvog”, and Peter Hovingh?. ! Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012; 2California Department of Fish and Game, San Joaquin Valley Southern Sierra Nevada Region, Sonora, CA 95370; 3Utah Museum of Natural History, Salt Lake City, Utah 84112-0050. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. Diatoms Taxonomy and Ecology: A Marriage of Necessity Proceedings of a Workshop 23-27 October 2002, Kulice, Poland Edited by J.P. Kociolek! and A. Witkowski2 !Diatom Collection, California Academy of Sciences, 875 Howard Street San Francisco, CA 94103, and 2Department of Paleooceanology, Institute of Marine Sciences, University of Szczecin, Waska 13, PI-71-415 Szczecin, Poland 93 Table of Contents JOHN PATRICK KOCIOLEK: Preface to Taxonomy and Ecology: A Marriage of Necessity. Proceedings of a Workshop 23—27 October, 2002, Kulice, Poland ................... JOHN PATRICK KOCIOLEK: Taxonomy and Ecology: Further Considerations .............. ROGER J. FLOWER: A Review of Diversification Trends in Diatom Research with Special Reference to Taxonomy and Environmental Applications Using Examples from Lake Br AatkealeanGdue: |SEWHETE Oe, sou. Sis EGR a BS ths eC eI ra eee Koto 3 ALEKO MIHO AND ANDRZEJ WITKOWSKI: Diatom (Bacillariophyta) Flora of Albanian @oastal) Wetlands) Taxonomy andiEcology::AvReview <.2-s.44-- «22ed sees ees EUGENE F. STOERMER: Marrying Taxonomy and Ecology: An Attempt.................. FRITHJOF A.S. STERRENBURG: Taxonomy and Ecology: An Inseparable Pair.............. GRAHAM J.C. UNDERWOOD: Microalgal (Microphytobenthic) Biofilms in Shallow TELSE MEYER, JULIA SAUER, ANDREAS OELTJEN, KLAUS WENDEROTH, JURGEN MARQUARDT, AND ERHARD RHIEL: Structural, Biochemical and Molecular Investigations on Wadden Seay Diatoms: Field Studies;and Laboratory Experiments. acne at - ese Davip M. WILLIAMS AND GERALDINE REID: The Sino-Siberian Distribution of Eunotia Clevei and its Relatives, from Lake Baikal to the Mekong Delta: Wie Wnion or laxonomybiogeosraphy.and ECologys 1. sce eee ae oe eee ae 95 a \ Participants in the Workshop Taxonomy and Ecology: A Marriage of Necessity held 23-27 October 2002 in Kulice, Poland PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 9, p. 97 April 29, 2005 Taxonomy and Ecology: A Marriage of Necessity Proceedings of a Workshop 23-27 October 2002, Kulice, Poland Preface John Patrick Kociolek !Diatom Collection, California Academy of Sciences, 875 Howard Street San Francisco, CA 94103; Email: pkociolek@ calacademy.org The workshop, “Diatom Ecology and Taxonomy: A Marriage of Necessity,” was convened by J.P. Kociolek and A. Witkowski, from 23-27 October 2002 at the University of Szczecin’s Conference Center in Kulice, Poland. The workshop was sponsored by the California Academy of Sciences (San Francisco), University of Szczecin (Poland), Alexander von Humboldt Foundation (Germany) and Carl von Ossietzky University (Oldenburg, Germany). A total of 48 participants from Albania, Belgium, Belarus, Croatia, Denmark, Germany, Hungary, Ireland, Macedonia, Poland, Russia, Spain, Sweden, Ukraine, United Kingdom and USA were in attendance. A number of invited speakers raised issues related to the workshop’s title, with regard to fos- sil and recent diatoms, and freshwater, estuarine and marine habitats. Viewpoints of those focused primarily on taxonomy and ecology were presented. The plenary presentations provided the spring- board for lengthy facilitated discussions. The areas of overlap and separation were discussed, and topics that might promote synergy for the two disciplines explored. The papers comprising this vol- ume represent most of the invited speakers at the workshop. The workshop could serve as a model for future discussions by diatomists across scientific disciplines. The organization of and ongoing support for the Workshop by faculty, staff, and students of the University of Szczecin was above reproach. Their efforts assured our workshop’s ultimate suc- cess, and the attendees were both aware of and greatly appreciated the hard work that went into dealing with the workshop’s many practical and impractical elements. ) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 10, pp. 99-106, 4 tables April 29, 2005 Taxonomy and Ecology: Further Considerations John Patrick Kociolek Diatom Collection, California Academy of Sciences 875 Howard Street, San Francisco, CA 94103; Email: pkociolek @ calacademy.org In this overview, points of the initial publication by Kociolek and Stoermer are reviewed and expanded upon, noting the gulf between the disciplines of taxonomy and ecology and the modest ways the two areas currently intersect. Some research areas for synergy are suggested. Previously published data on diatom community structure are assessed in terms of decreasing taxonomic resolution. The varying lev- els of taxonomic refinement are compared to the original data, and argue for the finest degree of taxonomic resolution. The final session of the workshop is summa- rized related to the desirability and implementation of on-line taxonomic resources, especially floras. The publication of our paper on the need for a marriage of diatom taxonomy and ecology (Kociolek and Stoermer 2001) prompted a wide range of comments and responses directed to us, some constructive, some not, and was one of the most requested either of us have ever published. It also prompted the gathering/workshop, which generated the series of papers included herein. The workshop was developed to foster dialogue on the general topic of describing and promoting ways for taxonomists and ecologists to collaborate. Despite some initial cultural hurdles, by the end of the workshop participation by the entire group was open, honest and constructive. The present report has three objectives: First, it reviews, amplifies and (hopefully) clarifies some of the points made in the earlier paper related to integrating taxonomic and ecological stud- ies. Second, an example is based on previously published data showing the relationship (and dependency) on ecology and relative to fine- and coarse-grained taxonomy. Lastly, one of the areas of extensive discussion that coalesced the opinions of many in the final workshop session is sum- marized and discussed; namely the potential and desirability of flexible, on-line taxonomic prod- ucts, particularly floras. TAXONOMY AND ECOLOGY-MORE DIFFERENCES THAN SIMILARITIES Tradition is one of the greatest barriers separating the disciplines of Ecology and Taxonomy. The number of strictly taxonomic (versus morphological/ultrastructural) papers in the diatom liter- ature is relatively modest, compared to the number of papers with ecology as a focal point. And the breadth of places for publications of ecological papers far exceeds the relatively smaller number of publication venues for taxonomic works. Taxonomic works rarely contain any specific ecological data-cursory summaries are usually the most offered. When included, ecological data have rarely been used to substantially forward our knowledge about the taxonomy of specific entities. Many ecological studies use taxonomy as a means to an end, but the species lists (usually containing many errors in names and authorities) usually suggest a continuation of previous approaches (and errors) rather than any serious analysis of taxonomic results (though many more diatoms are being seen by ecologists than taxonomists). Finer-grained taxonomy and the current state of flux in fresh- 99 100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 10 water diatom nomenclature appear to offer difficult hurdles for ecologists. The formal structures of our discipline help to perpetuate the gulf between taxonomists and ecologists. At meetings of the international society, oral presentations are arranged to separate presentations by ecologists and taxonomists; the two groups usually meet as they pass by each other going into/out of their respec- tive sessions. The divide between taxonomy and ecology is reflected not only in the outright absence of tax- onomists in the area of ecology, but in the approaches the two groups have to taxonomy. Although, of course, there are exceptions to the stereotypes/generalizations I offer here, I would suggest that the goal of most taxonomists is what I call “correct taxonomy,” that is trying to understand the mor- phological variability through the ontogeny of the cell and cell cycle (Kociolek and Williams 1986), what the circumscribed taxon should be called in the context of other taxa. Assignment of the taxon under study with in the Linnaean hierarchy not only offers an information storage- retrieval system (so that other data, including ecological data) can be compared and used in a wide range of studies, including comparative autecology, comparisons across space and time, alien species), but also predictive value so the hierarchy system should express the phylogenetic/evolu- tionary position of the taxon (Kociolek 1998). These typical taxonomic studies are usually organ- ized and presented in stylized formats and with jargon, which appear arcane to ecologists. The back-and-forth that can result from such studies suggest more uncertainty than certainty (recall the numerous exchanges related to the proposal of the genus Naviculadicta — e.g., Moser et al. 1995, Kociolek 1996). The result of these debates is a wait-and-see stance from ecologists using the tax- onomy, such that the impact of much of the primary literature is delayed unnecessarily. Even when the fervor of discussion has died down, and an approach has more or less been settled upon, some have found it difficult to embrace the consensus (e.g., Camburn and Charles 2000). To achieve their research agenda, ecologists, on the other hand, focus attention on “consistent” taxonomy, making sure that the entities they encounter in the light microscope can be reliably applied to their respective group, hopefully representing some “real” biological entity. Ecologists face the challenge to avoid “shoe-horning” specimens into already-established taxon names, and to ensure their “consistent” grouping have some biological reality. Another way taxonomists and ecologists are separated relates to their search for places to con- duct their research programs. Ecologists, for the most part, seem to be attracted to systems that have been in some way impacted, usually by the activities of human beings. A driver towards these systems and their related questions may be funding, as governments (locally, regionally, national- ly and internationally) seek understanding to possible impacts and remediation required. Taxonomists tend to seek out non-impacted situations to explore and conduct their research pro- grams. It is usually these places that support native and undescribed taxa (e.g., Moser et al. 1995, 1998; Metzeltin and Lange-Bertalot 1998), and it has been suggested that an understanding of the biogeography of freshwater diatoms has been clouded by the impacts of the human species on the distribution of species (Kociolek and Spaulding 2000). Due to human disturbance, taxonomists are less likely to encounter species that might bear on the questions they pose. Thus, an issue so basic as Where we work helps to divide taxonomists and most ecologists. It should be noted, however, that as ecologists seek to understand “pristine” or “unimpacted” situations, and the structure of communities found in those types of habitats, partnering with taxonomists in the investigation of those systems would be a wonderful setting for the marriage of taxonomy and ecology. A common concern of those using diatoms to estimate water quality is the (in)ability to iden- tify every individual encountered in their slides/counts. Many times the specimens are rare, or in cases of taxa where the entire population has nearly synchronized division, thus encountering many individuals, but who occupy a narrow range of variation. It is well documented that two or more KOCIOLEK: TAXONOMY AND ECOLOGY 101 distinct taxa may overlap morphologically, especially at the small end of the size range (Stoermer et al. 1986: Theriot and Ladewski 1986; Geitler 1932). Thus, although it may be impossible for the practicising ecologist to undertake studies to understand the full range of variation in a taxon (thus providing insights into the identity of the taxon), it must also be realized that closely related species share many similarities (hence the close evolutionary relationship), and thus it might be difficult to separate/identify isolated individuals. An analogy might be the difficulty to identify deciduous trees to species when leaves are off the tree, but easier when leaves and flowers are out. Thus, whereas the current state of ecological and taxonomic research is of two nearly inde- pendent fields, their interrelationships are at best utilitarian, with any dependency in terms of rela- tionships driven by taxonomists supporting ecologists with flora (rare, actually) or serving as “hired gun” identifiers of individual or groups of species. We believe that this separation of disciplines does not have a long, sustainable future. Integration of the two disciplines is a goal worth pursuing, with the payoffs of a more rigorous, robust enterprise (more students, more positions, research impacts and more funding) that approaches answers to questions posed by ecologists and taxonomists. Areas that are and should integrate these approaches include paleolimnology-done successfully, evolutionary ecology (espe- cially of ecologically pristine areas), conservation biology, co-evolution, and biogeography. TAXONOMIC RESOLUTION AND ITS IMPACTS ON ECOLOGICAL INTERPRETATIONS The interplay between taxonomy and ecology perhaps finds no closer relationship, the mar- riage is not more intimate, than the use of diatoms in the assessment of ecological conditions. A myriad of approaches has evolved since the early ideas of Kolkwitz and Marsson (1908) to apply diatoms to understanding freshwater ecology in particular, especially focused on human impacts (pollution). Excellent overviews of these appoaches can be found in Patrick and Roberts (1979), Cholnoky (1968) and Stoermer and Smol (2001). In quantitative approaches, early workers suggested robust analyses (robust in the sense of sta- tistics/mathematical models) required enumeration of thousands (in some cases tens of thousands) of valves to achieve reliable results (e.g., Patrick et al. 1954; Hohn 1961; Hohn and Hellerman 1963; Patrick 1968). Since the days of these extensive identification and enumeration methods, efforts were made to save time in the analysis of samples and to essentially reduce/minimize cost (because many of these studies were by now in the U.S. being funded by government agencies) but derive “correct” assessments of water conditions. This has led to a variety of approaches where counts are reduced to a certain number of valves (e.g., Stevenson and Pan 2001; 300—600/sample seems to be settled upon without too much debate) or until no new taxa have been encountered (e.g., Charles et al. 2002; assuming richness plays some role in the analysis/understanding of water quality). Although the effort has focused on reducing the number of valves to count (yet still achieving a correct assessment), few studies have looked at ways to reduce taxonomic resolution and still achieve a “correct” understanding of water quality. In other words, does all the “fuss” made by tax- onomists to identify and separate taxa at the level of species, variety and form (and this is being done with renewed vigor-e.g. Lange-Bertalot and Metzeltin 1996; Reichardt 1999 as good exam- ples) contribute in a substantitive way to our understanding of water quality? In the era when scientists were exploring myriads of ways to apply diatoms to water quality studies, complete data sets were often published (e.g., Hohn and Hellerman 1963; Patrick 1968; Patrick and Roberts 1979), as opposed to the summary statistics and data plots seen in most “mod- em’ analyses. 102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 10 To explore whether taxonomic resolution, i.e. coarse or fine-grained taxonomy, mattered in the interpretation of ecological data, we selected data sets published by Patrick (1968) on Darby Creek, Pennsylvania (USA). For each data set, we calculated species richness and Shannon-Wiener diver- sity, for three conditions, including the data set as presented (with identifications made by Patrick and finest taxonomic resolution presented in the paper at that time (what we have termed “ALL’’) and two levels of reduced taxonomic resolution. In one case, we subsumed all subspecific epithets into the species (“SPECIES”) and then all species and subspecific epithets subsumed into genera (“GENUS”). The richness and diversity calculations were then ranked for each of the 8 stations based on the three different levels of taxonomic resolution. We then made the assumption that the complete or “ALL” dataset, with the finest level of taxonomic resolution most closely representing the relative ecological conditions of the eight samples. We then compared these relative relation- ships with those derived from approaches with more coarse-grained taxonomies, to see how well they might serve as proxies for the finest-level of taxonomic resolution. We should note here that the finest-grained taxonomy applied in 1968 probably does not rep- TABLE |. Comparison of Rich- : Up eater resent the finer distinctions made today. ness and Shannon-Weiner Di- versity including all taxa. In Table | are listed the richness and diversity measures for the eight samples provided in Patrick (1968) for “ALL,” “SPECIES” and Sample Richness | Diversity | “GENUS.” The eight samples are then ranked from most to least rich, | _ and from most to least diverse in Table 2. The richness and diversi- Sample | | ¢ : 9 Mat 10s) 4.6555 | ty measures are provided for pennate taxa only in Table 3. Ranking SPECIES | 88 43523 | Of the pennate taxa measures is provided in Table 4. GENUS | 22 2.6942 The data suggest that the samples were relatively rich, in the pial | data including pennate and centric taxa, with total taxa numbers Sample 2 iS ALL | 104 | 46967 | ranging from 101 to 112 in the ALL samples, 84-98 taxa in the ee > 2 SPECIES samples and 19-23 taxa in the GENUS samples. Diversity ~— 7, ranged from 4.5170 to 4.9641 in ALL samples 4.2581 to 4.6687 in Samples | SPECIES samples and 2.4761 to 2.8527 in the GENUS samples. In the ALL 103 4.9641 peace leet cer cicicheatuavllicaleninn h SEEGIES a 4.6087 tankings, sample 8 was the richest in all calculations, w ereas sam- | GENUS | 20 2.8527 ple 6 ( ALL, SPECIES) and sample 7 (GENUS) were poorest in terms ISeaiaie ; —_— of taxa. Interestingly, in terms of diversity, sample 3 was most ye | Renan RAYG409 diverse in each of the three sample calculations, whereas sample 8 b it = — == — . . C 1 i | SPECIES | 95 4.366 | (the richest in terms of number of taxa) was the least diverse in all NU 93 D5 < 0 : GENUS 22/4 2! 25? three calculations. Assuming the ALL samples best approximated the Sample 5 | | P slag “true” con- | ALL 108 4.8087 TABLE 2. Ranking of samples by ALL, SPECIES only dui chee SPECIES 94 4.4895 and GENUS only for Richness and Shannon-Weiner S | GENUS 21. | 2.776 | Diversity with all taxa included. “#shared” shows the p E Cee — number of rankings that are in agreement with All taxa. rankings of ant ; 101 4.8705 | CLLenmess f | 6/05 Richness Richness | Richness | Diversity] Diversity | Diversity SPECIES | 85 4.5502 All Species Genus All __| Species | Genus _ | matched GENUS | 21 Agel aaa | exactly the | } 8 8 8 3 3 3} ix l | 4 = - Sample? i 7 7 5 6 5 ALL calcula ALL | 107 4.7037 5 5 l 5 eG tions, where- | SPECIES | 92 4.3687 7 7 5 LER PORE A bee? as GENUS | GENUS 19 2.6464 1 f g : | : | : THlancenead 2 2 2 Sample 8 3 3 3 ge ae rT only 50% of ALL 112 4.517 6 6 > 8 | 8s | 8....| the cankanes SPECIES 98 4.2581 | . ; t | For diversi- GENUS 23 2.4761 #shared with ALL 4 out of 8 |6 out of 8} 3 out of 8 KOCIOLEK: TAXONOMY AND ECOLOGY 103 + TABLE 3. Comparison of TABLE 4. Ranking of samples by ALL, SPECIES only ty, SPECIES Richness and Shannon-Weiner and GENusS only for Richness and Shannon-Weiner matched Diversity including pennate Diversity with pennate taxa only included. “#shared” 75q% of the taxa only. ee the number of rankings that are in agreement with rankings of taxa. Sample | Richness | Diversity FNAL but Sampled | Richness Richness | Richness | Diversity| Diversity | Diversity GREEN SUES ALL | 94. | 4.3976 All | Species | Genus All | Species | Genus matched ALL SPECIES | 79 | 4.0794 | | 8 8 8 3 3 6 in less than GENUS | 19 [2.2934 | | 4 4 4 6 6 3 40% of the | | | 5 5 1 5 5 5 Sample2_| | 7 7 5 7 5 i rankings. ALL os oa 2 D 5 2 4 7 In data SPECIES | 80 | 4.1681 | | r ai | 3 1 6 I 7 2 : : GENUS | 18 | 22619 | | i 3 5 7 , 7 including 7 | | AA eral | | 6 6 5 8 8 8 pennate taxa Sample 3 | ] ich ALL SS eee) Only DUCHESS SPECIES 79 | 4.389 | |#shared with ALL | 6 out of 8 |2 out of 8 4 out of 8} 3 outof 8) ranged from GENUS 17 2.4439 | 93-102 taxa Sample4 _| | ' in ALL, 79-88 in SPECIES and 16-20 in GENUS. Diversity ranged aeetiES c ae | from 4.2620 to 4.7224 in ALL, from 3.9695 to 4.3890 in SPECIES GENUS 0 1 1704 | and from 2.0455 to 2.4439 in GENUS. In the rankings, data for pen- — | nates only mirrored the total taxon scores, while sample 8 being the ample 5 | : : 6 : ALL [100 | 4.6403 | most rich, and sample 6 being least rich in ALL and SPECIES and SPECIES 86 4.2801 sample 7 least rich in the GENUS calculation. Likewise, sample 3 GENUS So EZ Z2 9 was the most diverse and sample 8 (the most species rich) was least Sample 6 | _ diverse. Order of ranking of samples in terms of richness, 6 of the 8 ani a | eure | rankings of SPECIES were the same as ALL, whereas only 2 of 8 were GENUS 118 257g. the same between GENUS and ALL. Order of ranking of samples in Sample? terms of diversity, as in pennate and centric taxa, showed less corre- ALL 99. | 44825 | Spondence between SPECIES and ALL (4 out of 8) and GENUS and SPECIES 83 4.0871 | ALL (3 out of 8). GENUS | 16 2.2709 | ; These data seem to suggest that even modest changes in taxo- Sample 8 nomic resolution can lead to large changes in the relative ranking of ALL 102 4.262 3 SSEaIES a8 | 3.9695 samples (up to 50% difference). In other words, reduced taxonomic GENUS 20 2.0455 | resolution does not provide accurate prediction of relative rankings of water condition. The surprising result of an inverse relationship between species richness and diversity suggest even the most common measures of water quality analysis may require further critical evaluation. Further analysis is needed, with robust statistical power, on the impacts of reduced taxonomic resolution on predicting the relative rankings of water conditions. CREATING MODERN TAXONOMIC TOOLS FOR A LARGE, INTERNATIONAL USER COMMUNITY The workshop discussed at length ways in which taxonomic information can best be conveyed to the broad community of diatomists, serving both taxonomists and ecologists (and others as well). Praise was evident for projects like the Siisswasserflora (Krammer and Lange-Bertalot 1986-1991), with its great taxonomic and geographic breadth, detailed taxonomic information, and incredible photo documentation. A second project also hailed by the workshop participants, though more restricted in scope, was the series on diatoms from the Baltic Sea (Snoetjs and co-workers, 104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 10 1993-1998). The project succinctly brings together illustrations with listings of important litera- tures and helpful comments into a common, useful format. It also represents a collaboration of sci- entists from different labs, countries and perspectives. Shortcomings of traditional floras include the lack of tying images or distributions explicitly to specimens in publicly accessible collections, uncertainty or lack of studies documenting syn- onymies, lack of detailed geographic summaries, the static nature of data and the high cost of the published volumes. Given the limited number of formally trained taxonomists and systematists worldwide (Kociolek and Stoermer 2001) and the increased possibilities for interaction and collaboration afforded by the internet, many workshop participants saw the opportunities and benefits of devel- oping an on-line flora. This concept has been discussed in part by Kociolek (accepted). Such a flora could also be linked to/integrated into other information systems that are already in place or in development that offer templates for achieving additional goals (offering the ability for users to provide comments and feedback, allowing the flora to create dialogue and be a dynamic entity for several possible communities; see one example dealing with the freshwater diatoms of south Florida (and the system in place at Academy of Natural Sscience, Phildelphia [ANSP] as current examples). The call for on-line tools such as floras was recently presented in Science (Wheeler et al. 2004). Information that can be an integral part of an on-line flora include name (linked to databases on nomenclature), description, important references (linked to on-line literature databases), veri- fied distributions (linked to collection/herbarium databases), reported distributions (linked to liter- ature databases), images (linked to image databases), types (linked to collection databases) and the person(s) responsible for the entry information. There is currently being developed enough infor- mation infrastructure available on-line such that an on-line flora is possible. It is time for members of the diatom community to work towards producing this much-needed tool—a tool that would serve both taxonomists and ecologists. LITERATURE CITED CHARLES, D., C. KNOWLES, AND R.S. Davis, EDS. 2002. Protocols for the Analysis of Algal Samples Collected as part of the U.S. Geological Survey National Water-Quality Assessment Program. Report No. 02-06. Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania, USA. 124 pp. CHOLNOKY, B.J. 1968. Die Okologie der Diatomeen in Binnengewdissern. J. Cramer, Lehre, Germany. 699 pp. GEITLER, L. 1932, Die Formwechsel der pennaten Diatomeen (Kieselalgen). Archiv fiir Protistenkunde 78:1-226. Houn, M. 1961. The relationship and population density in diatom populations from Silver Springs, Florida. Transactions of the American Microscopical Society 80:140—-165. Hon, M., and J. HELLERMAN. 1963. The taxonomy and structure of diatom populations from three eastern North American rivers using three sampling methods. Transactions of the American Microscopical Society 82:250-329. KocIoLek, J.P. 1996. Taxonomic instability and the creation of Naviculadicta Lange-Bertalot in Moser and Lange-Bertalot, a new, catch-all genus of diatoms. Diatom Research 11:223-224. KOcIOLEK, J. P. 1998. Does each genus of diatoms have at least one unique feature? Diatom Research 13:177-179. KOcIOLEK, J.P. (In press.) Some thoughts on the development of a diatom flora for freshwater ecosystems in the continental United States and a listing of Recent taxa described from U.S. freshwaters. Hydrobiologia. Festschrift for Rex Lowe. KOCIOLEK, J.P., AND S.A. SAULDING. 2000. Freshwater diatom biogeography. Nova Hedwigia 71:223-241. KOCIOLEK, J.P,. AND E.F. STOERMER. 2001. Taxonomy and ecology: A marriage of necessity. Diatom Research KOCIOLEK: TAXONOMY AND ECOLOGY 105 16:433—42. Kocio_ek, J.P,. AND D.M. WILLIAMS. 1987. Unicell ontogeny and phylogeny: Examples from the diatoms. Cladistics 3:274-284. Koikwitz, R., AND M. Marsson. 1908. Okologie der pflanzliche Saprobien. Berichte der Deutsche Botanischen Gesellschaft 26:505-519. KRAMMER K., AND H. LANGE-BERTALOT. 1986. Bacillariophyceae. Naviculaceae. 2/1. Jn: H. Ettl, J. Gerloff, H. Heynig, and D. Mollenhauer, eds., Siisswasserflora von Mitteleuropa . Gustav Fisher, Stuttgart, Germany. 876 pp. KRAMMER K., AND H. LANGE-BERTALOT. 1988. Bacillariophyceae. Bacillariaceae, Epithemiaceae, Surirellaceae. 2/2. In: H. Ettl, J. Gerloff, H. Heynig and D. Mollenhauer, eds., Siisswasserflora von Mittel- europa . Gustav Fisher, Stuttgart, Germany. 596 pp. KRAMMER K., AND H. LANGE-BERTALOT. 1991a. Bacillariophyceae. Centrales, Fragilariaceae, Eunotiaceae. 2/3. In: H. Ettl, J. Gerloff, H. Heynig, and D. Mollenhauer, eds.,.Siisswasserflora von Mitteleuropa. Gustav Fisher, Stuttgart, Germany. 576 pp. KRAMMER K., AND H. LANGE-BERTALOT. 1991b. Bacillariophyceae. Achnanthaceae. Kritische Erginzungen zu Navicula (Lineolatae) und Gomphonema. 2/4. Jn: H. Ettl, J. Gerloff, H. Heynig, and D. Mollenhauer, eds., Siisswasserflora von Mitteleuropa. Gustav Fisher, Stuttgart, Germany. 437 pp. LANGE-BERTALOT, H., AND D. METZELTIN. 1996. Indicators of Oligotrophy. 800 taxa representative of three ecologically distinct lake types. Carbonate, oligodystrophic, weakly buffered soft water. Iconographia Diatomologica 2:1-396. METZELTIN, D., and H. LANGE-BERTALOT. 1998. Tropical diatoms of South America I. About 700 predominate- ly rarely known or new taxa representative of the neotropical flora. Jconographia Diatomologica 5:1—695. Moser G., A. STEINDORF, AND H. LANGE-BERTALOT. 1995. Neukaledonien. Diatomeenflora einer Tropeninsel. Bibliotheca Diatomologica 32:\—340. Moser G., H. LANGE-BERTALOT, AND D. METZELTIN. 1998. Insel der Endemiten. Geobotanisches Phanomen Neukaledonien. Bibliotheca Diatomologica 38:1—464. PATRICK, R.M. 1968. The structure of diatom communities in similar ecological conditions. The American Naturalist 102:173-183. PATRICK, R.M., AND N.A. RoBerts. 1979. Diatom communities in the Middle Atlantic states, U.S.A. Some factors that are important for their structure. Nova Hedwigia, Beiheft 64:265—284. PaTRICK, R.M., M. HOHN, AND J. WALLACE. 1954. A new method for determining the pattern of the diatom flora. Notulae Naturae (259):1—12. REICHARDT, E. 1999. Zur Revision der Gattung Gomphonema. Die Arten G. affine/insigne, G. angustatum/micropus, G. acuminatum sowie gomphonemoide Diatomeen aus dem oberoligozién in Bohmen. [conographic Diatomologica 8:1—203. SNOEUS, P., ED. 1993. Intercalibration and Distribution of Diatom Species in the Baltic Sea, vol. 1. Opulus Press, Uppsala, Sweden. 129 pp. SNOEUS, P., AND S. VILBASTE, EDS. 1994. Intercalibration and Distribution of Diatom Species in the Baltic Sea, vol. 2. Opulus Press, Uppsala, Sweden. 125 pp. SNOEUS, P., AND M. Potapova, EDS. 1995. Intercalibration and Distribution of Diatom Species in the Baltic Sea, vol. 3. Opulus Press, Uppsala, Sweden. 125 pp. SNOEUS, P., AND X. KASPEROVICIENE, EDS. 1996. Intercalibration and Distribution of Diatom Species in the Baltic Sea, vol. 4. Opulus Press, Uppsala, Sweden. 125 pp. SNOEUS, P., AND N. BALASHOVA, EDS. 1998. Intercalibration and Distribution of Diatom Species in the Baltic Sea, vol. 5. Opulus Press, Uppsala, Sweden. 143 pp. STEVENSON, R.J., AND Y. PAN. 2001. Assessing environmental conditions in lakes and streams with diatoms. Pages 11-40 in E.F. Stoermer and J.P. Smol, eds., The Diatoms. Application for Environmental and Earth Sciences. Cambridge University Press, Cambridge, UK. 469 pp. STOERMER, E.F., T.B. LADEWSKI, AND J.P. KocIoLEK. 1986. Further observations on Gomphoneis. Pages 205-213 in M. Ricard, ed., Proceedings of the 8" Symposium on Living and Fossil Diatoms, Paris, 1984. STOERMER, E.F., AND J.P. SMOL, EDS. 2001. The Diatoms. Application for Environmental and Earth Sciences. Cambridge University Press, Cambridge, UK. 469 pp. 106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 10 THERIOT, E., AND T.B. LADEWSKI. 1986. Morphometric analysis of shape of specimens from the neotype of Tabellaria flocculosa (Bacillariophyceae). American Journal of Botany 73:224-229. WHEELER, Q.D., P.H. RAVEN, AND E.O. WILSON. 2004. Taxonomy: Impediment or Expedient? Science 303(5656):285. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11, pp. 107-128, 4 figs., 1 table. April 29, 2005 A Review of Diversification Trends in Diatom Research with Special Reference to Taxonomy and Environmental Applications Using Examples from Lake Baikal and Elsewhere Roger J. Flower Environmental Change Research Centre, University College London 26 Bedford Way, London WC1H OAP, UK; Email: rflower@ geog.ucl.ac.uk The study of diatoms has diversified strongly since its beginnings in the 18 centu- ry. Early work focused upon collecting material for taxonomic analysis but by the mid 19th century a strong global perspective had already developed. In the 20 cen- tury, use of diatoms in biostratigraphy, environmental change and ecology and in biogeography proliferated as did work on diatom ultra structures, life cycles, and systematics. Most recently, systematics and molecular genetics have sought to reveal diatom genealogy and to refine taxonomy. Similarly, diatoms are used increasingly as time-space indicators of water quality and there is renewed interest in global diversity patterns. Taxonomy underpins these applications but changing concepts can either diminish or enhance the value of diatoms to the environmental sciences. Some ways in which taxonomic analysis of diatom assemblages can benefit environ- mental research are discussed with reference to material from Lake Baikal and else- where. Because of their intricate siliceous cell walls and their global distribution in aquatic and ter- restrial environments, diatom microalgae have attracted the interest of naturalists and researchers alike since the 18" century. Nevertheless, it was mainly during the 20 century that the value of these organisms in ecological and, due to their resistant skeleton, in palaeo-environmental research became fully recognised. In recent decades, the major role of diatoms in global primary productiv- ity and the carbon cycle has been demonstrated (e.g., Round et al. 1990). At the same time, con- cern has increased about global biodiversity and the survival of species. Although microorganisms, diatoms are relevant to these issues. They are major contributors to global primary productivity and are abundant in all the earth’s major aquatic ecosystems, from the oceans, through wetlands to the alpine lakes. These ecosystems are all perceived as being threatened indirectly or directly by human activities, and time-space changes in the distribution of diatom taxa can provide valuable evidence about the nature and pace of global environmental change. There are several ways in which diatoms are particularly relevant to global change and biodi- versity issues. They are species diverse and, because of their well known ecological tolerances, they can indicate the nature of environmental change (climate, pollution, and habitat loss, for example). Diatom-environment data sets are now widely available whereby diatom assemblages can be used to predict nutrients, pH and salinity in freshwaters (see Stoermer and Smol 1999; Battarbee et al. 2001). However, with renewed interest in the ‘ecological quality’ of habitats, species monitoring and species comparisons using diatoms (especially in combination with other organismal groups) play an important role in the ecological assessment of water quality types (e.g., 107 108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 Coste et al. 1991). Consequently, diatoms are included within the recently introduced Water Framework Directive (WFD, see Foster et al. 2001) for the ecological management of European fresh and transitional waters (e.g., Kelly 2002). Long-term monitoring programmes are particular- ly useful for detecting trends in inter-annual species change, in relation to acidify for example (Monteith and Evans 2000). A special characteristic of the diatoms is their ability to form rich sedimentary archives, and this has been exploited in a variety of palaeo-environmental applications from marine biostratigra- phy to recent pollution. Sedimentary diatoms are also becoming increasingly useful in conserva- tion policy and management of water quality, particularly those issues focused upon in the WFD, where identification of reference conditions for impacted water bodies is required. Sedimentary diatoms, as well as other sub-fossil organism in dated sediment cores, can provide excellent eco- logical records of pre-impact conditions and can be used to identify modern reference lakes (Flower et al. 1997). The particular value of palaeolimnological techniques is the provision for pro- viding a variety of reference data where information from long-term monitoring and other sources are absent (Smol 2002). Application of diatom analysis to environmental issues rests fundamentally on adequate, pre- cise and harmonized taxonomic conventions and techniques. The recognition of morphotaxa in the environment and the unambiguous attribution of ecological optima and tolerances are the essence of diatom-based palaeolimnological reconstruction and of diatom ecology. Even without attribu- tion of these optima, the recognition of taxa remains of fundamental importance to other aspects of diatom research such as biogeography and systematics. Recent developments in diatom taxonomy regarding biogeographical limits for some diatom taxa and the re-definition of many taxa accord- ing to new taxon concepts (Mann 1999) have considerable implications for diatom based environ- mental studies. In the short-term, there is inevitable confusion concerning the new concepts (Stoermer 2001) but, if a more precise, comprehensive and appropriate taxonomic system arises, both diatom diversity appraisal and diatom ecology/palaeoecology will benefit. This paper pro- vides an overview of the diversification of diatom research and examines some issues, applications and implications of recent taxonomic revisions for assessing diatom diversity. DIVERSIFICATION TRENDS IN DIATOM RESEARCH Diatoms are readily observable in the light microscope and their occurrence was well known by the late 18" century (see Round et al. 1990), but it was not until the 19" century that awareness of the large diversity of micro-organisms developed as microscopical techniques improved. In the past 150 years or so, diatom research has diversified into several major fields ranging from ecolo- gy and micropalaeontology to systematics and molecular genetics. Although these diversification trends are indicated broadly in Figure 1, some of the perhaps less known recent applications are omitted (e.g., toxicology and archaeology, see Stoermer and Smol 1999). EARLY TAXONOMY.— The pioneering work of Ehrenberg, Greville and others during the early part of the 19" century (see Fig. 1) established diatoms as an important biological group and sev- eral classical works followed (e.g., Ehrenberg 1854). These works not only focused on describing species and erecting a taxonomic system for diatoms but the authors also were often very interest- ed in the provenance of diatom taxa. Ehrenberg, for example, corresponded extensively with Charles Darwin seeking diatomaceous material from exotic locations. He was also aware of the sig- nificance of the indicator value of marine and freshwater taxa in samples. Darwin sent numerous diverse samples including atmospheric ‘dust’ collected during the voyage of the Beagle (Darwin 1839) and the facial paint used by the indigenous people of Tierra del Fuego (see Burkhardt and FLOWER: TRENDS IN DIATOM RESEARCH 109 Smith 1987). A little later Cleve reported widely on diatoms from disparate global locations (e.g., Cleve 1878, 1894/5). Hence, even at this early stage of diatom research, there was much interest in the geographic distribution of taxa. The early emphasis was mainly on obtaining material for describing new taxa, and it was not until later in the 20" century that diatoms and biogeography began to receive systematic attention (see below). The first diatom catalogues appeared in the late 19 century (Habirshaw 1877; Schuett 1896), and more specific accounts of historical diatom stud- ies are available elsewhere (see Patrick 1982). BIOSTRATIGRAPHY.— The significance of diatoms in stratigraphic sequences was recognised in the latter part of the 19" century (e.g., Gregory 1854; Kitton 1870). This work developed into diatom micropalaeontology through the studies of Hanna (1927), Calvert (1929), Jouse (1939) and others on terrestrial deposits. In the first part of the 20" century, marine diatom biostratigraphy was mainly restricted to terrestrial deposits, but with the development of deep sea sediment coring tech- nology, palaeoceanography and diatom analysis rapidly expanded, firstly in the Pacific Ocean (Jousé 1960; Burckle 1972; Sancetta 1979; Barron and Baldauf (1995). The international Deep Sea Drilling Project and then the Ocean Drilling Programme opened up the world’s oceans to palaeo- ceanography and diatom biostratigraphic analyses of marine sediment cores. Such analyses have enabled chrono-biostratigraphies to be established and permitted the major taxon groups to be arranged into an evolutionary series according to geological time (e.g., Strelnikova 1990). Fresh- water diatom biostratigraphy also expanded and Bradbury (e.g., 1986) began examining continen- tal scale freshwater diatom deposits in exposures and in cores. In the 1990s, deep coring of Lake Baikal (the international Baikal Drilling Programme, BDP) was begun and has recovered an essen- tially continuous sedimentary diatom record for the past 8 million years (BDP-98 Members 2001). EcoLoGy.— The 19" century workers (e.g., Gregory 1854; Smith 1856; and Kitton 1870) can also be considered as the progenitors of diatom ecology through their observations and subjective inferences about environmental preferences of diatom taxa. The value of diatoms as ecological indicators of water quality became clearer during the 20" century as the central role of water chem- istry in influencing diatom taxa abundances in lakes and rivers was recognised. Firstly, the value of diatom taxa for indicating trophic gradients was established (Kolkwitz and Marson 1908), then for salinity (Kolbe 1928) and pH (Hustedt 1937-39). Environmental applications followed with diatoms being used to monitor river water quality (e.g., Patrick et al. 1954). From the 1970s, mul- tivariate methods were developed to quantify relationships among diatom taxa and environmental variables (e.g., Birks et al. 1990). Temperature was identified in the 1900s as an important descrip- tor of diatom abundances, but this was later discounted (see Patrick and Reimer 1966). Temperature has attracted renewed interest (e.g., Pienitz et al. 1995), but its value as a common predictor for diatom taxa remains questionable. The tight relationships of diatom taxa with their environment nevertheless makes these organisms highly suitable for ‘ecological’ monitoring of water quality (e.g., Kelly 2002) and of recovery responses following partial mitigation of water acidification (Monteith and Evans 2000). Irrespective of water quality relationships, diatom floris- tics alone can contribute to integrated biodiversity assessment of aquatic habitats (Flower 2001). PALAEOECOLOGY AND PALAEOLIMNOLOGY.— Nipkow (1920), working in Lake Zurich, prob- ably initiated the environmental reconstruction approach using sedimentary diatoms. The work of Deevey (1942) and Pennington (1943) then demonstrated the value of diatoms in sediment cores for palaeolimology. With the advent of precise sediment dating, diatoms became powerful tools for reconstructing recent limnological change (e.g., Battarbee 1978). However, it was multivariate analysis of contemporary diatom-water quality data sets that made palaeolimnological reconstruc- tions quantitative and numerically sound (e.g., Birks et al. 1990, Battarbee et al. 2001, Smol 2002). Robust models are now available for reconstructing a variety of environmental changes, including 110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 pH (Birks et al. 1990), salinity (Fritz et al. 1991) and nutrients (Bennion 1994). More recent devel- opments include the use of oxygen isotopes in diatom silica to infer lake level changes (Barker et al. 2001) and of diatoms to indicate lake enrichment by fish (Finney et al. 2000). On longer time- scales, changes in freshwater diatom species down long cores have enabled palaeoclimate infer- ences to be deciphered for several millions of years. For northern California, Bradbury et al. (1991) described climate driven floristic changes over the past 3 million years whereas the diatom record in Lake Baikal sediments now extends palaeoclimate information for more than 5 million years (Khursevich et al. 2001). LIFE-CYCLES AND EVOLUTION.— Diatom life-cycles interested Victorian naturalists such as GHK Thwaites and W. Smith and, in Germany, L. Rabenhorst; they described several types of reproduction strategies and spore formation (e.g., Smith 1856). Specific interest in life cycles and reproduction was energised by MacDonald (1869) and Pfitzer (1869) following their observations on division and cell size change. Cell division continued to receive much attention in the 20' cen- tury, especially from the cytological perspective (e.g., Geitler 1927; von Stosch 1958). Much about morphogenisis of the silica cell wall was understood by the 1980s (Volcani 1978). Towards the end of the 20" century, the significance of life cycle strategies and of resting stages for ecology was being emphasised (e.g., Jewson 1992). Understanding more about the detail of diatom sexuality has enabled the recognition of within species ‘demes’ that are reproductively isolated and sympatric (see Mann 1984, 1999). The significance of the species unit in evolution is currently under debate (ibid.). TAXONOMY AND SYSTEMATICS.— In the first part of the 20" century, taxonomic research increased as Hustedt (1909) began his remarkable half century of diatom publications. Floras spe- cific to more exotic locations also increased and included such regions as South America and Siberia (Frenguelli 1923, Skvortzow 1937). As the number of diatom taxa proliferated and nomen- clatural changes ensued, a recording system for diatom names and synonyms became necessary. Comprehensive catalogues, first by Mills (1933-35) and later by VanLandingham (1967-79) and Fourtanier and Kociolek (1999), followed. Major new freshwater diatom floras, which relied main- ly on photographs rather than on drawings, began to appear in the 1980s, largely due to the work of Krammer and Lange-Bertalot (e.g., 1986). Interestingly, this period (the last two decades of the 20 century) and the 1840s were the two most prolific for naming new diatom genera (Fourtainer and Kociolek 1999). The advent of electron microscopy (EM) made fine resolution of diatom ultra structures possible (e.g., Helmcke and Krieger 1951, 1953-77) and led the way to the recent taxo- nomic revisions and the descriptions of new taxa with more structural details. These improvements also enabled ideas about diatom systematics to advance (Simonsen 1971) and, using Hennig’s ideas about cladistic analysis of shared derived characters, taxa were arranged into clades and depicted on cladograms to indicate genealogy (Williams 1985). Advances in molecular biology and in par- ticular the polymerized chain reaction (PCR) for amplifying genetic base sequences, became avail- able by the late 1980s. This permitted the analysis of genetic material, rather than of morphologi- cal characters, to used to infer systematic relationships within the diatoms (Bhattacharya et al.1992), leading to new ideas about diatom phylogeny. NEW TAXON CONCEPTS.— With the rapid developments in ultra structure, molecular genetics and reproduction, it is unsurprising that taxon concepts began to change. The diatom species con- cept can be based on several views (see Theriot 1992, Mann 1999), but, suffice to say here, stud- ies using living and fossil taxa will necessarily always be at least partly based on morphological evidence. Concepts about non-species level taxa also began to change since the 1970s (e.g., Ross and Sims 1973, Lange-Bertalot and Simonsen 1978, Williams 1985, Round et al. 1990). The trend to describe new genera has expanded especially for biraphid taxa. The recent literature contains an ‘ FLOWER: TRENDS IN DIATOM RESEARCH 111 array of these new genera, but it is littered with invalid diagnoses, conflicting designations and idiosyncrasies associated with particular authors. Molecular genetics offers a way of establishing relationships amongst taxa and to some extent validating morphotaxa (Medlin 1997). Gene sequencing work on diatoms has helped in the validation of Aulacoseira taxa (cf. Edgar and Theriot 2002), but it is unlikely that the technique can be applied to all the controversial morphotaxa in the foreseeable future. REGIONAL FLORAS AND BIOGEOGRAPHY.— The proliferation of new names, especially in the past decade, has come at a time of renewed interest in diatom biogeography and biodiversity and in diatoms in remote areas and on islands. Whilst providing further evidence for widespread distri- butions of some taxa, the incidence of discontinuous distributions and endemism is becoming more widely appreciated. However, taxa with regionally restricted distributions were well known in the 19% century (see Cleve 1895/6). Endemism is a special case of restricted geographical distribution, and its high incidence in ancient Lake Baikal was recognised early in the 20 century (Dorogostaisky 1904; Skvortzow and Meyer 1928). Baikalian, Siberian (Palaearctic), relict and other endemic elements were then recognised in Baikal’s benthic diatom flora. Also notable for regional endemism are the recent floras of the Patagonian region (Rumrich et al. 2000) and espe- cially of the island of New Caledonia (Moser et al.1998). As long ago as Smith (1856), the wide- spread distribution of many diatom taxa as compared with higher plants was recognized, and because many diatom taxa are undoubtedly cosmopolitan, this view has persisted (Lund 2002). However, a minority of diatom taxa do appear to have restricted regional distributions, the signif- icance of which in systematics and biogeography is now being evaluated (Williams 1995; Kociolek and Spaulding 2000). The diversification of diatom research indicated in Figure | suggests that the main lines of development are diverging through time. This impression 1s only partly justified because current interests in diatom biogeography, biodiversity, ecology and palaeoecology are interrelated. Furthermore, morphotaxonomy provides the essential foundation for these linked lines of research. Hence, there is a necessity to improve ways in which taxonomic convention 1s achieved and dis- seminated to all investigators, researching or otherwise. Diatom diversity in time and in space has and will make major contributions to both environmental science and to evolutionary studies. One current challenge is to provide a firm validated taxonomic framework for achieving these aims. TAXONOMIC OBJECTIVES, PROTOCOLS AND INFORMATION DISSEMINATION The science of taxonomy is the search for natural order; or is it? According to S.J. Gould, quot- ed in Hakansson (2002), taxonomy is about genealogy and the search for natural order, but the International Code of Botanical Nomenclature (ICBN, Greuter et al. 2001) states that the role of taxonomy is simply to supply a means of referring to a taxon and indicating its rank. The binomi- al system, developed by Linnaeus and upon which the ICBN is founded, was conceived as an aide memoir for individual taxa and this usage persists, despite other suggested systems (cf. Forey 2002). The binomial names and the higher taxonomic levels can usefully indicate something about phylogeny and taxon group affinities, but it is the role of systematics to elucidate biological rela- tionships using homologous characters or genetic markers (Kitching et al. 1998). In much applied diatom research, the relationships amongst taxa are usually of less importance than the relationships of taxa (usually species level or below) with environmental variables. Nevertheless, typification and the unambiguous designation of diatom taxa remains of fundamen- tal importance in both systematics and environmental applications. Use of diatoms in time-space diversity studies is increasing and requires taxonomic consistency not only for attributional reasons Volume 56, No. 11 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 112 “svar JOofgns-v.QUI UdIMJOg UOHRI]IOv} PUB SUOTIL.IDUI JO da1Bap oy) Ajayenbape ayeorput jou SOOP WILISLIP OY} IY) PUL (GGG [OWS PUP IOLIAO]G 99S) DASNeYXe JOU SI SvoIR YOIRASAI WO Ip Jo Ave GY OOOT~ OUP IY) “Ir ‘stsayjUared UT payeoIpUT ore sddUSJayOI WOVRIP -UOU JN JURATAI puv JULIOdUUT [eIDAAS “1 ‘JeY} ION “SOLIN{US. OM) Ised ay} SULINP YOIRASaI WIP JO UONITJISIOAIP dU) UL Spud.) SuNKoIpur weAseIp ArewUUNs Y “| AUN 0661 0002 Je J9 punoy ye Ja YouWwny 9861 punoy F SWENII/\ pue jojejieg ¥ Jewel 4 SVHO1d IVNOIDSY i) { | { eZ6! I anv ua@suolU|S | | jojeyiag © Sld3ONOO NOXVLM3N | eee peeel | i a | 62-2961 weybupue7 ue, 495844 8 | 1 666) UueW AYOU/AH 200g uossueyey—: = GS-LS61L ; Nyon. 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It is therefore wise to consider ways in which diatom taxono- my can be made more user-friendly, less ‘messy’ (Mann 1999) and less ambiguous to promote con- sistency at both local and international levels. When a new diatom taxon is described, either at the variety (Flower 1989), species (Skvortzow 1937) or genus (Vyverman et al. 1998) level, typification should be relatively straight forward, assuming the author(s) have carefully checked the literature and relevant type material and obey the conventions (see Greuter et al. 2000). Formal typification is, therefore, regulated. In reality, however, the system is far from straight forward, and difficulties are often amplified when original investigations of particular taxa are translated into identification floras. There is currently a lack of coherence in diatom taxonomy that obstructs consistent and accurate species identifications. This is especially important for routine diatom counting within research consortia where light microscopy is principally used. Taxonomic inconsistencies amongst authors can obfuscate studies of both regional flora comparisons and species distribution patterns. There are several specific ways in which dis-harmonies can arise and these include: 1. The tendency to combine some taxa into rather broad species groups (e.g., Lange-Bertalot and Simonsen 1978) was reversed in the 1980s as many of the traditional species were subdivided into new genera (Williams and Round 1986; Round et al. 1990). This has led to conflicts in the literature (e.g., Kociolek 1998) and creating new genera by re-designation is currently on going (e.g., Round and Maidana 2001). New genera of course have merit where they better reflect different morphological groups but the ICBN definition of taxonomy suggests that some of the genus splitting effort may be questionable because what- ever legitimate binomial is used the taxon characteristics are still amenable to systematic analysis. 2. The recognition of sympatric taxa based mainly on sexually incompatible subgroups or demes (cf. Mann 1999) raises the issue of describing and typifying many more diatom taxa. The tendency to subdivide con- ventional species taxa began earlier (e.g., Knudson 1953; Koppen 1975). Steps towards increased taxo- nomic resolution are generally welcomed by those using diatoms as environmental indicators because it can improve discrimination of environmental change (Flower and Battarbee 1985). However, if a system is adopted whereby reproductively incompatible demes are designated as species (Mann 1999), the great multiplication in names for within groups of (sympatric) taxa, all characteristic of the same environmen- tal conditions, will contribute little more to understanding the role of environmental variables. 3. Increased taxonomic resolution and separating taxa by using details of ultra-structure is an important part of taxonomic research. However, when such details alone are used to discriminate taxa and especially at the generic level then difficulties for routine light microscopy can arise. Defining a taxon using single cri- teria, for example striae ultra structure (cf. Kobayasiella micropunctata [Germain] Lange-Bertalot) or sep- arating two genera (Cyclostephanos and Stephanodiscus) using ultra structure detail of the external rimo- portula (see Guerrero and Echenique 2001) is problematic and such ultra structure requirements led Camburn and Charles (2000) to abandon many taxonomic revisions in their work on the North American softwater diatom flora. 4. There is an increasing trend to use informal and/or (usually) inadequate descriptions of taxa in some floral and palaeoecological studies (e.g., Stevenson et al. 1991). Unverified book-form floras are extremely use- ful for opening up new areas for investigating diatom diversity (e.g., Rumrich et al. 2000) but they do not provide site-specific species inventories, and can obfuscate identifications by non-specialists. Informal designations can be a useful expedient, but lack of information can make widespread use of inadequately described taxa impossible (examples are many, such as Frustulia spec., Cyclotella kuetzingiana [agg. which without accompanying images are of questionable value). Elsewhere, Gould (1989) used the term ‘shoe-horning’ to describe the tendency to fit potentially new taxa into pre-existing descriptions; undoubt- edly this practise has caused difficulties in diatom identification and taxonomic comparisons especially amongst biogeographical regions (Williams 1995). 5. The existence of type specimens for all described taxa together with precisely catalogued synonymies should in a perfect world permit validation of problematic taxa. In practice, type specimen material is often inaccessible, unavailable, poorly conserved or is imprecisely identified. There are many cases where type 114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 material in diatom herbaria is not mounted on glass cover slips (especially the older collections) so mak- ing observation difficult. For more modern material, type slides are often lodged in collections without any indication as to which is the type specimen. In mixed material this can make selection of the type subjec- tive or at worst makes the preparation virtually useless for taxonomic validation purposes. The practical problems of making type material more amenable are now being tackled as tech- niques to display and characterise individual specimens are ever improving. However, some taxon characteristics will always be questioned and the extents to which current literature conflicts can be rectified is debatable. Taxonomic research cannot, nor should it, be carried out by consensus, but focusing conflicts in the primary research literature would help reduce ambiguity for taxono- my users. Where integration of diatom distributions in time-space is needed, neither molecular nor cytoplasmology is likely to resolve current problems. Neglecting the value of morphotaxa in favour of other taxonomic tools is probably not yet a practicable option in most environmental work. Applied diatom research is growing and there is a pressing need to introduce more consensus into standard taxonomy. For example, European protocols for diatom monitoring of rivers (e.g., Kelly 2002) advocate use of particular identification floras, but this standardisation attempt 1s subjective. Some steps to reconcile current difficulties could include: 1. Abandon the binominal species name concept and develop a uninominal system (e.g., Ereshefsky 2001), so inter alia eliminating confusions caused by generic changes 2. Abandon the binomial naming system and develop an iconographic numerical coding system as is used for chrysophyte cysts (see Duff et al. 1995) 3. Continue with the current system but introduce a consensus platform into taxonomy in a way that reduces ambiguities and makes diatom iconographs and regional floras more accessible to end-users. The latter step is in many ways preferable and would allow taxonomic research to continue its erratic individualistic course but permit progress towards a coherent world diatom flora. This flora, although very incomplete at first, could incorporate many existing works and be updated follow- ing decisions made through an authorised consensus forum. In reality, this practice has already been used to harmonise diatom taxonomy within several research consortia projects (Stevenson et al. 1991; Gell 1988; Camburn and Charles 2000). Internet and web site developments are begin- ning to make such harmonization procedures available on a wider scale. Nevertheless, local con- cepts and project driven agenda continue to dominate data sets. If AQC (analytical quality control) is to be achieved widely, a permanent over-arching central body or tightly co-ordinated group is needed. This would ideally result in a maintained web site database of images, descriptions and names for all known diatom taxa as well as provenance and environmental information. Appropriate recording of taxonomic decisions and descriptions should follow ICBN protocols (Greuter et al. 2001) and type specimens must remain the ‘gold standard’ in diatom taxonomy. Using the virtually limitless capacity of web-based data sets, combined with ease of updating, makes their use attractive and considerable progress has already been achieved. See the following: http://www.algaterra.org, for taxonomic baselines and original type material; http://www.calacademy.org/research/diatoms/, for cataloging diatom names and synonymies; http:/Avww.ualg.pt/adiac, for diatom image identification aids; http://www. geog.ucl.ac/ecrc/enclosed/darwin.htm, for diatom images from Lake Baikal; http:/>vww. geog.ucl.ac.uk/ukawmn, for monitoring diatom taxa and water chemistry; http://www.craticula.ncl.ac.uk8000/Eddi, for images and taxa with ecological optima. This latter site is particularly useful for applied diatom taxonomy users because it combines taxonomy, ecology and provenance data for application to diatom assemblages in sediment cores so that environmental reconstructions of water quality can be made (Battarbee et al. 2001a). FLOWER: TRENDS IN DIATOM RESEARCH 1S Diatoms are key organisms for monitoring natural water quality in Europe and elsewhere, but major taxonomic contradictions need to be reconciled. Taxonomy does suffer from a lack of glob- al organisation and planning generally (Alberch 1993), yet taxonomic consistency is a key neces- sity in diatom research (Kociolek and Stoermer 2001), and it is difficult to envisage how this can be achieved internationally without access to a coherent reference system. If a web-based global image database were developed, this would inter alia help harmonise diatom taxonomy (for inter- national monitoring programmes, biogeographic comparisons, etc.). However, any such interna- tional system must be (1) comprehensive, (2) have long-term maintenance/up-dating mechanisms, (3) incorporate consensus into taxonomic designations, (4) be of sufficient quality to supersede the use of book-based floras, (5) conservative regarding the verification and incorporation of new taxa from the primary literature, and (6) able to use type specimens where possible. Even if these con- ditions are met, some problems specific to web sites remain. The e-data are free of peer review (although authentication through consensus management could minimise this criticism), they require permanent specialist staff, and some institutional internet-servers can impart loss of trans- mitted image quality. Where on-line databases provide environmental data for particular taxa, revi- sions involving splitting taxa could invalidate any ecological optima. However, updating syn- onymy dictionaries could help maintain database integrity. DIATOM DIVERSITY Diatom diversity is an attempt to describe all the variability present in the group and as such includes an array of factors such as life forms, genetics and morphology. For evaluating time-space diversity aspects in diatom distributional studies, morphotaxon concepts are probably the most use- ful in the first instance. Despite difficulties in diatom taxonomic harmonisation (above), there are two main ways to consider the diversity of diatom morphotaxa, according to: 1. Spatial scales — these vary from local, through regional to global scales and are modified primarily by water quality, substrata and by biogeographic factors. 2. Temporal scales — these vary from diurnal and seasonal changes through inter-annual and millennial trends to evolution and species replacements over geological time (stratigraphic facies). The role of temporal and spatial scales on the occurrence and distribution of organisms has a large general literature (e.g., Rosenzweig 1995). Aquatic organisms received rather less attention although environmental change and fish speciation is well researched (Ruber et al. 1998) and Brooks (1950) made a careful global study of endemicity in lakes. Both studies demonstrate the importance of morphotaxonomy in comparing systems but the former showed some morphology groups diverged from mDNA phylogeny. Nevertheless, morphotaxonomy remains the initial tool to investigate diversity. Lake Baikal is remarkable for both speciation and endemicity (Kozhova and Izmesteva 1998), yet the benthic diatom diversity of Baikal requires thorough re-assessment (see Mann 1999, Flower et al. 2004). This lake can be used as an example for diatom diversity stud- ies from several aspects. LAKE BAIKAL AND TAXON DISTRIBUTIONS.— Lake Baikal has a large endemic component in its diatom flora. Most of the planktonic diatoms are endemic as are about half of the benthic taxa (according to Skwortzov 1937). This latter estimate will almost certainly need revising upwards (Mann 1999). Initial work on common shallow-water taxa around the lake (Flower et al. 2004) has not demonstrated any major distributional relationships associated with the three main sub-basins. Hence, these diatoms provide no evidence that the Lake Baikal sub-basins were isolated in the past (cf. fish populations in Lake Tanganyika [Ruber et al. 1998]). However, distributions of the large 116 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 conspicuous endemic taxa (mainly in the deep littoral) remain largely undescribed (Flower et al. 2004). The shallow water taxa appeared to be mainly cosmopolitan forms, but careful examination indicated that some of these too might have restricted distributions. Common Synedra vaucheriae (sensu Skvortzow) occurs in two characteristic forms in Lake Baikal. Similarly, in Lake Baikal, Hannaea arcus (sensu Skvortzow) seems to be a new species although it is not restricted to the lake (Bixby 2002). The more conspicuous deep water benthic endemic taxa in Baikal, e.g., Didymosphenia den- tata Dorog. are thought to be entirely restricted to the lake (Kociolek et al. 2000). A consideration of the lineage of gomphonemoid taxa suggests that this species evolved in Baikal (ibid.) but veri- fication by fossil records is needed. The fossil record can offer definitive evidence about past taxon distributions and immigrations (cf. Bradbury 1986). Being able to draw conclusions about the ori- gin of taxa has important bearings on interpreting modern distributions. This is why the Baikal Deep Drilling Project cores and the ~ 5 million years of sediment records are so valuable with many new species and even genera being described (Khursevich et al.2001). Although this palaeo- work so far indicates in situ evolution of endemic planktonic taxa (ibid.), benthic diatoms often comprise up to ~10% of the sediment record and their stratigraphic records need evaluating. However, the Baikal record is so extensive that it may be difficult to find comparable sedimentary diatom facies elsewhere for evidencing ideas about speciation. Fossil material from the Vitim Plateau (NE of Lake Baikal) is lodged at the Baikal Museum (see Flower et al. 2004) and casual examination has indicated the presence of several current Baikal endemic taxa (Aulacoseira baicalensis and Cyclotella baicalensis), indicating wider Siberian distributions of these taxa in Pleistocene lakes. The situation regarding fossil benthic taxa is largely unknown. LAKE BAIKAL AND DIATOM REGIONALITY.— Some common benthic diatom taxa in Baikal that appear to be rather different from the normal concept of the species form (e.g., Synedra vaucheri- ae) are probably restricted to the Baikal region. Cymbella stuxbergii and Eunotia clevei also occur in Baikal but have wider though still restricted distributions (Williams et al. 1999; Reid and Williams 2001). Other taxa were recognised by Skvortzow (Skvortzow and Meyer 1928; Skvortzow 1937) as having distributions restricted to Asia. However, many of the satellite lakes around Lake Baikal possess entirely different floras to that in Baikal, but there are important dif- ferences in water quality. Also, all the upland satellite lakes have only existed since the last glacia- tion. In Lake Bolshoie in the Khamar Daban Mountains above Lake Baikal, the surface sediment is dominated by the plankton diatom Pliocaenicus costatus v. sibirica (see Flower et al. 1998). This diatom genus is mainly known as a fossil, but the taxon is extant in a few sites in Siberia. With only a very few exceptions, the benthic diatom flora of Lake Bolshoie is otherwise very similar to olig- otrophic upland lakes in the UK. The unusual distribution of this planktonic taxon seems to be linked more with a limited dispersal and biogeography rather than with special water quality pref- erences. Regional issues regarding the relationships of Lake Baikal endemic diatoms with closely relat- ed taxa elsewhere can be examined using systematic analysis (cf. Williams 1985, 1996). This offers one way of placing Baikal’s endemic flora in an evolutionary context by defining lineages. Quantitative comparisons using morphological characters possessed by both the endemics and closely related taxa offers an alternative to using the fossil record. This approach has been applied to Stephanodiscus in the USA (Theriot 1992) and to benthic Tetracyclus spp. (Williams 1996), but its application to Lake Baikal taxa is perhaps premature. Here the first priority is to provide a thor- ough inventory of the benthic taxa within the lake, to revise previously described taxa, and to pro- vide an iconograph for the benthic taxa. LAKE BAIKAL AND MEASURING DIVERSITY.— Assessing diatom diversity in a large lake such ——— FLOWER: TRENDS IN DIATOM RESEARCH NIU as Baikal presents practical as well as theoretical difficulties. Size can be tackled by systematic sampling (cf. Flower et al. 2004), but measuring species diversity offers a variety of challenges. Biodiversity estimation has a considerable literature and an array of methods (Hawksworth 1995; Heywood 1995). Traditionally, a range of “biodiversity indices’ have been applied to multi-species sample populations and much information on species:area distributions has been generated (Rosenzweig 1995). Interestingly, however, diatom taxa do not seem to show the expected number and area relationship (Allen et al.1999) probably for reasons of water quality (Flower 2001). Yet assessing diatom diversity is important and may benefit from an approach other than application of the traditional indices. Refinements for assessing diversity by taxic measures, such as the ratio of genera to species (see Hawksworth 1995), and by phylogenetic methods to reconcile species num- bers with genealogical divergence (Williams 1996), are interesting concepts. However, like molec- ular genetics, using these techniques to address general issues of diatom occurrences and abun- dances in Lake Baikal and many other poorly described areas must inter alia await an adequate base in morphotaxonomy. No single diversity measure can capture all aspects of ‘diversity’, and at sites such as Baikal, measures that ignore the significance of endemicity are incomplete. However, getting taxonomy to the level where Baikal’s endemic taxa can be validated in routine diatom analysis remains a major challenge (Flower et al. 2004). Identifying any with in hot spot regions of Baikal for endemic taxa is not yet feasible. Estimates of species richness are, however, relatively easy to make, if consis- tency in morphotaxon identifications is maintained rigorously (ibid.). Partitioning taxa into rare and non-rare categories is an additional measure of ecological quality for a particular site or habi- tat (see Gaston 1994). Most diatom samples can be characterised by a few common species and a limited number of rare taxa (van Dam 1982), and for Baikal the rare taxon category often captures some of the conspicuous endemics. For shallow water epilithon samples, taxon abundance distri- butions (examples in Fig. 2) were plotted for 51 stations around Lake Baikal (see Flower et al. 2004). There are considerable variations between abundances of common and less common taxa in each sample. Rather than make assumptions about these distributions (cf. Rosenzweig 1995), it is instructive to consider the rare taxa alone. Rare taxa can be defined in various ways (Gaston 1994): A Station 3.3 O Station 4.1 ®@ Station 8.1 © Station 18.1 Abundance % (log scale) Number of taxa FiGURE 2. Relative abundance rank curves for diatom taxa in several shallow water epilithon samples selected from col- lections made from 51 sampling stations around Lake Baikal in 1997/98 (see Flower et al. 2004). 118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 they occur at very few sites (either locally or internationally); they are always found at low abun- dances; or they may be perceived as rare because of identification problems. For these Baikal sam- ples, the ‘rare’ taxa are those with frequencies of <1% in a fixed count of 300 valves (see Flower et al. 2004). The rare taxon data can be used to generate a simple index that places an emphasis on the pro- portion of rare taxa in samples. The relationship between diatom taxa occurrences and abundances is usually log-normal (van Dam 1982), and it is implicit in the log series of species-abundance dis- tributions that the total number of species in a sample is proportional to the number of uncommon or rare species (e.g., Pielou 1974). Nevertheless, the precise relationship between abundant and rare taxa may be of potential value in assessing the relative ecological value of a site, sub-sites or a group of sites. An index of diatom rarity is indicated below. Where n is the number of sub-samples and | indicates the population or region of interest, V = the total number of valves counted and z is the number of taxa present at frequency abundances of 1% or less. A simple diatom rarity index: n EV, i=l Diatom rarity index = 1- ( ) For sample | n n LV; + Ze: i= Jil This index should be <1 TABLE |. Diatom rarity index values (see text) calculated from diatom assuming rare taxa are taxon abundances in shallow water epilithon samples collected from always present and the term around Lake Baikal (1997/98) and summed to represent each shore zone ‘rare’ is subject to the count 1” each of Baikal’s three main basins (WSB=West shore South Basin, WMB = West shore Middle Basin, WNB= West shore North Basin, ENB = East shore North Basin, EMB = East shore Middle Basin and ESB = East shore South Basin). constraints imposed during sample analysis. There is, of course, a substantial number of taxa that are so rare that — gnoye Zone WSB WMB WNB ENB EMB _ ESB they wall not be captured by 025 021 036 046 Os Mose this modest count total. In fact, cumulative counting of 02? Oe Oe OF 0.08 0:60 one Baikal sample (34.1) 0.52 0.18 0.25 0.29 0.66 0.55 indicated that only above a 0.14 0.46 0.52 0.49 0.78 0.43 count total of 650 valves did Station scores ¢ 4g 0.49 0.49 0.36 0.60 (0.29 the count/taxon curve begin to approach the asymptote. 0.55 0.21 0.46 0.40 0.66 0.43 Nevertheless, by carefully Oe 082 oe Cee fixing the count total, the 0.32 0.46 0.32 index can be used to make 0.86 0.43 within lake comparisons dylan scores 039 031 047 2043 064. 048 amongst communities (Fig. 3) and amongst regions (Table 1). Increasing the standard count size will produce more taxa per sub-sample, but when a large number of samples required for comparison, the key factor is to keep the total count size constant. LAKE BAIKAL AND DIATOM BIOGEOGRAPHY.— The large number of endemic taxa in Lake Baikal is also of interest to ideas about ubiquity and the global distributions of micro-organisms FLOWER: TRENDS IN DIATOM RESEARCH IK) (cf. Finlay and Clarke 1999). Round (1981) indicated that freshwater algal taxa were more be widely distributed geographical- i ye ia | ly than marine taxa but that some | ee 19 freshwater diatoms (Asterionella, 1B B | Stephano-discus) were restricted I. 3 by water quality. Round (1981) I 2. also noted that distinctive distrib- richer z | utional patterns existed but that i studies were very incomplete. i > 2 | Diatom endemicity in Baikal I Sora offers a major challenge to ideas 1 peat 24, about ubiquity. The diatom flora a 2s | of Baikal is demonstrably able to a yx resist invasions and displacement 9 | by cosmopolitan species (which he ¥ ‘| generally predominate in small co, oe .| lakes around Baikal). The recent 1 if o{ fossil record shows continuous a 2 ean ay domination by endemic taxa I a7 ese | (Flower et al. 1996). Elsewhere | \ Rarity scale | yy 3 é Z hy 3 and against conservative ideas ar coy Se bd ; about diatom distributions, | wae 0.5 ; : pie me, les »— 6/46 1 34, regional diatom endemicity is Miyg+43 44 4s ery | 0 5 : . 41 37 x well recognised, in Australasia at @ 40 39 38% a | | both genus and species level A S0km (Tyler 1996) and on some ocean- mee Bus FiGureE 3. Diatom rarity index (see text) values as histograms calculated ic islands (Moser et al.1999). from shallow water epilithon samples from Lake Baikal (collected in 1997/98) There appear to be several from around the lake (see Flower et al. 2004). types of endemism in Baikal aris- ing from recent in situ speciation or preservation of relict taxa (neoendemics and palaeoendemics sensu Cronk 1992). Forms of Navicula lacus-baicali (Mann 1999) are probably recent whereas some species of Tetracyclus (Williams et al. 2002) have a substantial geological record. In the fos- sil record, Aulacoseira baicalensis seems to post date many endemic Stephanodiscus and Cyclotella taxa (Khursevich et al. 2001). Eunotia clevei and Pliocaenicus costatus are examples of taxa that are not endemic to Lake Baikal and its environs but nevertheless display distributions that are more restricted today than in the recent geological past. In some continental areas elsewhere, the impact of past glaciations were probably of paramount importance in obliterating centres of endemism and diminishing former wider distributions of ancient taxa. It is notable that much of southern Siberia, like most of Australasia, was not directly affected by major Pleistocene ice sheets. Those lakes that have persisted through major environmental changes are more likely to have retained ancient taxa (see Brooks 1950). Conversely, palaeoendemics seem absent in the post- glacial Great Lakes of North America. The contemporary endemic taxa of Lake Baikal seems large- ly restricted to the Baikal region and have been unable, possibly through poor dispersal mecha- nisms or lack of ecological niches, to expand elsewhere. Making floristic comparisons amongst different regions to delimit the distributions of particu- a i 120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES lar taxa can demonstrate clear biogeo- graphic patterns for some taxa (Williams 1995; Kociolek and Spaulding 2000). However, making quantitative compar- isons using communities is more difficult, not least because of differing taxonomic concepts and methodologies. Neverthe- less, this approach is needed to establish precise taxon based floristic distances between communities or assemblages (see Flower et al. 1997) and to explore the pro- portional mix of cosmopolitan and endem- ic or otherwise restricted taxa at a particu- lar site or location. For an extreme and preliminary quantitative comparison of two regional floras, Correspondence Analysis (see Jongman et al. 1987) of combined data was undertaken. These data comprised, (1) 51 samples of shallow water diatom epilithon samples from around Baikal and, (2) 28 samples of epilithon and surface sediment diatoms from the Falkland Islands (R. Flower, un- published) on the opposite side of the globe (Fig. 4). Sampled substrata were not constant but other sampling and counting variables and taxonomic concepts were consistent because both data sets were enumerated by the same analyst. The most noticeable feature of this initial and crude analysis is that the amount of floristic variation in all the Baikal samples is very much less than that displayed by the 28 individual water body samples on the Falkland Islands. Also, there were only seven taxa common to both data sets and included such cosmopolitan species as Cocconeis placentula, Navicula crypto- cephala and Staurosira construens. The major floristic differences between Baikal and the Falklands samples clearly do not offer an unequivocal test of a biogeographic hypothesis for diatom distributions. Diatom taxa are particularly good indicators of water quality and are used by palaeolimnologists essentially for this reason. We would, therefore, not +5.5 : NAQOBA* CY Q04A = NAOO7A* OPO001A* : Volume 56, No. 11 FRKO1A ACO16A FILR YHOO1A + x +: FROO2C* Seateas a : CM118A ioe _S17 | NAQO3A x FR9965 $26 lfaoo CM031A -4.0 -1.0 Fy ‘ #8,55003A FILG FILQ ISE P1005a FISC EU9891 EU1194 Bi Pi018B +2.5 FiGuRE 4. Correspondence analysis ordination of a combined data set of diatom taxa (shown as selected alpha-numeric taxon codes and crosses proportional to frequency abundances) and sites (shown as squares and coded with a bold S or FI prefix to indicate Siberian Lake Baikal or Falkland Islands sites). Note, floristic vari- ation is relatively small for the Baikal sites and that few taxa are common to both data sets (see text). FLOWER: TRENDS IN DIATOM RESEARCH ZI expect ubiquity in species distributions: acidiphilous or alkaliphilous taxa do not commonly occur in calcareous or in siliceous geological regions respectively. Falkland Islands’ inland waters were mainly acid. Hence ecological factors could explain most of the taxon differences in the two regions tested and very few, if any, of the Falklands Island diatoms are endemic (as on other glaciated islands, Foged 1964). On the other hand, the extremely basic rocks, unusual water qual- ity and geological time has produced many new species in New Caledonia (Moser et al.1999). From these few observations it is likely that the incidence of diatom endemicity is affected by a variety of factors including, (1) longevity of habitat, (2) extreme environments or environmental instability (palaeoclimate or geological change), (3) proximity to centres of evolution and, perhaps, (4) the nature of the community founder species. Evidence from numerical comparisons of inter-continental floras is scarce, but higher plant diversity (species number in disjunct genera) is higher in eastern Asia than in North America and seems mainly caused by physiographical heterogeneity in conjunction with climate change (Qian and Ricklefs 2000). Explanations of the spatial differences in diatom distributions must also involve climate, but heterogeneous geochemical conditions are probably more important locally. Causes of large scale differences in diatom distributions can be polarized into two main stand- points. One is based primarily on vicarience biogeography and the existence of regional floras aris- ing from evolutionary events. The other is based on an assumption that taxa have global distribu- tions that are punctuated by ecological conditions. Distributional patterns in the global diatom flora — at least for inland waters — seems to be the result of various combinations of these influences as modified by local ecology and sympatric events. However, a fuller picture of global patterns and trends in diatom diversity cannot be made until consistent floristic comparisons and time and space evaluations are undertaken for more biogeographic regions. SOME CONCLUSIONS Ecology, evolution, biogeography, systematics and palaeontology have drifted apart during the 20" century (Ricklefs and Schluter 1993) and a similar trend has occurred within diatom research. Recent major developments in the latter regarding species concepts, speciation, life cycles and molecular biology have all made important contributions but, for the foreseeable future, morpho- species concepts will dominate environmental studies where living and fossil taxa are used. Where concepts about diatom taxonomy and taxon designations have changed markedly, some ambigui- ties have arisen that give current cause for concern. Reconciling ever-finer resolution of taxa com- bined with nomenclatural revisions threaten to out-strip the capabilities of end-users (the ecolo- gists, palaeoecologists). Whilst debate is necessary, the current lack of coherence in diatom taxon- omy is now hindering some areas of applied diatom research, especially where light microscopy is routine. Introducing more consensus in diatom taxonomy at a level that is accessible to end users is advocated. Developing a more unified approach to accessing diatom type material and nomen- clature, regional floras and ecological information through the Internet medium is one promising way of improving taxon designations and identification. Diatom ecologists generally use the taxonomy available in published floras to identify taxa so that relationships between diatoms and measurable environmental variables can be discriminated. Diatom species distributions are not however always a predicable response to water quality and inter alia biogeographical factors can play an important role in the provenance of some taxa. The interplay of biogeographical and water quality factors as an influence on discontinuous distribu- tions of diatom taxa requires the attention of both ecologists and systematic taxonomists. Any ‘mar- riage of necessity’ between ecologists and taxonomists (sensu Kociolek and Stoermer 2001) that 1D PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 would help tackle the central issue of explaining diatom species distribution patterns on time-space scales that vary from ancient to modern (e.g., Khursevich et al. 2001) and local to continental (e.g., Fig. 4) is desirable. However, disentangling the causes of these patterns will require consideration of speciation processes, dispersal mechanisms, ecological tolerances, biological interactions and of environmental change. The existing extensive and essentially morphotaxonomic diatom reference system will doubtless endure as the foundation by which research proceeds and unifying and extending this database must facilitate progress. Despite some of the current difficulties in diatom taxonomy and its relationship with ecology, many endemic or regionally restricted diatom taxa are well defined and offer compelling evidence for the existence of biogeographic factors. They are an important aspect of global aquatic diversi- ty assessment and maintaining global biodiversity has become paramount as anthropogenic changes proceed at unprecedented rates. Sites such as ancient Lake Baikal are hot spots for aquat- ic diversity and the planktonic and benthic diatom communities are largely endemic or otherwise restricted. Diversity can be measured in a variety of ways and morphotaxa concepts are usually the primary tool to investigate species time-space changes in the environment. However, where taxo- nomic knowledge is incomplete, concepts about species richness and rare taxa can offer a pragmat- ic ways of appraising the composition of diatom communities. Such measures cannot substitute for species level inventories or assess the significance of genealogical relationships within communi- ties. Nevertheless, an optimal combination of these measures should help floristic comparisons and define community characteristics. Irrespective of taxonomic discrepancies, diatoms are increasing- ly important for setting baselines and references conditions and evidencing environmental change through multi-scale studies of their distributions and relationships. These are all essential parts of global biodiversity assessment in aquatic ecosystems. ACKNOWLEDGMENTS I wish to thank Patrick Kociolek and Andrzej Witkowski for organizing the Workshop on Ecology and Taxonomy (24-29 Oct. 2002, Poland) that enabled this paper to be presented. Also David Williams for discussions and contributions concerning some of the less uninteresting aspects of the manuscript. Galina Pomaskina (Limnological Institute, Irkutsk) and others all kindly helped assemble data from Lake Baikal. I am grateful to the referees and to Nina Strelnikova and mem- bers of the ECRC (UCL), especially Rick Battarbee and Viv Jones, for advice on referencing diver- sification in diatom research. John Birks, Gavin Simpson and Helen Bennion also contributed use- ful comments and art work was by Elanor McBane. This study was made possible by funding through the Darwin Initiative for the Survival of Species (DEFRA, UK) and the Foreign and Commonwealth Office (HMG, UK) REFERENCES ALBERCH, P. 1993. Museums, collections and biodiversity inventories. TREE 8: 372-375. ALLEN, A.P., T.R. WHITTIER, P.R. KAUFMANN, D.R. LARSEN, R.J. O° CONNER, R.M. HUGHES, R.S. STEMBERGER, S.S. Dixit, R.O. BRINKHURST, A.T. HERLIHY, AND S.G. PAULSEN. 1999. Concordance of taxonomic richness patterns across multiple assemblages in lakes of the northeastern United States. Canadian Journal of Fisheries and Aquatic Sciences 56:739-747. BATTARBEE, R.W. (1978) Observations on the recent history of Lough Neagh and its drainage basin. Philosophical Transactions of the Royal Society of London 281:303-345 BATTARBEE, R.W., V.J. JONES, R.J. FLOWER, N.G. CAMERON, AND H. BENNION. 2001. Diatoms. in Tracking Environmental Change Using Lake Sediments Pages 155-202 in J P. Smol, H.J.B. Birks, and W.M. Last, eds., Tracking Environmental Change Using Lake Sediments. Kluwer Publishers, Dordrecht, Netherlands. FLOWER: TRENDS IN DIATOM RESEARCH 13 BATTARBEE, R.W., S. JUGGINS, F. GASSE, N.J. ANDERSON, H. BENNION, N.G. CAMERON, D.B. RYVES, C. PAILLES, F. CHALIE, AND N. TELFORD. 2001a. European Diatom Database (EDDI). An Information System for Palaeoenvironmental Reconstruction. ECRC Research Report Number 81. University College London. 210 pp. BARKER, P.A., F.A. STREET-PERROTT, M.J. LENG, P.B. GREENWOOD, D.L. SwAIn, R.A. PERROTT, R.J. TELFORD, AND K.J. FICKEN. 2001. A 14000 year oxygen isotope record from diatom silica in two alpine lakes on Mt. Kenya. Science 292:2307-2310. BarRON, J.A., AND J.G. BALDAUF. 1995. Cenozoic maine diatom biostratigraphy and applications to paleocli- matology and paleocenography. Pages 107-118 in C.D. Blome, P.A. Whalen, and K.M. Reed, eds., Siliceous Microfossils. Paleontological Society Short Courses in Paleontolgy, 8. The Paleontology Society. UTC Publications, Knoxville, Tennessee, USA. BDP-98 MemBers. 2001. The new BDP-98 600m drill core from Lake Baikal: a key late Cenozoic sedimen- tary section in continental Asia. Quaternary International 80-8 1:19—36. BENNION, H. 1994. A diatom-phosphorus transfer function for shallow, eutrophic ponds in southeast England. Hydrobiologia 275/276:391-410. BHATTACHARYA, D., L. MEDLIN, P.O. WAINRIGHT, E.V. ARIZTIA, C. BIBEAU, S.K. STICKEL, AND M.L. SOGIN. 1992. Algae containing Chlorophylls a + c are paraphyletic: molecular evolutionary analysis of the Chromophyta. Evolution 46 (6):1801—1817. Birks, H.J.B., J.M. LIne, S. JuGGINS, A.C. STEVENSON, AND C.J.F. TER BRAAK. 1990. Diatoms and pH recon- struction. Philosophical Transactions of the Royal Society of London , Ser. B, 327:263-278. Brxpy, R.J. 2002. Endemism within the diatom genus Hannaea Patrick. Page 12 in M. Poulin, ed., Abstracts of the 17 International Diatom Symposium. Canadian Museum of Nature, Ottawa, Canada. BRADBURY, J.R. 1986. Continental diatom biostratigraphy and palaeolimnology. Pages 667—886 in M. Ricard, ed., Proceedings of the 8 International Diatom Symposium on Living and Fossil Diatoms. Koeltz, Koenigstein, Germany. BRADBURY, J.R. 1991. The late Cenozoic diatom stratigraphy and paleoclimatology of Tule Lake, Siskiyou Co. California. Journal of Paleolimnology 6:205—255. Brooks, J.L. 1950. Speciation in Ancient Lakes. Quarterly Review of Biology 25:30—60. BURKHARDT, F., AND S. SMITH, EDS. 1987. Correspondence of Charles Darwin. Vol. 3, 1844-46. Cambridge University Press, Cambridge, UK. CALVERT, R. 1929. Diatomaceous Earth. American Chemical Society Monograph 52: Chemical catalogue. American Chemical Society, New York, USA. 250 pp. CAMBURN, K.E., AND D.F. CHARLES. 2000. Diatoms of Low Alkalinity Lakes in the Northeastern United States. Special Publication 18. Scientific Publications, Academy of Natural Sciences of Philadelphia, USA. 152 pp. CLEVE, P.T. 1878. Diatoms from the West Indies Archipelago. Bihang till Kongliga Svenska-Vetenskaps- Academiens Handlingar 5:\—22. CLEVE, P.T. 1894/5. Synopsis of Naviculoid Diatoms. Kongliga Svenska-Vetenskaps-Academiens Handlingar 26:1—194, 27:1-219. Coste, M., C. Bosca, AND A. DautTA. 1991. Use of algae for monitoring rivers in France. Pages 75-88 in B. Whitton, E. Rott, and G Friedrich, eds., Use of Algae for Monitoring Rivers. Institiit fiir Botanik, Universitat Innsbruck, Austria. CRONK, Q.C.B. 1992. Relict floras of Atlantic islands-patterns assessed. Biological Journal of the Linnean Society, London 46:91—103 Darwin, C. 1839. Voyage of The Beagle. Henry Coburn, London, UK. 244 pp. DeevEY, E.S. 1942. Studies on Connecticut lake sediments. III The biostratigraphy of Linsley Pond. American Journal of Science 240:233-64, 313-38. DorocostTaisky, V. 1904. Materiaux por servir a l’algologie du lac Baikal et de son bassin. Bulletin de la Société impériale des naturalistes de Moscou 2:229-265. Durr, K., B. ZEEB, AND J.P. SMoL. 1995. Atlas of Chrysophycean Cysts. Kluwer Academic Publishers, Dordrecht, Netherlands. 179 pp. EpGar, S.M., AND E.C. Tuertor. 2000. A total evidence approach to inferring the phylogenetic relationships 124 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 within Aulacoseira. Page 33 in M. Poulin, ed., Abstracts of the 17 International Diatom Symposium. Canadian Museum of Nature, Ottawa, Canada. EHRENBERG, C.G. 1854. Zur Mikrogeologie. Das Erde und Felsen schaffende Wirken des unsichtbar kleinen selbstaendigen lebens auf der Erde. L. Voss, Leipzig, Germany. 374 pp. ERESHEFSKY, M. 2001. The Poverty of the Linnaean Hierarchy: A Philosophical Study of Biological Taxonomy. Cambridge University Press, Cambridge, UK. 328 pp. FINLAY, B.J., AND K.J. CLARKE. 1999. Ubiquitous dispersal of microbial species. Nature 400:828 FINNEY B.J., I. GREGORY-EVANS, J. SWEETMAN, M.D. DOUGLAS, AND J.P. SMoL. 2000. Impacts of climate change on Pacific salmon abundances over the past 300 years. Science 290:795-799. FLOWER, R.J. 1989. A new variety of Tabellaria binalis from several acid lakes in Galloway, Scotland. Diatom Research 4:21-23. FLOWER, R.J. 2001. Change, stress, sustainability and aquatic ecosystem resilience in North African wetland lakes during the 20" century: an introduction to integrated biodiversity studies within the CASSARINA Project. Aquatic Ecology 35:261—280 FLOWER, R.J., AND R.W. BATTARBEE. 1985. The morphology and biostratigraphy of Tabellaria quadriseptata in acid waters and lake sediments in the Galloway, southwest Scotland. British Phycological Journal 20:69-79. FLOWER, R.J., S. JUGGINS, AND R.W. BATTARBEE. 1997. Matching diatom assemblages in lake sediments cores and modern surface sediment samples: the implications for lake conservation and restoration with special reference to acidified systems. Hydrobiologia 344:27-40. FLoweR, R.J., A. MACKAY, J. BOYLE, AND J. DEARING. 1995. Recent environmental change in Lake Baikal. The Holocene 5:323—327 FLowER, R.J., S.P. OZORNINA, A. KUZMINA, AND F.E. RouND. 1998. Pliocaenicus taxa in modern and fossil material mainly from Eastern Russia. Diatom Research 13:39-62. FLOWER, R.J., G POMASKINA, E. RADIONOVA, AND D.M. WILLIAMS. 2004. Benthic diatoms in Lake Baikal: assessing and recording local and meso-scale patterns in diversity. Pages 69-92 in M. Poulin, ed., The 17*h International Diatom Symposium. Canadian Museum of Nature, Ottawa, Canada. Biopress Ltd., Bristol, UK. FoGeb, N. 1964. Freshwater diatoms from Spitsbergen. Tromso Museums Skrifter 11:1—204. Forey, P.L. 2002. A knotty problem of nomenclature. Nature 415:839. Foster, D., A. WooD, AND M. GRIFFITHS. 2001. The EC Water Framework Directive and its implications for the Environment Agency. Freshwater Forum 16:4—28. FOURTANIER, E., AND J.P. KOCIOLEK. 1999. Catalogue of the diatom genera. Diatom Research 14:1—190. FRENGUELLI, J. 1923. Diatomeas de Tierra del Fuego. Anales de la Sociedad Cientifica Argentina 96:225—263 Fritz, S.C., S. JuGGINs, S., R.W. BATTARBEE, AND D.R. ENGSTROM. 1991. Reconstruction of past changes in salinity and climate using a diatom-based transfer function. Nature 352:706—708. GasTON, K.J. 1994. Rarity. Chapman and Hall, London, UK. 244 pp. GEITLER, L. 1927. Die Reduktionsteilung und Copulation von Cymbella lanceolata. Archiv fiir Protistenkunde 58:465—S07. GELL, P.A. 1998. Diatoms, calibration sets and taxonomic quality control. Pages 16—20 in P. Newal, ed., The First Australian Diatom Workshop. Proceedings. Deakin University School of Ecology and Environment. Technical paper 1. GouLb, S.J. 1989. Wonderful Life The Burgess Shale and the Nature of History. Hutchinson Radius, London, UK. 347 pp. Grecory, W. 1854. Notice of the new forms and varieties of known forms occurring in the diatomaceous earth of Mull. Quarterly Journal of the Microscopical Society 2:90—100. GREUTER, W., J. MCNEILL, D.L. HAWKSORTH, AND E.R. BArRRIE. 2000. International Code of Botanical Nomenclature. Koeltz, Koenigstein, Germany. 253 pp. GUERRERO, J.M., AND R.O. ECHENIQUE. 2001. Cyclostephanos patagonicus sp. nov. a new freshwater diatom from Western Patagonia. Diatom Research 17:141-151. HABIRSHAW, F. 1887. Catalogue of the Diatomaceae with References to the Various Published Descriptions and Figures. R. Hitchcock, New York, USA. 272 pp. FLOWER: TRENDS IN DIATOM RESEARCH 125 HAKANSSON, H. 2002. A compilation and evaluation of species in the genera Stephanodiscus, Cyclostephanos, Cyclotella with a new genus in the family Stephanodiscaceae. Diatom Research 17:1—139 HALDEN, B. 1929. Kvartargeologiska diatomaceestudier belysande den postglaciala transgression a Svenska Vastkusten. Geologiska Foreningens I Stockholm Forhandlinger 51:311—366. HawkswortH, D.L., ED. 1995. Biodiversity Measurement and Estimation. Chapman and Hall, London, UK. 140 pp. Haywoop, V.H., ED. 1995. Global Biodiversity Assessment. Cambridge University Press, Cambridge, UK. 1140 pp. HELMCKE, J.G., AND W. KRIEGER. 1951. Electronenmikroskopische untersuchungen uber den kammerbau der Diatomeenmembran. Berichte der Deutsche. Botanische Gesellschaft 64:27-29. HELMCKE, J.G., AND W. KRIEGER. 1953-77. Diatomeenschalen im electronenmikroskopischen. Bild. Parts 1-10. J. Cramer, Weinheim, Germany. HustTepT, F. 1909. Swesswasser-Diatomeen Deutschlands. Stuttgart: Franckh’sche Verlagshandungen. 7Opp. HustTept, F. 1937-39. Systematische und Oekologische untersuchungen tiber den Diatomeen-Flora von Java, Bali, Sumatra. Archiv fiir Hydrobiologie, Suppls. 15:131-177 and 16:274—394. JEWSON, J.H. 1992. Size reduction, reproductive strategy and the life-cycle of a centric diatom. Philosophical Transactions of the Royal Society of London, Ser. B, 336:191—213 JONGMAN, R.H.G, C.F.J. TER BRAAK, AND O.F.R. VAN TONGEREN. 1987. Data Analysis in Community and Landscape Ecology. Pudoc, Wageningen, Netherlands. 299 pp. JouseE, A.P. 1939. Paleogeografia vodoemovy na osnove diatomovogo analisa.7rudi Verchnevolzskoy expedicii geografo-economichescogo Instituta LGU 4:1—86. (In Russian) Jouse, A.P. 1962. Stratigraficheskie i paleogeograficheskie issledovanija v cevero-sapadnoy chasti Tichogo okeana. Akademiya Nauk SSSR, Institut Okeanologii. [sdateljstvo Akademii Nauk SSSR Moskova 258:1—80. (In Russian) KetLty, M.G. 2002. Role of benthic diatoms in the implementation of the Urban Wastewater Treatment Directive in the River Wear, North-East England. Journal of Applied Phycology 14: 9-18. KHURSEVICH, G.K., E.B. KARABANOV, A.A. PROKOPENKO, D.F. WILLIAMS, M.I. KUZMIN, AND S.A. FEDENYA. 2001. Biostratigraphic significance of new fossil species of the diatom genera Stephanodiscus and Cyclotella from Upper Cenozoic deposits of Lake Baikal, Siberia. Micropalaeontology 47:1—24. KITCHING, I.J., P-L. Forty, C.J. HUMPHRIES, AND D.M. WILLIAMS. 1998. Cladistics. Oxford University Press. Oxford, UK. Kitton, F. 1870. On diatomaceous deposits from Jutland. Journal of the Quekett Microscopical Club 2: 98-99, 205-207. KocIoLek, J.P. 1998. Does each genus of diatoms have at least one unique feature? Diatom Research 13:177-179. Kocio_eK, J.P., R.J. FLOWER, AND G. REID. 2000. Valve structure of Didymosphenia dentata (Bacillariophyta): an endemic diatom species from Lake Baikal. Nova Hedwigia 71:113—120. KOCIOLEK, J.P., AND S.A. SPAULDING. 2000. Freshwater diatom biogeography. Nova Hedwigia 71:223-241. KOcIOLEK, J.P., AND E.F. STOERMER. 2001. Taxonomy and ecology: a marriage of necessity. Diatom Research 16:433-442. KOLBE, R.W. 1927. Zur Oekologie, Morphologie und Systematik der Brackwasser-diatomeen. Pflanzen- forschung 7:\-146. KOLKWITz, R., AND M. Marssen. 1908. Oekologie der pflanzlichen Saprobien. Berichte der Deutsche Botanische Gesellschaft 26: 505-519. KopPeNn, J.D. 1975. A morphological and taxonomical study of Tabellaria (Bacillariophyceae) from the north- central United States. Journal of Phycology 11:235-244. KOZHOVA, O.M., AND L.R. IZMESTEVA, EDS. 1998. Lake Baikal Evolution and Biodiversity. Backhuys, Leiden, Netherlands. 447 pp. KNUDSON, B.M. 1953. The diatom genus Tabellaria I. Taxonomy and morphology of the plankton varieties. Annals of Botany 17:131-155. KRAMMER, K., AND H. LANGE-BERTALOT. 1986. Bacillariophyceae. 1. Teil: Naviculaceae. In: H. Ettl, J. 126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 Gerloff, H. Heynig, and D. Mollenhauer, eds., Suesswasser Flora von Mitteleuropa. Gustav Fischer Verlag, Stuttgart, Germany. 876 pp. KRAMMER, K., AND H. LANGE-BERTALOT. 1991. Bacillariophyceae. 3. Teil: Centrales, Fragilariaceae, Eunotiaceae. Jn: H. Ettl, J. Gerloff, H. Heynig, and D. Mollenhauer, eds., Suesswasser Flora von Mitteleuropa.. Gustav Fischer Verlag, Stuttgart, Germany. 576 pp. LANGE-BERTALOT, H. 1990. Current biosystematic research on diatoms and its implications for the species concept. Limnetica 6:13-22. LANGE-BERTALOT, H., AND R. SIMONSEN. 1978. A taxonomic revision of the Nitzschia Lanceolatae Grunow. European and related extra-European fresh water and related taxa. Bacillaria 1:11-111. Lunp, J.W.G. 2002. Research Reports. Pages 43-46 in Seventieth Annual Report of the Freshwater Biological Station. Freshwater Biological Association, Ambleside, UK. MAcDonaLD, J.D. (1869) On the structure of the diatomaceous frustule and its genetic cycle. Annals and Magazine of Natural History, ser. 4, 3:1-8. MANN, D.G. 1984. Observations on copulation in Navicula pupula and Amphora ovalis in relation to the diatom species. Annals of Botany 54:429-438 MANN, D.G. 1999. The species concept in diatoms. Phycologia 38:437-495 MEDLIN, L.K. 1997. Can molecular techniques help define species limits. Diatom 13:19-23. MILLS, F.W. 1933-35. An Index to the Genera and Species of the Diatomaceae and Their Synonyms. Parts 1-3. Wheldon and Wesley, London, UK. MontEITH, D.T., AND C.D. EvaANs, EDS. 2000. UK Acid Waters Monitoring Network: 10 year Report. Analysis and Interpretation of Results 1988-1998. Department of the Environment, Transport and the Regions. ENSIS Publishing, London, UK. 362 pp. (down-loadable at hitp:/Avww.geog.ucl.ac.uk/ukawmn/) Moser, G., H. LANGE-BERTALOT, AND D. METZELTIN. 1998. Insel der Endemiten Geobotanisches Phanomen Neukaledonien. Bibliotheca Diatomologica 18. J. Cramer, Stuttgart, Germany. 464 pp. Nipkow, F. 1920. Vorlaufige Mitteilung ueber Untersuchungen des Schlammabsatzes im Zuerichsee. Schweizerische Zeitschrift fiir Hydrobiologie 1:100—120. PATRICK, R. 1982. The history of the science of diatoms in the United States of America. Pages 11—20 in D.G. Man, ed., Proceedings of the 7 International Diatom Symposium. Koeltz, Koenigstein, Germany. PATRICK, R., M.H. HOHN, AND J.H. WALLACE. 1954. A new method for determining the pattern of the diatom flora. Notulae Naturae 259:1—12. PATRICK, R., AND C.W. REIMER. 1966. The Diatoms of the United States. The Academy of Natural Sciences of Philadelphia, Monograph 13. 688 pp. PeILou, E.C. 1974. Population and Community. Ecology: Principles and Methods. Gordon and Breach Science, New York, USA. 424 pp. PENNINGTON, W. 1943. Lake sediments: the bottom deposits of the north basin of Windermere with special ref- erence to diatom succession. New Phytologist 43:1—27. Pritzer, E. 1869. Uber den Bau and die Zellteilung der Diatomeen. Botanische Zeitung 27:774-716. PIENITZ, R., J.P. SMOL, AND H.J.B. BirKs. 1995. Assessment of diatoms as quantitative indicators of past cli- mates change in the Yukon and Northwest Territories, Canada. Journal of Paleolimnology 13:21-49. QIAN, H., AND R.E. RICKLEFS. 2000. Large-scale processes and the Asian bias in species diversity of temper- ature plants. Nature 407:180—-182. Reip, G, AND D.M. WILLIAMS. 2001. On Eunotia clevei var. hispida Skvortzow from Lake Baikal. Pages 89-105 in A. Economou-Amilli, ed., Proceedings of the 16" International Diatom Symposium, Athens and Aegean Islands. University of Athens, Greece. RICKLEFS, R.E., AND D. SCHUTER. 1993. Species diversity: regional and historical influences. Pages 350-363 in R.E. Ricklefs and D. Schuter, eds., Species Diversity in Ecological Communities. University of Chicago Press, Chicago, Illinois, USA. ROSENZWEIG, M.L. 1995. Species Diversity in Space and Time. Cambridge University Press, Cambridge, UK. 436 pp. Ross, R., AND P.A. Sims. 1973. Observations on family and generic limits in the Centrales. Nova Hedwigia, 45:97-121. ROUND, F.E. 1981. The Biology of the Algae. Cambridge University Press, Cambridge, UK. 655 pp. FLOWER: TRENDS IN DIATOM RESEARCH WT) RouND, F.E., R.M. CRAWFORD, AND D.G. MANN. 1990. The Diatoms. Biology and Morphology of the Genera. Cambridge University Press, Cambridge, UK. 747 pp. ROUND, F-E., AND N.I. MAIDANA. 2001. Two problematic freshwater araphid taxa re-classified in new genera. Diatom 17:21-28. RUBER, L., E. VERHEYEN, C. STURMBAUER, AND A. MEYER. 1999. Lake level fluctuations and speciation in rock-dwelling ciclid fish in Lake Tanganyika, East Africa. Pages 225—240 in P.R. Grant, ed., Evolution on Islands. Oxford University Press, Oxford, UK. RumricH, U., H. LANGE-BERTALOT, AND M. RumricH. 2000. Diatomeen aus Patagonia. Jconographia Diatomologica 9:1-678. SANCETTA, C. 1979. Oceanography of the North Pacific during the last 18,000 years: evidence from fossil diatoms. Marine Micropaleontology 4:103—123. SCHUETT, F. 1896. Bacillariales (Diatomae). Die naturlichen Pflanzenfamilien. Verlag von W. Engelmann, Leipiz, Germany. SIMONSEN, R. 1979. The diatom system: ideas on phylogeny. Bacillaria 2:9-71. SKABICHEVSKI, A.P. 1936. Neue und interessante Diatomeen aus dem noerdlichen Baikalsee. Botanichevski Journal 21:705-719. SKABICHEVSKI, A.P. 1984. The genus Gomphonema Ag. (Bacillariophyta) in Lake Baikal. Novosti Sistematiki Nizshikh Rasteni 21:36-42 SkKvorTzow, B.V. 1937. Bottom diatoms from Olhon Gate of Baikal Lake, Siberia. Philippine Journal of Science 62:293-377. SKvorTzow, B.V,. AND C.I. MEYER. 1928. A contribution to the diatoms of Lake Baikal. Proceedings of the Sungaree River Biological Station 1:1—55. SMOL, J.R. 2002. Pollution of Lakes and Rivers A Paleoenvironmental Perspective. Arnold Publishers, London, UK. 280 pp. SmitH, W. 1856. A Synopsis of the British Diatomaceae. Vol. 2. J van Voorst, London, UK. 107 pp. STRELNIKOVA, N.I. 1990. Evolution of diatoms during the Cretaceous and Paleogene Periods. Pages 195—204 in H. Simola, ed., Proceedings of the 10 International Diatom Symposium. Koeltz, Koenigstein, Germany. STEVENSON, A.C., ET AL. 1991. The Surface Waters Acidification Project Palaeolimnology Programme: Modern Water Diatom/ Lake Water Chemistry Data-Set. ENSIS Ltd. Publishing, London, UK. 86 pp. STOERMER, E.F. 2001. Diatom taxonomy for palaeolimnologists. Journal of Paleolimnology |. 25: 393-398. STOERMER, E.F., AND J.P. SMOL, EDS. 1999. The Diatoms: Applications for the Environmental Sciences. Cambridge University Press, Cambridge, UK. 469 pp. StoscH, H.A. von. 1958. Entwicklungsgeschichtliche Untersuchungen an zentrischen Diatomeen. HI Die Spermatogenese von Melosira moniliformis Agardh. Archiv fiir Mikrobiologie 31:274—282. THERIOT, E.C. 1992. Clusters, species concepts and morphological evolution of diatoms. Systematic Biology 41:141-157. TyYLer, P.A. 1996. Endemism in freshwater algae. Hydrobiologia 336:19-32. VAN Dam, H. 1982. On the uses of measures of structure and diversity in applied diatom ecology. Nova Hedwigia 73:97-115. VANLANDINGHAM, S.L. 1967-79. Catalogue of the Fossil and Recent Genera and Species of Diatoms and Their Synonyms. Parts 1-8. J. Cramer, Lehre, and J. Cramer, A.R. Ganter Verlag KG, Vaduz, Germany. VOLCANI, B.E. 1978. Role of silicon in diatom metabolism and silicification. Pages 177—204 in G. Bendz and I. Lindqvist, eds., Biochemistry of Silicon and Related Problems. Plenum Press New York, USA. VYVERMAN, W., K. SABBE, D.G. MANN, C. KILROY, R. VYVERMAN, K. VANHOUTTE, AND D. HopGson. 1998. Eunophora gen. nov. (Bacillariophyta) from Tasmania and New Zealand. European Journal of Phycology 33:95-111. Witiiams, D.M. 1985. Morphology, taxonomy and inter-relationships of the ribbed araphid diatoms from the genera Diatoma and Meridion (Diatomaceae: Bacillariophyta). Bibliotheca Diatomologica 8:1—228. Wiiiiams, D.M. 1995. Diatom Biogeography: some preliminary considerations. Pages 311-319 in D. Marino and M. Montresor, eds., Proceedings of the 13 International Diatom Symposium. BioPress Ltd., Bristol, UK. 128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 11 WILLIAMS, D.M. 1996. Fossil species of the genus Tretracyclus (Bacillariophyta, ‘ellipticus’ group): morphol- ogy, interrelationships and the relevance of ontogeny. Philosophical Transactions of the Royal Society of London, Ser. B, 351:1759-1782. WILLIAMS, D.M., G. REID, R.J. FLOWER, AND N.E. VOTYAKOVA. 2002. New fossil diatom species of Tetracyclus (Bacillariophyceae) from the Miocene deposit of Tunka Ridge, Lake Baikal, Siberia, Russia. Diatom Research 17:437-443. WILLIAMS, P.H., C.J. HUMPHRIES, AND P.I. VANE-WRIGHT. 1991. Measuring biodiversity: taxonomic related- ness for conservation priorities. Australian Journal of Systematic Botany 4:665-679. WILLIAMS, R.M., M.B. EDLUND, AND E.F. STOERMER. 1999. Taxonomy and morphology of Cymbella stuxbergii from lakes in the Baikal Rift Zone. Diatom Research 14:381-392. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12, pp. 129-145, 1 fig., 2 pls., Appendix April 29, 2005 Diatom (Bacillariophyta) Flora of Albanian Coastal Wetlands Taxonomy and Ecology: A Review Aleko Miho! and Andrzej Witkowski ' Section of Botany, Department of Biology, Faculty of Natural Sciences, Tirana University, Albania. Email: amiho @icc-al.org; Department of Paleooceanology, Institute of Marine Sciences, University of Szczecin, Waska 13, PI-71-415 Szczecin, Poland. Email: witkowsk@ sus.univ.szczecin.pl The Albanian coast still features interesting and important lagoons and other wet- lands, characterised by high biodiversity. The present paper focuses on their diatom flora: taxonomy and ecology. The cheklist of hitherto identified taxa contains about 430 taxa observed in different coastal habitats, e.g., Butrinti, Karavasta, and Lezha. From the diatom structure, an evaluation of the ecological values and the trophic state of some coastal ecosystems can be made. In some of these (e.g., Butrinti), diatom and peridinophyte blooms, formed by potentially toxic species (e.g., Pseudonitzschia seriata, Prorocentrum micans, and P. minimum), were observed dur- ing the summer. The blooms were probably the result of the limited water exchange and were enhanced by other environmental factors (i.e., temperature, nutrients). Studies on the ecology and taxonomy of brackish water diatoms should always be interrelated with and supported by integrated environmental programs. In the existing programs dealing with coastal wetlands, the ecological aspect is always manifestly important. However, the ecological goals cannot be satisfactorily achieved if the need for a basic taxonomic approach is neglected, which is often the case. Recently, numerous geobotanical and ecological studies of the diatom flora of lagoons from regions with different climatic conditions have been made (e.g., Miho 1994; Danielidis 1980; Witkowski 1991; Bak et al. 2001; Trobajo 2003). Similarly to other Mediter- ranean countries, Albania features a high diversity of species and sensitive habitats, which are rather poorly known and remain exposed to a significant impact of human activity (NEA/AKM 1999). Biological zonation and/or the natural trophic/saprobic status of coastal lagoons can be estab- lished by studying the taxonomy and ecology of the diatom assemblages present (Miho 1994; Miho and Mitrushi 1999; Bak et al. 2001; Silvestre et al. 2002; Trobajo 2003). Such studies provide a substantial aid in exploration of biodiversity and functioning of food webs in coastal areas (Essink, 2003) and prove useful in assuring and strengthening further efforts towards their protection, restoration and sustainable use. Publications on microscopic algae in Albania began with sporadic taxonomic surveys by for- eign algologists, e.g., Forti (1902) in Shkodra Lake, Beck (1904) in Prespa Lake, Protic (1907) in brackish water habitats of Vlora, Krenner (1926) in the Drini River and Uherkovic (1963) in Shkumbini and some other rivers. Albanian studies were initiated in the 1980s. During 1987-1991, Miho (1994, 1996) surveyed the phytoplankton in Butrinti lagoon. These were expanded by research on the relationship between microalgae species composition and the trophic status of the habitats studied (e.g., Ohrid and Prespa lakes, Shkumbini and other rivers, Lezha and Vlora 129 130 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 lagoons, glacial lakes, e.g., Mihol1998a, 1998b; Miho and Mitrushi 1999; Miho and Dedej 1999; Miho and Lange-Bertalot 2001, 2003; Miho, Cake, and Carcani [in press]; Cullaj et al. 2003). Dedej (1994, 1995) provided also some preliminary data on the phytoplankton of the Durresi and Karavasta regions. Recently, Rakaj (2002), Rakaj and Kashta (1999) and Rakaj et al. (2000, 2001) have reviewed the Lake Shkodra phytoplankton. Some other publications, i.e., those by Hustedt (1945), Jurilj (1954) and Vilicic et al. (2002) are important taxonomic contributions, particularly with respect to diatoms. The present paper reviews the diatom flora of the Albanian coastal wetlands, and is an exam- ple of how taxonomy and ecology can be mutually supportive. In addition to the floristic compo- sition, information on the ecological and trophic status of various coastal wetlands is provided. GENERAL BACKGROUND Albania is a small country that extends between a high mountain range and the Adriatic Sea, bordering Montenegro and Kosovo to the north, the Republic of Macedonia to the east and Greece to the south (Fig. 1). The country features large wetland areas surrounded by evergreen hills along the Adriatic coast; in addition, almost two-thirds of the country’s surface area is covered by high mountain ranges (up to 2700 m) separated by valleys. The Albanian coast is about 427 km long, 273 km along the Adriatic Sea and 154 km along the Ionian Sea (Kabo 1990-91). The low accumulative Adriatic coast extends from the Buna/Bojana delta to Vlora Bay (Fig. 1) and is characterised by the presence of fluvial deltas, inlets and lagoons, beaches, coastal sand bars, rocky promontories, submerged caves, sandy dunes and extended inland bays. Estuaries of major rivers intersect the coast: Drini, Buna, Mati, Ishmi, Shkumbini, Semani, and Vjosa that run westwards from the country’s interior (Kabo 1990-91). The rivers are major water conduits not only for Albania, but also for the whole eastern Adriatic Sea. In their eastern sections they are tor- rential and erosive, whereas in the Western Coastal Plain they generally form wide meandering beds. In contrast, the Ionian coast is steep, mountainous and rocky, intersected only by a few small rivers, such as the Bistrica and Pavlla. The coast also features some small wetlands and islets. The region is characteristic in its high diversity of habitats and species. In addition, it supports active aquaculture and fisheries, while urbanisation and industrial activities are limited. Butrinti, a wet- land complex, unique in its biodiversity and aquaculture, is situated in the southern part (Fig. 1). COASTAL WETLANDS AND THEIR ECOLOGICAL VALUES Despite reclamation for agricultural purposes during the past decades, Albania still has about 109 km2 of coastal wetlands or lagoons. Kabo (1990-91) described the main hydrological charac- teristics of Albanian lagoons, and the ecology has been reviewed by Peja et al. (1996). The most important coastal wetlands are Karavasta, Narta, Lezha, and Patoku on the Adriatic, and Butrinti on the Ionian coast (Fig. 1). The lagoons extend along the coastline and are separated from the sea by rather narrow sandy spits, which continuously change in size and shape. They are generally characterised by brackish waters, being connected with the sea through one or more channels. Generally, wetlands display a large number of biotopes with enormous diversity of aquatic flora and fauna. Moreover, they are important especially for wintering of migratory birds (more than 70 species; NEA/AKM 1999). Some of these wetlands, e.g., Narta and Lezha, are partly pro- tected. Regarding richness in habitats, flora and fauna (especially globally threatened water birds, i.e., the Dalmatian pelican) Karavasta and Divjaka National Park area have recently been designat- ed Ramsar sites (Ramsar 1971, convention on wetlands [see <http://vww.ramsar.org/ MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS | 131 profiles_albania.htm>}]). Wetland areas have a very sub- stantial economic potential for tourism, as breeding and refuge habitats for fish, and as valuable S U F.Rof Yugoslavia) Noo Bajram Curr ‘4 f i ® aquaculture sites. According to a Kopliku a Peja et al. (1996), up to 6,000 kg = SS — es of fish has been annually harvest- ~ “Hh ol a j oe 5 ¥ | ed during periods of high produc- Ww pe Vau-Deies se uae tivity. ; Sy Ne 2 ( 3 Karavasta lagoon (surface vue Lezha ig s iS area 43 km?), situated between oan resheni Pest the Semani and Shkumbini deltas, is the largest of Albania. It is connected with the sea through three short channels. Divjaka for- Lalzi est (12 km?) spreads out in the northwest of the lagoon (Kabo, y. : Adriatic ie, 1990-91). It is a typically dune @ Kavaja Eibasani forest, bordered by brackish or S#4 ., Pegini at a z i Bu oe ae a freshwater, where pines (Pinus Karavasta eae E halepensis and P. pinea) grow up aapen © Lushnja { Sages in old dunes, mixed with shrubs, wena eC Pogradeci grasses or reeds near lagoon ‘ ce Kseors ast at a o es we is Oe om —s shores. The lagoon system of ‘ ip Beran ee ES) o Lezha is 10-15 km long and 3 Ae ee pie : e “ Narta cS Balishi AA LEY 7 km wide with a total area of 5 \___ Gorovoda fay about 22 km2, 11.6 km2 consist- Vlora Yn, Youth t = ; id ing of lagoons (Ceka, Merxhani, fae Sees ¥ = 5 Onikum ‘sTepelena x and Kenalla), 2 km? forests Ne pared (Vaina and Kune) and 7 km2 of ia mw wetlands. Their origin is related Greece to deltaic processes of the Drini ionian Ny River. Narta, situated in the southern part of the Vjosa delta, see % arport covers 42 km2 and connects with Al] Butint eee Pp the sea via two channels. All isncon bel tel Saal saa these lagoons are shallow and as enone reranecsmee fluvial in origin. Butrinti (16 Ficure 1: Location of Albanian coastal wetlands. km2) is one of the most interest- ing lagoons of tectonic origin. Due to its relatively great depth (averaging 14 m), its water column is permanently stratified. Its bottom is characterized by anaerobic decomposition. However, during the past few decades, the lagoon has been used intensively in aquaculture of mussels (Mytilus gal- loprovincialis) (up to 2,000 to 4,500 ton gross product/year). The wetlands are, however, very sensitive ecosystems that were under strong impact in the past due to the extensive agricultural reclamation and unsustainable industry. At present, the western lowland supports densely populated industrial centres, intensive agriculture and tourism. 3 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 Therefore, some riverine waters (i.e., of Ishmi, Tirana, Lana, Gjanica, etc.) are heavily loaded with urban and industrial sewage discharged directly to rivers that transport this load to the sea. Moreover, high levels of heavy metals have been found in Vlora and Durresi bays, Mati delta, etc. Petroleum industry in Fieri and Vlora also result in an adverse ecological impact on the Semani and Vjosa deltas and their related lagoons. Coastal dune forests are under pressure of touristic devel- opment. Also, the high rate of erosion caused by excessive woodcutting, overgrazing or firing in realtive shallow water basins, further increases the amount of suspended matter transported to the sea by the rivers (Cullaj et al. 2003; UNEP 2000). DIATOMS: TAXONOMIC APPROACH The survey of recent publications on Albanian waters, including the lakes on its borders, shows the number of diatom species to exceed 1 200. The most interesting area seems to be Lake Ohrid with about 550 species (Miho and Lange-Bertalot 2003), of which more than 100 are endemic or rarely occurring species. In their 1999-2000 study of Albanian freshwater diatom flora, Miho and Lange-Bertalot identified 900 taxa, including some whose distribution is poorly known (Miho and Lange-Bertalot 2001, 2003). Some of these (Aneumastus albanicus, A. rosettae, A. humboltianus, Navicula pseudoppugnata, N. parahasta N. hastatula, and Cymbopleura albanica, C. lura and C. lata var. lura) have recently been described as new to science in Lange-Bertalot (2001) and Krammer (2003), respectively. Altogether, more than 430 diatom taxa were identified in the brackish water coastal wetlands, either published or presented here for the first time. Of this group, 365 taxa are pennate and 65 are centrics. More than 160 taxa were found in Butrinti and about 115 in Karavasta, representing two ecosystems with the highest number of taxa. The checklist of the species identified is given in table 1. Despite the efforts to avoid synonyms or changed names of longer established taxa, some faxa might be recorded here twice under different names and some identifications may not be correct. Selected taxa (13 taxa in 25 figures), either the most abundant ones or those taxonomically inter- esting, are illustrated in Plates I and II. The data from this compilation represent the first step to shed light on the diatom flora of this Mediterranean area, hitherto unknown. About 240 taxa have already been published, whereas about 110 were not recorded before in this area; therefore, about 80 taxa have not been observed in Albanian samples examined previously. Vilicic et al. (2002) have already published a checklist of phytoplankton taxa in the Eastern Adriatic Sea, referring to the Croatian coast. In all, 888 taxa were reported, of which 518 were diatoms (330 pennates and 174 centrics). The data refer to naturally eutrophic areas (bays, estuar- ies), harbours, etc. However, only 118 taxa were also included in the Albanian checklist reported here. Probably, the difference in comparison with the Croatian checklist could be due to the fact that many of the Albanian samples were of periphyton. In addition, the various habitats differ in relation to their ecology. Protic (1907) provided the first data on some brackish habitats from the Albanian coast (in Orikumi and Narta: Vlora region). His list included 196 species, 107 of which were diatoms. When dealing with phytoplankton of Butrinti, Miho (1994, 1996) reported 90 species, 60 of which were diatoms. This was the first ecological approach to study an important habitat, at that time intensive- ly used for aquaculture. The diatom genus Nitzschia was represented by the highest number of species; however, centrics, especially Chaetoceros spp. and Cyclotella aff. choctawhatcheeana, were the most abundant. The difficulty in their identification was the principal taxonomic problem. As a matter of fact, more than 40 taxa in Table | are not fully determined. Other abundant species included Pseudonitzschia seriata, Nitzschia capitellata and Thalassionema nitzschioides. They MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS — 133 were accompanied by peridinophytes, e.g., Prorocentrum micans, P. minimum, Scrippsiella sp., Ceratium fusus, C. pulchellum. In the three Lezha lagoons (Ceka, Merxhani, Kenalla), about 90 phytoplankton species were recorded in July 1996 (Miho and Mitrushi 1999). A relatively high diversity was observed in Merxhani where Chaetoceros spp., Amphora holsatica, Nitzschia sigma, Entomoneis paludosa, Cerataulus turgidus, Melosira nummuloides, Grammatophora oceanica, Pleurosigma angulatum and Striatella unipunctata were observed as abundant species. In Ceka, an algal bloom was observed, dominated by Nitzschia reversa, Peridinium spp., Gonyaulax monacan- tha, and Prorocentrum minimum. Kenalla, a relatively deep pond close to the shallow lagoon of Merxhani, supported large populations of the centric diatom Chaetoceros muelleri, associated with abundant filamentous blue-green algae, e.g., Anabaenopsis circularis, Oscillatoria sp., and a small peridinophyte Gymnodinium sp. Occasional surveys were also carried out in other Albanian wetlands, 1.e., Saranda, Karavasta, Narta, Patoku, Viluni and Durresi (Lalzi) (Miho 1998b). In Karavasta, 65 diatom species were identified. In this locality, again, pennates represented by the genera Fragilaria and Cocconeis pre- vailed. In 1992-93, more than 75 diatoms were identified in the Orikumi wetland, where pennates also dominated. In a recent assessment of different habitats of Butrinti, Armura and Saranda Harbour, spanning the period of 1998-2000, over 200 epiphytic diatoms were identified. These included 28 centric species, represented by Cyclotella (6 species), Actinocyclus (5), and Chaetoceros (4), as well as 178 pennates, including, e.g., Nitzschia (25 species), Mastogloia (20), Amphora (14) and Navicula (12). The most abundant taxa included Cyclotella cf. choctawhatch- eeana, Achnanthes brevipes, Amphora cf. tomiakae, A. coffeaeformis, Bacillaria paxillifer, Cocconeis placentula, C. scutellum, Staurosira construens, Mastogloia crucicula, Nitzschia coarc- tata, N. constricta, Pleurosigma formosum and Thalassionema nitzschioides. Certain species, interesting either from an ecological or taxonomical point of view, are present- ed below. Some of them represent the dominant organism, whereas others are poorly known. Cyclotella aff. choctawhatcheeana Prasad (Plate 1: Figs. 1-9) was abundant in the phytoplank- ton of Butrinti (Miho 1994) where it was responsible for an autumn bloom. The species occurred as solitary cells or colonies and was present in the water column throughout the growing season. C. aff. choctawhatcheeana occurred in combination with various Chaetoceros species. In other Albanian lagoons (Orikumi, Karavasta) the distribution was scarce and represented by single cells only. Albanian specimens belong to the Cyclotella caspia group and are closely related to C. choctawhatcheeana Prasad; however, some characters differ from the descriptions given by Prasad (1991), Wendker (1991), and Hakansson et al. (1993). Unlike C. choctawhatcheeana, the Albanian specimens possess 1—3 fultoportulae in the middle; the transapical striae were radiate but finer, about 30-40/10uUm, and measured 5—15 um. Chaetoceros spp. (Plate 1: Figs. 10-14): the species belonging to the genus Chaetoceros occurred abundantly in the phytoplankton of the lagoons, especially in Butrinti, Orikumi, and Merxhani. However, due to their very fine structure, it was not easy to identify the most common species. Nevertheless, as shown by Miho (1994), Ch. wighamii Brightwell seems to be the most abundant, characterised by very small colony-forming cells with very fine setae. Actinocyclus cf. subtilis (Gregory) Ralfs (Plate 2: Fig. 2) was found in brackish and almost marine waters, e.g., Butrinti and Armura Bay (Saranda). Based on their overall morphology, our specimens resembled A. subtilis (Gregory) Ralfs. However, they differed from this in having a gen- erally larger valve diameter (exceeding 68 Um) and denser areolation of the striae (more than 16 puncta/10 um). The taxon appears to have morphologically distinct valves; according to the species description by Witkowski et al. (2000), A. subtilis valve diameter is smaller and the striae possess a coarser areolation (about 12/10 um). 134 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 2000x 7, ~men ar epnenrcsceenir meme NSS pies betes tibonils whe pe SS. ee A Re Pl re gh eaaeiteen = PLATE I. Figs. 1-9: Cyclotella aff. choctawhatcheeana Prasad; Figs. 10-14: Chaetoceros sp. diverse (cf. Ch. wighamii Brightwell); Fig. 7-9, SEM, bar = 3 um); Fig. 15: Pseudonitzschia seriata Cleve; Fig. 16: Navicula cf. besaren- sis Giffen; Fig. 17: Nitzschia reversa W. Smith. (Figs. 1-6, 11-16: 1500x; Figs. 7-9: SEM, bar=3 um; Fig. 10: 1100x; Fig. 17: 2000x.) MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS © 135 a ee. a a see — _—— — en | PLATE IL. Fig. 1: Actinocyclus cf. ochotensis Jouse; Fig. 2: A. cf. subtilis (Gregory) Ralfs; (900x) Fig. 3: Nitzschia granulata Grunow; Fig. 4: N. coarctata Grunow; Fig. 5: N. constricta (Kiitzing) Ralfs; Fig. 6: Campylodiscus cf. simulans Gregory; Fig. 6: Surirella cf. fluminensis Grunow. (Figs. 1, 3-7: 1500x; Fig. 2: 900; Fig. 8: 1200x.) 136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 Campylodiscus cf. simulans Gregory (Plate 2: Fig. 7) was found in Butrinti and Saranda Bay. The morphology seems to be intermediate between that of C. simulans Gregory (Schmidt’s Atlas: Tafel 17, Figs. 12-14) and C. thuretii var. baldjikiana Grunow (Plate 51, Figs. 16-21). Surirella ct. fluminensis Grunow (Plate 2: Fig. 8) is a poorly known species. The morphology appears to be similar to that of the specimen drawn in Schmidt’s Atlas (Plate 5: Fig. 6). Specimens were 44-62 um long and 23—27 um broad with 8 ribs/100 um. The axial area was relatively nar- row. This organisms were frequent in Karavasta and scarce in other Albanian lagoons. Nitzschia reversa W. Smith was found to form blooms in Ceka lagoon (Lezha) (Plate 1: Fig. 17). It is characterised by sigmoid valve shape (35—95 um long and 2.5—3.5 um broad) and a very fine striation (not resolvable in LM). It was smaller and finer than the specimens from the type pop- ulation characterized in Witkowski et al. (2000) or Proshkina-Lavrenko (1955). Pseudonitzschia seriata Cleve (Plate 1: Fig. 15) was very abundant in Butrinti lagoon. During late spring and summer of 1987 it was observed to form a bloom (Miho 1994). DIATOMS: ECOLOGICAL APPROACH Guelorget and Perthuisot (1984) have proposed a biological zonation system for lagoons, based on their ‘confinement’, a hydrological parameter linked mainly to exchange/renewal of the water. It can be based on benthic communities, where each community characterizes one of the six defined zones. Therefore, each zone has its own specific richness, density, biomass, productivity (including phytoplankton), etc. Hence, based on the composition of the microalgae, such zonation could be drawn up, and as a consequence a diagnosis of the biological and economic potential of a lagoon could even be made. Here, some preliminary insights into biological zones that prevail in Albanian lagoons are presented, based on the algal assemblages. Butrinti ranks among typical meromictic water bodies; therefore, optimum conditions for phy- tobenthos are to be found in some borderline locations, dominated by Enteromorpha prolifera (Miho 1994, 1996). The remaining part of the bottom is affected by anaerobic conditions, charac- terized by high rates of organic matter sedimentation and decomposition. Thus, the primary pro- duction is virtually due to the phytoplankton of the upper layers (S—7 m depth), which was domi- nated by one or two species of centric diatoms (often accounting for more than 90% of the primary producers). The seasonal pattern involved a very pronounced peak in spring (up to 44,000 cells/ml in March 1991) and a smaller one in the autumn. A phytoplankton decline, accompanied by a rel- ative increase in peridinophyte abundance, was recorded in the summer. Intensive growth of phytoplankton, mainly by the neritic forms of the centric diatoms men- tioned, shows that the upper layers of Butrinti mainly belong to zone 3. According to Dutrieux and Guelorget (1988) this zone is characterized by limited impact of hydrodynamic conditions; the organic matter content increases in both sediment and water, although the overall living conditions remain good. It represents a habitat favourable for shellfish reproduction, as evidenced by profu- sion of Mytilus galloprovincialis everywhere in Butrinti. However, being permanently stratified with anoxic bottom waters, the lagoon poses a potential risk to aquaculture. Centric diatoms were the most abundant in Orikumi (Miho and Dedej 1999). The phytoplank- ton structure seemed to be similar to that in Butrinti, but the cell density was much lower. However, the increase of peridinophytes was observed in summer as well. Taking the species composition into account, even Orikumi seems to represent predominantly zone 3, with similar features of sur- face waters in Butrinti, related to shellfish production. However, it is worth mentioning that the coastal wetland at Orikumi is exposed to one of the strongest human impacts among the Albanian lagoons. Its area was extremely reduced by land reclamation about 35 years ago. Consequently, its biological characteristics were significantly altered. MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS _ 137 To obtain a complete zonation view for each lagoon a more detailed approach of their physi- co-chemical and biological features would be necessary. A small number of phytoplankton samples from Narta, a large and very shallow lagoon (the depth ranges from 0.3 to | m), contained a few species only. This water body is characterized by a very limited connection with the sea and slow water exchange, belonging mainly to zone 6. After Dutrieux and Guelorget (1988) it does not sup- port intensive phytoplankton growth, and its use for intensive aquaculture is rather limited; there- fore, the lagoon could be suitable for traditional fishing. Pennate diatoms were most abundant in Karavasta. Unlike in other lagoons (Butrinti or Ceka), blooms or intensive growth of peridinophytes were not observed there. In agreement with Guelorget and Lefebvre (1993), zones 4 and 5 seem to prevail within the lagoon, characterised by a high productivity, mostly attributable to phytoplankton. Therefore, it may be used mainly in an extensive way for farming detritivorous fish species (e.g., mullet) or peneid shrimps (Dutrieux and Guelorget 1988). Similar characteristics seem to be typical of Patoku and Ceka (Lezha). As indi- cated by the diatom flora, dominated by centrics, Merxhani lagoon shows a relatively good water exchange with the sea, characteristic of zone 3. During April-June 1987, an abnormal bloom of Pseudonitzschia seriata was observed in Butrinti, followed by a bloom of Prorocentrum micans and P. minimum (Miho 1994), both known to produce toxins (Hallegraeff 1995). Algal bloom was observed also in Ceka (Lezha; spring-sum- mer 1992), where Nitzschia spp. or Prorocentrum spp. and Gonyiaulax spp. were dominant. This indicates conditions of stress, ensuing probably due to a combination of climate, the very limited water circulation in the lagoon and restricted water exchange with the sea. According to Marasovic (1989), the very high evaporation rate accompanied by increasing water temperature and salinity may contribute to the environmental stress as well. These critical conditions allow the presence of only a few species, with extremely dense growth. Parameters important in evaluation of the trophic status and other ecological features of a habi- tat may include the diversity index (Shannon and Weawer 1949) and/or dominance index (McNaughton 1967). In Butrinti, the diversity index was higher in marine habitats (varying from 1.9 to 3.4 in the littoral zone) than within the lagoon (ranging from 0.6 to 2.5; Miho 1994, 1996). In lagoon habitats, contributions of the two dominant species were very high (up to 92.4 % in Butrinti) and showed a strong seaward decrease (27.8% in Armura or 30.3% in Saranda). Generally, even the number of species was higher in the littoral marine habitats than in the lagoons, where the water exchange was limited. Armura, Butrinti (northern part), Karavasta, and Merxhani were the most distinct habitats in terms of species richness and trophic status. CONCLUDING REMARKS Albanian coastal wetlands, exemplified by the habitats studied in Butrinti, Armura, Karavasta, Orikumi and Merxhani show a high diversity of diatoms. The species composition found in some lagoons provides evidence of relatively good water exchange with the sea and favourable condi- tions of the natural environment. Stress conditions were observed in some other localities. Algal blooms were sporadically observed in Butrinti and Ceka, especially during the summer. The blooms were formed by pennate diatoms, e.g., Pseudonitzschia seriata and Nitzschia reversa, and by a few peridinophytes some of which are known to produce neurotoxins. Highly stressing con- ditions seem to prevail for an extended period of time in Narta and Kenalla pond (Lezha). The eutrophic status and related stressing conditions there are probably the result of limited water exchange with the sea. Moreover, the Albanian coast and the wetlands continue to be endangered by tourism, urban and industrial wastewater pollution, etc. 138 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 Potentially, the Albanian coast is a valuable environmental asset, but there is an urgent need of responsible protection and management. Taking appropriate measures in important and sensitive watershed areas will help to prevent damage to biodiversity and other natural values. Studies of the structure of diatom floras would provide additional information leading toward a better ecological approach of coastal ecosystems. ACKNOWLEDGMENTS This study was completed with the support from the Alexander von Humboldt Foundation to Dr. Miho. The authors are grateful to Prof. H. Lange-Bertalot, for using his laboratory facilities and library. Special thanks are to Manfred Ruppel, Botanical Institute, J.W. Goethe University, Frankfurt am Main for operating the Scanning Electronic Microscope and to D. Tase, Tirana University, who assisted preparation of the manuscript. Dr. Teresa Radziejweska for her help in correcting the English. LITERATURE CITED Bak, M., B. WAWRZYNIAK-WYDROWSKA, AND A. WITKOWSKI. 2001. Odra River discharge as a factor affecting species composition of the Szczecin Lagoon diatom flora, Poland. Pages 491-506 in R. Jahn, J.P. Kociolek, A. Witkowski and P. Compere, eds., Diatoms studies — Lange-Bertalot Festschrift. Gantner, Ruggell. Beck, G. 1904. Beitrag zur Flora des 6stlichen Albanian. Annales Naturhistorisches Hofmuseum, Wien. 78 pp. CuLLAJ A., A. Mino, B. BARAJ, A. HASKO, R. BACHOFEN, H. BRANDL, AND F. SCHANZ. 2003. Peliminary water quality report for some important Albanian Rivers. Journal of Environmental Protection and Ecology. B.EN.A. ed. (Special Issue):5—11. DANIELIDIS, D.B. 1991. A Systematic and Ecological study of Diatoms in the Lagoons of Messologhi, Aetoliko and Klessova. Ph.D. Dissertation. University of Athens, Athens, Greece. 320 pp. DebeJ, Z. 1994. On some new data of phytoplankton of Durres Gulf. Works of Biological Research Institute 9:67—73. (In Albanian) DepeJ, Z. 1995. Data on phytoplankton of Karavasta lagoon. Works of Biological Research Institute 10:26-30 (In Albanian) DUTRIEUX, E., AND O. GUELORGET. 1988: Ecological Planning: A Possible Method for the Choice of Aquacultural Sites. Ocean and Shorline Management 11:427-447. EssINK, K. 2003. Response of an estuarine ecosystem to reduced organic waste discharge. Aquatic Ecology 37:65-76. Forti, A. 1901—02. Primi appunti per uno studio sul phytoplancton del lago di Scutari d’ Albania (presentato da G.B.de Toni). Atti del Reale Instituto Veneto di Scienze, Lettere ed Arti. 61:704—708. GEULORGET, O., AND A. LEFEBVRE. 1993. Résultats de la campagne réalisée an avril 1993 sur trios lagunes Albanais: Butrinti, Karavasta et Patoku. Laboratoire d’ Hydrobiologie Marine, Université des Sciences et Techniques du Languedoc, France. GEULORGET, O., AND J.-P. PERTHUISOT. 1984. Indicateurs biologiques et diagnose ecologique dans le domaine paralique. Bulletin Ecologique 15:67—76. HAKANSSON, H., S. HAsDu, P. SNOEUS, AND L. LOGINOvA. 1993. Cyclotella hakanssoniae Wendker and its rela- tionship to C. caspia Grunow and other similar brackish water Cyclotella species. Diatom Research 8:333-347. HALLEGRAEFF, G.M., D.M. ANDERSON, AND A.D. CEMBELLA, EDS. 1995. Manual on Harmful Marine Microalgae. Intergovernmental Oceanographic Commission (IOC), Manuals and Guides No. 33. United Nations Educational, Scientific, and Cultural Organization (UNESCO), Paris, France. 551 pp. (See also: <http-://erf.org/education/erefs/Harmful_Algae.pdf>) Hustept, F. 1942-1945. Diatomeen aus Seen und Quellgebieten der Balkanhalbinsel. Archiv fiir Hydrobiolo- gie 40: 867-973, pls. 3143. MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS _— 139 JuriLJ, A. 1954. Flora i vegetacija diatomeja Ohridskog Jezera. Pages 100-190 in Prirodoslovna Istrazivanja, Knjiga 26, Jugoslavnaja Akademija, Zagreb. Kaso, M., ED. 1990-91. Gjeografia Fizike e Shgipérisé, Vol. | and Hl. Academy of Sciences. Geographic Centre, Tirana, Albania. KRENNER, J.A. 1926. Algae: Bacillariales. Né: Additamenta ad Floram Albanieae. Magyar Tudomdnyos Akad. Balkan-Kutatasainak Tudomdnyos Erdményoo (Budapest) 3:90—128 Marasovic, I. 1989. Phytoplankton and eutrophication of the Adriatic Sea. Pages 36—46 in Atti del Convegno ‘Lo stato di salute dell’Adriatico’ , Urbino, Italy: McNAuGHTON, S.J. 1967. Relationships among functional properties of California grassland. Nature 216: 168-169. Mino, A. 1994. Qualitative and Quantitative Data on Phytoplankton of Butrinti Lake (Saranda). Ph.D. Dissertation. Tirana University, Tirana, Albania. 145 pp. (In Albanian) Mino, A. 1996. Primary production and trophic state of Butrinti lake, Albania. Pages 37-43 in R.M.H. Tekke, ed., Proceedings of the International Seminar: Managment of Coastal Lagoons in Albania, May 29—June 4, 1994, Tirana, Albania. European Union for Coastal Conservation (EUCC), The Netherlands. Mino, A. 1998a. Contribution on microalgae of Ohrid, Macro and Micro Prespa lakes. Pages 102—107 in Proceedings of the International Seminar: Towards Integrated Conservation and Sustainable Development of Transboundary Macro and Micro Prespa Lakes, Korcha, 24—26 Oct. 1997, Korcha (Albania). Edition of Preservation and Protection of Natural Environment in Albania (PPNEA), Tirana, Albania. Mino, A. 1998b. Considerations on trophic state of Albanian coastal wetlands.. Pages 99-105 in Proceedings of the Third National Symposium of Geography: Applied Geomorphology, Environment and Coastal Tourism of Albania, October 16-18, 1998. Edition of Geographical Studies Center, Albanian Academy of Sciences, Tirana, Albania. (In Albanian) Mino A., A. CAKE, AND E. CARCANI. (In press.). Diatoms in the stomach content of barbel (Barbus meridion- alis) from Shkumbini River (Central Albania). Journal of Environmental Protection and Ecology. Balkan Environmental Association (B.EN.A.) ed. Sofia, Bulgaria. Mino A., AND H. LANGE-BERTALOT. 2001. Data on microflora of Albanian waters: diatoms from glacial lakes. Studime Biologjike 5—6/2001:234-236. Mino A., AND H. LANGE-BERTALOT. 2003. Considerations on biodiversity and trophic state of lake Ohrid (Albanian part) from a microscopic algae point of view. Journal of Environmental Protection and Ecology 4/3:543-549. Mino A., AND R. MITRUSHI. 1999. Phytoplanktonic data and trophic state of Lezha lagoons. Albanian Journal of Natural and Technical Sciences (AJNTS) (Academy of Sciences, Tirana, Albania) 1:69-76. Mino A., AND Z. DEDEJ. 1999. Microscopic algae form littoral habitats of Albanian Sourthern Adriatic. Studime Biologjike 2:112—123. NEA/AKM eb. 1999. Albania: Convention on Biological Diversity. Biodiversity Strategy and Action Plan (National Report), National Envrionmental Agency, Tirana, Albania. 100 pp. PEJA N., A. Vaso, A. MIHO, N. RAKAJ, AND J.L. CRIVELLI. 1996. Characteristics of Albanian lagoons and their fisheries. Fisheries Research 27:215—225. PRASAD, A.K.S.K., J.A. NIENOW, AND R.J. LivinGston. 1990. The genus Cycotella (Bacillariophyta) in Choctawahtchee Bay, Florida, with species reference to C. striata and C. choctawhatcheeana sp. nov. Phycologia 29:418-436. PROSHKINA-LAVRENKO, A.J. 1955. Diatomovie Vodorosli Planktona Cernovo Moria. Akademia Nauk, SSSR, Moscow-Leningrad, USSR. 222 pp. Protic, G. 1907. Beitrag zur Kenntnis der Algenflora Albaniens. Wissenschaftliche Mitteilungen aus Bosnien und Herzegovina 10:611-621. RAKAJS, M. 2002. Study on the Phytoplankton of Lake Shkodra (Floristic Composition, Seasonal Dynamics and Trophic State). Ph.D. Dissertation. Tirana University, Tirana, Albania. 154 pp. (In Albanian) RAKAJ M., F. HINDAK, AND A. HINDAKOvA. 2000. Phytoplankton species diversity of the Albanian part of Lake Shkodra in 1998-1999. Biologia, Bratislava 55:329-342. RakAJ, M., A. Mino, AND L. Kasuta. 2001. Data on diatoms (microalgae) from Shkodra Lake. Studime Biologjike 5—6/2001:237-241. 140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 ScumipT, A. 1874-1959. Atlas der Diatomaceen-Kunde. Band J and II. Leipzig and Berlin, Germany. SHANNON, C.E., AND W. WEAWER. 1949. The Mathematical Theory of Communication. University of Illinois Press, Urbana, Illinois, USA. 117 pp. SILVESTRE, F., B. BECK-EIHLER, W. DULEBA, AND J-P. DEBENAY. 2002. Modern benthic diatom distribution in a hypersaline coastal lagoon: the Lagoa de Araruama (R.J.), Brazil. Hydrobiologia 443:213-231. TROBAJO, R. 2003. Ecological analysis of periphytic diatoms in Mediterranean coastal wetlands (Emporda wetlands, NE Spain). Tesis Doctoral. Universidad de Girona, Girona, Spain. UHERKOVIC, G. 1963. Contributions a la connaisance du potamo-phytoplancton de quelques fleuves de 1’ Albanie. .Acta Botanica Academiae Scientiarum Hungaricae 9(1—2):161—172. UNEP, Eb. 2000. Post-Conflict Environmental Assessment — Albania. United Nations Environment Programme (UNEP), SMI (Distribution Services) Limited, Stevenage, UK. 80 pp WENDKER, S. 1991. Cyclotella hakanssoniae sp. nov. (Bacillariophyceae) — eine kleine Cyclotella-Art aus dem Schlei-Astuar (BRD). Nova Hedwigia 52:359-363. Viticic D., I. MARASOvIC, AND D. MroKovic. 2002. Checklist of phytoplankton in the eastern Adriatic Sea. Acta Botanica Croatica 61:57—91. (See also: <hitp://hirc. botanic. hr/actabot/acta6 1%281%29.html>) WirkowskI, A. 1991. Diatoms of the Puck Bay coastal shallows (Poland, Southern Baltic). Nordic Journal of Botany 11:689-701. WiTKowskI, A., H. LANGE-BERTALOT, AND D. METZELTIN. 2000. Diatom Flora of Marine Coasts. I. Iconographia Diatomologica 7:\—925. MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS 141 Appendix TABLE |. Checklist of Diatoms Found in Albanian Coastal Wetlands Key: A: Armura; B: Butrinti; C: Ceka; D: Durresi; K: Karavasta; Ke: Kenalla; L: Lalzi; Le: Lezha; M: Merxhani; N: Narta; O: Orikumi; P: Patoku; S: Saranda; VI: Vlora; V: Viluni (see Fig. 1). Note: The taxa labeled with “VI were reproted by Protic (1907), “D’ and partly “K’ by Dedej (1994, 1995), “A, B, C, L, Le, K, Ke, M. O, P, S, V’ by Miho (1994, 1996), Miho and Dedej (1999), and Miho and Mitrushi (1999); data for unlabeled taxa are from the unpublished notes of A. Miho. Bacillariophyceae: Centricae Actinocyclus cf. subtilis (Gregory) Ralfs [S] Actinocyclus ochotensis Jouse [A] Actinocyclus octonarius Ehrenberg (= A. ehrenbergii Ralfs) [B] Actinocyclus subtilis (Gregory) Ralfs [A, B, S] Actinoptychus senarius (Ehrenberg)Ehrenberg [B] Actinoptychus splendens (Shadbolt) Ralfs [M] Amphitetras antediluviana Ehrenberg [B, K, O] Asteromphalus heptactis (Brébisson) Ralfs [B, M] Asteromphalus robustus Castracane [B] Aulacoseira ambigua (Grunow) Simonsen [B] Aulacoseira granulata (Ehrenberg) Simonsen [B, K] Aulacoseira islandica (Mueller) Simonsen [L] Bacteriastrum furcatum Shadbolt [B, Ke] Bacteriastrium delicatulum Cleve [B; Ke] Biddulphia pulchella Gray [A, B] Cerataulina pelagica (Cleve) Hendey [K] Cerataulina bergonii (Peragallo) Schuett [B, D] Ceratulus turgidus (Ehrenberg) Ehrenberg [B, C, K, M, P] Chaetoceros affinis Lauder [B] Chaetoceros atlanticus Cleve [B, K] Chaetoceros atlanticus var. neapolitana (Schroeder) Schuett [B, K, M] Chaetoceros decipiens Cleve [B] Chaetoceros densus Cleve [B, Le] Chaetoceros diversus Cleve [B, D] Chaetoceros insignis Proshkina-Lavrenko [D] Chaetoceros lorenzianus Grunow [D] Chaetoceros muelleri Lemmermann [Ke] Chaetoceros peruvianus Brightwell [D] Chaetoceros subtilis Cleve [B] Chaetoceros teres Cleve [D] Chaetoceros tortissimus Grunow [B]| Chaetoceros wighamii Brightwell [B] Coscinodiscus cf. curvatulus Grunow var. minor[B]| Coscinodiscus nodulifer Schmidt [B, K] Cyclotella aff. choctawhatcheeana Prasad [B, O, K] Cyclotella cf. hustedti Jurilj [B] Cyclotella meneghiniana Kiitzing [B, Le, O, D] Cyclotella ocellata Pantocsek [B, O] Cyclotella radiosa (Grunow) Lemmermann [B, K] Cyclotella striata (Kiitzing) Grunow [B, P] Cymatosira lorenziana Grunow [S] Dactyliosolen blavyanus (Peragallo) Hasle [B, D, M, O] Dactyliosolen fragilissimus (Bergon) Hasle [B, D, K, M, O] Guinardia flaccida (Castracane) Peragallo [K] Guinardia striata (Stolterfoth) Hasle (=Rhizosolenia stolterfothii Peragallo) [B] Hemiaulus hauckii Grunow [B, D, M]| Hyalodiscus radiatus (O’Meara) Grunow [B, D, K, Le, O, P, V] Hyalodiscus scoticus (Kititzing) Grunow [B] Leptocylindrus adriaticus Schroeder [B, K, O] Leptocylindrus minimus Grunow [B] Melosira moniliformis (Mueller) Agardh var. moniliformis [B, Le, M, O, P, V] Melosira nummuloides Agardh [B, K, Ke, M, O, P] Melosira sol (Ehrenberg) Kiitzing [S] Melosira varians Agardh [B, K, V1] Odontella aurita (Lyngbye) Brébisson [B, K] Odontella mobiliensis (Bailey) Grunow [B, K] Porosira cf. pseudodenticula (Hustedt) Jouse [A] Proboscia alata (Brightwell) Sundstroem (=Rhizosolenia alata Brightwell) [B, D, K, M, O] Pseudosolenia calcar-avis (Schultze) Sundstroem [B, D] Rhizosolenia imbricata var. shrubsolei (Cleve) Schroeder [B] Skeletonema costatum (Greville) Cleve [K] Thalassiosira lacustris Grunow var. lacustris [B, K] Triceratium favus Ehrenberg [B] Triceratium reticulum Ehrenberg [S] Bacillariophyceae: Pennatae Achnanthes brevipes Agardh var. brevipes [B, K] Achnanthes brevipes var. angustata (Greville) Cleve [B, P] Achnanthes brevipes var. intermedia (Kiitzing) Cleve [B, K,M, O, P, VJ Achnanthes cf. groenlandica (Cleve) Grunow [S] Achnanthes delicatula (Kitzing) Grunow var. delicatula [B] Achnanthes exigua Grunow [B] Achnanthes exilis Kiitzing [V1] Achnanthes hungarica Grunow [K] Achnanthes lanceolata (Brébisson) Grunow agg. [B] Achnanthes longipes Agardh [B, K, M, P, V] Achnanthes minutissima Kiitzing [B, D, K, M, N, V1] Achnanthes separata Hustedt [A, S] Achnathes parvula Kiitzing [S] Adlafia minuscula Grunow (Lange-Bertalot) [Le] Amphipleura pellucida (Kiitzing) Kiitzing [B, D, K, V1] Amphora angularis Gregory [V1] Amphora angularis var. lyrata Van Heurck [V1] Amphora arenaria Donkin [A] Amphora cf. granulata Gregory [S] Amphora cf. tomiakae Witk., Lange-Bert. and Metz. [B, S] 142 Amphora coffeaeformis (Agardh) Kiitzing [B, C, K, O, P] Amphora commutata Grunow [V1] Amphora costata W. Smith [S] Amphora holsatica Hustedt [B, D, Le, O, P] Amphora inflata Grunow [K] Amphora kolbei Aleem [A, S] Amphora laevis Gregory [K] Amphora laevissima Gregory [S] Amphora lineolata Ehrenberg [B, K, Ke, M, V1] Amphora lybica Ehrenberg [B, D, K, O] Amphora marina (W. Smith) Van Heurck [A, B] Amphora ovalis (Kiitzing) Ktitzing [K, Le, O, VI] Amphora ovalis vat. affinis Kiitzing [V1] Amphora pediculus (Ktitzing) Grunow [B, K, Le, V1] Amphora pseudohyalina Simonsen [S] Amphora robusta Gregory [B, K, Le, P] Amphora salina W. Smith [V1] Aneumastus tusculus (Ehrenberg) Mann [Le, V1] Brachysira serians (Brébisson) Round and D.G. Mann [V]] Ardissonia crystallina (Agardh) Grunow (Agardh) Kiitzing [A, B] Ardissonia fulgens (Greville) Grunow [B, K, Le, P, S] Asterionellopsis glacialis (Castracane) Round [K] Bacillaria cf. socialis (Gregory) Ralfs [S] Bacillaria paradoxa Gmelin [B, D, K, Le, P] Berkeleya scopulorum (Brébisson) Cox [S] Caloneis amphisbaena (Bory) Cleve [B, K] Caloneis amphisbaena var. subsalina Van Heurck [V1] Caloneis bicuneata (Grunow) Wolle [B] Caloneis liber (W. Smith) Hendey [B] Caloneis silicula (Ehrenberg) Cleve agg. [B, D, K, V1] Caloneis westii (W. Smith) Hendey [B, K, P] Campylodiscus bicostatus W. Smith [B, K, O] Campylodiscus cf. simulans Gregory Campylodiscus clypeus Ehrenberg [B, K, O, V1] Campylodiscus echeneis Ehrenberg [B, K, Le, P] Campylodiscus levanderi Hustedt [B, K] Campylosira cymbelliformis (Schmidt) Grunow [V1] Cavinula cocconeiformis (Gregory) Mann and Stickle [B, K] Cocconeis costata Gregory [B, K, O] Cocconeis guttata Hustedt [S] Cocconeis molesta Kiitzing [K, Le] Cocconeis neodiminuta Krammer [B, K. O] Cocconeis pediculus Ehrenberg [B, O, V1] Cocconeis placentula (Ehrenberg) Hustedt agg. [V1] Cocconeis placentula Ehrenberg var. placentula [B, K, O] Cocconeis placentula var. euglypta (Ehrenberg) Cleve [B, Ke Le: P] Cocconeis placentula var. lineata (Ehrenberg) Van Heurck [Le] Cocconeis scutellum Ehrenberg [B, K, Le, P] Cocconeis pseudomarginata Gregory [B, A] Craticula cuspidata (Kiitzing) Mann [B, D, K, O] Craticula halophila (Granow) D.G. Mann [B, D, K, Le, V] Ctenophora pulchella (Ralfs ex Kiitzing) Williams and Round (=Fragilaria pulchella (Ralfs) Lange-Bertalot [B, Reka) Cylindrotheca cf. gracilis (Brébisson) Grunow [B, K, P] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 Cylindrotheca closterium (Ehrenberg) Reimann and Lewin (=Nitzschia closterium (Ehrenberg) W. Smith) [B, D, K, M, P] Cymatopleura elliptica (Brébisson) W. Smith [B, V1] Cymatopleura solea (Brébisson) W. Smith [B, V1] Cymbella affinis Kiitzing agg. [B, K, O, P] Cymbella ampicephala Naegeli var. amphicephala [B] Cymbella aspera (Ehrenberg) Peragallo [V1] Cymbella cistula (Ehrenberg) Kirchner agg. [V1] Cymbella cymbiformis Agardh var. cymbiformis [V1] Cymbella delicatula Kiitzing [V1] Cymbella ehrenbergii Kiitzing [V1] Cymbella helvetica Kiitzing [B, P, V1] Cymbella lanceolata (Ehrenberg) Kirchner [V1] Cymbella pusilla Grunow [V1] Cymbella tumida (Brébisson) Van Heurck [K, P, V1] Cymbella turgida (Gregory) Cleve [B] Delphineis cf. livingstonii Prasad [S] Denticula tenuis Kiitzing [V1] Diatoma anceps (Ehrenberg) Kirchner [V1] Diatoma ehrenbergi Kiitzing [Le] Diatoma hyemalis (Rith) Heiberg [V1] Diatoma tenuis Agardh (=D. elongatum (Lyngbye) Agardh) [O] Diatoma vulgaris Bory var. vulgaris [B, V1] Diatoma vulgaris var. tenuis Van Heurck [V1] Dimerogramma minor (Gregory) Ralfs [S] Diploneis cf. interrupta (Kiitzing) Cleve [S] Diploneis cf. littoralis (Donkin) Cleve [B, S] Diploneis ct. mirabilis Koenig [A] Diploneis cf. smithii var. dilatata (Peragallo) Terry Diploneis didyma (Ehrenberg) Cleve [B, Le, K, O, P] Diploneis domblittensis Cleve [B] Diploneis elliptica (Kiitzing) Cleve (=Navicula elliptica Kiitzing) [V1] Diploneis marginestriata Hustedt [B] Diploneis oblongella (Naegeli) Cleve-Euler [B] Diploneis ovalis (Hilse) Cleve [B, Ke] Diploneis smithii var. pumila (Grunow) Hustedt [B, K, P] Diploneis subovalis Cleve [B, M] Diploneis vetula (Schmidt) Cleve [A] Dploneis subcincta (Schmidt) Cleve [B] Encyonema caespitosum (Kiitzing) Grunow [V1] Encyonema prostratum (Berkeley) Cleve [V1] Encyonema ventricosum Kiitzing [B, V1] Encyonopsis minuta Krammer and Reichardt [S] Entomoneis paludosa (W. Smith) Reimer [B, K, Le, O, P] Entomoneis alata (Ehrenberg) Ehrenberg [B, S] Epithemia adnata (Kiitzing) Brébisson [B, K, O, V1] Epithemia smithii Carruthers [B] Epithemia sorex Kiitzing [B, K, O, V1] Epithemia turgida (Ehrenberg) Kiitzing [V1] Epithemia turgida var. granulata (Ehrenberg) Grunow [Le] Eunotia arcus Ehrenberg [V1] Eunotia cf. exigua (Brébisson) Rabenhorst [Le] Eunotia flexuosa (Brébisson) Kiitzing [V1] Eunotia pectinalis (Dillwyn) Rabenhorst [V1] Eunotia praerupta Ehrenberg [B, K, O] Fallacia versicolor (Grunow) Mann [S] MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS Fragilaria affinis Kiitzing [K] Fragilaria capucina Grunow var. capucina [V1] Fragilaria capucina vat. perminuta (Grunow) Lange- Bertalot [B] Fragilaria crotonensis Kitton [B, D, K] Fragilaria dilatata (Brébisson) Lange-Bertalot (=F. capita- ta (Ehrenberg) Lange-Bertalot) [V1] Fragilaria fasciculata (Agardh) Lange-Bertalot [B, K, Le, OEP Vi] Fragilaria investiens (W. Smith) A. Cleve [B] Fragilaria laevissima Oestrup [B, S] Fragilaria pinnata Ehrenberg gr. [B, K, O, P] Fragilaria ulna (Nitzsch) Lange-Bertalot agg. [B, Le, O, P] Fragilaria ulna (Synedra) var. longissima Van Heurck [V1] Fragilaria ulna (Synedra) var. oxyrhynchus (Kiitzing) Van Heurck [V1] Fragilaria (Synedra) ulna var. subaequalis Grunow [V1] Fragilaria virescens Ralfs [V1] Frustulia cf. weinholdii Hustedt [B] Gomphonema acuminatum Ehrenberg [Le, V1] Gomphonema angustatum (Kiitzing) Rabenhorst [V1] Gomphonema angustum Agardh [V1] Gomphonema constrictum Ehrenberg [V1] Gomphonema gracile Ehrenberg [Le] Gomphonema minutum (Agardh) Agardh agg. [B. K, O] Gomphonema olivaceum (Horn.) Brébisson var. olivaceum [B, K, O, V1] Gomphonema olivaceum var. calcareum (Cleve) Cleve [B, K, O] Gomphonema olivaceum var. staurophorum Pantocsek [B, D, K] Gomphonema parvulum (Kiitzing) Grunow [Le, V1] Gomphonema pseudotenellum Lange-Bertalot [Le] Gomphonema truncatum Ehrenberg [B, D, K, Le, O] Grammatophora angulosa Ehrenberg var. angulosa [B, Le] Grammatophora macilenta W. Smith [B, K, V] Grammatophora oceanica (Ehrenberg) Grunow var. ocean- ica [B, K, Le, N, O, P, V] Gyrosigma acuminatum (Kiitzing) Rabenhorst var. acumi- natum [B, K, Le, N, P, V] Gyrosigma attenuatum (Kiitzing) Rabenhorst [B, K, O] Gyrosigma arcuatum (Donkin) Sterrenburg Gyrosigma balticum (Ehrenberg) Rabenhorst [B, K, Le, N, O, P. VJ Gyrosigma cf. obscurum (W. Smith) Griffith and Henfrey [B] Gyrosigma cf. peisonis (Grunow) Hustedt [Le] Gyrosigma cf. wansbecki (Donkin) Cleve [Le] Gyrosigma compactum (Grev.) Cleve [B, S] Gyrosigma fasciola (Ehrenberg) Griff. and Henf. [B, K, Le, NODE. Vi) Gyrosigma Strigilis (W. Smith) Cleve [B, K, Le, P] Hantzschia amphioxys (Ehrenberg) Grunow [B, V1] Hantzschia amphioxys fo. capitata Hustedt [B] Licmophora cf. grandis (Kiitzing) Grunow [B, K] Licmophora cf. hyaline Kiitzing [B, K] Licmophora ehernberghii (Kiitzing) Grunow [B, K] Licmophora flabellata var. splendida W. Smith [K] Licmophora remulus Grunow [B, K, Le, N, O, P, V] 143 Licmophora dalmatica (Kiitzing) Grunow [V]] Licmophora flabellata (Carmichael) Agardh Licmophora gracilis (Ehrenberg) Grunow Licmophora paradoxa (Lyngbye) Agardh [A, S] Lyrella amphoroides D.G. Mann [S] Lyrella cf. clavata (Gregory) D.G. Mann [M] Lyrella lyra (Ehrenberg) Karayeva [B, K] Lyrella spectabilis (Gregory) D.G. Mann [A, S] Lyrella sulcifera (Hustedt) Witkowski [B, K, O, P, Le] Mastogloia angulata Lewis [B, K, O, P, Le] Mastogloia binotata (Grunow) Cleve [B] Mastogloia braunii Grunow [B, K, O, V1] Mastogloia cf. adriatica Voigt [B] Mastogloia cf. baldjikiana Grunow [B, K] Mastogloia cf. biocellata (Grunow) Nov. and Muft. [B] Mastogloia cf. labuensis Cleve [B] Mastogloia cf. recta Husted [B] Mastogloia crucicula (Grunow) Cleve [B, K, O] Mastogloia cuneata (Meister) Simonsen [B] Mastogloia dansei Thwaites [V1] Mastogloia grunowii A. Schmidt [B] Mastogloia ignorata Hustedt [B] Mastogloia lanceolata Thwaites [A] Mastogloia laterostrata Hustedt [B] Mastogloia ovalis A. Schmidt [A] Mastogloia paradoxa Grunow [B] Mastogloia pseudoexigua Cholonky [B] Mastogloia pseudolatecostata Yohn and Gibson [A] Mastogloia pumila (Grunow) Cleve [B] Mastogloia pusilla (Grunow) Cleve [B] Mastogloia robusta Hustedt [B] Mastogloia smithii Thwaites var. smithii [B] Mastogloia smithii var. lacustris Grunow [B, D, K, O] Mastogloia splendida (Gregory) Cleve [B] Mastogloia vasta Hustedt [B] Navicula agnita Hustedt [A] Navicula arenaria Donkin [B, S] Navicula cf. besarensis Giffen [S] Navicula cf. heimansii Van Dam et Kooyman [B] Navicula cf. indulgens Simonsen [B] Navicula cf. johanrossii Giffen [S] Navicula cf. subhamulata Grunow [Le] Navicula cincta (Ehrenberg) Kiitzing [V1] Navicula cluthensis Gregory var. cluthensis [P, V] Navicula cryptocephala Kiitzing [D, K, V1] Navicula cryptotenelloides Lange-Bertalot [B] Navicula dealpina Lange-Bertalot [V1] Navicula digitoradiata (Gregory) Ralfs [V1] Navicula distans (W. Smith) Ralfs [V1] Navicula duerrenbergiana Hustedt [B] Navicula erifuga Lange-Bertalot [A] Navicula granulata Bailey [C, Ke, V] Navicula gregaria Donkin [B, S, V] Navicula integra W. Smith [V1] Navicula monilifera Cleve [B, K] Navicula oblonga (Kiitzing) Kiitzing [B, K, O, V1] Navicula pavillardii Hustedt [B, C] Navicula peregrina (Ehrenberg) Kiitzing [V1] Navicula perminuta Grunow [B] 144 Navicula perrhombus Hustedt [B] Navicula phyllepta Kiitzing [B, K, O, P, Le] Navicula phylleptosoma Lange-Bertalot [B] Navicula pseudosilicula Hustedt [V1] Navicula radiosa Kiitzing [V, V1] Navicula recens (Lange-Bertalot) Lange-Bertalot [Le] Navicula rhynchocephala Kiitzing [B, V1] Navicula rolandii Lange-Bert. and Witk. [O] Navicula salinarum Grunow [B, K, O, V1] Navicula salinicola Hustedt [A, B, S] Navicula termes Ehrenberg [V1] Navicula tripunctata (Mueller) Bory [B, Le] Navicula veneta Kiitzing [B, K] Navicula viridula (Kiitzing) Ehrenberg [V1] Neidium iridis var. affinis Ehrenberg [V1] Neidium iridis var. amphirhynchus Ehrenberg [V1] Nitzschia acicularioides Hustedt [S] Nitzschia amphibia Grunow [B] Nitzschia angularis W. Smith [A, S] Nitzschia cf. filiformis var. conferta [Le] Nitzschia cf. graeffii Grunow [S] Nitzschia cf. plana W. Smith [B, S]} Nitzschia cf. prolongata Hustedt [B] Nitzschia cf. splendida Kiitzing [B, K] Nitzschia capitellata Hustedt [B, K, O] Nitzschia circumsuta (Bailey) Grunow [B, D, K, P, Le, V] Nitzschia clausii Hantzsch [B] Nitzschia coarctata Grunow [B, K, O, P, Le] Nitzschia communis Rabenhorst [V1] Nitzschia commutata Grunow [V]] Nitzschia compressa (Bailey) Boyer var. compressa [B, D, KAOsPs Le] Nitzschia constricta (Kiitzing) Ralfs [B, D, K, O, P, Le] Nitzschia denticula Grunow (=Denticula Kiitzingii Grunow) [B, K, O, P] Nitzschia dissipata (Kiitzing) Grunow [B] Nitzschia dissipata var. media (Hantzsch) Grunow [Le] Nitzschia distans var. tumescens Gregory [K] Nitzschia filiformis (W. Smith) Hustedt [A, B, S] Nitzschia flexoides Geitler [B] Nitzschia fossilis Grunow [Le] Nitzschia granulata Grunow [B, Le] Nitzschia hungarica Grunow [V1] Nitzschia incospicua Grunow [B] Nitzschia lacuum Lange-Bertalot [A, B, S] Nitzschia lanceola var. minutula Grunow [B] Nitzschia lanceolata W. Smith [B] Nitzschia linearis (Agardh) W. Smith [V1] Nitzschia littoralis Grunow [S] Nitzschia longissima (Brébisson) Ralfs [B, D, K, Le, V] Nitzschia macilenta Gregory [B] Nitzschia microcephala Grunow [Le] Nitzschia palea (Kiitzing) W. Smith [B, K, Le, V1] Nitzschia panduriformis Gregory [A] Nitzschia pararostrata (Lange-Bertalot) Lange-Bertalot [B] Nitzschia recta Hantzsch [B] Nitzschia reversa W. Smith [Le, O} Nitzschia scalpelliformis Grunow [V] Nitzschia sigma (Kiitzing) W. Smith [B, K, Le, V1] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 12 Nitzschia sigma var. sigmatiella Grunow [D, K, Le, P, V] Nitzschia sigmoidea (Nitzsch) W. Smith [V1] Nitzschia paleacea (Grunow) Grunow [V1] Nitzschia vitrea var. salinarum Grunow [V1] Opephora mutabilis (Grunow) Sabbe and Vyverman [B] Parlibellus berkeleyi (Kiitzing) Cox [A, S] Parlibellus cruciculoides (Brock.) Witk., Lange-Bert. and Metzeltin [S] Parlibellus protracta (Grunow) Witk., Lange-Bert. and Metzeltin [A, Le, S] Parlibellus rhombicula (Hustedt) Witk., Lange-Bert. and Metzeltin [S] Petroneis humerosa (Brébisson) D.G. Mann [B, K, M] Pinnularia appendiculata (Agradh) Cleve [V1] Pinnularia gibba Ehrenberg [V1] Pinnularia legumen (Ehrenberg) Ehrenberg [V1] Pinnularia major (Kiitzing) Rabenhorst [V1] Pinnularia microstauron var. brebissonii (Kiitzing) Mayer [V1] Pinnularia nobilis (Ehrenberg) Ehrenberg [V1] Pinnularia pulchra Oestrup Ehrenberg) [V1] Pinnularia tabellaria Ehrenberg [V1] Placoneis elginensis (Gregory) Cox [V1] Placoneis gastrum vat. signata (Hustedt) [B, K] Plagiotropis gibberula Grunow [S] Plagiotropis lepidoptera (Gregory) Kuntze [A, B, V] Pleurosigma angulatum (Quekett) W. Smith [B, K, Le, O, P, Vv) Pleurosigma cf. rostratum Hustedt [M] Pleurosigma elongatum W. Smith [B, D, K, Le, O, P, V] Pleurosigma formosum Peragallo [B, K, Le, O, P, V] Pleurosigma salinarum Grunow [S, Le] Pseudonitzschia delicatissima (Cleve) Peragallo [D] Pseudonitzschia seriata (Cleve) Peragallo [B, K, Le, O, P, Vv] Prerodictyon gemma (Ehrenberg) D.G. Mann [B, K, P] Rhabdonema adriaticum Kiitzing [B, K, Le, O, P, V] Rhoicosphenia abbreviata (Agardh) Lange-Bertalot [B, V1] Rhoicosphenia marina (W. Smith) M. Schmidt [B] Rhopalodia acuminata Krammer [A, S] Rhopalodia brebissonii Krammer [B] Rhopalodia cf. constricta (W. Smith) Krammer [A] Rhopalodia cf. gibberula (Ehrenberg) Mueller [B, K] Rhopalodia gibba (Ehrenberg) Mueller [B, K, Le, O, P, V1] Rhopalodia gibba var. minuta Krammer [A, S] Rhopalodia musculus (Kiitzing) Mueller [B, D, K, Le, O, P, vil Sellaphora bacillum (Ehrenberg) Mereschkowsky [V1] Sellaphora seminulum (Grunow) D.G. Mann [V1] Seminavis basilica Danielidis and D.G. Mann [A, B, S] Stauroneis acuta W. Smith [V1] Stauroneis anceps Ehrenberg [V1] Stauroneis gregori Ralfs [V1] Stauroneis phoenicenteron Ehrenberg [V1] Stauroneis salina W. Smith [V1] Staurosira brevistriata Grunow (=Fragilaria brevistriata Grunow, Pseudostaurosira brevistriata (Grunow) Williams and Round) [B, O] Staurosira construens Ehrenberg (=Fragilaria construens MIHO AND WITKOWSKI: DIATOM FLORA OF ALBANIAN COASTAL WETLANDS © 145 (Ehrenberg) Grunow) [B, S] Surirella splendida (Ehrenberg) Kiitzing [V1] Striatella unipunctata (Lyngbye) Agardh [B, K, Le,O,P,V] — Surirella striatula Turpin [B, C, K, O, P] Surirella biseriata Brébisson [V1] Synedra tabulata (Agardh) Kiitzing var. tabulata [B, K, Le] Surirella brebissonii Krammer and Lange-Bertalot [B, K, | Synedra undulata (Bailey) Gregory (=Toxarium undulatum O}] Bailey) [B, Le, O] Surirella cf. fluminensis Grunow [B, K, M, P] Tabellaria fenestrata (Lyngbye) Kiitzing [V1] Surirella cf. scalaris Giffen [A, S] Tabellaria flocculosa (Roth) Kiitzing [V1] Surirella fastuosa (Ehrenberg) Kiitzing [B, K, Le, O, P, V] Thalassionema nitzschioides (Grunow) Grunow [B, D, K, Surirella minuta Brebisson [V1] Le, O] Surirella ovalis Brébisson [B, K, O, V1] Trachyneis aspera (Ehrenberg) Cleve [A] Surirella spiralis Kiitzing [K, V1] Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 56, No. 13, pp. 146-155, 9 figs. April 29, 2005 Marrying Taxonomy and Ecology: An Attempt Eugene F. Stoermer Michigan Herbarium and School of Natural Resources and Environment, University of Michigan, Ann Arbor, MI 48109-1115;Email: stoermer@umich.edu Freshwater diatoms present an interesting challenge in an era when biodiversity is becoming a major concern. Although tremendously useful indicators of ecological conditions, past and present, lack of taxonomic knowledge limits the potential of eco- logical interpretation. At the same time the ecological studies that are carried out provide less than optimal feedback to the taxonomic literature. I suggest that appro- priate use of available computer-based technologies can integrate these fields to the benefit of both. I further outline the approaches taken in an early and primitive attempt to accomplish this goal, the benefits derived, and the mistakes made and inadequacies of our effort at that time. Thoughtful application of technologies now available has the potential to further integrate studies and expand eventual under- standing. The following is a discussion of an attempt to marry the fundamental approaches of systemat- ic practice to ecological studies. The tools and approaches used are, in retrospect, quite primitive, but there is an underlying logical framework that applies to all such endeavors, in taking on any problem at any time. | hope that discussing the way we attempted to solve problems common to any taxonomically based diatom study, what worked and what didn’t, and the mistakes made, will be of some value to current investigators. I should hasten to explain that the “we” in the previous paragraph is used advisedly. I am not a programmer, so much of the development and implementation was done by other people, better equipped to deal with the intricacies of programming than I. I thank the late Dr. Vincent Noble and Dr. Edward Johnston (Johnston and Stoermer 1976) for enlightening discussions of logical struc- tures appropriate for human — computer interactions. The initial programming was done by Dr. J.K.C. Huang and the system was brought to its most advanced state largely through the efforts of Theodore and Barbara Ladewski (Ladewski and Stoermer 1973: Sicko-Goad et al. 1977). Numer- ous helpful comments and suggestions were also made by many technical staff and students, which materially helped shape the project. THE PROBLEM In the mid 1960s I was a young investigator faced with the rather intimidating problem of investigating the algal flora of the Laurentian Great Lakes. At the time, severe eutrophication prob- lems were apparent in many regions of these lakes (Beeton 1965, 1969). Because of the Great Lakes’ tremendous value to the economies of the United States and Canada, considerable resources were available for studies related to water quality. Many of the practical problems that beset the lakes at that time were directly related to algae. Taste and odor problems caused by diatoms in the spring (Vaughn 1961, 1962) and cyanophytes in the summer and fall (Stoermer and Stevenson 146 STOERMER: MARRYING TAXONOMY AND ECOLOGY 147 1980; Bierman and Dolan 1981; Stoermer and Theriot 1985). Cladophora was a nuisance 1n many regions of the lakes (Wolfe and Sweeney 1980) and generally unpleasant obnoxious conditions were present in many areas. Lake Erie, in particular, became a cause célébe of the environmental activism of the day, and was widely reported in the common press to be a “dead lake.” This was somewhat problematic to biologists, as the actual problem was over-production, which eventually led to de-oxygenation of the bottom waters in certain areas of the lakes, creating so called “dead zones” where benthic invertebrates were periodically exterminated. In retrospect, the problems of the 1960s and 1970s were only the most recent in a long history of environmental catastrophes, such as epidemics of water-borne diseases (Beatty 1982; Bonner 1991) that devastated communi- ties that drew drinking water from the lakes. For example, the great cholera epidemic of 1854 was estimated to have killed five percent of the total population of the city of Chicago. Collapse of native fish stocks began soon after western settlement of the region (Smith 1972), and culminated in total extermination of some native stocks by 1950 (Beeton 1969) and introduction of many exot- ic fish species. One would rationally suppose such a valuable, but clearly damaged, ecosystem would have received careful and comprehensive study, especially considering the large number of well-known academic institutions in the region. Unfortunately, this was not the case. The ecological history of the Great Lakes, in many respects, provides a sterling example of precisely the wrong way to approach management of a large and complex ecosystem. Each successive crisis generated a wave of “directed research” centered on the apparent problem and to a lesser extent, if at all, on its root causes. “Charismatic vertebrates,” in this case fish, were the initial center of attention, and lesser attention and resources were devoted to the rest of the biota or to chemical and physical factors of the environment. In the case of diatoms, early (in the North American context) exploratory studies were carried out by J.W. Bailey in 1839, first mentioned in 1842 (Bailey 1842a, 1842b), and sent to C.G Ehrenberg, who more formally published them in his monumental works (Ehrenberg 1845, 1854). These collections are still maintained at the Museum fiir Naturkunde,