7 @755x NG SEPTEMBER 30, 2004 * VOLUME 55 * NUMBERS 13-25 o PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES CALIFORNIA ACADEMY or SCIENCES FOUNDED 1853 SAN FRANCISCO, CALIFORNIA Copyright © 2004 by the California Academy of Sciences All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage or retrieval system, without permission in writing from the publisher. SCIENTIFIC PUBLICATIONS Alan E. Leviton, Ph.D., Editor Gary C. Williams, Ph.D., Associate Editor Michele L. Aldrich, Ph.D., Consulting Editor COVER IMAGE On its return from the Galapagos expedition in November 1906, the Schooner Academy had in its hold an abundance of scientific specimens obtained by the scientists-crew during the 17-month long sojourn in the islands. Little did anyone realize that when the expedition departed San Francisco in June 1905 that the collections that the expedition would bring back to San Francisco would, in fact, form the nucleus of the Academy’s new research collections. Nearly all that had been gathered previously, dat- ing from the Academy’s founding in 1853, was destroyed in the earthquake and fire that engulfed the city of San Francisco on 18 April 1906. When the expedition returned, Academy staff proceeded to wall of a part of the severely damaged old museum building to serve as a temporary warehouse for its pre- cious collections. The photograph here shows the area in which the 258 tortoises brought back by the expedition were stored until the old museum building, which was slated for demolition a short time later, required that the specimens be moved to a new temporary site in a vacant downtown warehouse ISSN 0068-547X The Proceedings of the Califernia rteademy of Scieneed 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 Golden Gate Park, San Francisco, California 94118 U.S.A. Printed in the United States of America by Allen Press Inc., Lawrence, Kansas 66044 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Y’S WWISON IZ ~ Vis \ Volume 55, No. 13, pp. 255-299, 1 fig. / : ~y~- 0 7 AAR \ September 30, 2004 (OCT 07 2008 | a Annotated Listing of Diatom Types in Rabenhorst’s Die Algen Europa’s 1848-1882, and a Consideration of Their Nomenclatural Status J.P. Kociolek Diatom Collection, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103 USA; Email: pkociolek@calacademy.org Accumulation and dissemination of diatom collections during the 19** century were a significant means for diatomists to establish new taxa and communicate with the communi- ty of fellow researchers about the diversity of diatoms through space and time. The exsiccatae sets of Van Heurck (1882-1885), Cleve and Moller (1877-1882), Tempére and Peragallo (1889-1895; 1907-1915) and H.L. Smith (1876-1888) are relatively well known, but sets were also distributed by Wartmann and Schenk (1862-1882), M’Calla (1848) and Eulenstein (1867-1869). The materials distributed includ- ed at least some processed diatoms made into permanent slides or as raw material dried onto paper or mica. Another relatively well-known exsiccatae set, that followed more in the tradition of phy- cology, bryology, mycology and lichenology was distributed not as processed slides, but as raw material. This set, which included more K than 2500 numbers and is known as “Die Algen We YG. poe we Europas,” was developed and distributed by / y Ludwig Rabenhorst (Fig. 1) between 1848 and 1882 (Rabenhorst 1848-1860, 1861-1882). Material included came from all over Europe as well as from far away places such as India, Ficure 1. Ludwig Rabenhorst. From an engraving in Burma, New Zealand and Honduras. The set Flora Europaea Algarum (1864). Courtesy Department of 2 ; see Botany, National Museum of Natural History, Washington, formed the basis of numerous investigations by ¢ Rabenhorst and a wide range of collaborators, some immediately recognizable to diatomists (e.g., Grunow, Janisch, and Hilse) and others not so well known (Reinsch, Bleisch, Schwarz, and Hantzsch, after whom the genus Hantzschia was named). These sets were distributed widely, and may be found in herbaria across North America and the world. pad « SD WS fp ice 606. aa FEDS 256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 From these samples, Rabenhorst and colleagues observed what they believed to be new taxa. In some cases, the taxa were identified as such on the labels of the exsiccatae; in other cases, the material was the basis for separate scientific publications. In some cases in both paces, the exsic- catae and separate publications, were notices for new taxa. According to the International Code of Botanical Nomenclature (Greuter et al. 2000, Articles 32 and 42.2), distribution of exsiccatae sets satisfies the requirement of effective publication, but the sole listing of names as new taxa without illustration or description does not satisfy the requirement of valid publication. The presentation of names in the exsiccatae set of Rabenhorst was not consistent. In some cases, descriptions accom- panied the listing of the new taxon. In more rare situations, illustrations were included with the labels and descriptions. Frequently, the listing of names was without descriptions or illustrations. A similar situation to the listing of names only in exsiccatae was recently described for the exsic- catae slide set of Tempeére and Peragallo (Servant-Vildary et al. 2001). Recent taxonomic workers have also accessed the exsiccatae and described new taxa from the material (e.g., Krammer and Lange-Bertalot 1988; Krammer 2000). The Diatom Herbarium of the Academy of Natural Sciences of Philadelphia (ANSP) devel- oped a catalogue of the types contained in that collection (Mahoney and Reimer 1987, 1997). They listed taxa described from the Rabenhorst exsiccatae as types in ANSP. The approach taken to the Rabenhorst material was, apparently, to take the exsiccatae labels at face value, without consider- ation whether the name was validly published or not. The treatment in Mahoney and Reimer (1997) of new taxa in the Rabenhorst exsiccatae is uneven, attributing species to Rabenhorst where others are listed as the author, not indicating the place of valid publication, listing of taxa that have not been validly published as well as missing many species either validly published in the exsiccatae or elsewhere. Recently, Robert Edgar of the Farlow Herbarium has developed a website where indi- vidual packets with printed material can be accessed (http://www.huh.harvard.edu/diatom/pub- collfset.htm). Given the inconsistent way in which new taxa are treated in the Rabenhorst exsiccatae, and the superficial treatment in developing the type catalogue for ANSP, as well as the fact that the Rabenhorst exsiccatae contained type material for a wide range of diatom taxa, it was decided to present a detailed account of the types and new diatom names found in this interesting material. This work was also determined to be important, inasmuch as the number of catalogues of diatom types is relatively few (Mahoney and Reimer 1987, 1997; Williams 1988; Simonsen 1987; Kociolek et al. 1999; Servant-Vildary et al. 2001). This work should also facilitate a new planned catalogue of diatom species, similar to the catalogue of diatom genera published recently (Four- tanier and Kociolek 1999). METHODS AND APPROACH The resulting research was conducted on the Rabenhorst exsiccatae contained in the Farlow Herbarium at Harvard University. In the present publication, a listing of samples that serve as the type material for new diatom taxa is detailed. We count four new genera and 233 new taxa at the level of species or lower. This contribution is presented in four parts, as follows: Part 1 includes those names that have been published in the exsiccatae; herein they are distinguished whether they are validly or invalidly published. This should help direct students and researchers to those taxa of interest that may be new and and whose names require valid publication. For each sample, the fol- lowing information is provided: exsiccatae serial number, the year of its distribution, locality data, taxon name with authority, date of publication and page number, and then any notes on the publi- cation and taxonomic comments. Part 2 is an index of taxa and the exsiccatae number referenced KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S DY) to the first part for both validly and invalidly published names. Part 3 summarizes the data for each name referenced in the first two parts; the names are presented alphabetically. Lastly, in Part 4, data are provided with respect to place of publication, date, pagination, and nomenclatural status of each name. PART 1. TYPE MATERIAL OF DIATOM NAMES IN THE RABENHORST EXSICCATAE. Validly Published Names Exsiccatae Number: 383 Year Distributed: 1854 Locality Information: “Wasserfalles bei Schenpfenthal in Thiiringen” Taxon: Melosira roeseana Rabenhorst 1853, p. 13 Taxonomic Comments: Van Landingham (1971, p. 2241) indicates ““Rabenhorst 1852, #382” Exsiccatae Number: 403 Year Distributed: 1855 Locality Information: “Grosszschocker b. Leipzig in dem grossen Steinbruche, zwischen einem Oedogonium. 30. Aug. 1854” Taxon: Synedra interrupta Auerswald in Rabenhorst 1855 Taxonomic Comments: A brief description accompanies the exsiccatae. This is the basionym of the name Synedra splendens var. interrupta (Auerswald) Rabenhorst 1864a, p. 134 and S. ulna var. interrupta (Auerswald) M. Peragallo 1903. Exsiccatae Number: 505 Year distributed: 1856 Locality Information: “Wassermiihlen bei Bergamo” Publication Notes: A Latin description is included with the exsiccatae. See also Rabenhorst 1864a, p. 249, and Grunow 1860, p. 565. Taxon: Stauroneis rotaeana Rabenhorst 1856, p. 103, pl. XII, fig. 7 (without a description) Exsiccatae Number: 561 Year Distributed: 1857 Locality Information: “Mineralwassern zu Canstatt, Temp. von 16—-17° Reaum” Taxon: Falcatella zelleri Rabenhorst 1857a Taxonomic Comments: A brief description is included with the exsiccatae. Van Landingham (1969, p. 1646) indicates F’ zelleri was described in Rabenhorst (1863a, p. 40). This taxon is the basionym for Synedra zelleri (Rabenhorst) Rabenhorst (1864a, p. 139). Exsiccatae Number: 561 Year Distributed: 1857 Locality Information: “Mineralwassren zu Canstatt, Temp. von 16—17° Reaum” Taxon: Achnanthidium thermale Rabenhorst 1864a, p. 107 Exsiccatae Number: 604 Year Distributed: 1857 Locality Information: “In der Mergelgrube bei Peterwitz in der Nahe von Strehlen in Schlesien” Taxon: Cymbella helvetica f. silesiaca Rabenhorst 1857a Taxonomic Comments: A brief description is provided with the exsiccatae. Name not included in Van Landingham’s Catalogue. 258 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Exsiccatae Number: 611 Year Distributed: 1857 Locality Information: “Falaise” Taxon: Pinnularia falaiseana Krammer 1992, p. 113 Publication Notes: Krammer (1992) indicates that H.L. Smith (1876-1888) slide 285 represents type material for this taxon. Exsiccatae Number: 624 Year Distributed: 1857 Locality Information: “An feuchten Felsen beim Hirschensprung in Héllenthal (bei Freiburg im Br.)” Taxon: Gomphogramma rupestre (Kiitzing) A. Braun ex Rabenhorst 1853, p. 33 Taxonomic Comments: This is a new combination of Denticula thermales Beta rupestris Kiitzing 1849. This does not appear to represent type material. Exsiccatae Number: 625 Year Distributed: 1857 Locality Information: “. . .im Elbette .. .” Taxon: Nitzschia stagnorum Rabenhorst 1857a Taxonomic Comments: A Latin description is included with the exsiccatae. This name is the basionym of Nitzschia thermalis var. stagnorum (Rabenhorst) Rabenhorst 1864a. Exsiccatae Number: 642 Year Distributed: 1857 Locality Information: “An Leptomitus lacteus bei Stuttgart... . Taxon: Surirella intermedia Rabenhorst 1857b Taxonomic Comments: A brief description accompanies the exsiccatae. Surirella ovata var. inter- media (Rabenh) Rabenhorst 1864a, p. 57, is based on this name. bb) Exsiccatae Number: 682b Year Distributed: 1858 Locality Information: “Fossés pr. Falaise. De Brébisson” Taxon: Pinnularia viridula (Kiitzing) Rabenhorst 1853, p. 43. Taxonomic Comments: This is a new combination of Frustulia viridula Kiitzing 1833. This does not appear to represent type material. Exsiccatae Number: 683 Year Distributed: 1858 Locality Information: “Marais de bois, Falaise” Taxon: Navicula major var. crassa Brébisson ex Rabenhorst 1858a Taxonomic Comments: Van Landingham’s Catalogue (1975, p. 2658) lists the basionym of this taxon as “Pinnularia major var. crassa Rabenhorst 1848-1860; Rabenhorst 1864,” with the combination in Navicula made by M. Peragallo (1903). This name appears in the exsiccatae in Navicula; transfer to Pinnularia was effected by Rabenhorst (1864a, p. 210). Exsiccatae Number: 687 Year Distributed: 1858 Locality Information: “. . . se trouve en groupes assez épais sur les confervées. Falaise.” Taxon: Synedra aggregata Brébisson in Rabenhorst 1858a Publication Notes: Exsiccatae provides a description in Latin of this taxon. KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 259, Exsiccatae Number: 688 Year Distributed: 1858 Locality Information: “Dresden” Taxon: Himantidium dilatatum Wigand 1860, p. 43 Exsiccatae Number: 743 Year Distributed: 1858 Locality Information: “Auf Vaucheria clavata in der Ober-Léssnitz bei Dresden im sogen ‘ Walter’s Grund’” Taxon: Gomphonema capitatum var. gracile Rabenhorst 1858b Taxonomic Comments: A very brief description in German is included with the exsiccatae. Name not included in Van Landingham’s Catalogue. Exsiccatae Number: 761 Year Distributed: 1858 Locality Information: “Im Torfmoor am Titisee . . im badische Schwarzwalde” Taxon: Frustulia torfacea A. Braun ex Rabenhorst 1853, p. 50 Taxonomic Comments: Specific epithet listed as “torphacea” in exsiccatae. Exsiccatae Number: 765 Year Distributed: 1858 Locality Information: “Falaise” Taxon: Amphora ovalis f. nana Rabenhorst 1864a, p. 92. The exsiccatae entry says “Amphora ovalis (Ehrbg.) Ktz.” Taxonomic Comments: The index to Rabenhorst (1864a, p. 327) indicates the name as “var. nana.” Exsiccatae Number: 781 Year Distributed: 1859 Locality Information: “Toofmoor bei Pohlenz, Oct. 1858” Taxon: Nitzschia arcus Bulnheim 1859, p. 22 Exsiccatae Number: 801 Year Distributed: 1859 Locality Information: “Graben in Peissengrunde vor Neider-Hasslich bei Dresden” Taxon: Cylindrotheca Rabenhorst 1859a Publication Notes: This genus is described and figured in the exsiccatae. Exsiccatae Number: 801 Year Distributed: 1859 Locality Information: “Graben in Peissengrunde vor Neider-Hasslich bei Dresden” Taxon: Cylindrotheca gerstenbergei Rabenhorst 1859a. Publication Notes: This species is described and figured in the exsiccatae. Taxonomic Comments: This is the generitype for the new genus Cylindrotheca. Exsiccatae Number: 802 Year Distributed: 1859 Locality Information: “auf Elbschlamm bei Dresden” Taxon: Navicula reinickeana Rabenhorst 1859a Publication Notes: A short description is provided with the exsiccatae. See also Rabenhorst 1863a, p. 35 for a description of this taxon. 260 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Exsiccatae Number: 803 Year Distributed: 1859 Locality Information: “St. Gallen, in einem Brunnentrog im Klosterhof, den 15 January 1859” Taxon: Cymbella variabilis Wartmann in Rabenhorst 1859a Publication Notes: Description in German provided with the exsiccatae Exsiccatae Number: 811 Year Distributed: 1859 Locality Information: “aus einem Feldbrunnen bei Peterwitz bei Strehlen in Schlesien” Taxon: Campylodiscus punctatus Bleisch 1860, p. 29 Exsiccatae Number: 812 Year Distributed: 1859 Locality Information: “Strehlen in Schlesien, Ohlaufluss” Taxon: Stauroptera truncata Rabenhorst 1853, pl. 9, fig. 12 Exsiccatae Number: 841 Year Distributed: 1859 Locality Information: “In einer Spalte des Chlorischieferfelsens am Franzensberge in Briinn” Taxon: Navicula nodosa f. curta Rabenhorst 1864a, p. 207 Publication Notes: This name is listed as a “variety” in the index (p. 338). Rabenhorst lists Navicula quinquerodes Grunow 1860 as a synonym. It appears that Rabenhorst’s is a new name for Grunow’s taxon. Exsiccatae Number: 842 Year Distributed: 1859 Locality Information: “mit Sphagnum umgiirteten Tiimpfel auf dem Eulengebirge bei Reichenbach in Schlesien” Taxon: Pinnularia rabenhorstii Hilse 1860b, p. 82 Taxonomic Comments: Van Landingham (1978a, p. 3302) reports the specific epithet as “raben- horstiana (Hilse in Rabenhorst 1859; Hilse 1960 [sic]) M. Peragallo 1903 p. 723.” Exsiccatae Number: 847 Year Distributed: 1859 Locality Information: “Petersdorf bei Gleiwitz in Schlesien . . .” [Poland] Taxon: Pleurostauron acutum (W. Smith) Rabenhorst 1859c, Plate 1, fig. B, Plate 2, fig. F Taxonomic Comments: Rabenhorst 1864a (p. 259) indicates this is a new combination for Smith’s taxon. This does not appear to represent type material. Exsiccatae Number: 848f Year Distributed: 1859 Locality Information: “Um Gleiwitz in Schlesien .. . .” [Poland (Katowice) Gliwice] Taxon: Stauroneis janischii Rabenhorst 1859b Publication Notes: A description in German and figure are provided with the exsiccatae. Exsiccatae Number: 943 Year Distributed: 1860 Locality Information: “Dresden, in der Oberléssnitz in Mai 1859” Taxon: Nitzschia hantzschiana Rabenhorst 1860a, p. 40 Publication Notes: The exsiccatae also includes a brief description. It is also described in Grunow (1862, p. 576) KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 261 Exsiccatae Number: 944 Year Distributed: 1860 Locality Information: “in einem Bache bei Tharandt (Sachsen)” Taxon: Nitzschia clausii Hantzsch 1860b, p. 40 Publication Notes: The exsiccatae also includes a brief description. Descriptions of this taxon can also be found in Grunow (1862, p. 573, 559) and Rabenhorst (1863a, p. 48). Exsiccatae Number: 945 Year Distributed: 1860 Locality Information: “Oberléssnitz bei Dresden, in einem Wasserbassin auf Steinem im September 1859” Taxon: Nitzschia media Hantzsch 1860b, p. 40 Publication Notes: The exsiccatae also includes a brief description. Grunow (1862, p. 576) also provides a description for this taxon. Exsiccatae Number: 946 Year Distributed: 1860 Locality Information: “in einem Tiimpel bei Mortizburg bei Dresden im September 1859” Taxon: Nitzschia gracilis Hantzsch 1860b, p. 40 Publication Notes: The exsiccatae also includes a brief description. See Grunow (1862, p. 560, 575) for a description of this taxon. Exsiccatae Number: 947 Year Distributed: 1860 Locality Information: “Oberléssnitz bei Dresden auf Wurzeln und Steinem in einem Wasserbassin” Taxon: Navicula macrogongyla Rabenhorst 1860a, p. 40 Taxonomic Comments: The exsiccatae does not indicate the specific epithet of the new Navicula, but it is given in the publication. Exsiccatae Number: 949 Year Distributed: 1860 Locality Information: “In Graben um Dreden haufig von Gerstenberger” Taxon: Nitzschia communis Rabenhorst 1860b Publication Notes: See Grunow (1862, p. 561, 578) and Rabenhorst (1864a, p. 159) for descrip- tions of this taxon. Taxonomic Comments: There is reference to an effectively published illustration, (Hedwigia 1860, pl. 6, fig. 3). It appears from the text of the exsiccatae that this is to be a new name for a diatom identified by Gerstenberger (in Hedwigia) as Synedra frustulium Kitzing. Exsiccatae Number: 950 YearDistributed: 1860 Locality Information: “Am Galgenberge bei Strehlen in Schlesien” Taxon: Nitzschia minuta Bleisch in Rabenhorst 1860b Taxonomic Comments: A description accompanies the exsiccatae. See also Bleisch 1863, p. 78 and Grunow 1862, p. 578. Exsiccatae Number: 951 Year Distributed: 1860 Locality Information: “in einem einzigen Brunnen bei Strehlen an der Strasse nach Striege” 262 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Taxon: Pinnularia gibba f. curta Bleisch in Rabenhorst 1860b Taxonomic Comments: A brief description is provided with the exsiccatae. The exsiccatae packet lists this taxon as “Pinnularia gibba forma curta, varietas nova”. This name is not listed in Van Landingham’s Catalogue as either a variety or a form of P. gibba. Exsiccatae Number: 952 Year Distributed: 1860 Locality Information: “in einem Brunnen an der Strasse von Riegersdorf nach Strehlen” Taxon: Pinnularia medioconstricta Bleisch in Rabenhorst 1860b Publication Notes: A description is provided with the exsiccatae. See also Bleisch 1863, p. 81. Exsiccatae Number: 953 Year Distributed: 1860 Locality Information: “in einem Graben auf dem Riichen der Eule . . .October 1859” Taxon: Pinnularia hilseana Janisch in Rabenhorst 1860b Publication Notes: A description is provided with the exsiccatae. See also Hilse 1860b, p. 82. Exsiccatae Number: 954 Year Distributed: 1860 Locality Information: “Ohlaufluss bei Strehlen in Schlesien unter Oscillarien; im Juli 1859” Taxon: Pinnularia silesiaca Bleisch in Fresenius 1862 Publication Notes: See also Bleisch 1863, p. 81 and Bleisch in Rabenhorst 1863a, p. 38. Taxonomic Comments: A description is lacking in the exsiccatae. Exsiccatae Number: 958 Year Distributed: 1860 Locality Information: “im Zitiricher See bei der Badeanstalt” Taxon: Encyonema prostratum f. helvetica-robustior Rabenhorst 1860b Taxonomic Comments: The name is listed as “Encynonema prostratum forma helvetica robustior. ” A short description is provided, and reference to a previous description and figures qualifies as a valid publication for this name. Name not included in Van Landingham’s Catalogue. Exsiccatae Number: 962 Year Distributed: 1860 Locality Information: “Steinbriichen am Galgenberge bei Strehlen” Taxon: Stauroneis cohnii Hilse in Rabenhorst 1860b Taxonomic Comments: A brief description is included with the exsiccatae. See also Hilse 1860b, p. 83. Exsiccatae Number: 963 Year Distributed: 1860 Locality Information: “in altern Steinbriichen am Galgenberge bei Strehlen” Taxon: Stauroneis undulata Hilse 1860, p. 83 Taxonomic Comments: A brief description is included with the exsiccatae. Exsiccatae Number: 984 Year Distributed: 1860 Locality Information: “In einem Sumpfe in Kaitzer Grunde bei Dresden” Taxon: Nitzschia tryblionella Hantzsch in Rabenhorst 1860c Publication Notes: A description is included with the exsiccatae. KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 263 Exsiccatae Number: 1003 YearDistributed: 1861 Locality Information: “in einem Quellwasser bei Salem” Taxon: Achnanthidium jackii Rabenhorst 1861la Publication Notes: Latin description provided with the exsiccatae. This name is also proposed in Rabenhorst 1864a, p. 106, but the description differs significantly enough from the exsiccatae that two separate species might be described under the same name from the same material. Exsiccatae Number: 1021b Year Distributed: 1861 Locality Information: “[in den Mergelgruben von] Peterwitz bei Strehlen in Schlesien” Taxon: Epithemia goeppertiana Hilse 1860, p. 79 Publication Notes: This diatom is described in the exsiccatae under the number 1021 and listed under 1021b. Exsiccatae Number: 1022 Year Distributed: 1861 Locality Information: “von Holmachern zur Erlangung von Trinkwasser angelegten Rinne auf dem Riicken der Eule (bei Reichenbach in Schlesien)” Taxon: Cyclotella hilseana Rabenhorst 1864a, p. 33 Taxonomic Comments: This was originally listed with the exsiccatae as Cyclotella dubia Hilse in Rabenhorst, a name also published in Hilse 1860, p. 79. A note with the exsiccatae indicates “Ob die Pflanze gar ein neues Genus bildet, wie ich sast vermuthe, dtirste wohl die Zukunst Ichren.” Exsiccatae Number: 1023 Year Distributed: 1861 Locality Information: “Pfingsten 1860 auf dem Riicken der Eule in einer h6he von 3000 F” Taxon: Pinnularia sudetica Hilse 1860, p. 82 Taxonomic Comments: This diatom is described in the exsiccatae. Van Landingham (1978a, p. 3324) lists this taxon as “Navicula sudetica Hilse 1861 p. 82; Hilse in Rabenhorst 1861—1879 No. 1023” and “Pinnularia sudetica (Hilse) M. Peragallo 1903, p. 725”. Krammer (2000) des- ignated this material as the lectotype of Hilse’s taxon. Exsiccatae Number: 1024 Year Distributed: 1861 Locality Information: “Brunnenabfluss von Katschwitz bei Strehlen” Taxon: Synedra campyla Hilse 1860, p. 84 Publication Notes: A brief description is also provided with the exsiccatae. Exsiccatae Number: 1025 Year Distributed: 1861 Locality Information: “im Ziegengrunde bei Strehlen” Taxon: Pinnularia ovalis Hilse 1860, p. 82 Publication Notes: A description is also provided with the exsiccatae. Exsiccatae Number: 1026 Year Distributed: 1861 Locality Information: “in den Mergelgruben von Gross-Lauden bei Strehlen” Taxon: Epithemia intermedia Hilse 1860, p. 76 264 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Publication Notes: A description is also provided with the exsiccatae. Taxonomic Comments: This name is an earlier homonym of Epithemia intermedia Fricke 1904 Exsiccatae Number: 1041 Year Distributed: 1861 Locality Information: “Dresden, im artefischen Brunnen” Taxon Fragilaria mesolepta Rabenhorst 1861b Publication Notes: Exsiccatae provides Latin description. Exsiccatae Number: 1045 Year Distributed: 1861 Locality Information: “in Graben bei Canstatt” Taxon: Surirella suevica Zeller in Rabenhorst 1861b Publication Notes: Exsiccatae provides Latin and German descriptions. See Krammer & Lange- Bertalot 1987, p. 89. Exsiccatae Number: 1046 Year Distributed: 1861 Locality Information: “Gegend von Marburg” Taxon: Himantidium striatum Wigand 1860, p. 43 Exsiccatae Number: 1046 Year Distributed: 1861 Locality Information: “Gegend von Marburg” Taxon: Synedra arcuatum Wigand 1860, p. 43 Taxonomic Comments: This name is an earlier homonym for Synedra arcuatum Cleve-Euler 1953, p. 68. Exsiccatae Number: 1065 Year Distributed: 1861 Locality Information: “Bochla pr. Grossenhain (Saxoniae) in fossis pratorum” Taxon: Cymbella naviculiformis Auerswald ex Heiberg 1863 Publication Notes: No description provided in exsiccatae. Taxonomic Comments: See the interpretation of the nomenclature of this taxon by Reimer (pp. 31-32 in Patrick & Reimer 1975). Exsiccatae Number: 1066 Year Distributed: 1861 Locality Information: “Um Leipzig, in der Elster und im Teiche bei Barneck” Taxon: Pleurosigma gracilentum Rabenhorst 1861b Publication Notes: A short description is provided with the exsiccatae. See also Rabenhorst 1863a, p. 617 and Rabenhorst 1864a, p. 240. Taxonomic Comments: Van Landingham (1978a, p. 3375) lists this as “Rabenhorst 1861—1879, No. 166”. Exsiccatae Number: 1084 Year Distributed: 1861 Locality Information: “St. Gallen, in einem Brunnentrog des Zweibriicher Tobels” Taxon: Odontidium hiemale f. rotundata Rabenhorst 1864a, p. 116 Taxonomic Comments: This taxon was listed with the exsiccatae as “Odontidium mesodon Kitz. KOCIOLEK: DIATOM TYPES IN RABENHORST?’S DIE ALGEN EUROPA’S 265 form!” Van Landingham (1978a, p. 3154) indicates the basionym of this taxon is O. rotundatum Rabenhorst 1853 (p. 34, plate 2, fig. 4c), although this name is used in the figure legends for plate 2, fig. 4c, the listing of this taxon name on page 34 is under O. hyemale. The name O. rotundatum Rabenhorst is a later homonym of O. rotundatum (Ehrenb.) Kiitz.1849. Exsiccatae Number: 1088 Year Distributed: 1861 Locality Information: “in einem kleinen Teiche bei St. Gallen (Schweiz)” Taxon: Epithemia rabenhorstii Wartmann in Rabenhorst 1861c Publication Notes: A brief description is provided with the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 1089 Year Distributed: 1861 Locality Information: “bei St. Gallen in einem kleinen Teiche auf Miihlegg” Taxon: Epithemia zebra var. intermedia Wartmann in Rabenhorst 1861c Publication Notes: A brief description is provided with the exsiccatae. Taxonomic Comments: This name is also presented in Rabenhorst 1864a, p. 65. This name is not included in Van Landingham’s Catalogue. It is an earlier homonym for the combination made by Hustedt (1934). Exsiccatae Number: 1101 Year Distributed: 1861 Locality Information: “Am Elbufer im Gehege bei Dresden” Taxon: Pleurosigma scalproides Rabenhorst 1861d Publication Notes: Description and illustration provided in exsiccatae. Exsiccatae Number: 1104 Year Distributed: 1861 Locality Information: “Elbufer im grossen Gehege bei Dresden” Taxon: Diatoma gracillimum Hantzsch in Rabenhorst 1861d Publication Notes: Description and illustration are provided in the exsiccatae. Taxonomic Comments: The specific epithet was originally listed as “gracillinum’”, but this should probably be considered a typographical error. Van Landingham’s Catalogue (1969, p. 1295) lists the specific epithet as “gracillimum”’. Exsiccatae Number: 1106 Year Distributed: 1861 Locality Information: “Walther’s Grund in der Oberléssnitz” Taxon: Pinnularia peracuminata Krammer 2000 Exsiccatae Number: 1107 Year Distributed: 1861 Locality Information: “Dresden, im artefischen Brunnen” Taxon: Cocconema pachycephalum Rabenhorst 1861d Publication Notes: Description and illustrations provided in exsiccatae. Rabenhorst (1863a, p. 616) had created the superfluous name Cocconema robustum for this taxon. Krammer (2002) has recently renamed this taxon Cymbella hantzschiana Krammet. Exsiccatae Number: 1114 Year Distributed: 1861 266 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Locality Information: “Oberléssnitz bei Dresden” Taxon: Nitzschia tumida Hantzsch in Rabenhorst 186le Publication Notes: A specimen is figured with the exsiccatae. Mahoney & Reimer (1997) indicate this species is described in Rabenhorst 1864a (p. 153). Exsiccatae Number: 1161 Year Distributed: 1861 Locality Information: “Strehlen in Schlesien . . . Jordansmiihle” Taxon: Pleurostaurum lineare Hilse 1860b, p. 83 Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1164 Year Distributed: 1861 Locality Information: “Im Marmorbruch zu Prieborn bei Strehlen in Schlesien” Taxon: Nitzschia perpusilla Rabenhorst 1861f Publication Notes: See also Rabenhorst 1864a, 159. Taxonomic Comments: A brief Latin description is included with the exsiccatae. Van Landingham (1978a, p. 3096) lists this taxon occurring in exsiccatae number 1163. Exsiccatae Number: 1166 Year Distributed: 1861 Locality Information: “in einem Graben bei Teppendorf in Schlesien” Taxon: Cymbella maculata var. major Rabenhorst 1864a, p. 80 Taxonomic Comments: According to Rabenhorst in comments accompanying the exsiccatae, this diatom was initially identified as Cymbella lunula (Ehrenb.) Rabenhorst 1853, p. 23. Van Landingham (1969, p. 1210) suggests the name Cymbella lunula Hilse in Rabenhorst was cre- ated in this exsiccatae number, but there is no indication from the exsiccatae that this combina- tion was made. Exsiccatae Number: 1167 Year Distributed: 1861 Locality Information: “Aus der Eule in einer HGhe von gegen 3000 Fuss, im October 1860” Taxon: Eunotia minuta Hilse 1860, p. 79 Publication Notes: A description is also provided with the exsiccatae. Exsiccatae Number: 1183 Year Distributed: 1861 Locality Information: “im stadtischen granitnen RGhrtroge zu Strehlen in Schlesien” Taxon: Stauroneis goeppertiana Bleisch ex Rabenhorst 1861g Publication Notes: A description is provided with the exsiccatae. See also Bleisch 1863, p. 81. ExsiccataeNumber: 1201 Year Distributed: 1861 Locality Information: “Dresden, Prinz Georg’s Garten 1861” Taxon: Cymatopleura nobilis Hantzsch 1860a, p. 36 Publication Notes: Illustrations provided with exsiccatae. Exsiccatae Number: 1203 Year Distributed: 1861 Locality Information: “Krippengrund 1.d. sachs. Schweiz” KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 267 Taxon: Pinnularia rupestris Hantzsch ex Rabenhorst 1861h Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1208 Year Distributed: 1861 Locality Information: “in einem Quelle beim Géthe-Hauschen auf dem Giickelhahn in Thiiringer Walde” Taxon: Surirella thuringiacea Hantzsch in Rabenhorst 1861h Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1246 Year Distributed: 1862 Locality Information: “in einem Bach in Héskler bei Ziirich” Taxon: Cocconema variabile Cramer ex Rabenhorst 1862a Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1246 Year Distributed: 1862 Locality Information: “in einem Bach in Héskler bei Ztirich” Taxon: Cymbella vulgata Krammer 2002, p. 55 Exsiccatae Number: 1248 Year Distributed: 1862 Locality Information: “Miihlegg bei St. Gallen (Schweiz)” Taxon: Encyonema maximum Wartmann ex Rabenhorst 1862a Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1261 Year Distributed: 1862 Locality Information: “bei Gerbersdorf in Schlesien” Taxon: Cymbella minuta Hilse in Rabenhorst 1862b Publication Notes: See also Hilse 1863, p. 66. Taxonomic Comments: Krammer (1997, p. 54) points out the error made by Reimer (in Patrick & Reimer 1975, p. 42) about the exsiccatae number where this taxon was published. Exsiccatae Number: 1263 Year Distributed: 1862 Locality Information: “in einem Graben bei Hussinetz bei Strehlen” Taxon: Pinnularia viridis var. coerulescens Hilse in Rabenhorst 1862b Publication Notes: See also Hilse 1863, p. 68. Taxonomic Comments: A brief description is included in the exsiccatae. This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 1266 Year Distributed: 1862 Locality Information: “in einem Graben im Dorfe Hussinetz bei Strehlen” Taxon: Nitzschia thermalis var. minor Hilse in Rabenhorst 1862b Publication Notes: See also Hilse 1860, p. 67. Taxonomic Comments: Van Landingham (1978a, p. 3128) indicates this taxon was published by Hilse in 1860. 268 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Exsiccatae Number: 1267 Year Distributed: 1862 Locality Information: “in einem Graben hinter dem Walde von Dobergast bei Strehlen in Schlesien” Taxon: Nitzschia kuetzingiana Hilse in Rabenhorst 1862b Taxonomic Comments: Nitzschia kuetzingii Rabenhorst 1864 is the valid name for this taxon. Exsiccatae Number: 1268 Year Distributed: 1862 Locality Information: “in einem Graben am Teiche von Hussinetz bei Strehlen” Taxon: Stauroneis anceps f. gracilis Rabenhorst 1864a, p. 247 Exsiccatae Number: 1283 Year Distributed: 1862 Locality Information: “Dresden, in Prinz Georgs Garten” Taxon: Nitzschia recta Hantzsch in Rabenhorst 1862c Publication Notes: A brief diagnosis is provided in the exsiccatae. Taxonomic Comments: Van Landingham gives year of publication as “1861-1879”. Exsiccatae Number: 1283 Year Distributed: 1862 Locality Information: “Dresden, in Prinz Georgs Garten” Taxon: Navicula rotundata Hantzsch in Rabenhorst 1862c Publication Notes: A brief description is provided in the exsiccatae. Exsiccatae Number: 1301 Year Distributed: 1862 Locality Information: “am obersten Gradirhause Sulza” Taxon: Homoeocladia bulnheimiana Rabenhorst 1862d Publication Notes: A description of this taxon is provided with the exsiccatae. The species is also described in Rabenhorst 1863a, p. 617 and Rabenhorst 1864a, p. 167. Exsiccatae Number: 1321 Year Distributed: 1862 Locality Information: “Dresden, im artefischen Brunnen” Taxon: Cocconema nanum Hantzsch in Rabenhorst 1862e Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1322 Year Distributed: 1862 Locality Information: “Strehlen bei Dresden, in einem Graben” Taxon: Sphenella naviculoides Hantzsch in Rabenhorst 1862e Publication Notes: A description is provided with the exsiccatae. 292 Exsiccatae Number: 1323 Year Distributed: 1862 Locality Information: “Dresden, im Abfliisse des artefischen Brunnens” Taxon: Synedra amphirhynchus var. undulata Rabenhorst 1862e Publication Notes: See also Rabenhorst 1864a, p. 134. KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 269 Exsiccatae Number: 1381 Year Distributed: 1862 Locality Information: “Ruisseaux et fossés. Falaise” Taxon: Surirella crumena Brébisson ex Kiitzing 1849 Taxonomic Comments: Rabenhorst 1864a, p. 57 says No. 1381 is “specimen authen.”” Krammer in Krammer & Lange-Bertalot (1987, p. 96) designates this material as the lectotype for the name of the species. Exsiccatae Number: 1385 Year Distributed: 1862 Locality Information: “Bei Leipzig” Taxon: Encyonema auerswaldii Rabenhorst 1853a, p. 24 Exsiccatae Number: 1386 Year Distributed: 1862 Locality Information: “San Giuliano prope Genuam, ad sax leg. Dufour’ Taxon: Homoeocladia dufourii DeNotaris in DeNotaris & Baglietto 1871 9 Exsiccatae Number: 1403 Year Distributed: 1862 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlamm aus dem ostindis- chen Archipel” Taxon: Climacosphenia indica Hantzsch 1863, p. 18 Exsiccatae Number: 1403 Year Distributed: 1862 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlamm aus dem ostindis- chen Archipel” Taxon: Synedra pulcherrima Hantzsch 1863, p. 19 Exsiccatae Number: 1403 Year Distributed: 1862 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlarmm aus dem ostindis- chen Archipel” Taxon: Synedra formosa Hantzsch 1863, p. 19 Exsiccatae Number: 1403 Year Distributed: 1862 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlamm aus dem ostindis- chen Archipel” Taxon: Joxarium rostratum Hantzsch 1863, p. 19 Exsiccatae Number: 1403 Year Distributed: 1862 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlamm aus dem ostindis- chen Archipel” Taxon: Mastogloia interrupta Hantzsch 1863, p. 20 Exsiccatae Number: 1403 Year Distributed: 1862 270 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlamm aus dem ostindis- chen Archipel” Taxon: Nitzschia formica Hantzsch 1863, p. 20 Taxonomic Comments: Figure legend on p. 22 says “Tryblionella formica”’, and Hantzsch indicates this species is identical with Nitzschia tryblionella in Rabenhorst exsiccatae 984. Exsiccatae Number: 1403 Year Distributed: 1862 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlamm aus dem ostindis- chen Archipel” Taxon: Eupodiscus minutus Hantzsch 1863, p. 20 Exsiccatae Number: 1403 Year Distributed: 1862 Locality Information: “Ostindische Diatomeen. Von Algen und Meeresschlamm aus dem ostindis- chen Archipel” Taxon: Cocconeis heteroidea Hantzsch 1863, p. 20 Exsiccatae Number: 1405 Year Distributed: 1862 Locality Information: “im Polenzthale (sachs. Schweiz)” Taxon: Synedra bicurvata Biene ex Rabenhorst 1863b Publication Notes: There is a reference to an illustration (Nitzschia arcus Bulnheim in Hedwigia II, Tafel II, fig. 1) to validate the name. This taxon was described in Rabenhorst 1864a, p. 129. Exsiccatae Number: 1421 Year Distributed: 1863 Locality Information: “Kraussnitz bei Grossenhain in Sachsen, in einem moorigen Wiesengraben” Taxon: Surirella saxonica Auerswald in Rabenhorst 1863b Publication Notes: A Latin description is provided with the exsiccatae. See also Rabenhorst 1864a, Oy DS). Exsiccatae Number: 1441 Year Distributed: 1863 Locality Information: “aus eine Bache unweit Sennerei unterhalb Casaccio auf dem Kukmanier” Taxon: Cymbella elegans Cramer 1863, p. 65 Exsiccatae Number: 144] Year Distributed: 1863 Locality Information: “aus eine Bache unweit Sennerei unterhalb Casaccio auf dem Kukmanier” Taxon: Fragilaria |Fragillaria in exsiccatae] undulata Cramer 1863, p. 65 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “Meeralgen von Honduras” Taxon: Amphitetras cruciata Janisch & Rabenhorst 1863, p. 4 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “[Meeralgen] an den Kiiste von Honduras” KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S Poy) Taxon: Campyvlodiscus rabenhorstianus Janisch in Janisch & Rabenhorst 1863, p. 6 Publication Notes: This name was published earlier by Grunow (1862). Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “|Meeralgen] an den Kiiste von Honduras” Taxon: Amphitetras parvula Janisch & Rabenhorst 1863, p. 4 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “[Meeralgen] an den Ktiste von Honduras” Taxon: Climacosphenia linearis Janisch & Rabenhorst 1863, p. 6 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “[Meeralgen] an den Ktiste von Honduras” Taxon: Cocconeis kirchenpaueriana Janisch & Rabenhorst 1863, p. 7 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “[Meeralgen] an den Kiiste von Honduras” Taxon: Cocconeis flexella Janisch & Rabenhorst 1863, p. 7 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “[Meeralgen] an den Kiiste von Honduras” Taxon: Denticella ventricosa Janisch & Rabenhorst 1863, p. 8 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “[Meeralgen] an den Kiiste von Honduras” Taxon: Navicula bleischiana Janisch & Rabenhorst 1863, p. 9 Exsiccatae Number: 1481 YearDistributed: 1863 Locality Information: “[Meeralgen] an den Kiiste von Honduras” Taxon: Navicula janischiana Rabenhorst in Janisch & Rabenhorst 1863, p. 10 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “[Meeralgen] an den Kiiste von Honduras” Taxon: Navicula marina Janisch & Rabenhorst 1863, p. 10 Taxonomic Comments: Non Navicula marina Ralfs in Pritchard 1861 Exsiccatae Number: 1481 Year Distributed: 1863 Locality Information: “|Meeralgen] an den Kiiste von Honduras” Taxon: Synedra gomphonema Janisch & Rabenhorst 1863, p. 13 Exsiccatae Number: 1481 Year Distributed: 1863 272 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Locality Information: “[Meeralgen] an den Kiiste von Honduras” Taxon: Jessella hyalina Janisch & Rabenhorst 1863, p. 13 Exsiccatae Number: 1489 Year Distributed: 1863 Locality Information: “Brunnen bei Weiselwitz bei Strehlen” Taxon: Surirella diaphana Bleisch ex Rabenhorst 1863c Publication Notes: Short description provided with exsiccatae. See also Bleisch 1863, p. 78 and Rabenhorst 1864a, p. 55. Exsiccatae Number: 1492 Year Distributed: 1863 Locality Information: “Brunnen an der Miihle zu Toeppendorf bei Strehlen in Schlesien” Taxon: Diatoma elongatum var. abbreviatum Rabenhorst 1864a, p. 122 Exsiccatae Number: 1501 Year Distributed: 1863 Locality Information: “Am einem Blumentopfe, melcher in einem Warnhause langere Zeit in einem Napfe mit Wasser gestanden” Taxon: Epithemia minuta Hantzsch in Rabenhorst 1863d Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1503a Year Distributed: 1863 Locality Information: “In einer Cisterne, . . . in dem Chloritschieferfelsen des Franzensberges in Briinn” Taxon: Nitzschia dubia var minor Grunow 1862, p. 568 Exsiccatae Number: 1503a Year Distributed: 1863 Locality Information: “In einer Cisterne, . . . in dem Chloritschieferfelsen des Franzensberges in Briinn” Taxon: Nitzschia constricta f. minor Rabenhorst 1864a, p. 153 Exsiccatae Number: 1561 Year Distributed: 1863 Locality Information: “In dem kalten reissenden Wasser der Aare bei Interlaken” Taxon: Synedra frigida Zeller in Rabenhorst 1863e Publication Notes: A description is provided with the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 1651 Year Distributed: 1864 Locality Information: “Am Ziegenberg bei Husiant [sic ? = Hussinetz] in Schlesien” Taxon: Nitzschia bleischi Janisch in Bleisch 1863, p. 78 Publication Notes: This taxon is illustrated in the exsiccatae. Taxonomic Comments: Van Landingham (1978a, p. 3019) gives the publication as Rabenhorst “1848-1860”. Exsiccatae Number: 1700 Year Distributed: 1864 KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 2S Locality Information: “Dresden” Taxon: Pinnularia parvillisima Krammer 2000 Exsiccatae Number: 1701 Year Distributed: 1864 Locality Information: “In stagnis turfosis propre Neukirch Lusatiae superioris legit M. Rostock” Taxon: Cymbella cuspidata f maxima Rabenhorst 1864d Publication Notes: A brief description is provided in Latin. The name was presented originally as Cymbella cuspidata var. naviculiformis f. maxima. Exsiccatae Numbers: 1711 and 1712 Year Distributed: 1864 Locality Information: “An Pfahlen des Hafendammes von Ostende, griine incrustirende Ueber- zuge. Juli 1861. Grunow coll.” Taxon: Nitzschia incrustans Grunow 1862, p. 566 Publication Notes: Krammer & Lange-Bertalot (1988, p. 246) discuss the lectotype of this taxon. Exsiccatae Number: 1716 Year Distributed: 1864 Locality Information: “Incrustirend auf Pflanzenstengeln in Nildelta” Taxon: Amphora tumidula Grunow in Rabenhorst 1864a, p. 175 Publication Notes: A Latin description accompanies the exsiccatae. Exsiccatae Number: 1718 Year Distributed: 1864 Locality Information: “Timsah See (bittersalzhaltig) bei Ismailia, Nildelta” Taxon: Nitzschia schweinfurthii Grunow in Rabenhorst 1864a, p. 175 Publication Notes: A Latin description and “nov. spec.” accompany this name. This is the basionym of Nitzschia obtusa var. schweinfurthii (Grunow) Grunow in Cleve & Grunow 1880. Exsiccatae Number: 1725 Year Distributed: 1864 Locality: Austria, “In einem kleinen See am Ufer der Donau im Prater bei Wien” Taxon: Surirella gracilis (W. Smith) Grunow 1862 Publication Notes: See Krammer & Lange-Bertalot (1988, p. 189, plates 136, figures 1—4). Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia formica var. elongata Grunow 1865, p. 3 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia formica var. genuina Grunow 1865, p. 3 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia formica var. intermedia Grunow 1865, p. 3 274 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia formica var. bigibba Grunow 1865, p. 3 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia ventricosa var. elongata Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia camelus var. genuina Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia camelus var. didymodon Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia camelus var. denticulata Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia indica Grunow 1865, p. 5 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia indica var. ventralis Grunow 1865, p. 5 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia major var. sexundulata Grunow 1865, p. 5 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia major var. undulata Grunow 1865, p. 16, pl. 1, fig. 8 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia major var. octoundulata Grunow 1865, p. 5, pl. 1, fig. 8 Exsiccatae Number: 1727 Year Distributed: 1864 KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S POU Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia pectinalis var. undulata f. biundulata Grunow 1865, p. 4, pl. 1, fig. 5a Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “insel Banka (Ostindien)” Taxon: Eunotia pectinalis var. undulata f. quadriundulata Grunow 1865, p. 4, pl. 1, fig. 5b [PK confirmed] Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia pectinalis var. undulata f. octoundulata Grunow 1865, p. 4, pl. 1, fig. Sc Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Desmogonium rabenhorstianum Grunow 1865, p. 6 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Encyonema gerstenbergeri Grunow 1865, p. 9 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Gomphonema turris var. apiculatum Grunow 1865, p.10 Exsiccatae Number: 1765 Year Distributed: 1865 Locality Information: “In aquis frigidis prope Insbruck” Taxon: Odontidium alpigenum A.J. Kerner ex Rabenhorst 1865b Publication Notes: A Latin description is provided with the exsiccatae. Exsiccatae Number: 1802 Year Distributed: 1865 Locality Information: “im Brunnentroge des Geh6ftes in Sedlitiz bei Strehlen” Taxon: Cymbella silesiaca Bleisch in Rabenhorst 1865c. Publication Notes: A description is provided with the exsiccatae. Exsiccatae Number: 1804 Year Distributed: 1865 Locality Information: “Westgalizien, Jeziorki bei Chrzanow” Taxon: Nitzschia schliephackeana Grunow ex Rabenhorst 1864a, p. 324 Publication Notes: See also Rabenhorst 1866c, p. 56. Exsiccatae Number: 1806 Year Distributed: 1865 Locality Information: “Diatomaceen-Erde, Nottingham Maryland, U.S. (Bermuda Tripel aut.)” Taxon: Heliopelta ehrenbergii Eulenstein ex Rabenhorst 1865c 276 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Publication Notes: Though no formal description is offered, reference to previous descriptions, here in the form of three previous descriptions of Ehrenberg, suffice to validate this name. The name is, however, illegitimate inasmuch as Eulenstein should have used one of Ehrenberg’s epithets. Exsiccatae Number: 1891 Year Distributed: 1866 Locality Information: “In fossis humidis sylvae Sebaldianae (Reichsforst) prope Neuhof (Norim- bergae) in Franconia” Taxon: Nitzschia |Nitschia on exsiccatae label] franconica Reinsch in Rabenhorst 1866a Taxonomic Comments: A Latin description is part of the exsiccatae. Exsiccatae Number: 1891 Year Distributed: 1866 Locality Information: “In fossis humidis sylvae Sebaldianae (Reichsforst) prope Neuhof (Norim- bergae) in Franconia” Taxon: Nitzschia [Nitschia on exsiccatae label] franconica var. serpentina Reinsch in Rabenhorst 1866a Taxonomic Comments: A Latin description is part of the exsiccatae. Exsiccatae Number: 1895 Year Distributed: 1866 Locality Information: “in Tuberi fluminis locis leniter fluentibus infra Rubromontum in Franc” Taxon: Cocconeis striolata f. substriolata Reinsch in Rabenhorst 1866a Publication Notes: A description and illustrations accompany the exsiccatae. Exsiccatae Number: 1912 Year Distributed: 1866 Locality Information: “In per sylvis ductis fossis sempiterne humidis sylvae Sebaldianae (Reichsforst) infra ‘Kalkreuth’ in Franconia” Taxon: Melosira pfaffiana Reinsch in Rabenhorst 1866b Publication Notes: A Latin description and illustrations accompany the exsiccatae. See also Reinsch 1866, p. 11. Exsiccatae Number: 1933 Year Distributed: 1866 Locality Information: “Weisengraben unter Eis bei Eibenstock” Taxon: Denticula quadrata Kuntze in Rabenhorst 1866c Publication Notes: A description is provided with the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 1934 Year Distributed: 1866 Locality Information: “einer Bucht des salzigen See’s bei Halle” Taxon: Amphora bulnheimii Rabenhorst 1866c Publication Notes: A brief diagnosis is provided with the exsiccatae. Exsiccatae Number: 1998 Year Distributed: 1867 Locality Information: “Moler, . . . aus der Insel Mors in nérdlichen Jiitland” Taxon: Triceratium heibergianum Grunow 1866, p. 145 KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 277 Taxonomic Comments: Van Landingham (1978b, p. 4047) considers this diatom conspecific with T. heibergii Grunow in Van Heurck 1883. Exsiccatae Number: 1998 Year Distributed: 1867 Locality Information: “Molér, . . . aus der Insel Mors in nérdlichen Jiitland” Taxon: Triceratium jensenianum Grunow 1866, p. 145 Exsiccatae Number: 1998 Year Distributed: 1867 Locality Information: “Molér, . . . aus der Insel Mors in nérdlichen Jtitland” Taxon: Stephanogonia danica Grunow 1866, p. 146 Exsiccatae Number: 1998 Year Distributed: 1867 Locality Information: “Molér, . . . aus der Insel Mors in nérdlichen Jiitland” Taxon: Sceptroneis gemmata Grunow 1866, p. 146 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Podosphenia remulus Grunow 1867, p. 2 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Asterionella bleakeleyi var. notata Grunow 1867, p. 2 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Synedra undosa Grunow 1867, p. 4 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Synedra cuneata Grunow 1867, p. 5 Taxonomic Comment: Non Synedra cuneata Ehrenberg 1834 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Striatella intermedia Grunow 1867, p. 6 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Striatella lindigiana Grunow 1867, p. 6 Exsiccatae Number: 2000 Year Distributed: 1867 278 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Plagiodiscus Grunow & Eulenstein in Grunow 1867, p. 8 Publication Notes: This is the original publication of the genus name Plagiodiscus. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Plagiodiscus nervatus Grunow 1867, p. 8 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Berkeleya fusidium Grunow 1867, p. 17 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Nitzschia affinis f. major Grunow 1867, p.18 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Nitzschia kolaczekii Grunow 1867, p.18. Publication Notes: See also Grunow 1877, p. 173. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Bacillaria tropica Grunow 1867, p.19 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Rhoicosigma Grunow 1867, p. 19 Publication Notes: This is the original publication of the genus name Rhoicosigma. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Rhoicosigma reichardtianum Grunow 1867, p.19 Taxonomic Comment: This is equivalent to Rhoicosigma reichardtii Grunow 1877, p. 181. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Rhoicosigma reichardtianum var. constrictum Grunow 1877, p. 181. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Amphora decussata Grunow 1867, p. 23 KOCIOLEK: DIATOM TYPES IN RABENHORST?’S DIE ALGEN EUROPA’S 279 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Amphora hemisphaerica Grunow 1867, p. 24 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Amphora cymbelloides Grunow 1867, p. 24 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Navicula reticulata Grunow 1867, p. 26 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Navicula triundulata Grunow 1867, p. 27 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Rtickreife von Neu-Granada” Taxon: Schizostauron Grunow 1867, p. 28 Publication Notes: This is the original publication of the genus name Schizostauron. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Schizostauron lindigianum Grunow 1867, p. 28 Publication Notes: See also Grunow 1877. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Licmophora gracilis var. borealis Grunow 1867, p. 34 Taxonomic Comments: This name is actually a new combination for Rhipidophora borealis Kiitzing 1844 and should be presented as Licmophora gracilis var. borealis (Kiitzing) Grunow 1867; the Rabenhorst exsiccatae does not appear to be original Kiitzing material. Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: [sthmia lindigiana Grunow & Eulenstein in Grunow 1867, p. 29 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: The name was originally offered as “Triceratium (orbiculatum Shadbolt?) elongata” Gru- now 1867, p. 31. Validly published names from this are Triceratium elongatum Grunow 1867 or Triceratium orbiculatum var. elongatum (Grunow) Grunow 1877 280 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Exsiccatae Number: 2000 Year Distributed: 1867 Locality Information: “von Sargassum aus der Riickreife von Neu-Granada” Taxon: Licmophora gracilis var. constricta Grunow 1867, p. 34 Exsiccatae Number: 2005 Year Distributed: 1867 Locality Information: “Dives” [= Dives-sur-Mer, in Calvados] Taxon: Eupodiscus roperii Brébisson in Rabenhorst 1867 Publicaton Notes: A description is provided in the exsiccatae. See also Brébisson 1870, p. 41. Taxonomic Comments: The name is proposed as “roperii” though the correct spelling is “roperi.” This a nomen novum for Coscinodiscus ovalis Roper 1858, required for its transfer to Eupodiscus since priority exists for Eupodiscus ovalis Norman 1861. The Rabenhorst material does not appear to be original material for any of these names. Exsiccatae Number: 2012 Year Distributed: 1867 Locality Information: “Sallenelles (Calvados)” Taxon: Surirella ovata var. marina Brébisson in Rabenhorst 1867 Publication Notes: A description is provided with the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 2061 Year Distributed: 1868 Locality Information: “Wiesengraben bei Quartschen” Taxon: Cymatopleura marchica Hermann in Rabenhorst 1868 Publication Notes: A brief description is provided wth the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 2150 Year Distributed: 1870 Locality Information: “de la baie du M. St. Michel, pres d’ Avranches” Taxon: Amphiprora arenaria Brébisson in Rabenhorst 1870 Publication Notes: A description is provided with the exsiccatae. Van Landingham (1967, p. 160) gives question marks next to this name. He attributes it to Rabenhorst, but then indicates “Dec 217 according to Chase in Habirshaw (1885)” and “No. 2150 according to Mills (1934, p. 904)”; the entry actually occurs in Mills 1933, p. 129. Exsiccatae Number: 2171 Year Distributed: 1870 Locality Information: “Hab. ad plantas submersas Galliae, prope Falaise, imprimis ad folia Sphagnorum [sic]” Taxon: Peronia erinacea Brébisson & Arnott in Kitton 1868, p. 16 Taxonomic Comments: This is a nomen novum for Gomphonema fibula Brébisson ex Kiitzing 1849, p. 65 (=Peronia fibula (Brébisson ex Kiitzing) Ross). Peronia erinacea is an illegitimate name. Exsiccatae Number: 2172 Year Distributed: 1870 Locality Information: “Hab. in aquis submarinis Galliae: Dives, Trouville” Taxon: Navicula humerosa Brébisson ex Wm. Smith 1856, p. 93 KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 281 Exsiccatae Number: 2222 Year Distributed: 1870 Locality Information: “Ad thermas insulae Inarimes (Ischia)” Taxon: Denticula elegans f. [gamma] valida Pedicino 1867, p. 7 Publication Notes: No description accompanies the exsiccatae. Exsiccatae Number: 2261 Year Distributed: 1872 Locality Information: “Meeresgrundprobe, Carral bei Valdivia” Taxon: Navicula seminulum var. fragilarioides Grunow in Van Heurck 1880, plate 14, fig. 10 Exsiccatae Number: 2264 Year Distributed: 1872 Locality Information: “Auf Algen beim Leuchtthurme von Livorno” Taxon: Cocconeis pseudomarginata var. intermedia Grunow 1868, p. 13 Exsiccatae Number: 2264 Year Distributed: 1872 Locality Information: “Auf Algen beim Leuchtthurme von Livorno” Taxon: Cocconeis pellucida var. minor Grunow 1868, p. 13 Exsiccatae Number: 2312 Year Distributed: 1873 Locality Information: “Calcutta” Taxon: Nitzschia kurzeana Rabenhorst 1873 Publication Notes: A Latin description is provided with the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Reference is given to N. kurzii Rabenhorst ex Cleve & Moller #78. Exsiccatae Number: 2331 Year Distributed: 1873 Locality Information: “Im Felsen am Meeresufer bei Akyab (Haupstadt der Provinz Arracan)” Taxon: Podosira kurzii Zeller 1873, p. 175. Exsiccatae Number: 2357 Year Distributed: 1873 Locality Information: “Rangoon” Taxon: Eunotia kurzeana Grunow in Moller 1881 Taxonomic Comments: The entry in Rabenhorst (1873) as Eunotia kurziana is an invalid name because it was published without a description. Possible orthographic variant of E. kurzeana Grunow in Moller 1881. Exsiccatae Number: 2380 Year Distributed: 1874 Locality Information: “Kiiste von Japan” Taxon: Licmophora divisa f. japanica Rabenhorst 1874 Publication Notes: A brief description accompanies the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 2382 Year Distributed: 1874 282 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Locality Information: “Salzwasser Graben bei Kaschan (Pers. med.)” Taxon: Nitzschia [Nitzchia on label] parvula f. elongata-gracilis Hausknecht in Rabenhorst 1874 Publication Notes: A short description is provided with the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 2391 Year Distributed: 1874 Locality Information: “Grundprobe von Whampoa” Taxon: Triceratium sinense Schwarz 1874, p. 163 Exsiccatae Number: 2391 Year Distributed: 1874 Locality Information: “Grundprobe von Whampoa” Taxon: Triceratium whampoense Schwarz 1874, p. 163 Exsiccatae Number: 2481 Year Distributed: 1877 Locality Information: “Vera Cruz (zwischen Sertularien)” Taxon: Navicula lobata Schwarz in Rabenhorst 1877 Publication Notes: A Latin description and illustration are provided with the exsiccatae. Exsiccatae Number: 2483 Year Distributed: 1877 Locality Information: “Vera Cruz” Taxon: Navicula splendens Schwarz in Rabenhorst 1877 Publication Notes: A description is provided with the exsiccatae. This taxon is illustrated in the mate- rials accompanying No. 2481. Taxonomic Comments: This name is not listed in Van Landingham’s Catalogue Exsiccatae Number: 2560 Year Distributed: 1878 Locality Information: “vom Mathuri-Pass auf Neu-Zeeland, c. 3000; hoch ..” Taxon: Amphora berggrenii Cleve 1881, p. 4 Invalidly Published Names Exsiccatae Number: 1041 Year Distributed: 1861 Locality Information: “Dresden, im artefischen Brunnen” Taxon Fragilaria hantzschiana Gerstenberger in Rabenhorst 1861b Publication Notes: Exsiccatae lacks a description. Exsiccatae Number: 1249 Year Distributed: 1862 Locality Information: “Mitteloppaquelle auf dem Leiterberg im Gesenke (Mahren)” Taxon: Melosira distans f. articulis longioribus Rabenhorst 1862a Taxonomic Comments: Name not in Van Landingham’s Catalogue. No description or illustration associated with the exsiccatae. It is not clear whether Rabenhorst intended this to be a new taxon. Exsiccatae Number: 1489 Year Distributed: 1863 KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 283 Locality Information: “Brunnen bei Weisselwitz bei Strehlen” Taxon: Amphora abbreviata Bleisch in Rabenhorst 1864a, p. 55 Publication Notes: No description provided or reference to previous descriptions with exsiccatae. Van Landingham’s Catalogue (1967, p. 187) considers this name validly published in the exs- iccatae, but no description and/or illustration is provided. Exsiccatae Number: 1700 Year Distributed: 1864 Locality Information: “Dresden” Taxon: Pinnularia gibba f. minor Rabenhorst 1864b Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia pectinalis var. undulata f. triundulata Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia pectinalis var. undulata f. quinqueundulata Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia pectinalis var. undulata f. septemundulata Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Eunotia pectinalis var. undulata f. novemundulata Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Euvnotia pectinalis var. undulata f. sexundulata Grunow 1865, p. 4 Exsiccatae Number: 1727 Year Distributed: 1864 Locality Information: “Insel Banka (Ostindien)” Taxon: Gomphonema dichotomum var. affine f. minor Grunow 1865, p. 10 Exsiccatae Number: 1954 Year Distributed: 1867 Locality Information: “In den Torfgruben von Jentendorf und Quitzdorf, Kr. Rothenburg” Taxon: Frustulia saxonica f. turfacea Hilse Publication Notes: No description, illustrations or reference to other descriptions are provided with the exsiccatae. Taxonomic Comments: Van Landingham (1971, p. 1848) equates this taxon with Frustulia tur- facea A. Braun in Rabenhorst 1853, p. 50, yet no such relationship is indicated on the label exs- iccatae number 1954. 284 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Exsiccatae Number: 2025 Year Distributed: 1867 Locality Information: “Falaise” Taxon: Synedra lunaris var. elongata Brébisson in Rabenhrost Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Publication Notes: No description is provided in the exsiccatae. Exsiccatae Number: 2213 Year Distributed: 1870 Locality Information: “Eisenbahn-Ausstich bei Breslau” Taxon: Achnanthidium lanceolatum var. major Rabenhorst 1870 Publication Notes: No description accompanies the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 2223 Year Distributed: 1870 Locality Information: “Ad algas majores maris Neapolitani circa insulan Caprearum” Taxon: Cocconeis parthenopaea N.A. Pedicino in Rabenhorst 1870. Publication Notes: No description accompanying the exsiccatae. Exsiccatae Number: 2234 Year Distributed: 1870 Locality Information: “An der Ems unweit Emden” Taxon: Berkeleya dillwynii var. sericeum Eiben in Rabenhorst 1870 Publication Notes: No description accompanies the exsiccatae. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 2263 Year Distributed: 1872 Locality Information: “An der Kiiste der Insel S. Paul in der Stidsee, Expedition Novara” Taxon: Cocconeis grevillei var. minor Grunow in Rabenhorst 1872 Publication Notes: No description accompanies the exsiccatae. Exsiccatae Number: 2353 Year Distributed: 1873 Locality Information: “Calcutta, Balliaghatta Canal und saltlakes” Taxon: Nitzschia obtusa var. kurziana Rabenhorst 1873 Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. =~ Exsiccatae Number: 2353 Year Distributed: 1873 Locality Information: “Calcutta, Balliaghatta Canal und saltlakes” Taxon: Navicula calcuttensis Grunow in Rabenhorst 1873 Exsiccatae Number: 2353 Year Distributed: 1873 Locality Information: “Calcutta, Balliaghatta Canal und saltlakes” Taxon: Pleurosigma kurzianum Grunow in Rabenhorst 1873 Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 285 Exsiccatae Number: 2358 Year Distributed: 1873 Locality Information: “Rangoon-lake” Taxon: Nitzschia rangoonensis Grunow Publication Notes: No description accompanies the exsiccatae. Label is printed as 2558. Taxonomic Comments: This name is not included in Van Landingham’s Catalogue. Exsiccatae Number: 2551 Year Distributed: 1878 Locality Information: “Quilly-Le-Vicomte (France) Dr. Ch. Manoury” Taxon. Nitzschia palea f. curta Grunow Publication Notes: Van Landingham (1978, p. 3089) indicates this name is published in Cleve & Moller 1878, No. 124, but this name is invalidly published in that exsiccatae set because it lacks a decription. 286 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 PART 2. TAXONOMIC NAME REGISTER (A) VALIDLY PUBLISHED NAMES Taxon Name Exsiccatae Taxon Name Exsiccatae Achnanthidium jackii Rabenhorst 1003 Epithemia minuta Hantzsch 1501 Achnanthidium lanceolatum var. major Rabenhorst 2213 Epithemia rabenhorstii Wartmann 1088 Achnanthidium thermale Rabenhorst 561 Epithemia zebra var. intermedia Wartmann 1089 Amphiprora arenaria Brébisson 2150 Eunotia camelus var. denticulata Grunow 1727 Amphitetras cruciata Janisch & Rabenhorst 1481 Eunotia camelus var. didvymodon Grunow 1727 Amphitetras parvula Janisch & Rabenhorst 1481 Eunotia camelus var. genuina Grunow M2) Amphora berggrenii Cleve 560 Eunotia formica var. bigibba Grunow 1727 Amphora bulnheimii Rabenhorst 1934 Eunotia formica var. elongata Grunow 1727 Amphora cymbelloides Grunow 2000 Eunotia formica var. genuina Grunow 1727 Amphora decussata Grunow 2000 Eunotia formica var. intermedia Grunow 1727 Amphora hemisphaerica Grunow 2000 Eunotia indica Grunow WZ Amphora ovalis f. nana Rabenhorst 765 Eunotia indica var. ventralis Grunow 1727 Amphora tumidula Grunow 1716 Eunotia kurzeana Grunow in Moller 2357 Asterionella bleakeleyi var. notata Grunow 2000 Eunotia major var. undulata Grunow 1727 Bacillaria tropica Grunow 2000 Eunotia major var. sexundulata Grunow 1727 Berkeleya fusidium Grunow 2000 Eunotia major var. octoundulata Grunow 1727 Campylodiscus punctatus Bleisch 811 Eunotia minuta Hilse 1167 Campylodiscus rabenhorstianus Janisch in Janisch & Rabenhorst 1481 Eunotia pectinalis var. biundulata Grunow 1727 Climacosphenia indica Hantzsch 1403 Eunotia pectinalis var. undulata f. triundulata Grunow WZ Climacosphenia linearis Janisch & Rabenhorst 1481 Eunotia pectinalis var. undulata f. octoundulata Grunow 1727 Cocconeis flexella Janisch & Rabenhorst 1481 Eunotia ventricosa var. elongata Grunow 1727 Cocconeis heteroidea Hantzsch 1403 Eupodiscus minutus Hantzsch - 1403 Cocconeis kirchenpaueriana Janisch & Rabenhorst 1481 Eupodiscus roperi Brébisson 2005 Cocconeis pellucida var. minor Grunow 2264 Falcatella zelleri Rabenhorst 561 Cocconeis pseudomarginata var. intermedia Grunow 2264 Fragilaria mesolepta Rabenhorst 1041 Cocconeis striolata f. substriolata Reinsch 1895 Fragilaria undulata Cramer 1441 Cocconema nanum Hantzsch 1321 Frustulia torfacea A. Braun ex Rabenhorst 761 Cocconema pachycephalum Rabenhorst 1107 Gomphogramma rupestre (Kiitzing) A. Braun ex Rabenhorst 624 Cocconema variabile Cramer 1246 Gomphonema capitatum var. gracile Rabenhorst 743 Cyclotella dubia Hilse 1022 Gomphonema turris var. apiculatum Grunow 1727 Cyclotella hilseana Rabenhorst 1022 Heliopelta ehrenbergii Eulenstein 1806 Cylindrotheca Rabenhorst 801 Himantidium dilatatum Wigand 688 Cylindrotheca gerstenbergei Rabenhorst 801 Himantidium striatum Wigand 1046 Cymatopleura marchica Herrmann 2061 Homeocladia bulnheimiana Rabenhorst 1301 Cymatopleura nobilis Hantzsch 1201 Homoeocladia dufourii DeNotaris in DeNotaris & Baglietto 1386 Cymbella cuspidata f. maxima Rabenhorst 1701 Isthmia lindigiana Grunow & Eulenstein 2000 Cymbella elegans Cramer 1441 Licmophora divisa f. japonica Rabenhorst 2380 Cymbella hantzschiana Krammer 1107 Licmophora gracilis f. constricta Grunow 2000 Cymbella helvetica f. silesiaca Rabenhorst 604 Licmophora gracilis f. borealis Grunow 2000 Cymbella lunula (Ehrenberg) Rabenhorst 1166 Mastogloia interrupta Hantzsch 1403 Cymbella maculata var. major Rabenhorst 1166 Melosira pfaffiana Reinsch 1912 Cymbella minuta Hilse 1261 Melosira roeseana Rabenhorst 383 Cymbella naviculiformis Auerswald ex Heiberg 1065 Navicula bleischiana Janisch & Rabenhorst 1481 Cymbella silesiaca Bleisch 1802 Navicula humerosa Brébisson in Wm. Smith 2172 Cymbella variabilis Wartmann 803 Navicula janischiana Rabenhorst 1481 Cymbella vulgata Krammer 1246 Navicula lobata Schwarz 2481 Denticella ventricosa Janisch & Rabenhorst 1481 Navicula macrogongyla Rabenhorst 947 Denticula elegans var. valida Pedicino 2222 Navicula major var. crassa Brébisson 683 Denticula quadrata Kuntze 1933 Navicula marina Janisch & Rabenhorst 1481 Desmogonium rabenhorstianum Grunow in Rabenhorst 1727 Navicula nodosa f. curta Rabenhorst 841 Diatoma elongatum var. abbreviatum Rabenhorst 1492 Navicula reinickeana Rabenhorst 802 Diatoma gracillinum Hantzsch 1104 Navicula reticulata Grunow 2000 Encyonema auerswaldii Rabenhorst 1385 Navicula rotundata Hantzsch 1283 Encyonema gerstenbergeri Grunow 1727 Navicula seminulum var. fragilarioides Grunow 2261 Encyonema maximum Wartmann 1248 Navicula splendens Schwarz 2483 Encyonema prostratum f. helvetica-robustior Rabenhorst 958 Navicula triundulata Grunow 2000 Epithemia goeppertiana Hilse 1021 Nitzschia affinis f. major Grunow 2000 Epithemia intermedia Hilse 1026 Nitzschia arcus Bulnheim 781 KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S Taxon Name Nitzschia bleischi Janisch Niteschia clausii Hantzsch Nitzschia communis Rabenhorst Nitzschia constricta f. minor Rabenhorst Nitzschia dubia var. minor Grunow Nitzschia formica Hantzsch Nitzschia franconica Reinsch Nitzschia franconica var. serpentina Reinsch Nitzschia gracilis Hantzsch Nitzschia hantzschiana Rabenhorst Nitzschia incrustans Grunow Nitzschia kolaczekii Grunow Nitzschia kurzeana Rabenhorst Nitzschia kutzingiana Hilse Nitzschia media Hantzsch Nitzschia minuta Bleisch Nitzschia parvula f. elongata-gracilis Hausknecht Nitzschia perpusilla Rabenhorst Nitzschia recta Hantzsch Nitzschia schliephackeana Grunow Nitzschia schweinfurthii Grunow Nitzschia stagnorum Rabenhorst Nitzschia thermalis var. minor Hilse Nitzschia tryblionella Hantzsch Nitzschia tumida Hantzsch Odontidium alpigenum A.J. Kerner Odontidium hiemale f. rotundata Rabenhorst Odontidium rotundatum Rabenhorst Peronia erinaceae Brébisson & Arnott Pinnularia falaiseana Krammer Pinnularia gibba var. curta Bleisch Pinnularia hilseana Janisch Pinnularia medioconstricta Bleisch Pinnularia ovalis Hilse Pinnularia parvillisima Krammer Pinnularia peracuminata Krammer Pinnularia rabenhorstii Hilse Pinnularia rupestris Hantzsch Pinnularia silesiaca Bleisch in Fresenius Pinnularia sudetica Hilse Pinnularia viridis var. coerulescens Hilse Pinnularia viridula (Kiitzing) Rabenhorst Plagiodiscus Grunow & Eulenstein Plagiodiscus nervatus Grunow Pleurosigma gracilentum Rabenhorst Pleurosigma scalproides Rabenhorst Pleurostauron lineare Hilse Podosira kurzii Zeller Taxon Name Achnanthidium lanceolatum var. major Rabenhorst Amphora abbreviata Bleisch in Rabenhorst Berkeleya dillwynli var. sericeum Eiben Cocconeis grevillei var. minor Grunow Cocconeis parthenopaea Pedicino Eunotia pectinalis var. undulata f. quinqueundulata Grunow Eunotia pectinalis var. undulata f. septemundulata Grunow Eunotia pectinalis var. undulata f. novemundulata Grunow Eunotia pectinalis var. undulata f. quadriundulata Grunow Eunotia pectinalis var. undulata f. sexundulata Grunow Fragilaria hantzschiana Gerstenberger Exsiccatae 1651 944 949 1503 1503 1403 1891 1891 946 943 1711 (1712) 2000 2312 1267 945 950 2382 1164 1283 1804 1718 625 1266 984 1114 1765 1084 1084 2171 611 951 953 952 1025 1700 1106 842 1203 954 1023 1263 682b 2000 2000 1066 1101 1161 2331 (B) INVALID NAMES Exsiccatae 2213 1489 2234 2263 2223 1727 1727 1727 1727 1727 1041 287 Taxon Name Exsiccatae Podosphenia remulus Grunow 2000 Rhoicosigma Grunow 2000 Rhoicosigma reichardtianum Grunow 2000 Rhoicosigma reichardtianum vat. constrictum Grunow 2000 Sceptroneis gemmata Grunow 1998 Schizostauron Grunow 2000 Schizostauron lindigianum Grunow 2000 Sphenella naviculoides Hantzsch 1322 Stauroneis anceps f. gracilis Rabenhorst 1268 Stauroneis cohnii Hilse 962 Stauroneis goeppertiana Bleisch 1183 Stauroneis janischii Rabenhorst 848 Stauroneis rotaeana Rabenhorst 505 Stauroneis undulata Hilse 963 Stauroptera truncata Rabenhorst 812 Stephanogonia danica Grunow 1998 Striatella intermedia Grunow 2000 Striatella lindigiana Grunow 2000 Surirella crumena Brébisson ex Kutzing 1381 Surirella diaphana Bleisch 1489 Surirella gracilis Grunow 1725 Surirella intermedia Rabenhorst 642 Surirella ovata var. intermedia Rabenhorst 642 Surirella ovata var. marina Brébisson 2012 Surirella saxonica Auerswald 142] Surirella suevica Zeller 1045 Surirella thuringiacea Hantzsch 1208 Synedra aggregata Brébisson ex Rabenhorst 687 Synedra amphirhynchus var. undulata Rabenhorst 1323 Synedra arcuata Wigand 1046 Synedra bicurvata Biene 1405 Synedra campyla Hilse 1024 Synedra cuneata Grunow 2000 Synedra formosa Hantzsch 1403 Synedra frigida Zeller 1561 Synedra gomphonema Janisch & Rabenhorst 1481 Synedra interrupta Auerswald 403 Synedra pulcherrima Hantzsch 1403 Synedra splendens var. interrupta Rabenhorst 403 Synedra undosa Grunow 2000 Tessella hyalina Janisch & Rabenhorst 1481 Toxarium rostratum Hantzsch 1403 Triceratium (orbiculatum Shadbolt?) elongatum Grunow — 2000 Triceratium heibergianum Grunow 1998 Triceratium jensenianum Grunow 1998 Triceratium sinense Schwarz 2391 Triceratium whampoense Schwarz 2391 Taxon Name Exsiccatae Frustulia saxonica f. turfacea Hilse 1954 Gomphonema dichotomum var. affine f. minor Grunow 1727 Melosira distans f. articulis-longioribus Rabenhorst 1249 Navicula calcuttensis Grunow 2353 Nitzschia obtusa var. kurziana Rabenhorst 2353 Nitzschia palea f. curta Grunow 2551 Nitzschia rangoonensis Grunow 2358 Peronia erinacea Brébisson & Arnott in Kitton 2171 Pinnularia gibba f. minor Rabenhorst 1700 Pleurosigma kurzianum Grunow 2353 Synedra lunaris var. elongata Brébisson 2025 288 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 PART 3. TAXONOMIC STATUS OF NAMES PRESENTED AS NEw OR TYPIFIED IN RABENHORST EXSICCATAE Achnanthidium jackii Rabenhorst 1861a, no. 1003. The name is also proposed in Rabenhorst 1864a, p. 106, but the description differs significantly from the exsiccatae that two separate species might be described under the same name from the same material. Achnanthidium jackii Rabenhorst 1864a, p. 106. The name is also proposed in Rabenhorst 1861, no. 1003, but the description differs significantly from that of 1864 that two separate species might be described under the same name from the same material. Later homonym. Achnanthidium lanceolatum var. major Rabenhorst 1870, no. 2213. Invalidly published name. No description. Achnanthidium thermale Rabenhorst 1864a, p. 107. Amphiprora arenaria Brébisson in Rabenhorst 1870, no. 2150. Amphitetras cruciata Janisch & Rabenhorst 1863, p. 4; pl. 1, fig. 5. Amphitetras parvula Janisch & Rabenhorst 1863, p. 4; pl. 1, fig. 4. Amphora abbreviata Bleisch in Rabenhorst 1863c, no. 1489. Invalidly published name. No description. Amphora berggrenii Cleve 1881, p. 4, pl. 4, fig. 3. Name (with no description) in Cleve & MOller 1878, no. 90, and in Rabenhorst 1878, no. 2560. Amphora bulnheimii Rabenhorst 1866c, no. 1934. Amphora cymbelloides Grunow 1867, p. 24. Amphora decussata Grunow 1867, p. 23. Amphora hemisphaerica Grunow 1867, p. 24. Amphora ovalis f. nana Rabenhorst 1864a, p. 92. In index p. 327 as “var. nana”. Amphora tumidula Grunow in Rabenhorst 1864a, p. 175. Also described in no. 1716 (Rabenhorst 1864d). Asterionella bleakeleyi var. notata Grunow 1867, p. 2. Bacillaria tropica Grunow 1867, p. 19 (as Bacillaria (paradoxa var.?) tropica). Berkeleya dillwynii var. sericeum Eiben in Rabenhorst 1870, no. 2234. Invalidly published name. No descrip- tion. Berkeleya fusidium Grunow 1867, p. 17. Campylodiscus punctatus Bleisch 1860, p. 29; pl. 5, figs. 1-6. Campylodiscus rabenhorstianus Janisch in Janisch & Rabenhorst 1863, p. 6; pl. 1, figs. 6-7. Illeg. Nom. superfluous (C. ecclesianus Greville and C. fenestratus Greville listed as synonyms). Campylodiscus rabenhorstii Janisch ex Grunow 1862, p. 435. Also in Rabenhorst 1863a, p. 6, as Campylodiscus rabenhorstianus Janisch & Rabenhorst. Climacosphenia indica Hantzsch in Rabenhorst 1863a, p. 18; pl. 5, fig. 1. Climacosphenia linearis Janisch & Rabenhorst 1863, p. 6; pl. 2, fig. 2. Cocconeis flexella Janisch & Rabenhorst 1863, p. 7; pl. 1, fig. 11. Cocconeis grevillei var. minor Grunow ex Rabenhorst 1872, no. 2263. Invalidly published name. No descrip- tion. Cocconeis heteroidea Hantzsch in Rabenhorst 1863, p. 21; pl. 6 A, fig. 10. Cocconeis kirchenpauperiana Janisch & Rabenhorst 1863, p. 7; pl. 1, fig. 9. Cocconeis parthenopoea Pedicino in Rabenhorst 1870, no. 2223. Inv. Cocconeis pellucida var. minor Grunow 1868, p. 13; pl. 1, fig. 7. Cocconeis pseudomarginata var. intermedia Grunow 1868, p. 13; pl. 1, fig. 6. Cocconeis striolata f. substriolata P. Reinsch in Rabenhorst 1866a, no. 1895. Description and illustration with exsiccatae. See also Reinsch 1866, p. 150. Cocconema nanum Hantzsch in Rabenhorst 1862e, no. 1321. KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 289 Cocconema pachycephalum Rabenhorst 1861d, no. 1107, [4] figs. Cocconema variabile Cramer in Rabenhorst 1862a, no. 1246. Cyclotella dubia Hilse in Rabenhorst 1861a, no. 1022. Description also in Hilse 1860, p. 79. Cyclotella hilseana Rabenhorst 1864a, p. 33. New name for Cyclotella dubia Hilse in Rabenhorst 1861. Cylindrotheca Rabenhorst 1859a, no. 801. Cylindrotheca gerstenbergeri Rabenhorst 1859a, no. 801. Cymatopleura marchica Hermann in Rabenhorst 1868, no. 2061. Cymatopleura nobilis Hantzsch 1860a, p. 36. Illustration in Rabenhorst 1861h, no. 1201. Cymbella cuspidata f. maxima Rabenhorst 1864d, no. 1701. Cymbella cuspidata f. maxima Rabenhorst 1868, p. 414 (as [Cymbella cuspidata var. naviculiformis| f. max- ima). Name previously published as Cymbella cuspidata f. maxima in Rabenhorst 1864d, no. 1701. Cymbella cuspidata var. naviculiformis (Auerswald in Rabenhorst) Rabenhorst 1864a, p. 78. Basionym: Cymbella naviculiformis Auerswald in Rabenhorst 1861b. Cymbella elegans Cramer in Rabenhorst 1863a, p. 65. Cymbella hantzschiana Krammer 2002, p. 47. Cymbella helvetica f. silesiaca Rabenhorst 1857a, no. 604. Cymbella lunula (Ehrenberg) Rabenhorst 1853, p. 23; pl. 7, fig. 11. Basionym: Cocconema lunula Ehrenberg 1843. Cymbella maculata var. major Rabenhorst 1864a, p. 80. Rabenhorst cites as a synonym: “Cymbella lunula Hilse in Rabenhorst Alg. no. 1166”. Previously identified by Rabenhorst as Cymbella lunula (Ehr.) Rabenhorst in no. 1166 (1861). Cymbella minuta Hilse in Rabenhorst 1862b, no. 1261. Cymbella naviculiformis Auerswald in Rabenhorst 1861b, no. 1065. Invalidly published name. No descrip- tion. Cymbella naviculiformis Auerswald ex Heiberg 1863, p. 108; pl. 1, fig. 2. Cymbella silesiaca Bleisch in Rabenhorst 1865c, no. 1802. Cymbella variabilis Wartmann in Rabenhorst 1859a, no. 803. Cymbella vulgata Krammer 2002, p. 55. Denticella ventricosa Janisch & Rabenhorst 1863, p. 8; pl. 2, fig. 11. Denticula elegans var. valida Pedicino 1867, p. 7; pl. 1, figs. 42-45. Denticula quadrata Kuntze in Rabenhorst 1866c, no. 1933. Desmogonium rabenhorstianum Grunow 1865, p. 6; pl. 1, fig. 1. Diatoma elongatum var. abbreviatum Rabenhorst 1864a, p. 122. (= Diatoma elongatum v. ‘gamma’ Smith 1856). Diatoma gracillimum Hantzsch in Rabenhorst 1861d, no. 1104b + figs. a—b (‘gracillinum’ ) Encyonema auerswaldii Rabenhorst 1853, p. 24; pl. 7, Encyonema : fig. 2. Encyonema gerstenbergeri Grunow 1865, p. 9; pl. 1, fig. 11. Encyonema maximum Wartmann ex Rabenhorst 1862a, no. 1248. Encyonema prostratum f. helvetica-robustior Rabenhorst 1860b, no. 958. Epithemia goeppertiana Hilse 1860, p. 79 (Géppertiana). Also in Rabenhorst 1861, no. 1021. Epithemia géppertiana Hilse in Rabenhorst 1861, no. 1021. Epithemia intermedia Hilse 1860, p. 76. Also described in exsiccatae no. 1026a (Rabenhorst 1861). Epithemia intermedia Hilse in Rabenhorst 1861, no. 1026a. Epithemia intermedia Wartmann in Rabenhorst 1861c, no. 1089. Error cit. for Epithemia zebra var. interme- dia Wartmann in Rabenhorst. Epithemia minuta Hantzsch in Rabenhorst 1863d, no. 1501. 290 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Epithemia rabenhorstii Wartmann in Rabenhorst 1861c, no. 1088. Epithemia zebra var. intermedia Wartmann in Rabenhorst 1861c, no 1089. Eunotia camelus var. denticulata Grunow 1865, p. 4; pl. 1, fig. 6d. Listed in Van Landingham’s Catalogue (1969) as a new combination of Himanthidium denticulatum Brébisson in Kiitzing. Treated here as the name of a new species because Grunow expressed doubt (query) regarding the synonymy. Eunotia camelus var. didvmodon Grunow 1865, p. 4; pl. 1, fig. 6c (as Eunotia camelus var. ‘beta’ didymod- on). Eunotia camelus var. genuina Grunow 1865, p. 4; pl. 1, figs. 6a—b (as Eunotia camelus Ehrenb. var. ‘alpha’ genuina). Eunotia formica var. bigibba Grunow 1865, p. 3; pl. 1, figs. 2f-g (as Eunotia formica var. ‘delta’ bigibba). Eunotia formica var. elongata Grunow 1865, p. 3; pl. 1, figs. 2a—b (as Ewnotia formica Ehr. var. ‘alpha’ elon- gata). Eunotia formica var. genuina Grunow 1865, p. 3; pl. 1, figs. 2c—d (as Eunotia formica var. ‘beta’ genuina). Eunotia formica vat. intermedia Grunow 1865, p. 3; pl. 1, fig. 2e (as Eunotia formica var. ‘gamma’ interme- dia). Eunotia indica Grunow 1865, p. 5; pl. 1, figs. 7a—b. Eunotia indica var. ventralis Grunow 1865, p. 5; pl. 1, fig. 7¢ (as Eunotia indica var. ‘beta’ ventralis). Eunotia kurzeana Grunow in MOller 1881, Typ. pl. 1.2.2. Eunotia kurziana Grunow in Rabenhorst 1873, no. 2357. Invalidly published name. No description. Probable orthographic variant of Eunotia kurzeana Grunow. Eunotia kurziana Grunow in Cleve & Moller 1878, no. 129. Invalidly published name. No description. Eunotia major var. octoundulata Grunow 1865, p. 5; pl. 1, fig. 8. Illustrated as Eunotia major var. undulata. Eunotia major var. sexundulata Grunow 1865, p. 5. Invalidly published name. No description. See Ewnotia major var. undulata Grunow 1865. Eunotia major var. undulata Grunow 1865, p. 16; pl. 1, fig. 8, p. 5. Described in text p. 5 as var. sexundula- ta and var. octundulata. Eunotia minuta Hilse in Rabenhorst 1861f, no. 1167. Eunotia minuta Hilse 1860, p. 79. See also Rabenhorst 1861, no. 1167. Eunotia pectinalis f. biundulata Grunow 1865, p. 4; pl. 1, fig. 5a (as Eunotia pectinalis var. undulata f. biun- dulata). Invalidly published name. No description. Eunotia pectinalis f. octoundulata Grunow 1865, p. 4; pl. 1, fig. Se (as Eunotia pectinalis var. undulata f. octoundulata). Invalidly published name. No description. Eunotia pectinalis f. quadriundulata Grunow 1865, p. 4; pl. 1 (as Eunotia pectinalis var. undulata f. quadri- undulata). Invalidly published name. No description. Eunotia pectinalis f. quinqueundulata Grunow 1865, p. 4 (as Eunotia pectinalis var. undulata f. sexundulata). Invalidly published name. No description. Eunotia pectinalis f. septemundulata Grunow 1865, p. 4 (as Eunotia pectinalis var. undulata f. septemundu- lata). Invalidly published name. No description. Eunotia pectinalis f. sexundulata Grunow 1865, p. 4 (as Eunotia pectinalis var. undulata f. sexundulata). Invalidly published name. No description. Eunotia pectinalis var. triundulata Grunow 1865, p. 4. Invalidly published name. No description. Eunotia ventricosa var. elongata Grunow 1865, p. 4; pl. 1, fig. 4. Eupodiscus minutus Hantzsch in Rabenhorst 1863a, p. 21; pl. 6A, fig. 9. Eupodiscus roperi Brébisson in Rabenhorst 1867, no. 2005 (‘roperii’). New name for Coscinodiscus ovalis Roper 1858 (priority for Eupodiscus ovalis Norman 1861). Eupodiscus roperi Brébisson 1870, p. 41 (‘roperii’). Previously published in Rabenhorst 1867, no. 2005. Falcatella zelleri Rabenhorst 1857a, no. 561. Falcatella zelleri Rabenhorst 1864a, p. 139. Inv. Cited pro syn. Synedra zelleri Rabenhorst. Fragilaria hantzschiana Gerstenberger in Rabenhorst 1861b, no. 1041. Inv. In syn. Fragilaria mesolepta Rabenhorst. KOCIOLEK: DIATOM TYPES IN RABENHORST?’S DIE ALGEN EUROPA’S 291 Fragilaria mesolepta Rabenhorst 1861b, no. 1041. Also in Rabenhorst 1863a, p. 30. Fragilaria undulata Cramer 1863, p. 65; pl. 12, fig. 7. Frustulia saxonica f. turfacea (A. Braun ex Rabenhorst) Rabenhorst 1864a, p. 227 (as var. in index p. 333). Basionym: Frustulia torfacea A. Braun ex Rabenhorst 1853. Frustulia saxonica f. turfacea Hilse ex Rabenhorst 1867, no. 1954. Invalidly published name. No description.. Perhaps this is just a citation error for Frustulia saxonica var. torfacea (Braun ex Rabenhorst) Rabenhorst 1864. Frustulia torfacea A. Braun ex Rabenhorst 1853, p. 50; pl. 7, Frustulia: fig. 2. Epithet spelled “torphacea” in exsiccatae no. 761. Gomphogramma rupestre (Kiitzing) Braun in Rabenhorst 1853, p. 33, 67; pl. 9, fig. 1. Basionym: Denticula thermalis var. rupestris Kiitzing 1849. Gomphonema capitatum var. gracile Rabenhorst 1858, no. 743. Gomphonema dichotomum f. minor Grunow 1865, p. 10 (as Gomphonema dichotomum var. affine f. minor). Inv. Nomen nudum. Gomphonema turris var. apiculatum Grunow 1865, p. 10; pl. 2, fig. 12. Heliopelta ehrenbergii Eulenstein ex Rabenhorst 1865c, no. 1806. Illeg. Nomen novum. H. metii Ehrenb., H. leeuwenhockii Ehrenb. and H. euleri Ehrenb. given as synonyms. Himantidium dilatatum Wigand 1860, p. 43; pl. 7, fig. 9. Himantidium striatum Wigand 1860, p. 43; pl. 7, fig. 10-12. Homoeocladia bulnheimiana Rabenhorst 1862d, no. 1301. Also described in Rabenhorst 1863a, p. 617 and Rabenhorst 1864a, p. 167. Homoeocladia dufourii De Notaris in De Notaris & Baglietto 1871, Erbar. Critt. it. No. 764. Isthmia lindigiana Grunow & Eulenstein in Grunow 1867, p. 29. Licmophora divisa f. japonica Rabenhorst 1874, no. 2380. Licmophora gracilis f. borealis (Kiitzing) Grunow 1867, p. 35. Basionym: Rhipidophora borealis Kiitzing 1844. Licmophora gracilis f. constricta Grunow 1867, p. 34. Mastogloia interrupta Hantzsch in Rabenhorst 1863a, p. 20; pl. 6A, figs. Sa—b. Melosira distans f. articulis-longioribus Rabenhorst 1862a, no. 1249. Invalidly published name. No illustra- tion or description. Melosira pfaffiana Reinsch in Rabenhorst 1866b, no. 1912. Also in Reinsch 1866. Melosira roeseana Rabenhorst 1853, p. 13; pl. 10, supplemente: fig. 5. Melosira roeseana Rabenhorst 1854, no. 383. Description previously published. Navicula bleischiana Janisch & Rabenhorst 1863, p. 9; pl. 2, fig. 10. Navicula calcuttensis Grunow in Rabenhorst 1873, no. 2353. Invalidly published name. No description. Navicula humerosa Brébisson ex W. Smith 1856, p. 93. Navicula janischiana Rabenhorst in Janisch & Rabenhorst 1862, p. 10; pl. 2, fig. 15. Navicula lobata Schwartz in Rabenhorst 1877, no. 2481. Description in Latin and illustration are provided. Navicula macrogongyla Rabenhorst 1860a, p. 40; pl. 6, fig. 10. Navicula major var. crassa Brébisson in Rabenhorst 1858a, no. 683. Invalidly published name. No descrip- tion. Name of taxon validly published as Pinnularia major var. crassa Rabhenhorst 1864a, p. 210. Navicula marina Janisch & Rabenhorst 1863, p. 10; pl. 2, fig. 16. Illeg. Non Ralfs 1861. Navicula nodosa f. curta Rabenhorst 1864a, p. 207. As ‘var.’ in index p. 338. Syn: Navicula quinquenodis Grunow 1860. New name for Grunow’s taxon. Navicula reinickeana Rabenhorst 1859a, no. 802. Brief description provided. Description also in Rabenhorst 1863a, p. 35. 292 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Navicula reticulata Grunow 1867, p. 26. Navicula rotundata Hantzsch in Rabenhorst 1862c, no. 1283b. Navicula seminulum vat. fragilarioides Grunow in Van Heurck 1880, pl. 14, fig. 10. Navicula splendens Schwartz in Rabenhorst 1877, no. 2483. Illustration with no. 2481. Navicula triundulata Grunow 1867, p. 27. Nitzschia affinis f. major Grunow 1867, p. 18. Nitzschia arcus Bulnheim 1859, p. 22; pl. 2, fig. 1. Nitzschia bleischii Janisch in Bleisch 1863, p. 78. Nitzschia bleischii Janisch in Rabenhorst 1864c, No. 1651. Nitzschia clausii Hantzsch 1860b, p. 40; pl. 6, fig. 7. Illustration only. Brief description with exsiccatae no. 944 (1860). Description in Grunow 1862, p. 573, 559. Nitzschia communis Rabenhorst 1860b, no. 949. Reference to a published illustration (Hedwigia Il, pl. 6, fig. 3) is given. Described in Grunow 1862, p. 561, 578 and Rabenhorst 1864a, p. 159. Nitzschia constricta f. minor Rabenhorst 1863a, p. 153. Nitzschia dubia var. minor Grunow 1862, p. 568; pl. 28/12, figs. 24a—b. Nitzschia formica Hantzsch in Rabenhorst 1863a, p. 21; pl. 6 A, fig. 8. Nitzschia franconica Reinsch 1866, p. 26; pl. 1, figs. la, c-e. See also Rabenhorst 1866a, no. 1891. Nitzschia franconica var. serpentina Reinsch 1866, p. 27; pl. 1, figs. 1b-c. See also Rabenhorst 1866a, no. 1891. Nitzschia gracilis Hantzsch 1860b, p. 40; pl. 6, fig. 8. Illustration only. Brief description in exsiccatae no. 946. Description in Grunow 1862, p. 560, 575. Nitzschia hantzschiana Rabenhorst 1860a, p. 40; pl. 6, fig. 6. Brief description in exsiccatae no. 943. Description in Grunow 1862, p. 576. Nitzschia hantzschiana Rabenhorst in Grunow 1862, p. 576. Nitzschia incrustans Grunow 1862, p. 566; pl. 28/12, figs. 21a-f. Nitzschia kolaczekii Grunow 1867, p. 18. Spelled kolaizeckii in 1877. Nitzschia kolaizeckii Grunow 1877, p. 173. Orth. var. of Nitzschia kolaczekii Grunow 1867. Nitzschia kuetzingiana Hilse in Rabenhorst1862b, no. 1267. Nitzschia kuetzingii Rabenhorst 1864a, p. 160 [ref. 775]. Valid name for Nitzschia kuetzingiana Hilse in Rabenhorst 1862b, no. 1267. Nitzschia kurzeana Rabenhorst 1873, no. 2312. Nitzschia media Hantzsch 1860b, p. 40; pl. 6, fig. 9. Illustration only. Brief description in exsiccatae no. 945. Description in Grunow 1862, p. 576. Nitzschia minuta Bleisch in Rabenhorst 1860b, No. 950. Description also in Bleisch 1863, p. 78 and Grunow 1862, p. 578. Nitzschia minuta Bleisch 1863, p. 78. Nitzschia obtusa var. kurziana Rabenhorst 1873, no. 2353. Invalidly published name. No description. Nitzschia obtusa var. schweinfurthii (Grunow) Grunow in Cleve & Grunow 1880, p. 92. Basionym: Nitzschia schweinfurthii Grunow in Rabenhorst 1864. Nitzschia palea f. curta Grunow ex Rabenhorst 1878, no. 2551. Invalidly published name. No description. Nitzschia parvula f. elongata-gracilis Hausknecht in Rabenhorst 1874, no. 2382 (‘elongata gracilis’). Nitzschia perpusilla Rabenhorst 1861f, no. 1164. Also in Rabenhorst 1864a, p. 159, and Hilse 1863, p. 67. Nitzschia perpusilla Rabenhorst in Hilse 1863, p. 67. Previously validly published in Rabenhorst 1861f. Nitzschia rangoonensis Grunow ex Rabenhorst 1873, no. 2358. Invalidly published name. No description. Nitzschia recta Hantzsch ex Rabenhorst 1862c, No. 1283. Nitzschia schliephackeana Grunow ex Rabenhorst 1864a, p. 324. Nitzschia schweinfurthii Grunow in Rabenhorst 1864a, p. 175. Nitzschia stagnorum Rabenhorst 1857a, no. 625. Nitzschia thermalis var. minor Hilse in Rabenhorst 1862b, no. 1266. Name also in Hilse 1863, p. 67. Nitzschia thermalis var. stagnorum (Rabenhorst) Rabenhorst 1864a, p. 154. Basionym: Nitzschia stagnorum Rabenhorst 1857. KOCIOLEK: DIATOM TYPES IN RABENHORST’S D/E ALGEN EUROPA’S 298, Nitzschia tryblionella Hantzsch in Rabenhorst 1860c, No. 984. New name for 7ryblionella gracilis Smith 1853 (priority for Nitzschia gracilis Hantzsch 1860). Nitzschia tumida Hantzsch in Rabenhorst 186le, No. 1114. Illustration only. Description in Rabenhorst 1864a, pe 153. Odontidium alpigenum Kerner ex Rabenhorst 1865b, no. 1765. First appeared as nomen nudum in Hedwigia 4, p. 56 (1864). Odontidium hiemale (hyemale) f. rotundata (Rabenhorst) Rabenhorst 1864a, p. 116. Basionym: Odontidium rotundatum Rabenhorst 1853. Odontidium rotundatum Rabenhorst 1853, p. 34; pl. 2, fig. 4 c. Illeg. Non (Ehrenberg) Kiitzing 1849. The epi- thet appears in Rabenhorst 1853, on p. 34 (under O. hiemale) with an unclear status (new species of Odonthidium or new subdivision of O. hiemale?). Nevertheless, the name Odontidium rotundatum is defi- nitely validly published with the illustration and plate caption. Peronia erinacea Brébisson & Arnott ex Kitton 1868, p. 16. Illeg. Gomphonema fibula Brébisson ex Kiitzing 1849 cited as synonym. Name illegitimate, correct names for this taxon: Gomphonema fibula Bréb. ex Kiitzing, Peronia fibula (Bréb. ex Kiitzing) Ross. Pinnularia falaiseana Krammer 1992, p. 113-114, 172; pl. 41, figs. 5-8, 10-11. Pinnularia gibba var. curta Bleisch 1860, no. 951 (as Pinnularia gibba forma curta, varietas nova). Descrip- tion also in Rabenhorst 1863b, p. 39 (as Pinnularia gibba b. curta). Pinnularia gibba f. minor Rabenhorst 1864b, no. 1700. Invalidly published name. No description. Pinnularia hilseana Janisch in Rabenhorst 1860b, no. 953. See also Hilse 1860, p. 82. Pinnularia major var. crassa Rabenhorst 1864a, p. 210. Pinnularia medioconstricta Rabenhorst 1860b, no. 952 (‘medio constricta’). See also Bleisch 1863, p. 81. Pinnularia ovalis Hilse 1860, p. 82. Pinnularia ovalis Hilse in Rabenhorst 1861b, no. 1025. Pinnularia parvillisima Krammer 2000, p. 95. Pinnularia peracuminata Krammer 2000, p. 157. Pinnularia rabenhorstii Hilse 1860, p. 82. Pinnularia rupestris Hantzsch in Rabenhorst 1861h, no. 1203. Pinnularia silesiaca Bleisch in Fresenius 1862, p. 68; pl. 4, figs. 25—29. Pinnularia silesiaca Bleisch in Rabenhorst 1863a, p. 38. The exsiccatae (no. 954) does not provide a descrip- tion, but refers to an illustration (Hedwigia Il, pl. 4 figs. 3-4). See also Bleisch in Fresenius 1862. Pinnularia sudetica Hilse 1860, p. 82. Pinnularia sudetica Hilse in Rabenhorst 1861, no. 1023. Pinnularia viridis var. coerulescens Hilse in Rabenhorst 1862b, no. 1263. Also in Hilse 1863, p. 68. Pinnularia viridula (Kiitzing) Rabenhorst 1853, p. 43, 69; pl. 6, fig. 39. Basionym: Frustulia viridula Kitzing 1833 (1834). Plagiodiscus Grunow & Eulenstein in Grunow1867, p. 8. Plagiodiscus nervatus Grunow 1867, p. 8. Pleurosigma gracilentum Rabenhorst 1861b, no. 1066. Also in Rabenhorst 1863a, p. 617 and Rabenhorst 1864a, p. 240. Pleurosgima kurzianum Grunow ex Rabenhorst 1873, no. 2353. Invalidly published name. No description. Pleurosigma scalproides Rabenhorst 1861d, no. 1101 + fig. Also in Rabenhorst 1863a, p. 41, and Rabenhorst 1864a. Pleurostaurum acutum (W. Smith) Rabenhorst 1859c, p. 17; pl. 1, fig. B, pl. 2, fig. F. Basionym: Stauroneis acuta W. Smith 1853. Pleurostaurum lineare Hilse 1860, p. 83. Valid as the name of a new taxon. Could also be treated as a new combination of Stauroneis linearis Ehrenberg. Pleurostaurum lineare Rabenhorst 1861f, no. 1161. Valid as the name of a new taxon. Could also be treated as a new combination of Strauroneis linearis Ehrenberg. 294 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 13 Pleurostaurum lineare (Ehrenberg) Rabenhorst 1863a, p. 43. Basionym: Stauroneis linearis Ehrenberg 1843. Podosira kurzii Zeller 1873, p. 175 (= p. 2). Podosphenia remulus Grunow 1867, p. 2. Rhoicosigma Grunow 1867, p. 19. Rhoicosigma reichardtianum Grunow 1867, p. 19. Rhoicosigma reichardtii Grunow 1877, p. 181. Orth. var. of Rhoicosigma reichardtianum Grunow 1867. Rhoicosigma reichardtii var. constrictum Grunow 1877, p. 181. Sceptroneis gemmata Grunow 1866, p. 146. Schizostauron Grunow 1867, p. 28. Schizostauron lindigianum Grunow 1867, p. 28. Schizostauron lindigii Grunow 1877, p. 181; pl. 195, fig. 17. Orth. var. for Schizostauron lindigianum Grunow 1867. Sphenella naviculoides Hantzsch in Rabenhorst 1862, no. 1322. Stauroneis anceps f. gracilis Rabenhorst 1864a, p. 247. Stauroneis cohnii Hilse in Rabenhorst 1860b, no. 962. Stauroneis goeppertiana Bleisch ex Rabenhorst 1861g, No. 1183. Also in Bleisch 1863, p. 81. Stauroneis janischii Rabenhorst 1859b, No. 848f, fig. f. Stauroneis rotaeana Rabenhorst 1856, p. 103; pl. 13, fig. 7. Illustration only. For descriptions see Grunow 1860, p. 565 and Rabenhorst 1864a, p. 249. Latin description also included in exsiccatae no. 505 (1856). Stauroneis undulata Hilse 1860, p. 83. Brief description also in the exsiccatae Rabenhorst 1860b, no. 963. Stauroptera truncata Rabenhorst 1853, p. 49; pl. 9, fig. 12. Stephanogonia danica Grunow 1866, p. 146. Striatella intermedia Grunow 1867, p. 6. Striatella lindigiana Grunow 1867, p. 6. Surirella crumena Brébisson ex Kiitzing 1849, p. 38. In same publication, cited pro syn. Cyclotella menegh- iniana ‘beta’ major (p. 19). Surirella diaphana Bleisch in Rabenhorst 1863c, no. 1489. Also in Bleisch 1863, p. 78. Surirella gracilis (W. Smith) Grunow 1862, p. 144; pl. 7/10, fig. 11. Basionym: 7ryblionella gracilis W. Smith 1853. Surirella intermedia Rabenhorst 1857b, no. 642. Brief description provided. Described in Latin in Rabenhorst 1864a, p. 57 as Surirella ovata b. intermedia. Surirella ovata var. intermedia (Rabenhorst) Rabenhorst 1864a, p. 57, 345 (as Surirella ovata b. intermedia on p. 57, as var. intermedia on p. 345). Basionym: Surirella intermedia Rabenhorst 1857. New combina- tion if basionym is valid; valid name of a new taxon otherwise. Surirella ovata var. marina Brébisson in Rabenhorst 1867, no. 2012. Surirella saxonica Auerswald in Rabenhorst 1863b, no. 1421. Also described in Rabenhorst 1864a, p. 53. Surirella suevica Zeller in Rabenhorst 1861b, no. 1045. Surirella suevica Zeller ex Grunow in Schmidt et al. 1875, pl. 23, figs. 56-60. Name already published in Rabenhorst 1861b, no. 1045. Surirella thuringiaceae Hantzsch in Rabenhorst 1861h, no. 1208. Synedra aggregata Brébisson ex Rabenhorst 1858a, No. 687. Synedra amphirhynchus var. undulata Rabenhorst 1862a, no. 1323. A diagnosis is provided with the exsic- catae. Also Rabenhorst 1864a, p. 134. Synedra arcuata Wigand 1860, p. 44; pl. 7, fig. 15. KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 295 Synedra bicurvata Biene ex Rabenhorst 1863b, No. 1405. No description in exsiccatae, but reference to an illustration (Hedwigia II, pl. 2, fig. 1). Described in Rabenhorst 1864a, p. 120. Synedra bicurvata Biene ex Rabenhorst 1864a, p. 120. Synedra campyla Hilse 1860, p. 84. Synedra campyla Hilse in Rabenhorst 1861, no. 1024a. Synedra cuneata Grunow 1867, p. 5. Illeg. Non Ehrenberg 1834. Synedra formosa Hantzsch in Rabenhorst 1863a, p. 19; pl. 5, fig. 3. Synedra frigida Zeller in Rabenhorst 1863e, no. 1561. Synedra gomphonema Janisch & Rabenhorst 1863, p. 13; pl. 2, fig. 6. Synedra interrupta Auerswald in Rabenhorst 1855, No. 403. Synedra lunaris var. elongata purissima A. de Brébisson ex Rabenhorst 1867, no. 2025. Invalidly published name. No description. Synedra pulcherrima Hantzsch in Rabenhorst 1863a, p. 19; pl. 5, fig. 2. Synedra splendens var. interrupta (Auerswald in Rabenhorst) Rabenhorst 1864a, p. 134. Basionym: Synedra interrupta Auerswald in Rabenhorst 1855. Synedra undosa Grunow 1867, p. 4. Synedra zelleri Rabenhorst 1864a, p. 139. New combination of Falcatella zelleri? Tessella (Tessela) hyalina Janisch & Rabenhorst 1863, p. 13; pl. 2, fig. 13. Toxarium rostratum Hantzsch 1863, p. 19; pl. 5, fig. 4. Triceratium elongatum Grunow 1867, p. 31 (as Triceratium (orbiculatum var?) elongatum). Triceratium heibergianum Grunow 1866, p. 145. Triceratium jensenianum Grunow 1866, p. 145. Triceratium sinense Schwarz 1874, p. 163. Triceratium whampoense Schwarz 1874, p. 163. ACKNOWLEDGMENTS [| want to take this opportunity to thank Dr. Elizabeth Foutanier for her extensive work on the taxonomic status of taxa considered herein; also, Dr. Michele L. Aldrich who perused the entire manuscript in its penultimate typeset format. With a keen editorial eye, she uncovered both textu- al and formatting inconsistences that I had missed, thus giving me an opportunity to make changes before committing the whole to print. Drs. Robert Edgar and David Williams reviewed the paper in its penultimate form and offered many suggestions for its improvement. I am indebted to both for the care they took in their reviews. And, last but not least, thanks are due Dr. J. Thomas Dutro, Jr., U.S. Geological Survey [ret.], National Museum of Natural History, Washington, D.C., who generously assisted in arranging with the Department of Botany at the Museum to make available their copy of Rabenhorst’s 1864 edition of Flora Europaea Algarum from which I secured the copy of Rabenhorst’s portrait that is reproduced here. To all of the above, my sincere thanks with the caveat that I alone take responsibility for any errors of commission or omission that still exist. LITERATURE CITED BLEISCH, M. 1860. Ueber zwei Campylodiscus (C. costatus, C. spiralis). Hedwigia 2:29, pl. 5. BLeIscu, M. 1863. Uber einige in den Jahren 1856-1862 in der Gegend von Strehlen gefundene Diatomeen. Abhandlungen Schlesische Gesellschaft fur Vaterldndische Kulture, Abtheilung. fiir Naturwissenschaften und Medicin. Breslau 75-84. BREBISSON, A. de. 1870. Notes on some French Diatomaceae. Presented to the Club by Alph. de Brebisson, Corr. Mem. 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Monograph No. 13, Academy of Natural Sciences of Philadelphia. 213 pp. PeDICINO, N.A. 1867. Pochi studi sulle Diatomee viventi presso alcune terme dell’isola d’Ischia. Afti Accademia delle scienze fisiche e matematiche di Napoli 3:20, 2 pls. PepicINo, N.A. 1867. Pochi studi sulle Diatomee viventi presso alcune terme dell’Isola d’Ischia. Rediconto delle adunanze e de’lavori della Reale Accademia delle Scienze (Fisiche e Matematiche) di Napoli 6: 70-72. PERAGALLO, M. 1903. Le Catalogue général des Diatomées. Pp. 472-973. Clermont-Ferrand. PRITCHARD, A. 1861. A History of Infusoria, Living and Fossil: Arranged According to Die Infusionsthierchen of C.G. Ehrenberg; Containing Colored Engravings, Illustrative of All the Genera, and Descriptions of All the Species in that Work, with Several New Ones; to which is Appended an Account of those Recently Discovered in the Chalk Formations. xii. Edition IV, revised and enlarged by J.T. Arlidge, W. Archer, J. Ralfs, W.C. Williamson and the author. Whittaker and Co., London. xiii + 968 pp., 40 pls. RABENHORST, L. 1853. Die Siisswasser-Diatomaceen (Bacillarien) fiir Freunde der Mikroskopie. Eduard Kummer, Leipzig. 72 pp., 9 pls. RABENHORST, L. 1855. Die Algen Sachsens resp. Mittel-Europa’s. Decas 50-51. RABENHORST, L. 1856. Erklarung der Tab. XII. Hedwigia 1:103. RABENHORST, L. 1857a. Die Algen Sachsens resp. Mittel-Europa’s. Decas 61—62. RABENHORST, L. 1857b. Die Algen Sachsens resp. Mittel-Europa’s. Decas 63-64. RABENHORST, L. 1858a. Die Algen Sachsens resp. Mittel-Europa’s. Decas 69-70. RABENHORST, L. 1858b. Die Algen Sachsens resp. Mittel-Europa’s. Decas 73-74. 298 RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. 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In Aryptogamen-Flora von Sachsen, der Ober-Lausitz, Thuringen und Nord-Bohmen, mit Beruchsichtigung der benachbarten Lander. Leipzig. 653 pp. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. complectens, 359 pp. RABENHORST, L. RABENHORST, L. RABENHORST, L. RABENHORST, L. 1863b. Die Algen Europa’. Decas 143-144. 1863c. Die Algen Europa’. Decas 149-150. 1863d. Die Algen Europa’s. Decas 151-152. 1863e. Die Algen Europa’. Decas 157-158. 1864a. Flora Europaea Algarum aquae dulcis et submarinae. Sectio I. Algas diatomaceas cum figuris generum omnium xylographice impressis. Apud Eduardum Kummerum, Lipsiae. 1864b. Die Algen Europa’s. Hedwigia 3:175—176. 1864c. Die Algen Europa’. Decas 165-166. 1864d. Die Algen Europa’. Decas 170-171. 1865a. Flora Europea algarum aquae dulcis et submarinae. Sectio II. Algas phycochromaceas complectens. Lipsiae. Apud Eduardum Kummerum. 319 pp. RABENHORST, L. RABENHORST, RABENHORST, RABENHORST, RABENHORST, RABENHORST, RABENHORST, 1865b. 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Die Algen Europa’. Decas 248-249. 1878. Die Algen Europa’. Decas 250-251. KOCIOLEK: DIATOM TYPES IN RABENHORST’S DIE ALGEN EUROPA’S 2S) REINSCH, P.F. 1866. Die Algenflora des mittleren Theiles von Franken (des Keupergebietes mit den angren- zenden Partien des jurassischen Gebietes) enthaltend die vom Autor bis jetzt in diesen Gebieten beobachteten Susswasseralgen und die Diagnosen und Abbildungen von ein und funfzig vom Autor in diesem gebiete entdeckten neuen Arten und drei neuen Gattungen. Abhandlungen der Naturhistorischen Gesellschaft zu Nurnberg. Nurnberg 1866., Vol: 2 Issue: 2 Halfte. 238 pp., 13 pls. SayRE, G. 1969. Cryptogamae exsiccatae :an annotated bibliography of published exsiccatae of algae, lich- enes, hepaticae, and musci. Memoirs of the New York Botanical Garden, vol. 19, nos. 1-3. SCHUMANN, J. 1867. Die Diatomeen der Hohen Tatra. Verhandlungen kaiserlichkoniglische zoologisch botanische Gesellschaft in Wien 17:1—102. SCHWARZ, D. 1874. Grundproben aus den chinesischen Gewassern gesammelt von R. Rabenhorst. Hedwigia 13:161—166. SERVANT- VILDARY, S., E. FOURTANIER, J.P. KOCIOLEK, M. MACDONALD, AND A. MIETTE. 2001. Nomenclatural issues, types and conserved material from Tempere and Peragallo’s Diatomees du Monde Entier, Edition. Diatom Research 16:363-398. SIMONSEN, R. 1987. Atlas and Catalogue of the Diatom Types of Friedrich Hustedt. 3 vols. J. Cramer, Berlin. S25 pp: 772 pls. SmiTH, H.L. 1876-1888. Species Diatomacearum Typicae. Geneva, New York. [Exsiccatae] 750 slides. SmiTH, W. 1856. Synopsis of the British Diatomaceae, vol. 2. John Van Voorst, London. 107 pp., pls. 32-60, 61-62,,A-E. TEMPERE, J., AND H. PERAGALLO. 1889-1895. Diatomées Collection. Hy-Tribout. St. Paul. 304 pp. TEMPERE, J., AND H. PERAGALLO. 1907-1915. Diatomées du Monde Entier, 2°4 ed. Arcachon (Gironde). 480 pp. VAN HEuRCK, H. 1880. Synopsis des Diatomées de Belgique. Atlas. Ducaju & Cie., Anvers. VAN HEurRCK, H. 1881. 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Neidium through Rhoicosigma. J. Cramer, Vaduz. Pages 2964-3605. VAN LANDINGHAM, S.L. 1978b. Catalogue of the Fossil and Recent Genera and Species of Diatoms and Their Synonyms Part VII. Rhoicosphenia through Zygoceros. J. Cramer, Vaduz. Pages 3606-4241. WARTMANN, F.B., AND B. SCHENK. 1862-1882. Schweizerische Kryptogamen unter mitwirkung mehrer Botaniker. Centuries I-XI, fascicles I-X VII, numbers 1-900. Sankt Gallen (Scheitlin & Zollikofer), Hottingen. WIGAND, ALBERT. 1860. Bemerkungen iiber einige Diatomaceen. Hedwigia 2:41—46. WILLiAMs, D.M. 1988. An illustrated catalogue of the type specimens in the Greville diatom herbarium. Bulletin of the British Museum (Natural History), Botany Series, 18:1—148. ZELLER, G. 1873. Algae collected by Mr. S. Kurz in Arracan and British Burma, determined and systematical- ly arranged by G. Zeller. Journal of the Asiatic Society of Bengal 42:175—-193. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14, pp. 300-337, 11 pls. September 30, 2004 A SEM Study of the Diatom Genus Porodiscus Greville; Morphology of the Species and Comparison with Related Genera N.I. Strelnikova!, E. Fourtanier’, J.P. Kociolek?, and J.A. Barron? ! Department of Botany, Biological Faculty, St. Petersburg State University, Universitetskaya Emb. 7/9, St. Petersburg 199034, Russia; Email: diatomspb@mail.ru. 2 Diatom Collection, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103, USA; Email: efourtanier@ calacademy.org, pkociolek(@calacademy.org. 3 United States Geological Survey, MS910, 345 Middlefield Rd, Menlo Park, California 94025, USA; Email: jbarron@usgs. gov. We carried out SEM studies on the poorly known middle Eocene marine diatom genus Porodiscus Greville. Species examined include, Porodiscus splendidus Greville, P. nitidus Greville, P. conicus Greville, P- oblongus Greville. Selected species in the related genera Coscinodiscus Ehrenberg, Craspedodiscus Ehrenberg and Annellus Tempére are also considered. Two new combinations, Porodiscus splendidus var. excentricus (Olshtinskaya) Olshtinskaya n. comb. and Porodiscus splendidus var. corniger (Brun) Fourtanier n. comb. are proposed, and we designate Porodiscus nitidus Greville as the lectotype of Porodiscus Greville. Porodiscus resembles both Coscinodiscus and Craspedodiscus in the presence of locular areolae, which possess external cribra and internal foramina, and the presence of marginal rimoportulae. Porodiscus has unique characters, however, such as spines occurring in some taxa, which are not present in Coscinodiscus or Craspedodiscus. Porodiscus possesses a dis- tinctive central cavity (or “sack”), similar to that of Craspedodiscus; however, in Porodiscus, the central cavity is very steep (tube-like) and well defined, being bor- dered by a hyaline rim. Craspedodiscus species usually do not have such hyaline rim (except for some tropical Miocene forms of Craspedodiscus coscinodiscus) and lack a well-defined central depression. Porodiscus Greville 1863 is a small, poorly known genus of fossil centric marine diatoms closely allied to Coscinodiscus Ehrenberg from which it differs by the presence of a deep concav- ity at the center of the valve. The genus has received little attention by taxonomists, and the only SEM observations to date are those of Sims (1989) on a species identified as Porodiscus nitidus var. armatus Rattray from Joe’s River, Barbados. Sims’ specimens were not sufficiently well pre- served to observe the shape of the rimoportulae and the nature of the cribrum. Her observations, however, confirm the affinity of Porodiscus with Coscinodiscus, which was first noted by Greville (1863). We present here new information on the morphology of this intriguing genus based on detailed SEM observations and reexamination of Greville’s original slides. Our interest in the genus first arose after the observation under SEM of Porodiscus splendidus (2?) from a sample taken from DSDP Site 206, Core 16, Section 5 (Tasman Sea), a specimen with spines. A review of the literature indicated that P. splendidus Greville was the most cited and the most widely distributed species of Porodiscus. Illustrations of the reported specimens, however, showed both specimens with spines (Gombos 1983) and specimens without spines (Fenner 1985). This prompted us to reexamine the type material of Greville for that species and to consider the other Species in the genus. 300 ee STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 301 HISTORICAL PERSPECTIVE Greville (1863:63) described the genus Porodiscus from samples prepared by Johnson from the sediments of the Cambridge Estate in Barbados (middle-late Eocene according to the strati- graphic scheme of Mitchell-Thome 1979). He originally included five species: P. nitidus, P. ele- gans, P. major, P. conicus and P. oblongus (P. ovalis in plate caption), but did not designate a gener- itype. Subsequently, Greville (1865:46) described Porodiscus splendidus from the sediments of the Springfield Estate, Barbados, whose precise stratigraphic position is not known. Greville (1865) noted that P. splendidus was not found in the Cambridge Estate sediments. Greville (1863) described the genus in these words: “Frustules free, disciform, composed of two discs, united by an intermediate, ring-like zone; discs very convex, minutely radiato-cellulate or punctate, with a conspicuous central pseudo-opening.” Greville continued: “This genus is evident- ly closely allied to Coscinodiscus, differing chiefly in the remarkable pore-like pseudo-opening, which is not a mere blank circular space produced by the absence of cellulation at the apex, but a well-defined, concave, apparent orifice, provided with a thickened margin.” Greville added that “all the species described from Barbados had very convex valves, and puncta [areolae] arranged in radial rows. In nearly all the species certain of the radial rows start from the margin and continue to the valve apex, dividing the valve into perceptible sections [fascicles]. Surface is either plane or armed with variously arranged minute spines.” The types of Greville’s Porodiscus are illustrated by Williams (1988:44—-45, pl. 52, figs. 1-9, pl. 53, fig. 5). Castracane (1886) described P. stolterfothii, which he considered an extant species, from the tropical area of the Pacific Ocean. Grove and Sturt (1887) described two other species of Porodiscus: P. hirsutus and P. interruptus, from Oamaru (New Zealand, upper Eocene). Rattray (1890) described Porodiscus spiniferus, P. splendidus var. marginata, P. major var. densa, and P. nitidus var. armata from Barbados. Porodiscus corniger Brun (1896) was also described from Barbados from the Chimborazo sediments (unknown age) and Mount Hillaby (late Eocene). Porodiscus calyciflos Yempere and Brun in Brun and Tempére, 1889, is known from a Neogene limestone of Yedo, Yedo Bay, Japan, and is also known as living in the Sandwich Islands (Brun and Tempere 1889). Prema and Desikachary (1989) described Porodiscus minor and Porodiscus venkataraminii from the Oligocene of the Indian Ocean. The generic placement of these species has been questioned. Mann (1907:237), noted that P calyciflos, P. interruptus, P. hirsutus (which he placed in synonymy) have nothing in common with Greville’s genus. Jousé in Proschkina-Lavrenko (1949:33) created the illegitimate name Pseudopodosira pileiformis as a synonym of P. calyciflos. Various authors (e.g., Gombos 1982; Prema and Desikachary 1989) have also recognized the similarity of Porodiscus with Craspedodiscus Ehrenberg, a closely related genus with a central depression. Grunow 1881 in Schmidt (pl. 66, figs. 7-9) transferred Porodiscus oblongus Grev. to Craspedodiscus oblongus. Gombos (1982:232) transferred Porodiscus splendidus Grev. to Craspedodiscus splendidus. Based on our observations of original illustrations and descriptions, we recognize that the genus Porodiscus in Greville’s original sense currently contains the species of Greville (P. conicus, P. elegans, P. oblongus, P. nitidus, P- major, P. spendidus), Rattray’s taxa (P. major var. densa, P. nitidus var. armata, P. splendidus var. marginata), P. corniger Brun, and P. stolterfothii Castracane. As Grunow suggested (Grunow in Schmidt 1881, pl. 66, fig. 6), we believe that Craspedodiscus ovalis Grunow belongs to Porodiscus splendidus Greville. The specimen illustrated by Grunow is slightly oval and has a well defined depressed central area bordered by a rim. All these species are the focus of this paper. We agree with Mann (1907:237) that P. calyciflos, P. interruptus, and P. hirsutus do not belong 302 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 in Porodiscus. We also doubt that Porodiscus minor and Porodiscus venkataraminii Prema and Desikachary (1989) belong to Porodiscus, as these species are only slightly depressed at the cen- ter with the central area that is not surrounded by a hyaline rim. As argued by Mann (1907), Poro- discus coronarius (Mann) Mills 1934 (based on Melosira coronaria Mann) does not belong in Porodiscus. Specimens identified by Jousé (1954:128, pl. 3, fig. 1-2) as Porodiscus pliocenicus (an invalid name) do not belong in Porodiscus and were later placed in the genus Cosmiodiscus by Jousé (1961). MATERIALS AND METHODS The following slides were examined: British Museum slides BM 10602. Coll. Deby. L.H. no. 714. Barbados, Diatomaceae selected. Porodiscus splendidus. BM 10603. Coll. Deby. Named by Greville no. 10. “L.H. no. 715. Barbados Diatomaceae selected. Asterolampra affinis.” BM 10604. Coll. Deby. Named by Greville no. 19. “L.H. no. 716. Barbados. Diatomaceae selected. Porodiscus splendidus.” BM 10605. Coll. Deby. Named by Greville no. 17. “L.H. no. 714. Barbados. Diatomaceae selected. Porodiscus splendidus.” BM 10399. Coll. Deby. “L.H. no. 505. Springfield Estate. Barbados. Diatomaceae selected. Porodiscus splen- didus”’. BM 2045. Coll. Greville. “Barbados Earth. Porodiscus ovalis”. BM 2745. Coll. Greville. “Bridgewater. Barbados. Johnson. Porodiscus conicus.” BM 2744. Coll. Greville. “Cambridge. Barbados. Johnson. 1863. Porodiscus nitidus.” BM 2751. Coll. Greville. “Bridgewater. Barbados. Johnson. Porodiscus nitidus — Holotype.” BM 2812. Coll. Greville. “Cambridge. Barbados. Johnson.1862. Porodiscus conicus.” BM 2844. Coll. Greville. “Cambridge. Barbados. Johnson.1863. Porodiscus major — type.” CAS Collection, Barbados CAS 611174. Joe’s River. St. John District. 13°13'N, 59°32'W. CAS 611205, CAS 611235. Cambridge Estate, Adams Hill, St. James District. CAS 611193. Springfield. St. Peter District. CAS 611223, CAS 611226. Chimborazo village. Collection N. Strelnikova Sample CAS 624819, DSDP Leg 21, 206—16—5, 111—113 cm, Tasman Sea (Middle Eocene). Collection A.P. Olstinskaya Sample 74163, Ukraine, Settlement Staroverovka, Kievskaya Formation, middle-upper Eocene. Coll. L.H. Robinson (slides) At CAS, received from the New York Botanical Garden. NOTES ON COLLECTIONS The history of collections from Barbados of the British Museum and the Diatom Collection of CAS is very interesting and is described by J.-H. Robinson (1934, 1936, 1941). She indicates the absence of good information about the exact locality and the geological position of the samples that Dr. Davy collected in 1846 and sent to R.K. Greville under the name “Barbadoes Earth”: “more than two hundred and fifty of these are from Barbados fossil deposit and more than two-thirds of these are cited as “Cambridge Estate” in slides communicated by C. Johnson Esq. From one sam- ple of earth received by Mr. Johnson, the majority of the new species were obtained. Although it was never definitely ascertained where this good sample came from, Dr. Greville seems to have Ne STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 303 been under the impression that it was picked up on the Cambridge Estate” (Robinson 1934:3). The samples from Barbados at CAS were received from J.H. Robinson. She provided infor- mation about the localities Joe’s River Estate (Robinson 1941, V, p. 1), Springfield (Robinson, 1941, VI, p. 84), and Cambridge Estate (Robinson 1941, VII, p. 181) and gave a list of the species she observed in these localities. MICROSCOPY Specimens were photographed and examined under light microscopy (Leica DMRB) and by scanning electron microscopy (SEM) (Leo 1450VP). SYSTEMATIC ACCOUNT Porodiscus splendidus Greville, 1865: 46, pl. V, fig. 5. (Figs. 1—7, 9-10, 24-33) Synonyms (taxonomic): Craspedodiscus ovalis Grunow in Schmidt, 1881, Atlas Diat., Taf. 66, fig. 6. Porodiscus splendidus var. marginatus Rattray, 1890:672. This species was described by R.K. Greville from the Springfield Estate, Barbados. Picked valves of this species are arranged among the selected valves of different species on the type slide from the British Museum. Williams (1988:45) indicates that the lectotype is on slide BM 10602 (L.H. no. 714). Greville wrote on the slide the words “Diatomaceae selected. Porodiscus splen- didus.” This species was also selected and marked by Greville on slides: BM 10604 (L.H. no 716, named by Greville no. 19) and BM 10605 (L.H. no. 717, named by Greville no. 17). All slides men- tioned above are from Deby’s collection and have no locality information. Slide BM 10399 (L.H. no. 505), however, also from Deby’s collection bears the notes: “Springfield Estate. Barbadoes. Diatomaceae selected. Porodiscus splendidus.” OCCURRENCE.— In the material from Barbados at CAS, Porodiscus splendidus was found at all localities in Barbados (Springfield, Joe’s River, Cambridge and Chimborazo) but more often in the Joe’s River material. Porodiscus splendidus is the Porodiscus species that is the most often cited in contemporary literature (Glezer et al. 1965; Glezer 1969, 1974; Glezer and Jousé 1974; Olshtynskaya 1976; Jousé et al. 1977; Gombos 1983; Fenner 1985; Strelnikova 1992). The distribu- tion of P. splendidus (based on the literature) indicates that it is confined to middle Eocene sedi- ments. However, the accuracy of this distribution is in question as the species occurrence may be poorly documented and identification inaccurate. Our SEM and light microscope (LM) observations add more detail on the valve morphology of this species. DESCRIPTION.— We measured all the valves (a total of 6 specimens) of Porodiscus splendidus in Greville’s slides (BM 10602, BM 10399, BM 10604, BM 10605, BM 2844), and a total of 50 valves from Barbados sediments in the CAS collection, mostly from Joes’s river (CAS 611174 [1345], slides 581128-581131), but also from Springfield (CAS 611193 [3368], slide 581122), Cambridge (CAS 611205 [3142, 3143, 3144], slides 581107, 581108, 581109; CAS 611235 [3375, 3376], slides 581111-581113) and Chimborazo (CAS 611226 [3088, 3089, 3090], slides 58116-58118; CAS 611223 [3570], slide 581101). The valves are convex, sometimes almost hemispherical, round or slightly oval 30-91 um in diameter. A deeply concave cavity (4-18 um in diameter), which is sometimes a little eccentric in outline, is positioned in the center of the valve. The central concave part or “apparent orifice” (or ‘sack’ ) juts into the frustule to different depths (6-16 um). The ratio diameter of valve/diameter of 304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 center ranges between 3 and 12. There is no apparent correlation between the diameter of the cen- ter and the depth of the “sack.” Usually, the center is surrounded by a hyaline ring crest, but this ring is sometimes absent (Fig. 26). Areolae are arranged in radial rows 4—5 (sometimes 6) in 10 um. Rattray (1890:672) noted that areolae are a little larger at semi-radius, however, this is not con- firmed by our observations. Areolae loculate, locula pear-shaped, diameter of foramen about | um. Velum not observed. Rimoportulae arranged on the margin of the valve at distances of 1 to 3-4 are- olae from the margin (Figs. 28-29), with 3-4 areolae between rimoportulae. Rimoportulae open externally as small round holes (Fig. 26). Internally, they appear as simple small round holes, but it is not clear from our observations whether they are broken. Valves have no spines, only some protuberances around the areolae (Figs. 24-25). Girdle band is high (12-18 um) with a group of vertical lines separated by narrow hyaline lines, 14-18 pores in 10 um in a line (Figs. 31—33). These observations are consistent with Greville’s original description, and Rattray’s subsequent description (Rattray 1890:672). COMMENTS.— Craspedodiscus ovalis Grunow (= Porodiscus splendidus var.?) (Schmidt, 1881, Atlas Diat., Taf. 66, Fig. 6) from Springfield is a slightly oval specimen identical in shape and structure with P splendidus Greville. Rattray (1890:672) treated Craspedodiscus ovalis Grunow as a variety of Porodiscus splendidus and named it Porodiscus splendidus var. marginata. Based on our observations of round and slightly oval valves of P. splendidus in BM and CAS mate- rials, and Greville’s original description of P splendidus (“disc circular, occasionally broadly oval”) we treat here Craspedodiscus ovalis Grunow (= Porodiscus splendidus var. marginata Rattray) as a synonym of Porodiscus splendidus Greville. Porodiscus splendidus var. corniger (Brun) Fourtanier, new comb. (Figs. 37-40) n Basionym: Porodiscus corniger Brun, 1896:242, pl. XIX, fig. Porodiscus corniger Brun (1896:242, pl. XIX, fig. 15) (Figs. 37-40) was described from Chimborazo, Mount Hillaby, as a very rare form. Brun noted its similarity with P. splendidus, the two taxa differing only by the presence of spines in P. corniger. The valves of Porodiscus in the Chimborazo material are very rare. Only one frustule of P corniger was observed in the Chimborazo material (CAS 611226); it was mixed with several forms without spines. As the two types are identical in structure and dimensions and differ only by the presence of spines, we believe that P. corniger should be treated as a variety of P. splendidus. Since we did not observe a contin- uum of morphological expressions between specimens with spines and specimens without spines, We are recognizing P. splendidus var. corniger and P. splendidus var. splendidus as two separate taxonomic entities. Porodiscus splendidus var. excentricus (Olshtinskaya) Olshtinskaya, new comb. (Figs. 8, 11-12, 34-36) Basionym: Porodiscus nitidus var. excentricus Olshtinskaya, 2001:452, pl. I, figs. 1-3. Olshtinskaya (2001) described a new diatom taxon from the upper Eocene sediments of the Pristen village, Ukraine, as a variety of Porodiscus nitidus (Porodiscus nitidus var. excentricus). We studied specimens from Staroverovka (coll. A.P. Olshtinskaya) near the type locality of her taxon and in the same geological interval (Kievskaya Formation, middle to upper Eocene). Olshtinskaya’s taxon differs from Porodiscus splendidus only by the eccentric position of the “sack.” In the material from Barbados, there are a few forms with an eccentric “sack.” In the STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 305 Ukrainian material, however, the eccentric type is dominant, and the external surface of Ukrainian forms are smooth, without small bumps. Porodiscus nitidus Greville, 1863, p. 65, pl. IV, fig. 4. (Figs. 13-16, 41-56) Synonyms: Porodiscus nitidus var. armatus Rattray, 1890:672-673, pl. III, fig. 17. Porodiscus spiniferus Rattray, 1890:674, pl. Il, fig. 19. Greville described P. nitidus from the Cambridge Estate. Two slides in the Greville collection (slides BM 2751 and BM 2744) bear the indication “Porodiscus nitidus.” Slide BM 2751 is a smear slide and, according to Williams (1988), is the probable holotype slide for P. nitidus. The slide bears no locality information besides “Barbadoes. Johnson Coll. Greville.” P. nitidus is also present in slide BM 2812. Greville (1863:65) stated in his original description of P. nitidus, “Disc convex, unarmed, the longest lines of puncta single (not in pairs), alternating with two or three series of shorter ones; puncta distinct, all of them becoming much more minute towards the margin. Diameter .0026. Barbadoes deposit, from Cambridge Estate; C. Johnson Esq. . . . It is a scarce species.” Porodiscus nitidus var. armatus Rattray (1890:672—673) was also described from the Cambridge Estate. Rattray noted the presence of spines (“spines acicular, about .01 mm., long, sometimes shorter, inserted about */ of radius from center”). It appears from Rattray’s description and illustration that the main difference between P. nitidus var. armatus and the nominal variety is the presence of spines. The two varieties are otherwise identical in valve structure and dimensions. Porodiscus spiniferus Rattray (1890:674) was also described from the Cambridge deposit. It appears from Rattray’s description that P. spiniferus is similar to P. nitidus var. armatus, but it has more robust spines, more evident fascicles, and possibly more dome-shaped valves. Although Greville 1863 was clear in his original description of Porodiscus nitidus that spines were not present (“unarmed”), the slides in his collection (BM 2751, BM 2744) feature both spec- imens with spines and without spines. Our observations indicate that there is a continuum between specimens lacking spines, specimens with minute spines, and specimens with robust spines. There is also a great variability in the appearance of the fascicules, from very faint to evident. Despite three taxa having being described for this material, our observations suggest a continuum of mor- phological expressions within a single taxon, and support the synonymy between Porodiscus nitidus var. nitidus, P. nitidus var. armatus and P. spiniferus. Our description of P. nitidus is supported by the observation of 12 specimens (on 11 slides) from Barbados from slides at the British Museum (BM 2744, BM 2751, BM 2745, BM 2812) and at CAS (CAS 61174[1345], Joe’s River, slide 581129; CAS 611235[3376], Cambridge, slide 581112; CAS 611226[3089], Chimborazo, slide 581117). We found the best preserved specimens of P. nitidus in DSDP Leg 21, Sample 206-16-5, 111-113 cm, Tasman Sea (CAS 624819), from which we studied 25 valves in LM and SEM. DESCRIPTION.— Valves circular, convex, 40-118 um in diameter, with a deeply concave cen- tral part, round or very slightly oval, 5—7 um in diameter that is either surrounded by a hyaline ring or lacks a ring. Ratio valve diameter/diameter of center 8-20. Areolae loculate with external cribrum and internal foramen, 5—8 in 10 um. Loculae cylindrical. Areolae arranged in radial rows, 5—8 in 10 um in a row, organized in fascicles, separated by hyaline lines. Fascicles are usually faint, sometimes more distinct. The hyaline lines and fascicles may be more distinct internally. Valve sur- face smooth or with spines. There is great variability in the size of the spines from barely notice- 306 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 able to very long. Spines are situated on the hyaline lines at mid-radius or closer to the margin (74 of radius from center according to Rattray’s original description) (Figs. 41-42). Rimoportulae are positioned in the hyaline lines at the distance of 1 areola from the valve margin (Fig. 56). They con- sist of small round holes externally, and small round or oval holes internally. It is possible, howev- er, that they are eroded. Cribrum consists of several pores. In well preserved cribra there are 5—7 marginal pores surrounding a single (but sometimes 2-3) central pore (Figs. 52-55). Rattray (1890:672) noted that the areolae were more dense near the margin; however, this is not confirmed by our observations. COMMENTS.— Sims (1989:366, figs. 48-52) published SEM photos on specimens identified as P. nitidus var. armatus Rattray from Joe’s River, Barbados (material from CAS collection: A.L. Brigger’s samples, presented to British Museum by R.W. Holmes). Her specimens have spines, but unlike the specimens we observed, there are several spines on one hyaline ray. Spines nearest to the center (ca '/; radius) are long and thick, spines closer to the margin are of moderate size. The presence of more than one concentric row of spines in P. nitidus was not observed by us and not mentioned by Rattray in the description of P. nitidus var. armatus or P. spiniferus. This suggests that the specimens illustrated by Sims may belong to a different taxon. Further study of her popu- lation is needed to determine if it represents a separate taxonomic entity. The main differences between P. nitidus and P. splendidus are: 1) The ratio of valve diameter to diameter of the concave center, which is smaller for P. splen- didus (3—9) and larger for P. nitidus (9-18). The diameter of the center is therefore larger in P. splendidus (5—18 um) than in P. nitidus (4-6 um); 2) Areolae are larger in P. splendidus (4—6, commonly 5 in 10 um) than in P. nitidus (S—8, com- monly 6—8 in 10 um); 3) Areolae are arranged in radial lines in both species, but P. nitidus has slightly more evident fasciculae, which are separated by hyaline lines that are more distinct internally; 4) Spines are absent in P. splendidus, they are often (but not always) present in P. nitidus. Porodiscus conicus Greville, 1863: 65, pl. IV, fig. 3. (Figs 17, 18, 642) Synonym: Porodiscus elegans Greville, 1863:64, pl. IV, fig. 1. This species was described by Greville from Cambridge Estate, Barbados. We studied it only from the type slides of the British Museum as we did not find any specimen in the CAS Collection material from Cambridge, Barbados. This species has not been reported since Greville’s work. A complete frustule is present in slide BM 2812, the holotype according to Williams (1988). We also found this species on slides BM 2744 and BM 2745. We did not find definite specimens of this species in the CAS Collection. DESCRIPTION.— Greville’s description is short: “Small; disc conical, unarmed, with an obtuse- ly truncate apex; radiating lines of puncta extremely minute. Diameter .0014” [36 um].” In the comments to this description Greville writes — “The smallest of the species hitherto discovered, and occurring not infrequently in perfect frustules. The length of the connecting zone is consider- able, and that of the entire frustule, when both valves are symmetrical, about .0040” [ 98 um]. The valve is decidedly conical, but obtusely truncate at the top when seen in profile. It hardly ever hap- pens that the valves are equal in the same specimen. Indeed, I do not think that I have seen a sin- gle example perfectly symmetrical, one valve being almost always considerably shorter than the other. The length of the connecting zone gives the frustule a cylindrical appearance.” STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 307 The diameter of the specimen illustrated by Greville is 45 um. Rattray (1890) restudied Greville’s and Johnson’s material and added to Greville’s description: “Diameter .025 to .0525 mm. Major axis of frustule from .0625 to .0875 mm. Surface a more or less elongate regular cone, trans- versely truncate at the extremities, the opposite valves of a frustule of unequal height. Central space? Markings obtusely angular or subareolate, 6 in .01 mm, subequal; rows radial, straight, non- fasciculate; secondary oblique decussating rows evident, from the truncated ends of the cone a few short tapering clear lines, distinct. Girdle cylindrical, from .025 to .0325 mm, broad; a narrow band at each extremity, hyaline, the intervening portion clouded with diffuse parallel lines.” In Greville’s slides, we found the valve diameter of Porodiscus conicus to range between 40-43 um, the height of conical valves to be 30-35 um, the height of the second valve to be 16 um, and the girdle band to be 27—32 um in width. Areolae on the valves are arranged in radial rows, 7-8 in 10 um in a row, 8 rows in 10 um. On the shorter valve 5—6 rows of areolae alternate with thin hyaline lines. COMMENTS.— Greville (1863:64) described Porodiscus elegans from Cambridge and other localities. “Disc very convex, unarmed, divided into compartments by pairs of the radiating lines of very minute puncta, extending from the margin to the centre. Diameter .0020” [51 um] to .0033” [84 um].” The specimen illustrated by Greville has a diameter of 77 um. Greville (1863:64) com- mented: “This species is distinguished by the disc being divided into numerous compartments, by pairs of radiating lines of puncta, very distinctly seen under a moderately magnifying power, and at the same time being quite destitute of spines. It is the most frequent species, three or four valves sometimes occurring in a single slide. The connecting zone is rarely seen in situ.” Rattray (1890:673) added: “Circular. Diameter .0625 to .095 mm. Surface rounded and dome-shaped. Central space circular, 0075 mm. broad, sharply defined, hyaline. Markings obtusely angular or subareolate, decreasing gradually from the central space outwards, around the central space 6, near the border 10 to 12, in .01 mm; rows radial, straight; fasciculi distinct; interspaces minute, largest around the central space. Girdle cylindrical, .03 mm, broad, in a frustule, .06 mm in diameter; a narrow hyaline band at each extremity; the interval minutely punctate; at subregular intervals nar- row hyaline straight lines at right angles to the edges of the valve. The fasciculi are bounded by two adjacent radial rows, somewhat more conspicuous than the intervening rows. In one of the valves in Greville’s collection it is possible to trace downwards from the central space a cylindri- cal siliceous tube which is of sufficient length to have passed to a plane corresponding in position to the edges of the valve.” Our measurements of the specimen from slide BM 2745, the holotype of Porodiscus elegans according to Williams (1988) are: diameter of valve 45 um, diameter of cen- ter 8 um, areolae in a row 7 in 10 pm. Porodiscus conicus and P. elegans are found on the same slides (BM 2812 and BM 2745). The non-conical valve of P. conicus seems identical to valves attributed to P elegans. They both have a fasciculate arrangement of areolae and the same areolae density. Given that the two morphologies occur together as part of a single frustule, we believe that P conicus and P. elegans are conspecif- ic. We treat the two names as synonyms and retain P. conicus for the name of the species, since it was originally applied to a complete frustule. Porodiscus major Greville, 1863: 64, pl. 4, fig. 2. Porodiscus major was described from Cambridge Estate, Barbados, from a slide communicat- ed by C. Johnson (Greville 1863:64). The description was later amended by Greville (1865:46-47). We did not study this species as we did not find any additional specimens besides the valve on slide BM 2844 (holotype?) that was illustrated by Williams (1988, pl. 52, figs 5-6). Based on our obser- vations of slide BM 2844, we believe that this species belongs in Porodiscus. 308 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 Porodiscus oblongus Greviile, 1863: 65, pl. 4, fig. 5. (Figs. 22-23, 57-61) Nomenclatural synonyms: Porodiscus ovalis Greville, 1863, explan. pl. 4, fig. 5 (alternate name). Craspedodiscus oblongus (Greville) Grunow in Schmidt, 1881, explan. pl. 66, Figs. 7-9. Greville (1863:65) described Porodiscus oblongus from “Barbadoes deposit” and illustrated it as P. ovalis (Greville 1863, explan. pl. 4, fig. 5). Slide BM 2045 bears the note “Coll. Greville. Barbadoes earth” and the name ‘“Porodiscus ovalis.”” Williams (1986) cites this slide as possibly the holotype. This species is not rare in the British Museum slides. On slides BM 10602 and BM 10399, valves of P. oblongus were selected (picked) by Greville. Greville’s original description is short (Greville 1863:65): “Disc elliptical-oblong; pseudo- opening large. Long diameter about .0028” [71 um]. He also added, “The pseudo-opening is very large, the radiating lines of granules are less crowded, and the granules themselves larger than in any of the preceding species”. Greville (1865:46) adds to the description: “The diameter of the disc is .0030” [77 um]. Radiating cellules about 8 in .001.” The specimen figured in Greville’s original illustration (Greville 1863, fig. 5), has a diameter of 70-35 um, and there are about 5 areolae in 10 um on a row. Rattray (1890:674) added to this description: “Subacutely elliptical. Major axis .05 mm long, about 2.5 times minor. Surface sloping gradually downwards from edge of central space. Central space roundly elliptical, with major axis corresponding in direction to minor axis of valve. Markings angular, decreasing regularly and somewhat rapidly from central space to border; around the central space 4.5 at border 10 in .01 mm; rows radial, substraight. Border narrow, hyaline.” DESCRIPTION.— We measured five valves of PR. oblongus from slides BM 2045, BM 1062, BM 1063, BM 10399, and five valves from sample CAS 611193[3368], Springfield (slide 581122). Valves oval, length (S4—78 um) two times larger than width (26-37 um), with round or slight- ly elliptical center (9-11 tm diameter). Areolae organized in rows radiating from the center, 5—6 areolae in 10 um. Locula with velum better preserved on the elongated ends of the valves (Fig. 58), foramen about | um in diameter (Fig. 60). Velum consisting of a central cribral pore surrounded by 4-6 marginal cribral pores (Figs. 57-58). The central depression (“sack”) is not bordered by a real hyaline ring, but the areolae walls surrounding the depression form a little boundary around it that appears like a ring. Areolae in the depression are smaller than on the valve. Rimoportulae arranged along the margin of the valve, with more distance from the margin in the middle of valve (2-3 areolae from margin) and closer to the margin (1 areola from margin) at the ends of the valve. Rimoportulae opening externally as round small holes, internally also as holes, but may be broken (Fig. 60). COMMENTS.— Castracane (1886) described a similar species P. stolterfothii from the equato- rial part of the Pacific Ocean as a living species; it differs from P oblongus by a more rhombic shape. Also similar to P. oblongus is Coscinodiscus oblongus Greville (1866a:4, figs. 9-10) from Springfield Estate, Barbados. Both species (P. oblongus and C. oblongus) are present in British Museum slides BM 2045, BM 10605, BM 10604, BM 10603, BM 20602 and BM 10399. The main difference between the two taxa is a deeper (steeper) cavity in Porodiscus oblongus. Porodiscus oblongus Greville from Springfield is illustrated in Schmidt’s Atlas (Schmidt 1881, Taf. 66, figs. 7-9) as Craspedodiscus oblongus (Grey.) Grunow [in Schmidt] (a valid new combination). Schmidt also illustrates from Springfield Coscinodiscus oblongus Greville (Taf. 66, figs. 10-11) for which he suggests a generic placement in Craspedodiscus (“unbedingt zu Craspedodiscus zu rechnen”’). Schmidt, however, did not validly publish the new combination, which seems to have been first published by Hanna (1931:194). STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 309 Craspedodiscus oblongus (Greville) Hanna (= Coscinodiscus oblongus Greville) and Craspedodiscus oblongus (Greville) Grunow in Schmidt (= Porodiscus oblongus Greville) are sim- ilar species; however, Craspedodiscus oblongus (Greville) Grunow in Schmidt belongs to Porodiscus, whereas Craspedodiscus oblongus (Greville) Hanna belongs to Craspedodiscus and had been renamed Craspedodiscus ellipticus by Gombos (1982). Fenner (1985) refers to two species, Craspedodiscus ellipticus Gombos (= Coscinodiscus oblongus Greville) and Craspedodiscus oblongus (Greville) Grunow (= Porodiscus oblongus Greville). Many works (e.g., Barron et al. 1984; Strelnikova 1992) cite Craspedodiscus oblongus (Grev.) Grunow from Middle Eocene sediments, which corresponds to Coscinodiscus oblongus Greville (= Craspedodiscus ellipticus Gombos). This species has a wide distribution in the Middle Eocene in tropical and non- tropical regions. Porodiscus oblongus Grey. has a narrower distribution limited to the tropical region. We found it only in samples from Barbados. Craspedodiscus ellipticus Gombos, 1982, p. 231, figs 13, 14. (Figs. 19, 21, 62, 63) Nomenclatural synonyms: Coscinodiscus oblongus Greville, 1866a:4, pl. 1, figs 9, 10. Craspedodiscus oblongus (Greville) Hanna, 1931:194 (illegitimate name, later homonym). Craspedodiscus oblongus (Greville) Grunow in Schmidt, 1881, caption Taf. 66, figs. 10-11 (invalid name). Greville’s original description is short “Disc more or less oblong, having the center depressed, and an umbilicus containing a number of subremote granules; surface filled up with radiating gran- ules, which diminish in size next the umbilicus and towards the margin, where they resemble minute puncta. Length .0028” to .0050” [71 to 127 um].” It was described from Springfield. DESCRIPTION.— We measured several valves from slide BM 10603 and slide CAS 581122 (CAS 611193[3368]). We found that the valves, characterized by their elongated shape, are 56—67 um in length, near- ly 30 um in width, with areolae arranged in rows radiating from a concave center, 6 areolae in 10 um on a row. The concave center is oval, with diameter 10—12 x 6—8 tm. Rimoportulae arranged along the margin, opening externally as a small round holes, and internally also as round holes (they may be broken, however). CoMMENTS.— Williams (1988, pl. 28, figs 1-4) illustrated two specimens that he referred to Coscinodiscus oblongus Greville. Only the specimen illustrated in figures 3—4 belongs to Craspedodiscus ellipticus Combos (= Coscinodiscus oblongus Greville). The other specimen (figs. 1-2), which has a round center, belongs to Porodiscus oblongus Greville. Craspedodiscus umbonatus Greville, 1866b:79, pl. 8, fig. 15. (Figs. 65-70, 74) Craspedodiscus umbonatus Greville was described from “Cambridge Estate Barbados deposits in slides communicated by C. Johnson Esq.” The original description is short: “Disc hexagonally cellulate, the border nearly equal to half the radius, the centre rather sharply umbon- ate.” Greville further commented: “Distinguished at once by its umbonate centre. Cellules near the margin of the border 8 in .001”.” Williams (1988, pl. 33, fig. 7) illustrated the holotype from slide BM 5469. DESCRIPTION.— We did not study the holotype slide, but this species was common on slides from the CAS Collection, especially from material from Joe’s River (CAS 611174[1345], slides 581129, 581130). 310 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 The valves are circular, diameter 42-74 um, convex, sometimes hemispherical with deeply concave center, 7-10 um diameter. The ration of the valve diameter to the diameter of the center is 6-8. Areolae are arranged in radial rows, 4—5 in 10 um. External cribrum consists of one or sev- eral central pores surrounded by marginal pores. Areolae are a little smaller on the concave center. Rimoportulae are arranged around the margin, opening externally as small round holes, sometimes one of them situated in smaller cavities (Fig. 74) on the margin. They open internally as simple round holes; however, they may be broken. Valves have no spines. COMMENTS.— This species has been reported only from Barbados. Craspedodiscus umbona- tus differs from Porodiscus only by the absence the hyaline ring around the cavities and the little depression for one rimoportula at the margin (Fig. 74). TYPIFICATION OF THE GENUS PORODISCUS WITH PORODISCUS NITIDUS GREV. In the original publication of Porodiscus, Greville included five species (P. nitidus, P. elegans, P. major, P. conicus, and P. oblongus = P. ovalis in the plate caption), but he did not designate a type. The most common species of Porodiscus, P. splendidus Grev. is not a candidate as generitype inasmuch as it was not one of the original species. We designate here Porodiscus nitidus Grev. as the lectotype of genus Porodiscus Greville. Although not as common as P. splendidus, P. nitidus has been reported in various fossil localities, and it is a better choice than any of the other four species that have never been observed other than on Greville’s original slides. The holotype of P nitidus is on slide BM 2751 (Bridgewater, Barbados). DISCUSSION Coscinodiscus Ehrenberg, Craspedodiscus Ehrenberg, Porodiscus Greville, and Annellus Tempere are closely allied genera in the family Coscinodiscaceae; they differ only by the structure and shape of the cavity in the valve center. Based on the shapes and arrangements of the rimopor- tulae, Craspedodiscus and Porodiscus are similar. Stratigraphically, Coscinodiscus is the oldest of these genera (Cretaceous). Craspedodiscus apparently separated from Coscinodiscus during the early Paleocene. Porodiscus separated from Coscinodiscus or Craspedodiscus only during the Middle Eocene, whereas Annellus is restricted to the Miocene. The Cretaceous genus Pomphodiscus Barker and Meakin has locular areolae, and a central cavity that is reminiscent of some species of Porodiscus and Craspedodiscus. The central cavity in Pomphodiscus is a chamber that is formed by an inflation and separation of the basal siliceous layer into two layers. The shape and the central position of the rimoportulae, however, places this genus in the family Stellarimaceae (Nikolaev and Harwood 2000). Even more distant in the classification, but with a central chamber, is the Cretaceous genus Benetorus (family Stictodiscaceae). The uvular process of the Early Cretaceous genus Archaegladiopsis (family Archaegladiopsidaceae) is also reminis- cent of the cavity of Porodiscus. The uvular process, centrally located is a funnel-shaped depres- sion with a round opening in the valve exterior and a uvula-shaped intrusion on the inside, without visible openings (Nikolaev and Harwood 1997). We believe that the higher level classification of these fossil genera is still valid (i.e., we are not suggesting that these genera are closely related), and that their superficial similarities are due to parallel evolution. The following section will concentrate on the morphological differences between Porodiscus and Craspedodiscus and between Annellus and Craspedodiscus, because it is presently difficult to separate them. STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE Syl Craspedodiscus Ehrenberg We recognize three informal divisions among the diatoms that have been referred to Craspedodiscus. The classical concept of Craspedodiscus is represented by C. coscinodiscus (Figs. 81-86) and is characterized by a large internal depression that has a diameter that is at least one half of the entire valve. The central depression typically has a nearly flat bottom and has areolae that are reduced in size compared to those of the outer valve surface (Figs. 84-85). The transition from the outer valve surface to this internal depression is abrupt, often close to 90° (Figs. 82-85). Similar to Porodiscus, a hyaline rim occurs at this transition (Figs. 82-85) in most Miocene forms. The edge of the central depression is rough, as if a weakly silicified covering, now missing, was present over a chamber in that species. Such covering was never observed in Craspedodiscus. The majority of diatoms that have been referred to Craspedodiscus comprise a second group, represented by C. elegans, the lectotype of Craspedodiscus (see Ross in Farr et al. 1979). Diatoms of this group have an internal depression that is less distinct, characterized by a more gradual tran- sition from the valve’s mantle and by areolae that are not distinctly different from those on the valve’s margin. Some Oligocene diatoms that have traditionally been referred to Craspedodiscus coscinodiscus, but are better assigned to Craspedodiscus barronii Bukry (Figs. 76-77), have gen- tly sloping central depressions and could be referred to this group. Other taxa, including Craspedodiscus undulatus Gombos, C. elegans Ehrenberg and C. moelleri A. Schmidt, have undu- lated valve surfaces, reminiscent of the undulated valve surfaces of certain species of Actinocyclus and Cestodiscus. Diatoms of this group appear to be closely related to Coscinodiscus, and it is not clear why they should be separated from it. Indeed, Coscinodiscus excavatus (Fig. 92) might be placed in this group, although no formal transfer to Craspedodiscus has been proposed. We do not, however, wish to discuss the taxonomic implications of transferring Craspedodiscus elegans and other diatoms of this group to Coscinodiscus in the present paper. A third group of taxa that have been referred to Craspedodiscus are similar to Porodiscus in that they possess a distinct internal depression that is typically confined to only the centermost por- tion of the valve. This group is represented by Craspedodiscus ellipticus Gombos (= Coscinodiscus oblongus Greville) (Figs. 62-63) and C. umbonatus Greville (Figs. 65-70). As in Porodiscus, the central depression is deeply invaginated with a concave central valvar surface. The transition from the outer valve surface to the interior depression, however, is not as abrupt as in Porodiscus, and no hyaline ring is present. Valves of Craspedodiscus ellipticus, with their sloping central depres- sion (Figs. 62-63), clearly differ from valves of Porodiscus oblongus (Figs. 57-61) with their cen- tral depression more circular and sloping more abruptly. On his Figure 17, however, Gombos (1982) illustrates a diatom, which he refers to Craspedodiscus oblongus (Greville) Schmidt, which has a circular internal depression like that of P oblongus, but it also possesses a gradual or sloping transition from the outer valve surface, as in C. ellipticus. This specimen may be transitional between P. oblongus and C. ellipticus. Annellus Tempére The genus Annellus is restricted to the late-early and early-middle Miocene, separated by at least 15 million years from the Eocene records of Porodiscus. At present, Annellus is monospecif- ic, with A. californicus Tempere, as the only species. As discussed by Ryde (1962), Annellus is closely related to Miocene forms of Craspedodiscus coscinodiscus. Ryde (1962) made a detailed morphologic study of Annellus californicus Tempere, noting that the valve was “formed of an outer cylinder inturned at one end (and) deeply invaginated to produce within the first, a co-axial cylinder closed at the lower end.” He mentioned that the “inner cylinder 32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 often tends to bulge somewhat and its length is equal to the depth of the valve mantle,” appearing as a dome when viewed from the inside of the valve. Ryde (1962) stated that the diameter of the areolae on the valve surface of the inner cylinder is very much reduced in size compared to those of the valve mantle and the surface consists of rather fragile, diaphanous silica that is often not pre- served. Ryde (1962) further commented on the rather abrupt transition from the inner valve man- tle to the inner cylinder. Based on comparison with the internal valve surface of Craspedodiscus coscinodiscus, Ryde (1962) chose to transfer Annellus californicus Tempere to Craspedodiscus californicus (Tempére) Ryde. We feel, however, that the high mantle of Annellus californicus, which results in valves often being orientated in valve view on slides, is a unique feature to Annellus, distinguishing it from the Craspedodiscus coscinodiscus group. Annellus also lacks the hyaline rim, which is present in typical forms of Craspedodiscus coscinodiscus (Figs. 82—86). Barron (1983) argued that the shallower mantle of his new species, Craspedodiscus rhydei, war- ranted placing it into Craspedodiscus rather than Annellus; however, C. rhydei (Fig. 91) clearly lacks a hyaline ring and closely resembles valves of Annellus californicus (Figs. 87-88). CONCLUDING REMARKS ON PORODISCUS GREVILLE Porodiscus resembles both Coscinodiscus and Craspedodiscus with the presence of locular areolae with external cribrum and internal foramina, and marginal rimoportulae. It has unique char- acters, however, such as spines (in some species), which are not present in Coscinodiscus nor in Craspedodiscus. The central cavity, which also characterizes Craspedodiscus, seems to be of a dif- ferent nature in Porodiscus, where it is very steep (tube-like) and well defined, being bordered by a hyaline rim. Craspedodiscus species usually do not have such hyaline rims (except for some trop- ical Miocene forms of Craspedodiscus coscinodiscus) and lack a well-defined central depression. The hyaline rim in these forms may be the edge of a broken outer surface of a chamber. Miocene forms of Craspedodiscus coscinodiscus and Annellus californicus, which have a well defined cen- tral depression, typically have a central depression that covers at least half of the diameter of the valve. This character, and the separation in geologic time of more than 15 million years between the Eocene forms of Porodiscus and the Miocene forms of Craspedodiscus coscinodiscus and Annellus, are arguments in support of Porodiscus being retained as a valid taxonomic entity. ACKNOWLEDGMENTS We thank Dr. David Williams of the British Museum who made possible the study of the type slides of R. Greville. The work was supported by Russian Fund for Fundamental Research (RFFI) Grant 00-04-49333 and the Diatom Collection of the California Academy of Sciences. We are also thankful to D. Ubick, Scott Serata, M. Potapova, and E. Ruck for assistance in the preparation of this article, Dr. David Harwood for his review of the manuscript and helpful suggestions, and Dr. Michele L. Aldrich, who read the near final version of the manuscript and saved us from memor- alizing a number of grammatical and other baby blunders. LITERATURE CITED BARRON, J.A. 1981. Late Cenozoic diatom biostratigraphy and paleoceanography of the middle latitude east- ern North Pacific, DSDP Leg 63. Jnitial Reports, Deep Sea Drilling Project 63:507—538. BARRON, J.A. 1983. Latest Oligocene through early middle Miocene diatom biostratigraphy of the eastern tropical Pacific. Marine Micropaleontology 7:487-—515. STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 313 BarRON, J.A., D. BUKRY, AND R.Z. Poore. 1984. Correlation of the middle Eocene Kellogg Shale of northern California. Micropaleontology 30(2):138—170. Brun, J. 1896. Diatomées Mioceénes. Le Diatomiste 2(24):229-260, pl. 19-24. Brun, J., AND J. TEMPERE. 1889. Diatomées fossiles du Japon. Espéces marines et nouvelles des calcaires argileux de Sendai et de Yedo. Mémoires de la Société de Physique et d'Histoire Naturelle de Genéve 30(9):3-75. CASTRACANE, F. 1886. Report on the Diatomaceae collected by H.M.S. Challenger during the years 1873-1876. Report on the scientific results of the voyage of H.M.S.Challenger during the years 1873-76. Botany, Vol. 2. 178 pp. Farr, E.R., J.-A. LEUSSONK, AND Y. STAFLEU. 1979. Index Nominum Genericorum (Plantarium) Supplement. Regnum Vegetabile 113. 126 pp. FENNER, J. 1985. Late Cretaceous to Oligocene planktonic diatoms. Pages 713-762 in H.M. Bolli, J.B. Saunders, and K. Perch-Nielsen, eds., Planktonic Stratigraphy. Cambridge University Press, Cambridge. GLEZER, Z.1. 1969. Late Eocene complexes of diatoms, silicoflagellates and ebriidea of the south-western part of the Turgai depression. Proceedings of the VSEGEI (All-Union Geological Institute), new series, V. 130. Biostratigraphy (Biostratigraficheskii Sbornik), 4:67—84. GLEZER, Z.I. 1974. Diatoms of the Paleogene. Pages 109-146 in Glezer et al., eds., The Diatoms of the USSR. Fossil and Recent. Publishing House Nauka, Leningrad Branch, 1. GLEZER, Z.]., AND A.P. JousE. 1974. Diatoms and silicoflagellates of the Eocene of equatorial Atlantic. Pages 49-63 in Micropaleontology of Seas and Oceans. Nauka, Moscow, USSR. GLeZER, Z.I., V.YU. ZOSIMOVICH, AND M.N. KLYUSHNIKOV. 1965. Diatoms of the Paleogene sediments of the Severny Donez River basin, and their stratigraphic position. Paleontologicheski sbornik 2(2):73—84. Gompos, A.M., JR. 1982. Early and middle Eocene diatom evolutionary events. Bacillaria 5:225—242. Gomsos, A.M. 1983. Middle Eocene diatoms from the South Atlantic. Initial Reports of the Deep Sea Drilling Project 71:565—581. GREVILLE, R.K. 1863. Descriptions of new and rare diatoms. Series [X. Transactions of the Microscopical Society of London, new series 11(3): 63—76, pls IV—V. GREVILLE , R.K. 1865. Descriptions of new and rare diatoms. Series XVI. Transactions of the Microscopical Society of London, new series 13:43—57, pls V—VI. GREVILLE , R.K. 1866a. Description of new and rare diatoms. Series X VIII. Transactions of the Miscroscopical Society of London, n.s., 14:1—9, pls. I-I. GREVILLE , R.K. 1866b. Description of new and rare diatoms. Series XIX. Transactions of the Miscroscopical Society of London, n.s., 14:77—86, pls. VIH-IX. Grove, E., AND G. Sturt. 1887a. On a fossil marine diatomaceous deposit from Oamaru, Otago, New Zealand. Part III. Journal of the Quekett Microscopical Club, series 2, 1887, 3(18):63—78, pls 5-6. GROVE, E., AND G. StTurT. 1887b. On a fossil marine diatomaceous deposit from Oamaru, Otago, New Zealand. Appendix. Journal of the Quekett Microscopical Club, series 2, 1887, 3(19):131—148, pls 10-14. HANNA, G D. 1931. Diatoms and silicoflagellates of the Kreyenhagen shale. Mining in California. California Division of Mines, Report of the State Mineralogist 27(2):187-201. HANNA, G D. 1932. The diatoms of Sharktooth Hill, Kern county, California. Proceedings of the California Academy of Sciences, ser. 4, 20(6):161—263. Jouse , A.P. 1949. The genus Pseudopodosira Jousé. In: Proschkina-Lavrenko, ed., Diatomovy Analiz, Botanicheskii Institut im V.L. Komarova Akademii Nauk S.S.S.R. Gosudarstvennoe Izdatelystvo Geologicheskoi Literatury, Moskva-Leningrad 2(1):238 pp. Jouse, A.P. 1954. On diatoms of Tertiary age in bottom sediments of the Far Eastern Seas. Proceedings of the Institute of Oceanography, Academy of Sciences of the USSR 9:119—135. [In Russian] Jouse, A.P. 1961. Diatomeae marinae Mioceni et Plioceni ex oriente extremo. Botanicheskie Materialy okdela Sporovykh Rastenii (Notulae Systematicae e Sectione Cryptogamica Instituti Botanici Nomine VL. Komarovii Academiae Scientiarum URSS) 14:59—70, 3 pls. Jouseé, A.P. editor-in-chief. 1977. Atlas of Microorganisms in Bottom Sediments of the Oceans (Diatoms,Radiolarians, Silicoflagellates, Coccoliths) M. “Nauka’’. 196 pp. Mann, A. 1907. Report on the diatoms of the Albatross voyages in the Pacific Ocean, 1888-1904. 314 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 Contributions from the United States National Herbarium 10(5):221-443, 11 pls. MITCHELL-THOME, R.C. 1979. Pre-Quaternary stratigraphy of the Atlantic island. Geologische Rundschau 68(2):495—522. NIKOLAEY, V.A., AND D.M. HARwoop. 2000. Morphology and taxonomic position of the Late Cretaceous diatom genus Pomphodiscus Barker & Meakin. Micropaleontology 46(2):167-177. NIKOLAEY, V.A., AND D.M. HARwoop, D.M. 1997. New process, genus and family of Lower Cretaceous diatoms from Australia. Diatom Research, 12(1):43-53. OLSHTYNSKAYA, A.P. 1976. Diatom algae of the stratotypic section of the Tishkinskaya sediments. Geologichesky Zhournal 36(4):147-151. PREMA, P., AND T.V. DESIKACHARY. 1989. New and interesting fossil diatoms from the Indian Ocean. Phykos 28(1-2):127-131. Ratrray, J. 1890. A revision of the genus Coscinodiscus, Ehrb., and of some allied genera. Proceedings of the Royal Society of Edinburgh 16:449-692, 3 pls. RoBINSON, J.H. 1934. The study of the organic remains in the oceanic beds of Barbados with special reference to the diatoms. Journal of the Barbados Museum and Historical Society 1(2):|5 pp.] ROBINSON , J.H. 1936. The occurrence and distribution of the diatoms in theoceanic beds of Barbados. II. Journal of the Barbados Museum and Historical Society 4(1):[4 pp.]. ROBINSON , J.H. 1941. The occurrence and distribution of the diatoms in theoceanic beds of Barbados. III-IV. Journal of the Barbados Museum and Historical Society 4(4):[4 pp.]. Rybe, J.W. 1962. A study of Annellus californicus Tempere. Journal of the Royal Microscopical Society 80(3):243-246. Sims, P.A. 1989. Some Cretaceous and Paleogene species of Coscinodiscus: a micromorphological and sys- tematic study. Diatom Research 4(2):351-371. STRELNIKOVA, N.I. 1992. Diatoms of the Paleogene. St. Petersburg University, St. Petersburg. 312 pp. WILLIAMS, D.M. 1988. An illustrated catalogue of the type specimens in the Greville diatom herbarium. Bulletin of the British Museum (Natural History) 18(1):1—148, 74 pls. STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE PLATES S15 316 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 1 Light microscopy. Figures 1-7, 9-10. Porodiscus splendidus Greville. Figure 1. Lectotype (internal view), BM 10602. Figures 2, 3. BM10399. Figure 4. CAS 611235 (Cambridge). Figure 5. CAS 611174 (Joe’s River). Figure 6. CAS 611174 (Joe’s River). Figure 7. BM 2844. Figure 9. CAS 611235 (Cambridge). Figure 10. CAS 611193 (Springfield). Figures 8, 11, 12. Porodiscus splendidus var. excentri- cus (Olsht.) Olshtinskaya new comb. Figure 8. Olst. Collection, Ukraine, Staroverovka. Figure 11. CAS 611174 (Joe’s River). Figure 12. CAS 611174 (Joe’s River). Scale bars = 10 um. SY DIATOM GENUS PORODISCUS GREVILLE STRELNIKOVA ET AL. 318 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 2 Light microscopy. Figures 13-16. Porodiscus nitidus Greville. Figure 13. Valves with spines. BM 2751. Figure 14. Valves with no spines. BM 2751. Figure 15. Valves with spines. BM 2812. Figure 16. Valves with no spines. CAS 611174 (Joe’s River). Figure 17. Porodiscus conicus Greville. (non-conical valve; Synonym: Porodiscus elegans Greville.) BM 2745. Figure 18. Porodiscus conicus Greville. BM 2812. Holotype. Figures 19, 21. Craspedodiscus ellipticus Gombos. Figure 19. BM 1063. Figure 21. CAS 611193 (Springfield). Figure 20. Coscinodiscus excavatus Greville. BM 2745. Figures 22—23. Porodiscus oblongus Greville. Figure 22. BM 2045. holotype? Fig 23. CAS 611193 (Springfield). Scale bars = 10 um. STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE + * ‘oe ii Pe, rf ‘e ene) > i fs ae 4 ney 7 * r ae ks oh ae . 320 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 3 Scanning electron microscopy (SEM). Figures 24-30. Porodiscus splendidus Greville. Figures 24-26. External views. Figure 24. CAS 611226 (Chimborazo). Figure 25. CAS 711174 (Joe’s River). Figure 26. CAS 611193 (Springfield). Figures 27-30. Internal views. Figures 27,28. Internal “sack”. CAS (611193) Springfield. Figure 29. Broken “sack”. Figure 30. Broken valve. Figures 29-30. CAS 611174 (Joe’s River). Scale bars = 10 um. STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE Coerere OOF a; s@ we 02 ges o- 8¢e 99 af 798 a, 745 p88 » a revetese ay al ihe POL 06%,"5! ‘ (a @eea @eetes LoS) i) 32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 14 | PLATE 4 SEM. Figures 31—33. Porodiscus splendidus Greville. Valves with connection band. Figure 31. External view. Figures 32, 33. Internal view. CAS Joe’s River 611174. Figures 34-36. Porodiscus splendidus var. excentricus (Olsht.) Olshtinskaya new comb. Figure 34. External view. Figures 35-36. Internal views. Figures 34, 36. Staroverovka. Figure 35. CAS 611174 (Joe’s River). Scale bars = 10 um. 323 A ET AL.: DIATOM GENUS PORODISCUS GREVILLE STRELNIKO\ 324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 5 SEM. Figures 37-40. Porodiscus splendidus var. corniger (Greville) Fourtanier n. comb. Different positions of one frustule. Figure 40. “Sack” sur- rounded by hyaline ring. CAS 611226 (Chimborazo). Figures 41-42. Porodiscus nitidus Greville. External views; valves with spines. CAS 611174 (Joe’s River). Scale bars = 2 um (Figure 40), 10 um (Figures 37-39, 41-42). N oa) 1 GENUS PORODISCUS GREVILLE A ET AL.: DIATOM STRELNIKOV @ Oe, * of SR@e@eee ee ee © Feageeec 326 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 6 SEM. Figures 43-50. Porodiscus nitidus Greville. Figures 43-46. External views. Valves with velum. Figures 47-50. Internal views. Figures 43-44. Valves without spines. Figures 47, 49. Valves with spines. Figure 49. Rimoportulae at the ends of hyaline lines. Figures 44, 46. CAS 611226 (Chimborazo). Figures 45—S0. CAS 611174 (Joe’s River). Scale bars = 10 um. AN foe) DIATOM GENUS PORODISCUS GREVILLE ABT AL: STRELNIKOV e s Te 2% rw - ames: * ie 328 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 7 SEM. Figures 51—56. Porodiscus nitidus Greville. CAS 624819, Tasman Sea, DSDP Sample 206-16-5, 111-113 cm. Figures 51-55. External views. Figures 51, 54. Valves with spines. Figure 52. “Sack” surrounded by hyaline ring. Figure 53. The bases of broken spines, rimoportulae on the margin of valve on hyaline line under the spines. Figure 55. Velum. Figure 56. Internal view, rimoportulae. Scale bars = 25 um (Figure 51), 15 um (Figure 54), 6 um (Figure 53), 4 um (Figure 52), 3 um (Figures 55—56). STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 330 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES HI ) | Volume 55, No. 14 © PLATE 8 SEM. Figures 57-61. Porodiscus oblongus Greville. CAS 611193 (Springfield). Figures 57-58. External views. Velum and “sack” surrounded by hya- line ring. Figures 59-61. Internal views. Figures 59, 61. “Sack” and broken “sack.” Figure 60. Rimoportulae on the margin of valve. Figures 62-63. Craspedodiscus ellipticus Gombos. CAS 611193 (Springfield). Figure 62. Internal view. Rimoportulae on the margin of the valve. Figure 63. External view. Rimoportulae on the margin. Figure 64. Porodiscus conicus Greville ? (non-conical valve; Syno- nym: Porodiscus elegans Greville.) CAS 611235 (Chimborazo). Scale bars = 5 um (Figure 60), 10 um (Figures 58—59, 61-63) (aa) ioe) DIATOM GENUS PORODISCUS GREVILLE A ET AL. STRELNIKO\ 332 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 9 SEM. Figures 65—70. Craspedodiscus umbonatus Greville. Figures 65—66. External views. Figure 65. Velum. Figures 67-70. Internal views. Figures 67, 69. “Sack”. Figures 68, 70. Rimoportulae on the valve margin. Figures 71-72. Coscinodiscus sp. Figure 71. External view. Figure 72. Internal view. Rimoportulae on the valve margin. Figures 65-72. CAS 611174 (Joe’s River). Scale bars = 1 um (Figures 68, 70), 10 um (Figures 65—67, 69, 71—72) lag) ioe) DIATOM GENUS PORODISCUS GREVILLE ASAE: STRELNIKOV 334 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 10 SEM. Figure 73. Coscinodiscus sp. External view. Figure 74. Craspedodiscus umbonatus Greville. Rimoportulae on the margin. Figures 73-74. CAS 611174 (Joe’s River). Figure 75. Coscinodiscus radiatus Ehrenberg with slightly depressed center and rimoportulae on the valve face. Staroverivka (CAS 624820). Figures 76-77. Craspedodiscus barronii Bukry. Figure 76. Internal view. Rimoportulae on the margin. Figure 77. External view. Depressed center. CAS 611226 (Chimborazo). Figures 78-80. Coscinodiscus excavatus Greville. Figures 78-79. External view. Figure 79. Separate rimoportulae on the valve. Figure 80. Internal view. CAS 618793. Barbados, lower Oligocene. Scale bars = 1 um (Figure 74), 10 um (Figures 73, 75-77), 7 um (Figure 79), 60 um (Figures 78, 80) ioe) A ET AL.: DIATOM GENUS PORODISCUS GREVILLE STRELNIKOV a : ees © ow 336 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 14 PLATE 11 Figures 81—86. Craspedodiscus coscinodiscus Ehrenberg. (CAS 621025, Trinidad). Figure 81. Internal view. Figures 82—85. External view. Figures 81-85. SEM. Figure 86. LM. Figure 87. Annellus californicus var. hannai Barron. Holotype USNM 304194, sample Mf5208, southern California Continental Borderland. SEM. Figures 88-90. Annellus californicus var. californicus Tempére. Figures 88-89. Valve view. Figure 90. Mantle view. Sample Mf 5192, southern California Continental Borderland. Figure 88. SEM. Fig 89. 90. LM. Figure 91. Craspedodiscus rhydei Barron. DSDP 77B-28-2, 28-30 cm. SEM. Figure 92. Coscinodiscus excavatus Greville. Inside view. SEM (CAS 621025, Trinidad). Scale bars = 20 um (Figures 81—82, 84), 10 um (Figures 83, 85-91), 100 um (Figure 92) STRELNIKOVA ET AL.: DIATOM GENUS PORODISCUS GREVILLE 839) Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15, pp. 338-357, 6 pls. September 30, 2004 Mesodictyopsis Khursevich, Iwashita, Kociolek and Fedenya A New Genus of Diatoms in the Class Coscinodiscophyceae (Bacillariophyta) from Upper Miocene Sediments of Lake Baikal, Siberia G.K. Khursevich!?, J.P. Kociolek?, T. Iwashitat, S.A. Fedenya!, M.I. Kuzmin5, T. Kawai®, D.F. Williams?, E.B. Karabanov2>, A.A. Prokopenko2.’, and K. Minoura‘ ! Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk 220141, Republic of Belarus; Email: khurs@ns.igs.ac.by. * Baikal Drilling Project, Department of Geological Sciences, University of South Carolina, Columbia, SC 29208, USA. 3 Diatom Collection, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103, USA. 4 Institute of Geology and Paleontology, Faculty of Science, Tohoku University, Sendai 980, Japan. > Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk 664033, Russia. © National Institute for Environmental Studies, Tsukuba 305, Japan. 7 United Institute of Geology, Geophysics and Mineralogy, Siberian Branch of Russian Academy of Sciences, Novosibirsk 630090, Russia A new freshwater genus Mesodictyopsis, studied with light (LM) and scanning elec- tron microscopy (SEM), has been distinguished in the class Coscinodiscophyceae (Bacillariophyta) from Upper Miocene sediments of Lake Baikal. It is diagnosed by a combination of the following morphological features: a) location of the areola cribrum inside the loculus; b) arrangement of areolae on the valve face in uniseriate striae or in narrow or broad fascicles subdivided by hyaline strips or costae; c) var- ious positions of rimoportula(e) on the valve surface; d) presence of marginal fulto- portulae with three satellite pores; e) presence of valve face fultoportulae. Six new species are described. Another species, Stephanodiscus dzhilindus Khursevich, from the Dzhilinda basin of the Transbaikal area, appears to also share these same fea- tures. We propose to transfer this species to Mesodictyopsis. Mesodictyopsis is distin- guished from its close ally, Mesodictyon Theriot and Bradbury, in the structure and placement of fultoportulae and position of rimoportula(e). Key words: Bacillariophyta, Coscinodiscophyceae, diatoms, new genus, Mesodictyopsis, Late Miocene, Lake Baikal. Lacustrine sediments of long continuous sections from Lake Baikal are important archives for studying the evolution of freshwater centric diatoms. Plio-Pleistocene diatom succession is well documented in the sedimentary record of Lake Baikal from BDP 96-1 and BDP 96-2 cores (Grachevy et al. 1998: Khursevich et al. 1998, 2000, 2001la—c). The above-mentioned sections, as well as the new 600-m long drill core BDP-98, have recovered a continuous diatom record of the past 10 Ma (BDP-Members 2000) and show that the Pleistocene time interval is characterized by active speciation within the genus Stephanodiscus Ehrenberg. The Pliocene epoch is distinguished by the appearance, speciation and extinction of the genus Stephanopsis Khursevich & Fedenya. The Late Miocene differs by the appearance, speciation and extinction of a group of species whose 338 KHURSEVICH ET AL.: NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL 589 features do not allow placement into any previously described genus. The description of this new diatom genus, Mesodictvopsis, including six species from the Upper Miocene deposits of Lake Baikal, is given in the present report MATERIALS AND METHODS The 600 m long borehole BDP-98 was drilled by the Russian company “Nedra Enterprise” during winter 1998 using the Baikal-2000 drilling complex placed on a barge. The drilling was per- formed on the Academician Ridge of Lake Baikal at 53°44'48” N and 108°24'34"” E in water depth of 333 m. Continuous samples were collected to a depth of 600 m. Total core recovery was over 95%. Sediments consist of alternating biogenic diatomaceous ooze and terrigenous clay intervals. The age model for the BDP-98 section is based on the correlation of event/reversal boundaries with the reference polarity time scale (Cande and Kent 1995). Proceeding from paleomagnetic investi- gations and analysis of the Earth’s orbital frequencies in the climatic record of the core, the sedi- ments at a depth of 600 m have a preliminary date of 10.3 Ma (BDP-Members 2000). The Upper Miocene deposits occur in the BDP-98 core at the depth interval between approximately 600 and 221 m according to the preliminary age model for this core. Permanent diatom slides were prepared with identical volumes of material according to the method described in Grachev ef al. (1997). Diatoms were enumerated along vertical and horizon- tal transects of permanent slides in 50 or 200 fields of view depending upon diatom concentration. The distributions of total diatoms, as well as of dominant individual planktonic diatom genera including Mesodictvopsis, were quantitatively determined for the interval of core depth between 600 and 221 m. Elsewhere, ranked abundances (mln valves per gram of dry sediments) were used. Specimens were examined using oil immersion light microscopy (Ergaval brightfield, NA=1.25 in IGS, Belarus; DMRB with DIC optics, NA=1.4 in CAS) and scanning electron microscopy (JEOL JSM-35C in IGS, Belarus and JEOL JSM-6330F in IGPS, Sendai, Japan). To describe the new fos- sil diatom species, the terminology recommended by Ross et al. (1979) was used. The structural elements on the valve were measured using the procedure of Anonymous (1975) on not less than 30 complete diatom valves of each new taxon. RESULTS Genus Mesodictyopsis Khursevich, Iwashita, Kociolek, and Fedenya, gen. nov. DESCRIPTION: Frustules low-cylindrical with few intercalary bands. Valves circular, 3.0—91.5 um in diameter, more or less flat; occasionally the central area is markedly concave or convex. Areolae loculate, having a cribrum on the inside of each loculus and a foramen on both the inter- nal and external valve surfaces. Areolae arranged in radial rows of unequal length or in narrow or broad fascicles divided by hyaline strips or costae. Mantle not separated from the valve face by a sharp angle, but curved outwards; 1—5 um in wide, with areolae closer and smaller than those on the valve face; the outermost row of areolae larger. On the valve surface one rimoportula, occupy- ing various positions (at the center, near the center, in the submarginal zone of the valve face, on the valve face/mantle junction or on the mantle) or several rimoportulae (up to 7) forming a ring on the boundary between the valve face and mantle. Internal opening of the rimoportulae is a slightly raised slit; external opening is a pore or small tube. Fultoportulae on the valve face with three, rarely two or four, satellite pores internally, represented by small external apertures. Marginal fultoportulae with three satellite pores on the internal valve surface, appearing as small openings or tubes at the base of hyaline strips externally. A ring of spines is almost always present at the valve face/mantle junction. 340 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15 TYPE SPECIES.— Mesodictyopsis academicus Khursevich, lwashita, Kociolek, and Fedenya, Sp. nov. TYPE LOCALITY.— The underwater Academician Ridge of Lake Baikal, Russia, borehole BDP-98 (53°44'48"N; 108°24'34”E). Abundant in the Upper Miocene deposits. COMMENTS.— The new genus differs from Mesodictyon Theriot and Bradbury (Theriot and Bradbury 1987) in several aspects. The medial velum of Mesodictyopsis (a clean siliceous plate within loculate areola) is of a type different from that seen in Mesodictyon (a distinct medial cribrum in areola in Mesodictyon). In Mesodictvopsis. the position of rimoportulae is highly vari- able, while in Mesodictyon the rimoportula is always found on the valve mantle, behind the mar- ginal fultoportula internally. Mesodictvon is known to have occurred from 12—11 Ma to 7 Ma in the western USA, Peru, France, Bulgaria, Belarus, and in Lake Baikal (Mesodictvon nativus has the age range ca. up to 6.5 Ma). Mesodictvopsis is endemic to Lake Baikal region with the age range from ca.7—5.1 Ma (the end of Late Miocene-earliest Pliocene). In Lake Baikal there is a long, con- tinuous sedimentary record where there are several endemic freshwater centric diatom genera (Concentrodiscus, Stephanopsis) which replace each other from bottom to top and have some com- mon inherited morphological features (such as the various inconstant position of rimoportula on the valve surface including some taxa with the location of rimoportulae in or near the center of the valve). This situation is wanting in Mesodictvon species. Lake Baikal has very high levels of endemism, not only of species, but also genera (among many organisms including diatoms). The extinct endemic genera Mesodictvopsis and Stephanopsis, for example, have marginal fultoportu- lae with three satellite pores and the valve face fultoportulae with three satellite pores. We know many closely related centric genera with external cribra (Actinocyclus, Cestodiscus, Cosmiodiscus and others), many genera with internal cribra (Thalassiosira, Concentrodiscus, Stephanodiscus and others), and now we recognize several genera with a medial velum. We have recognized six new species in this genus; these are described below. Mesodictyopsis academicus Khursevich, Iwashita, Kociolek, and Fedenya, sp. nov. (Plate 1, Figs 1-3, 6; Plate 2, Figs 1-6. Plate 1, figure | is of the holotype.) DESCRIPTION.— Valves circular, 9.5—50.0 um in diameter. Areolae 10—25 in 10 um along the valve radius, arranged in radial, uniseriate striae or extending from the center and becoming bi- or triseriate fascicles in the submarginal zone of the valve face. Hyaline strips or costae divide unise- riate striae or fascicles from each other and continue from the valve face to the mantle, (6)8—16 in 10 um. A single rimoportula, usually positioned in the valve center or near the center. Valve face fultoportulae, 2—14, each with three satellite pores, situated within the central area in radial areolar rows replacing 1-4 areolae per row. Valve mantle (up to 2 um.in height) with 2-5 small areolae in a vertical row, with 36-40 rows of areolae in 10 um. Marginal fultoportulae with three satellite pores, (6)8-16 in 10 um, located at the base of each hyaline strip, rarely every second or third one. A ring of irregularly located spines may be present at the valve face/mantle junction. Ho.LoryPe.— Slide No. 3495a, BDP-98, core 198-1 (18 cm), deposited at the Institute of Geo- logical Sciences, National Academy of Sciences of Belarus, Minsk, Belarus. IsoryPe.— Slide No. 3495b, BDP-98, core 198-1 (18 cm), deposited at the Institute of Geo- chemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TYPE MATERIAL.— Housed in the G. K Khursevich Collection at the Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk, Belarus; Baikal Drilling Project Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia, and in Diatom Collection, California Academy of Sciences, San Francisco, USA. KHURSEVICH ET AL.: NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL 34] TYPE LOCALITY.— The underwater Academician Ridge of Lake Baikal, Russia, borehole BDP-98, depth 334-461 m from sediment surface. Mesodictyopsis medius Khursevich and Iwashita, sp. nov. (Plate 1, Fig. 11; Plate 3, Figs. 1-6. Plate 1, Fig. 11 is of the holotype.) DESCRIPTION.— Valves circular, more or less flat, 11.0-24.5 tm in diameter. Areolae 14-16(20) in 10 um along the valve radius, arranged in radial rows of unequal length separated by hyaline strips, 14-18 in 10 um. A single rimoportula situated in the valve center and surrounded by a hyaline ring. Valve face fultoportulae (from 3 to 9), each with three satellite pores, form an irreg- ular ring at a distance of '/—'/ from the valve center. Valve mantle (1—2 um in height) has 2—4 small areolae in a vertical row (35—40 areolar rows in 10 tm near the edge of mantle) and a marginal ring of fultoportulae with three satellite pores, 8—12(14) in 10 um, located at the base of each or every second or third hyaline strip. Spines may be present at the valve face/mantle junction. Ho.otyPe.— Slide No. 108616, BDP-98, core 231-1 (91 cm) deposited at the Institute of Geology and Paleontology, Graduate School of Science, Tohoku University, Sendai, Japan. TYPE MATERIAL.— Housed in the IGPS Collection at the Institute of Geology and Paleontology, Graduate School of Science, Tohoku University, Sendai, Japan; Baikal Drilling Project Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TYPE LOCALITY.— The underwater Academician Ridge of Lake Baikal, Russia, the borehole BDP-98, depth 403-424 m from sediment surface. COMMENTS.— This species differs from Mesodictvopsis academicus by the presence of non- fasciculated striae of areolae only and valve face fultoportulae forming an irregular ring in the mid- dle of the valve radius. Mesodictyopsis peculiaris Khursevich, Kociolek, and Fedenya, sp. nov. (Plate 1, Fig. 13; Plate 4, Figs. 1-2. Plate 1, Fig. 13 is of the holotype.) DESCRIPTION.— Valves circular, flat or slightly convex, 3.0—9.5 um in diameter. Areolae, up to 20 in 10 um along the radius, located in single radial rows divided by hyaline strips, 15—20 in 10 um. Not all rows of areolae reach the valve center; hence the center not densely structured. One fultoportula with two satellite pores occurs near the valve center. A single rimoportula occurs in the submarginal zone of the valve face, between two closely located marginal fultoportulae. 4—6 fulto- portulae with three satellite pores on the mantle; among which two are placed close to one anoth- er, the rest widely spaced. Valve mantle shallow (up to 1 um high), consisting of 1-2 small areo- lae in a vertical row. Spines positioned at every hyaline strip at the valve face/mantle junction; often broken. Ho.otype.— Slide No. 3495a, BDP-98, core 198-1 (18 cm), deposited at the Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk, Belarus. IsoryPE.— Slide No. 3495b, BDP-98, core 198-1 (18 cm), deposited at the Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TYPE MATERIAL.— Housed in the GK. Khursevich Collection at the Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk, Belarus; Baikal Drilling Project Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia, and in Diatom Collection, California Academy of Sciences, San Francisco, USA. Type LOCALITY.— The underwater Academician Ridge of Lake Baikal, Russia, borehole BDP-98, depth 328-377 m from sediment surface. 342 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15 Comments: This species differs from Mesodictyopsis medius by the smaller size of valves, as well as the specific location of a single rimoportula and marginal fultoportulae on the valve sur- face. Mesodictyopsis singularis Khursevich, Iwashita, and Fedenya, sp. nov. (Plate 1, Figs. 7, 10, 14-15; Plate 4, Figs. 3-5. Plate 1, Fig. 7 is of the holotype.) DESCRIPTION.— Valves circular, more or less flat, 4.0-17.2 um in diameter. Areolae 12-16 (20) in 10 um along the radius, grouped in uniseriate radial striae of unequal length, separated by hyaline strips, 14-18 in 10 um. One-several fultoportulae, each with three satellite pores, found near the valve center. A single rimoportula occurs just inside the ring of marginal fultoportulae. Marginal fultoportulae with three satellite pores, (6)8—10 in 10 um, located at the base of each or every second, rarely every third hyaline strip on the mantle. Valve mantle (up to 1 um high) perfo- rated by 1-3 small areolae in a vertical row (up to 40 rows of areolae in 10 um). Small pointed spines usually present at every hyaline strip on the boundary between the valve face and mantle. Ho.LortypeE.— Slide No. 3040a, BDP-98, core 174-1 (68 cm), deposited at the Institute of Geo- logical Sciences, National Academy of Sciences of Belarus, Minsk, Belarus. IsoType.— Slide No. 3040b, BDP-98, core 174-1 (68 cm), deposited at the Institute of Geo- chemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TYPE MATERIAL.— Housed in the G.K. Khursevich Collection at the Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk, Belarus, and Baikal Drilling Project Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TYPE LOCALITY.— The underwater Academician Ridge of Lake Baikal, Russia, borehole BDP-98, depth 304-452 m from sediment surface. COMMENTS.— This species differs from Mesodictyopsis peculiaris by the more dense place- ment of marginal fultoportulae. Mesodictyopsis baicalensis Khursevich, [washita, Kociolek, and Fedenya, sp. nov. (Plate 1, Figs. 4-5, 8, 17-18; Plate 5, Figs. 1-6, Plate 6, Figs 1-6. Plate 1, Fig. 5 is of the holotype.) DESCRIPTION.— Valves circular, with markedly concave or convex central area, 15.5-91.0 um in diameter. Areolae, 11—18(20) in 10 um along the valve radius, arranged in radial uniseriate uneven striae within the central area becoming bi- multiseriate fascicles (up to 5 areolar rows ) in the submarginal zone of the valve face. Fascicles of areolae divided by hyaline strips or costae, 5- 7 in 10 um, continuing from the valve face to the mantle. | to 7 valve face fultoportulae, each with three satellite pores (rarely one fultoportula may have four satellite pores), occur in the valve cen- ter or near the center replacing 1-4 areolae per row. Rimoportulae (1—7) located in the upper part of the valve mantle a little below the spine insertion; appear as short tubes, externally, and slight- ly raised slit oriented variously, internally. Rimoportulae may be positioned both on hyaline strips and in radial areolar rows. Valve mantle (up to 5 um high) with 2—8 small areolae in a vertical row (30-40 rows of areolae in 10 um) and a ring of marginal fultoportulae located at the base of each or every second, rarely every third, hyaline strip. Marginal fultoportulae having three satellite pores internally and small tubes externally. Conic spines are spaced irregularly at the valve face/mantle junction. HoLotyPe.— Slide No. 2810a, BDP-98, core 150-1 (18 cm), deposited at the Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk, Belarus. KHURSEVICH ET AL.: NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL 43 1S) IsotyPE.— Slide No. 2810b, Bdp-98, core 150-1 (18 cm), deposited at the Institute of Geo- chemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TYPE MATERIAL.— Housed in the GK. Khursevich Collection at the Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk, Belarus; Baikal Drilling Project Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia, and in Diatom Collection, California Academy of Sciences, San Francisco, USA. TyPE LOCALITY.— The underwater Academician Ridge of Lake Baikal, Russia, borehole BDP-98, depth 259-302 m from sediment surface. COMMENTS.— This species differs from Mesodictvopsis singularis mainly by the larger size of valves, as well as by the presence of distinct areolar fascicles towards the margin of the valve face. Mesodictyopsis similis Khursevich and Fedenya, sp. nov. (Plate 1, Figs. 9, 16; Plate 4, Figs. 6—7. Plate 1, Fig. 16 is of the holotype.) DESCRIPTION.— Valves circular, flat or slightly convex, 3.5—-10.0 um in diameter. Areolae, 15-20 in 10 um along the radius, arranged in short radial uniseriate striae extending from '/ to about ‘> of the radius from the valve margin to the center. Hyaline strips separating uniseriate stri- ae from each other 15—20 in 10 um. Central area hyaline except the presence of a single valve face fultoportula with two satellite pores. One rimoportula occurs at the same level with marginal ful- toportulae having three satellite pores. Valve mantle shallow (up to | um high) with one areolae placed in a vertical row. Spine occurrence and placement difficult to determine, usually broken. Ho.orype.— Slide No. 2525a, BDP-98, core 142-1 (94 cm), deposited at the Institute of Geo- logical Sciences, National Academy of Sciences of Belarus, Minsk, Belarus. IsoTyPE.— Slide No. 2525b, BDP-98, core 142-1 (94 cm), deposited at the Institute of Geo- chemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TYPE MATERIAL.— Housed in the GK. Khursevich Collection at the Institute of Geological Sciences, National Academy of Sciences of Belarus, Minsk, Belarus, and Baikal Drilling Project Collection, Institute of Geochemistry, Siberian Branch of Russian Academy of Sciences, Irkutsk, Russia. TyPE LOCALITY.— The underwater Academician Ridge of Lake Baikal, Russia, borehole BDP-98, depth 254-304 m from sediment surface. COMMENTS.— This species differs from Mesodictyopsis peculiaris and M. singularis by the different location of a single rimoportula on the valve surface. DISCUSSION There are now two freshwater genera (Mesodictyon and Mesodictyopsis) of the family Stephanodiscaceae Makarova which have the medial cribrum within the loculate areolae. This fea- ture is the main diagnostic character of these genera. Mesodictyon has no valve face fultoportulae and possesses marginal fultoportulae with two satellite pores, while Mesodictvopsis has not only marginal fultoportulae with three satellite pores, but also valve face fultoportulae. In addition, the rimoportula(e) is typically on the mantle outside of the ring of marginal fultoportulae in Mesodictyon (Theriot 1990). As far as the species of Mesodictyopsis are concerned, they can be separated into three morphological groups with respect to the rimoportula(e) position on the valve surface. One morphological group of Mesodictyopsis is composed of species characterizing by the location of a single rimoportula at the valve face/mantle junction or in the submarginal zone of the 344 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15 valve face (M. singularis, M. peculiaris). The second group is based on the placement of a single rimoportula in the valve center or near the center (IZ. academicus, M. medius). Finally, the third morphological group is distinguished by the position of rimoportula(e) on the valve mantle (Z baicalensis, M. similis). In general, species of Mesodictyopsis were found in the sediments of BDP-98 section in the interval of depth 461—254 m. According to the preliminary age model for the BDP-98 drill core (BDP-Members 2000), the appearance, development and extinction of Mesodictyopsis in the ancient basin proceeded during the Late Miocene period from ~ 8.5 to 6.0 Ma. In our opinion, the species Stephanodiscus dzhilindus Khursevich, which occurs in the Miocene deposits within the Dzhilinda depression of the Transbaikal area (Khursevich 1994) should also be transferred to the genus Mesodictyopsis. Although cribra were not observed in spec- imens of Stephanodiscus dzhilindus, presence of funnel-like areolae, marginal fultoportulae with three satellite pores and a single valve face fultoportula suggest that Mesodictyopsis is its best generic placement. Hence, the new combination is proposed: Mesodictyopsis dzhilindus (Khursevich) Khursevich. BASIONYM.— Stephanodiscus dzilindus Khursevich (Khursevich 1994: Morphology and tax- onomy of some centric diatom species from the Miocene sediments of the Dzhilinda and Tunkin hollows. Pages 271-272 and fig. 3 in J.P. Kociolek, ed., Proceedings of the 11 International Diatom Symposium, San Francisco. Memoirs of the California Academy of Sciences no. 17). The extinct genus Mesodictyopsis is characteristic both of the Upper Miocene sediments from Lake Baikal and the Transbaikal area. ACKNOWLEDGMENTS This work was supported by National Scientific Foundation of USA (NSF) grants EAR-93- 1720401, EAR-96-14770 and the Siberian Branch of Russian Academy of Sciences. Cores for this study were made possible through the Baikal Drilling Project (BDP) supported by NSF and International Continental Scientific Drilling Program (ICDP), the Russian Academy of Sciences, the Russian Ministry of Geology, the Science and Technology Agency (STA) of Japan. Authors wish to express their gratitude to B.N. Khakhaev, L.A. Pevzner and V.V. Kochukov of the Nedra Drilling Enterprise for organizing and providing the deep drilling of Lake Baikal sediments. We also thank A.N Gvozdkov and N. Soshina of the Institute of Geochemistry for technical assistance with the BDP-98 core and smear slide preparation. REFERENCES ANONYMOUS. 1975. Proposal for a standardization on diatom terminology and diagnoses. Nova Hedwigia 53:323-354. BDP-Mempers. 2000. Paleoclimatic record in the late Cenozoic sediments of Lake Baikal (600-m deep- drilling data). Russian Geology and Geophysics 41:3-32. CANDE, S.C., AND D.V. KENT. 1995. Revised calibration of the geomagnetic polarity time scale for the late Cretaceous and Cenozoic. Journal of Geophysical Research 100:6093-6095. GRACHEV, M.A., E.V. LikHosHwWay, S.S. VoROBYOVA, O.M. KHLYSTOV, E.V. BEZRUKOVA, E.V. VEINBERG, E.L. GOLDBERG, L.Z. GRANINA, E.G. KORNAKOVA, F.I. LAzo, O.V. LEvinA, P.P. LETUNOVA, P.V. OTINOV, V.V. PrroG, A.P. FEpotTov, S.A. IASKEVICH, V.A. BOBROV, F.V. SUKHORUKOV, V.I. REZCHKOV, M.A. FEDORIN, K.Z. ZOLOTAREV, AND V.A. KRAVCHINSKY. 1997. Signals of the paleoclimates of the Upper Pleistocene in the sediments of Lake Baikal. Russian Geology and Geophysics 38:957-980. KHURSEVICH ET AL.: NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL 345 GRACHEV, M.A., S.S. VoRoByovaA, E.V. LIKHOSHWAY, E.L. GOLDBERG, G.A. ZIBOROVA, O.V. LEVINA, AND O.M. KutysToy. 1998. A high-resolution diatom record of the paleoclimates of East Siberia for the last 2.5 My from Lake Baikal. Quaternary Science Reviews 17:1101—1106. KHURSEVICH, G.K. 1994. Morphology and taxonomy of some centric diatom species from the Miocene sedi- ments of the Dzhilinda and Tunkin hollows. Pages 269-280 in J.P. Kociolek, ed., Proceedings of the 11th International Diatom Symposium, San Francisco. Memoirs of the California Academy of Sciences 17. KHURSEVICH, G.K., S.A. FEDENYA, E.B. KARABANOV, A.A. PROKOPENKO, D.F. WILLIAMS, AND M.I. KuzMIN. 2001a. Late Cenozoic diatom record from Lake Baikal sediments. Pages 451—460 in A. Economou-Amilli, ed., Proceedings of the 16th International Diatom Symposium. Athens, University of Athens, Greece. KHURSEVICH, G.K., E.B. KARABANOV, A.A. PROKOPENKO, D.F. WILLIAMS, M.I. KUZMIN, S.A. FEDENYA, A.N. GVOZDKOV, AND E.V. KERBER. 2001b. Detailed diatom biostratigraphy of Baikal sediments during the Brunhes chron and climatic factors of species formation. Russian Geology and Geophysics 42:108—129. KHURSEVICH, G.K., E.B. KARABANOV, A.A. PROKOPENKO, D.F. WILLIAMS, M.I. KUZMIN, S.A. FEDENYA, AND A.N. Gvozpkov. 2001c. Insolation regime in Siberia as a major factor controlling diatom production in Lake Baikal during the past 800,000 years. Quaternary International 80-81:47-58. KHURSEVICH, G.K., E.B. KARABANOV, D.F. WILLIAMS, M.I. KUZMIN, AND A.N. GvOZDKOV. 1998. Pliocene- Pleistocene geochronology and diatom biostratigraphy of bottom deposits of Lake Baikal: new results of deep drilling cores. Pages 27—28 in Abstracts of the International Symposium “Paleoclimates and the Evolution of Paleogeographic Environments in the Earths Geological History.” Petrozavodsk. KHURSEVICH, G.K., E.B. KARABANOV, D.F. WILLIAMS, M.I. KUZMIN, AND A.A. PROKOPENKO. 2000. Evolution of freshwater centric diatoms within the Baikal rift zone during the late Cenozoic. Pages 146-154 in Koji Minoura, ed., Lake Baikal: A Mirror in Time and Space for Understanding Global Change Processes. Elsevier, Amsterdam. Ross, R., E.J. Cox, N.I. KARAYEVA, D.G. MANN, T.B.B. PADDOCK, R. SIMONSEN, AND P.A. Sims. 1979. An emended terminology for the siliceous component of the diatom cell. Nova Hedwigia 64:513-533. THERIOT, E. 1990. New species of Mesodictyvon (Bacillariophyta: Thalassiosiraceae) in Late Miocene lacus- trine deposits of the Snake River basin, Idaho. Proceedings of the Academy of Natural Sciences of Philadelphia 142:1-19. THERIOT, E., AND J.P. BRADBURY. 1987. Mesodictyon, a new fossil genus of the centric diatom family Thalassiosiraceae from the Miocene Chalk Hills Formation, western Snake River Plain, Idaho. Micropaleontology 33:356—367. 346 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15 Plate 1 Light Microscopy Species of Mesodictyopsis from the Upper Miocene sediments of BDP-98 drill core. Scale bar in fig. | =10 um for all figures. 1-3, 6. Mesodictyopsis academicus. Fig. 1. Holotype specimen, BDP-98 core, 198-1, 18 cm. 4-5, 8, 17-18. Mesodictvopsis baicalensis. Fig. 5. Holotype specimen, BDP-98 core, 150- 1, 18 cm. 7, 10, 14-15. Mesodictvopsis singularis. Fig. 7. Holotype specimen, BDP-98 core, 174-1, 68 cm. 9, 16. Mesodictyvopsis similis. Fig. 16. Holotype specimen, BDP-98 core, 142-1, 94 cm. 11-12. Mesodictvopsis medius. Fig. 11. Holotype specimen, BDP-98 core, 231-1, 91 cm. 13. Mesodictvopsis peculiaris Holotype specimen, BDP-98 core, 198-1, 18 cm. | j | | LLye® — < — < faa) = y, < — = S x on N 5 Z fa O = e = A aa Z, aa S g 2 Ss fo Z — a0 > aa fe = > LU N ee T } KHU 348 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES — Volume 55, No. 15 Plate 2 Scanning Electron Microscopy Mesodictyopsis academicus Figs. 1, 3-4. External views of the valve surface with small openings of valve face fulto- portulae near the valve center (arrows); scale bars = 10 um. Figs. 2, 5—6. Internal views of the valve surface with a single rimoportula and valve face fultoportulae with three satellite pores near the valve center (arrows). Scale bars = 10 um in Figs. 2-6; scale bar = 1 um in Fig. 5. 349 NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL RSEVICH ET AL. KHL e* eee « ° 5 o © c v x o SSOc00G0+ CO90CH6C ‘300 eis) 350 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 15 Plate 3 Scanning Electron Microscopy Mesodictyopsis medius Figs. 1-2, 4, 6. Internal views of the valve surface with a single rimoportula in the valve center and an irregular ring of valve face fultoportulae with three satellite pores located at the distance '/;—'/, from the valve center (arrows). Fig. 3. Fragment of internal view of the valve surface with a cribrum inside of loculus (arrow). Fig. 5. External view of the valve surface with a ring of valve face fultoportulae openings (arrows), scale bars =] um. — < 4 < on aa) y < — = © x i, N =) Z fa OC) 2 2 < A = Z jaa O © = S aa) Z KHURSEVICH ET AL B52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15 Plate 4 Scanning Electron Microscopy 1-2. Mesodictyopsis peculiaris 1. External view of the valve surface with a single opening of the valve face fultoportula near the center (arrow); scale bar = 2 um. 2. Internal view of the valve surface with a single rimoportula in the submarginal zone of the valve face (black arrow) and one valve face fultoportula with two satellite pores near the center (arrow); scale bar = | um. 3-5. Mesodictvopsis singularis 3, 5. External view of the valve surface with one opening of the valve face fultoportula near the center (arrow); scale bars = 1 um. 4. Internal view of the valve surface with a single valve face fultoportula with three satel- lite pores (arrow) and a ring of marginal fultoportulae with three satellite pores; scale bar = | um. 6-7. Mesodictvopsis similis 6. External view of the valve surface with a wide hyaline central area and one opening of the valve face fultoportula near the center (arrow); scale bar = 2 um. 7. Internal view of the valve surface with a single rimoportula located on the same level with marginal fultoportulae (arrows); scale bar = 2 um. ee Soe 353 NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL KHURSEVICH ET AL.: 354 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15 Plate 5 Scanning Electron Microscopy Mesodictyopsis baicalensis 1, 34. External views of the valve surface with a ring of conic spines at the valve face/man- tle junction and a single opening of the valve face fultoportula near the center (arrows); scale bars = 10 um in Fig. 1, 1 um in Figs. 3-4 2, 5—6. Interval view of the valve surface with a marginal ring of rimoportulae (arrows); scale bar =10 um Fig. 5, 1 um in Figs. 2, 6. KHURSEVICH ET AL.: NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL 355 ane sat! ¥ ty pene fatten eV Oy Ota Bey, 2g ow eoe0 Sb eo® te ®ow %o ® a ete ten o oe as «Se e aes eee eS. 356 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 15 Plate 6 Scanning Elelctron Microscopy Mesodictyopsis baicalensis 1, 3. External views showing irregularly-placed, elongate, conical spines at the valve face/mantle junction and short, thin openings of fultoportulae; scale bars = 10um in Fig. 1, 1 um in Fig. 3. 2, 4-6. Internal valve views showing pattern of areolae, ring of many marginal fultoportu- lae and central fultoportulae with three satellite pores (Fig. 6); scale bars = lum in Figs. 2, 4, 6 and 10 um in Fig. 5. eens ities KHURSEVICH ET AL.: NEW MIOCENE DIATOM GENUS FROM LAKE BAIKAL 3)5)]) Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16, pp. 358-372, 1 fig. September 30, 2004 The Pattern of the Lateral-line System on the Caudal Fin of Perccottus glenii Dybowski, 1877 (Teleostei: Odontobutidae), with Comments on the Arrangement of the Lateral-line System on the Caudal Fin of Gobioidei Harald Ahnelt and Josef Géschl Institute of Zoology, Department of Comparative Anatomy and Morphology, University of Vienna, Althanstrasse 14, A-1090 Vienna, Austria; Email: harald.ahnelt@Qunivie.ac.at The pattern of the lateral-line system on the caudal fin is surveyed in 120 gen- era and 200 species of gobioid fishes. In the majority of Gobioidei this sensory sys- tem is arranged in two general patterns on the caudal fin: Pattern 1 with four later- al lines, one transversal and three longitudinal, and Pattern 2 with three lateral lines, one transversal and two longitudinal. These lateral lines are formed by rows of free (superficial) neuromasts, except in the Rhyacichthyidae. In these most-basal Gobioidei, the longitudinal lateral lines are a combination of canals and free neuro- masts. The transversal lateral line in Gobioidei (including Rhyacichthyidae) is always formed by free neuromasts. This transversal row is discontinuous, consisting of a few short parts (plesiomorphic), or it is continuous and relatively long (apomor- phic). In some species, it is indistinct and difficult to separate from the longitudinal rows. Pattern 1 of the lateral-line system on the caudal fin is plesiomorphic for Gobioidei. As relatively rare deviations from these two patterns, an increase or a decrease in the number of longitudinal neuromast rows occurs in some Gobiidae. Perccottus glenii is the only known species of basal Gobioidei (including the Eleotridae) characterized by a specialized lateral-line pattern on the caudal fin. On each side of this fin, only two neuromast rows are developed: a discontinuous trans- versal row on its base and a long longitudinal row along its midline. This is the most specialized character state within the Odontobutidae. This research provides a fur- ther indication that the Odontobutidae are not monophyletic. The free neuromasts of the lateral-line system of Gobioidei are arranged in series of character- istic rows and aggregations on the head, the trunk and the caudal fin. Lateral-line canals are absent on the trunk and on the caudal fin except in the Rhyacichthyidae, the most basal Gobioidei (Miller 1973; Hoese and Gill 1993). The pattern of the cephalic lateral-line system has been widely used in the classification of Gobioidei (e.g., Sanzo 1911; Ijin 1930; Miller 1986; Takagi 1988; Akihito et al. 2000; Larson 2001), but detailed information on the neuromast pattern on the trunk and on the caudal fin is rare. In most species the free neuromasts on the body are tiny and hard to detect. They are also easy damaged or abraded during collecting or preservation. This is especially true for the caudal fin. In recent years more attention has been given to the neuromast pattern on the trunk and on the caudal fin (e.g., Ahnelt and Duchkowitsch 2001; Shibukawa et al. 2001; Ahnelt and Scattolin 2003). Gobioid fishes are characterized by two general patterns of the lateral-line system of the cau- 58 Uo AHNELT AND GOSCHL: CAUDAL FIN LATERAL-LINE SYSTEM IN GOBIOIDEI 359 dal fin: Pattern 1, with a transversal and three longitudinal lateral lines; Pattern 2, with a transver- sal and two longitudinal lateral lines (Ahnelt et al. 2000; Shibukawa et al. 2001; Ahnelt and Géschl 2003). The combination of lateral-line canals and free neuromasts is only known in the Rhyacichthyidae. In all other Gobioidei, only free neuromasts form the lateral-line system on the caudal fin. In some species, the transversal row is indistinctly developed and/or difficult to sepa- rate from the longitudinal rows. As a deviation from pattern 1, the number of longitudinal neuro- mast rows in some Gobiidae is increased (Mortara 1918) or decreased (Miller 1963). Here we describe the reduced pattern of the lateral-line system on the caudal fin of the odon- tobutid fish Perccottus glenii Dybowski, 1877. Compared with the plesiomorphic arrangement of the free neuromasts on the caudal fin of the other basal Gobioidei (Rhyacichthyidae, 7erateleotris, Odontobutidae and Eleotridae), P. glenii is characterized by a specialized arrangement of these neu- romasts, otherwise known only in two miniature Gobiidae. MATERIALS AND METHODS NOMENCLATURE.— Odontobutidae: According to Shibukawa et al. (2001) and Miller (2003), the intra-familial relationships of the Odontobutidae are unsolved and a redefinition of the Odontobutidae is needed. In the definition of the Odontobutidae, we follow, therefore, Hoese and Gill (1993) who recognize three genera within this family, Micropercops, Odontobutis and Perccottus. Rhyacichthyidae: Watson and Péllabauer (1998) did not assign Protogobius Watson and Péllabauer, 1998 to a gobioid family. In the phylogenetic tree of Akihito et al. (2000a), based on mitochondrial DNA, Rhyacichthys and Protogobius form a single cluster and were, therefore, united as Rhyacichthyidae by Shibukawa et al. (2001). In the definition of the Microdesmidae and the Ptereleotridae, we follow Thacker (2003). Institutional abbreviations follow Leviton et al. (1985) except for: CMN(FI) = Canadian Museum of Nature; IZUW = Institute of Zoology, University of Vienna; IECB = Institute of Ecology and Conservation Biology, University of Vienna. The following preserved specimens were examined (collection number, number of specimens, sex, SL in mm, sampling site). Length of specimens is given in standard length. The sex was deter- mined by the shape of the urogenital papillae: longer and more pointed in males, shorter and wider in females. MATERIAL EXAMINED Odontobutidae: Perccottus glenii: NMH 2003.26.1; 1 female, 76.8 mm; Hungary, Bodrog River, Satoraljaujhely. NMH 2003.27.1; 1 female, 85.3 mm; Hungary, Tisza River, Mindszent. NMH 2003.28.1; 1 male, 65.7mm; Hungary, Bodrog River, Satoraljaujhely. NRM 23932; 2 juve- niles, 25.9 — 26.8 mm; Russia, Amur River, 5 km E Troitskoye. USNM 105188; 1 male, 64.1 mm; Russia, Sakhalin Island, Khanka Lake. USNM 077008; 1 female, 91.6 mm; China, Sungari River near its junction with the Amur. IECB uncatalogued; | male, 101.8 mm and | female, 72.6 mm; Ukraine, Pond at Komarno, near River Dniestr. IECB uncatalogued; | male, 79.6 mm and | female, 104.5 mm; Ukraine, Pond at Komarno, near River Dniestr. Rhyacichthyidae: Protogobius attiti Watson & Péllabauer, 1998: NMW 94266, | male, 67.7 mm, New Caledonia, South Province, Trou Bleu River. NMUW 94267, 1 female, 55.5 mm, New Caledonia, Sotuh Province, Fausse Yaté River. Rhyacichthys aspro (Valenciennes, 1837): CAS 138655, 1 spm., sex ?, 98.8 mm, Philippines, Mindanao, Tagaloan River. CAS 51696, 1 spm., sex?, 102.2 mm, Philippines, Luzon, Mantugil River, Villar. NUW 45968, 1 female, 97.2 mm; Indonesia, Java, Semarang. NMW 82972, 1 male, 115.2 mm; Indonesia, Moluccas. NUW 82990, | male?, 121.7 mm, Philippines. 360 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16 COMPARATIVE MATERIAL Within the families, the subfamilies and the genera are listed alphabetically. The pattern of free neuromasts of the lateral-line system on the caudal fin is as follows: four rows (one transversal Iet and three longitudinal Ied, lem and Iev) (Fig. 1 A—B) or three rows (one transversal Iet and two lon- gitudinal Ied and Iem) (Fig. 1C—D). Row Iet is continuous at the base of the caudal fin or discon- tinuous, as two, three (generally), or four short rows anterior to each longitudinal row (Fig. 1). Species with two longitudinal rows are marked with an asterisk. Species with more than three lon- gitudinal rows are marked with an §. Species with one longitudinal row (lem) are marked with #. Number in parentheses is number of investigated specimens and not necessarily identical with the number of specimens in the lot. Odontobutidae: Micropercops swinhonis (Gunther, 1873), UMMZ 167389, USNM 336883 (12); Odontobutis aurarmus* Vidthayanon, 1995, USNM 325486, UMMZ 223284 (6); Odontobutis obscura* (Temminck & Schlegel, 1845), CAS 32827, CAS 28154, MNHN 6778, MNHN 6481, MNHN 1987-1213, USNM 264892, USNM 864893, USNM 086965, UMMZ 70284, UMMZ 142619 (25); Odontobutis potamophila* (Giinther, 1861), BMNH 1918.11.12.16—-18 (2). Eleotridae: Butinae: Bostrychus africanus (Steindachner, 1879), BMNH 1989.1.6: 23-29 (3); Bostrichthys zonatus (Weber, 1907), NMW 58166 (1); Butis butis (Hamilton, 1822), CAS 205539, NMW 22482 (5); Butis koilomatodon (Bleeker, 1849), BMNH 1988.2.8: 1-8 (4); Kribia nana* (Boulenger, 1901), CAS 64486, BMNH 1918.11.12: 16-18, BMNH 1948.1.14: 291-313 (4); Kribia kribensis* (Boulenger, 1907), NMW 78428 (2); Milveringa veritas Whitely, 1945, BMNH 1982.2.17: 1-9 (3); Ophiocara porocephala (Valenciennes, 1837), BMNH 1980.10.10: 260—263 (4); Oxyeleotris fimbriata (Weber, 1907), BMNH 1983.8.2: 147-148 (1); Oxveleotris heterodon (Weber, 1907), NMW 57206, NMW 57206 (2); Oxyeleotris lineolatus (Steindachner, 1867), BMNH 1983.8.2: 203-210 (4); Oxyeleotris marmorata (Bleeker, 1852), CAS 66251 (2); Oxyeleotris urophthalmoides (Bleeker, 1853), CAS 49456 (1); Pogoneleotris microps Weber, 1907, NMW 31456 (1); Zyphleotris pauliani Arnoult, 1959, BMNH 1981.111.9: 19-20 (2). Eleotrinae: Batanga lebretonis (Steindachner, 1870), NMW 22275, NMW 76245, NMW 76407, NMW 58729 (11); Bunaka gyrinoides (Bleeker, 1853), BMNH 1983.8.2: 211-214 (3); Dormitator latifrons (Richardson, 1844), CAS 54392 (2); Eleotris sp., NUW 94987 (1); Eleotris amblyopsis (Cope, 1871), NMW 85500 (6); Eleotris coxi (Krefft, 1864), NMW 78548 (1); Eleotris daganen- sis Steindachner, 1870, NMW 22488 (1); Eleotris fusca (Schneider & Forster, 1801), NMW 89288 (1); Eleotris gyrinoides Bleeker, 1853, NMW 80754 (2); Eleotris picta Kner, 1863, SIO 59-358- 58B (1); Eleotris sandwicensis Vaillant & Sauvage, 1875, NMW 78678, SIO 61-425-58A (7); Eleotris senegalensis (Steindachner, 1870), NMW 78553, NMW 85266 (3); Eleotris vittata Dumeril, 1861, BMNH 1956.9.6: 51, BMNH_ 1985.3.18: 191-192 (3); Gobiomorphus basalis (Gray, 1842), BMNH 1964.12.21: 17 (1); Gobiomorphus gobioids (Valenciennes, 1837), NMW 22508 (1); Gobiomorus dormitor Lacepede, 1800, BMNH 1982.8.19: 2021—2040 (4); Guavina guavina (Valenciennes, 1837), NVW 14644, NMW 22531 (4); Hypseleotris sp., NUW 30966 (1); Hypseleotris compressa (Krefft, 1864), NMW 22511, NMW 22520, NMW 58165 (6); Mogurnda mogurnda (Richardson, 1844), BMNH 1983.8.2: 4 (2); Mogurnda nesolepis (Weber, 1907), NMUW 22517 (1); Mogurnda variegata Nichols, 1951, BMNH 1983.8.2; 73-74 (2); Ophieleotris aporos (Bleeker, 1854), NMW 22524, NMW 22537, NMW 78607, NMW 83359 (7); Philypnodon grandi- ceps (Krefft, 1864), NMW 22534 (1). Gobiidae: Amblyopinae: Amblyopus caeculus Karoli, 1882, NMW 5779 (2); Taenioides buchanani (Day, 1873), NMW 76500 (1); Taenioides cirratus (Blyth, 1860), NMW 94598 (1). ee AHNELT AND GOSCHL: CAUDAL FIN LATERAL-LINE SYSTEM IN GOBIOIDEI 361 Gobiinae: Acentrogobius caninus (Valenciennes, 1837), NMW 28808-28811 (3); Acentrogobius frenatus (Giinther, 1861), NMW 30452-30453 (2); Acentrogobius masoni (Day, 1873), NMW 33926 (1); Acentrogobius simplex (Sauvage, 1880), NMW 29964 (6); Amblygobius albimaculatus (Riippel, 1830), NMW 28722-28727 (4); Amblygobius byonensis (Richardson, 1844), NMW, 33924, NMW 58800 (4); Amblygobius sphynx (Valenciennes, 1837), NMW 30102-30105 (3); Amoya sp., NUVW 16165, NMW 30757 (2); Anatirostrum profundorum* (Berg, 1927), CMNFI 1999-0023 (4); Asterropteryx semipunctatus Riippell, 1830, NMW 22516, NMW 78225, NMW 88504 (6); Bathygobius cotticeps (Steindachner, 1879), NMW 30439 (1); Bathygobius fuscus (Riippell, 1830), NMW _ 88321(5); Bathygobius soporator (Valenciennes, 1837), NMW 33927-33928, NMW 60391 (4); Benthophilus stellatus* (Sauvage, 1874), IZUW uncatalogued (2); Cafjrogobius caffer (Giinther, 1874), NMW 28819-28820 (2); Callogobius sclateri (Steindachner, 1879), NMW 30901 (1); Chasar bathybius (Kessler 1877), CMNFI uncatalogued (4); Chromogobius quadrivittatus (Steindachner, 1863), NMW 30639, NMW 30657, NMW 86084, NMW 88486 (4); Chromogobius zebratus (Kolombatovic, 1891), NMVW 29613, NMW 86082 (3); Corcyrogobius liechtensteini (Kolombatovic, 1891), NMW 37536-37543, NMW 78460; NMW 94458 (4); Deltentosteus collonianus§ (Risso, 1820), NMVW 28826-28830, NMW 29087-29100, NMW 30342, NMW 86693 (6); Deltentosteus quadrimaculatus§ (Valenciennes, 1837), NMW 28845-28849, NMW 37479-37484, NMW 30667 (5); Didogobius kochi Van Tassell, 1988, IZUW uncatalogued (2); Didogobius splechtnai Ahnelt & Patzner, 1995, NMW 92804, NMW 9280S, NMW 94451 (4); Economidichthys pygmaeus (Holly, 1929), MCN 106376—394 (2); Elacatinus sp., SIO uncatalogued (1); Elacatinus multifasciatus (Steindachner, 1876), NMW 76239 (1); Evermannia longipinnis (Steindachner, 1879), NMW 76420 (2); Eviota distigma Jordan & Seale, 1906, NMW 88173 (4); Favonogobius reichi (Bleeker, 1853), NMW 33897 (1); Gammogobius steinitzi Bath, 1971, SMF 11071, NMW 94452-94456 (4); Glossogobius celebius (Valenciennes, 1837), NMW 83363 (1); Glossogobius giuris (Hamilton, 1822), NMW 30543-30544 (2); Gobiodon citrinus (Riippell, 1838), NMW 30973 (5); Gobiosoma bosc (Lacepede, 1800), SIO 67- 277-59 (2); Gobius ater Bellotti, 1888, NMW 28551-28552, NMW 29041 (2); Gobius bucchichi Steindachner, 1870, NMW 88436, NMW 88491 (6); Gobius cobitis Pallas, 1814, NUW NMW 87092, NMW 88463 (6); Gobius cruentatus Gmelin, 1789, NUW 28986-28982 (2); Gobius fallax Sarato, 1889, NMW 28792, NMW 77932 (2); Gobius gasteveni Miller, 1974, MNCN 73576-73579, MNCN 79218-79219 (3); Gobius geniporus Valenciennes, 1837, NMW 87065 (1); Gobius niger Linnaeus, 1758, MNCN 74278-74282, IZUW uncatalogued (4); Gobius paganellus Linnaeus, 1758, NMW 80592, NMW 84937, NMW 94582, NMW 94583 (6); Gobius rouli De Buen, 1828, NMW 94277 (2); Gobius vittatus Vinciguerra, 1883, IZUW uncatalogued (1); Gobiusculus flavescens* (Fabricius, 1779), NMW 30693 (3); Hetereleotris vulgaris (Klunzinger,1871), NMW 31020, NMW 78237, NMW 88177 (12); Istigobius diadema (Steindachner, 1876), NMW 29171 (1); Knipowitschia croatica* Mrakovic, Kerovec, Misetic & Schneider, 1996, NMW 93978 (6); Knipowitschia caucasica* (Berg, 1916), LZUT uncatalogued (2); Knipowitschia pannizae (Verga, 1841), IZUW uncatalogued (4); Knipowitschia punctatissi- ma* (Canestrini, 1864), IZUW uncatalogued (5); Lebetus guilleti# (le Danois, 1913) NMW 94589-94592 (4); Lesueurigobius friesii (Malm, 1874), MNCN 78580-78583; NMW 76296, NMW 88348 (4); Lesueurigobius suerii (Risso, 1810), NMW 88351, NMW 88550 (2); Lophogobius cyprinoides (Pallas, 1770), NMW 30807-30808 (2); Lythrypnus cobalus Bussing, 1990, CAS 205778, SIO 72-97 (2); Lythrypnus dalli (Gilbert, 1890), CAS 118455, SIO 63-174- S9C (6); Lythrypnus gilberti (Heller and Snodgrass, 1903), CAS 39236 (4); Lythrypnus pulchellus Ginsburg, 1938, CAS 18097 (3); Lythrypnus rhizophora (Heller and Snodgrass, 1903), CAS 50078 (5); Lythrypnus zebra (Gilbert, 1990), CAS 25388, SIO H50-40-59A (6); Mauligobius maderensis 362 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16 (Valenciennes, 1837), NMW 87554 (1); Mesogobius batrachocephalus (Pallas, 1814), NMW | 11397 (3); Microgobius gulosus (Girard, 1858), NMW 87269 (3); Millerigobius macrocephalus (Kolombatovic, 1891), IZUW uncatalogued (2); Myersinia filifer (Valenciennes, 1837), NMW 86810 (1); Neogobius cyrius (Kessler, 1874), NMW 94327 (4); Neogobius eurycephalus (Kessler, 1874), NMW 92808 (2); Neogobius fluviatilis (Pallas, 1814), NUW 94446 (4); Neogobius gorlap Ijin, 1949; CMNFI 1993-0161, CMNFI 1993-0162 (3); Neogobius gymnotrachelus (Kessler, 1857), NMW 88596, NMW 94583 (6); Neogobius kessleri (Giinther, 1861), IZUW uncatalogued, NMW 93977, NMW 94560 (6); Neogobius melanostomus (Pallas, 1814), NMW 60200 (1); Neogobius ratan (Nordmann, 1840), NMW 33910 (1); Neogobius syrman (Nordmann, 1840), CMNFI 1970-0544 (1); Odondebuenia balearica (Pellegrin and Fage, 1907), MNCN 107992—108042 (6); Padogobius bonelli (Bonaparte, 1846), IZUW uncatalogued (5); Padogobius nigricans (Canestrini, 1867), IZUW uncatalogued (2); Paragobiodon echinocephalus (Rippel, 1830), NMW 29196 (4); Paragobiodon modestus (Regan, 1908), NMW 87478 (4); Paragobiodon xanthosomus (Bleeker, 1852), NUW 87477, NMW 87479 (2); Pomatoschistus adriaticus* Miller, 1873, NMW 28647-28649, NMW 28670 (5); Pomatoschistus kneri (Steindachner, 1861), NMW 37772-37776 (1); Pomatoschistus marmoratus* (Pallas, 1810), NMW 87359 (3); Pomatoschistus microps (Kroyer, 1838), IZUW uncatalogued (3); Priolepis sp., NUW 88507 (2); Priolepis nuchi- fasciata (Giinther, 1873); NMW 33908 (1); Proterorhinus marmoratus (Pallas, 1814), NMW 60240, NMW 79755, NMW 94561, NMW 94595 (7); Rhinogobiops nicholsii (Bean, 1882), CAS 135015, CAS 135016, CAS 27627 (6); Speleogobius trigloides# Zander and Jelinek, 1976, NMW 75826-75827 (1); Thorogobius macrolepis (Kolombatovic, 1891), NMW 37421-37422, NMW 94331, NMW 94332 (7); Thorogobius ephippiatus (Lowe, 1839), NMW 86591 (1); Trimma sp., NMW 89332 (1); Valenciennea sexguttat (Valenciennes, 1837), NMW 83959 (1); Vanneaugobius dollfusi Brownell, 1978, NMW 87961, NMW87962 (6); Zebrus zebrus (Risso, 1827), NMW 86097, NMW 88472, NMW 88492, IZUW uncatalogued (7); Zosterisessor ophiocephalus (Pallas, 1814), IZUW ucatalogued (1). Gobionellinae: Acanthogobius flavimanus (Temminck and Schlegel, 1845), CAS 36971, CAS 42487, CAS 52003, CAS 21367, CAS 213686, NMW 30454-30455 (28); Aphia minuta§ Risso, 1827, NMVW 31463, NMW 88030, NHRM 46688 (10); Arcygobius baliurus (Valenciennes, 1837), NMW 30278-30279, NMW 82114 (4); Awaous giuris (Hamilton, 1822), NMW 30096 (1); Awaous tajasica (Lichtenstein, 1822), NMW 91265, NMW 91277 (6); Brachygobius xanthozonus* (Bleeker, 1849), NMW 12685, NMW 30152-30156 (4); Cepola striata Bloch and Schneider, 1801, NMW 28742-28747 (3); Clariger cosmurus* Jordan and Snyder, 1901, IZUW uncatalogued (2); Chaenogobius gulosus (Sauvage, 1882), IZUW uncat- — alogued, NMW 30676 (4); Clevelandia ios* (Jordan and Gilbert, 1882), CAS 15476, SIO 47-73- 59D (5); Ctenogobius sagittula* (Giinther, 1861), CAS 55185 (3); Euclogobius newberryi* (Girard, 1856), CAS 31768, SIO 62-278-59A (6); Eutaeniichthys gilli* Jordan and Snyder, 1901, IZUW uncatalogued (2); Evorthodus lyricus (Girard, 1858), NMW 83107, NMW 88633 (3); Gillichthys mirabilis* Cooper, 1864), CAS 22071, CAS 79616 (6); Gillichthys seta* (Ginsburg, 1938), CAS 26055, SIO 67-133-59 (6); Gobiopterus cf. chuno§ (Hamilton, 1822), IZUW uncata- logued (5); Gymnogobius castaneus (O’Shaughnessy, 1875), IZUW uncatalogued, NMW 78031(3); Gymnogobius urotaenia* (Hilgendorf, 1879), NMW 29508 (1); Jlypnus_ gilberti* (Eigenmann & Eigenmann, 1891), CAS 24169, CAS 26896 (6); /ypnus luculentus* (Ginsburg, 1938), CAS 214242, SIO 62-235-59A (6); Lepidogobius lepidus* (Girard, 1858), CAS 19894, CAS 25383, CAS 53228 (6); Lethops connectens* Hubbs, 1926, SIO H46-46, SIO H51-239 (6); Leucopsarion petersii§ Hilgendorf, 1880, FAKU 103091 (5); Luciogobius guttatus* Gill, 1859, FAKU 102257—102258 (2); Mugilogobius poeyi (Steindachner, 1867), NMW 30608 (1); Oxyurichthys papuensis (Valenciennes, 1837), NUW 29935-29938 (3); Oxvurichthys tentacularis AHNELT AND GOSCHL: CAUDAL FIN LATERAL-LINE SYSTEM IN GOBIOIDEI 363 (Valenciennes, 1837), NMW 12688 (1); Pseudaphya ferreri* (de Buen and Fage, 1908) NMW 37424-37429, NMW 37431-37433 (7); Quietula guaymasiae* (Jenkins & Evermann,1889), CAS 26056, CAS 55187 (6); Quietula y-cauda* (Jenkins and Evermann, 1889), CAS 11497, CAS 200221 (6); Rhinogobius sp*., NMW 89618 (3); Stigmatogobius sadanundio (Hamilton, 1822), NMW 81598 (1); Stigmatogobius sella (Steindachner, 1881), NMW 30107-30108 (1); Synechogobius ommaturus (Richardson, 1845), NMW 34199-34200, NMW 34206-34208, NMW 81876 (4); Triaenopogon barbatus (Giinther, 1861), CAS 161208 (2); Tridentiger bifasciatus* Steindachner, 1881, CAS 82361 (3); Tridentiger trigonocephalus* (Gill, 1859), CAS 15405 (1); Typhlogobius californiensis* Steindachner, 1879, CAS 200223, CAS 211664 (5). Oxudercinae: Apocryptes bato (Hamilton, 1822), CAS 89289 (2); Boleophthalmus boddarti (Pallas, 1770), CAS 140028, IZUW 2419 (4); Boleophthalmus pectinirostris (Linnaeus, 1758), NMW 78444 (2); Periophthalmus barbarus (Linnaeus, 1766), IZUW 2418, NMW 31321 (2). Sicydiinae: Sicydium sp.. NMW 31434 (1); Sicyvdium plumieri (Bloch, 1786), NMW 91574, NMW 31411-31414 (6); Sicyopterus cynocephalus (Valenciennes, 1837), NMW 31405-31406 (1); Sicvopterus sarasini Weber and Beaufort, 1915, NMW 94340 (2); Stiphodon elegans (Steindachner, 1879), NMW 81318 (3). Kraemeriidae: Kraemeria samoensis Steindachner, 1906, NMW 83668 (2). Microdesmidae: Microdesmus dipus Giinther, 1864, BMNH 1967.12.30: 1-3 (1). Ptereleotridae: Nemateleotris decora Randall and Allen, 1973, BMNH 1983.3.25: 893-895 (1); Ptereleotris microlepis (Bleeker, 1856), BMNH 1983.3.25:892 (2). TERMINOLOGY OF THE LATERAL-LINE SYSTEM OF THE CAUDAL FIN Sanzo (1911) was the first to develop a terminology for the longitudinal rows of neuromasts on the caudal fin of gobioid fishes. The species he investigated have three longitudinal rows of neu- romasts developed on each side of the caudal fin: one an elongation of the median trunk lateral line, and ventral and dorsal to it, and two accessory rows. Sanzo named these rows ‘lateral caudal’ (Ic) and they differed, from dorsal to ventral, between Ie, Ie’ and Ic’’. The number of lateral lines on the caudal fin in Gobioidei may be secondarily reduced to one or two or increased to eight neuromast rows (Mortara 1918; Ahnelt and Géschl 2003; Scattolin and Ahnelt, unpublished), making it diffi- cult to identify homologous rows. Therefore, an alternative classification of the lateral-line system was proposed by Ahnelt et al. (2000) and Ahnelt and Duchkowitsch (2001) reflecting the position of the rows on the caudal fin: Ied (lateral caudal dorsal), lem (lateral caudal medial) and lev (lat- eral caudal ventral). Three longitudinal lateral lines are plesiomorphic for Gobioidei (Springer 1983). In Gobioidei with two longitudinal rows, the Ied and Iem are developed and the lev is absent (e.g., Ahnelt et al. 2000; Ahnelt and Géschl 2003). In species with one longitudinal row (lem), both accessory rows (Iced and Iev) are absent. The characteristic pattern of the lateral-line system on the caudal fin, led separated from lem by three and Iem from lev by two fin rays, allows the identification of the three longitudinal neu- romast rows in species with more than three rows developed (Fig. 1F). The transversal row (Ict) on the base of the caudal fin was included by Sanzo (1911) as the last transversal row in the lateral medial trunk series. These series of generally short rows run along the midline of the trunk and are associated with the trunk lateral-line canal in Rhyacichthys, Protogobius and Terateleotris (Wongrat and Miller 1991; Shibukawa et al. 2001). Such short trans- versal rows are also found on each first canal carrying scales on the caudal fin of R. aspro. Lateral- line canals on the trunk and on the caudal fin are absent in the other Gobioidei. The neuromast row Ict differs from the medial trunk series (Im) as it is distinctly longer, seemingly because it was orig- 364 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16 inally associated with the three lateral-line canals of the caudal fin. In many advanced taxa the three parts unite to form a long, continuous transversal row, exceeding the longitudinal rows dorsally and ventrally and often nearly extending over the entire caudal fin base (Fig. 1D—F). Generally, Iet is separated from the last Im row by a larger gap than the Im rows on the caudal peduncle from each other. The row Ict may be reduced to three neuromasts, each anterior to one longitudinal neuromast row. In many species, these neuromasts are still identifiable due to their larger size and a gap to the neuromasts of the longitudinal rows (Fig. 1). These neuromasts of Iet are in some taxa incorporat- ed in the origin of the longitudinal neuromast rows and, therefore, only separable if they are of a larger size. The incorporation of Ict in the longitudinal lateral-line rows and its secondary loss is seemingly apomorphic. The neuromast rows on the caudal peduncle and on the caudal fin of gobioid fishes are inner- vated by the ramus lateralis posterius (Sanzo 1911; Wongrat and Miller 1991). According to the nomenclature of Coombs et al. (1988), let (at the of caudal fin) is formed by secondary replace- ment neuromasts. Secondary replacement neuromasts, derived from free neuromasts associated with pores of the lateral-line canals (e.g., the trunk canal of Protogobius, Rhyacichthys and Terateleotris, and the origin of canals on the caudal fin of Protogobius and Rhyacichthys), form transverse rows along the former course of a canal. The longitudinally arranged neuromasts on the caudal fin (led, lem, lev) form rows along the direction of deleted canals and are, therefore, pri- mary replacement neuromasts, not secondary accessory neuromasts as postulated by Wongrat and Miller (1991). RESULTS LATERAL-LINE SYSTEM OF THE CAUDAL FIN OF PERCCOTTUS GLENI.— The lateral-line system on the trunk and caudal fin is formed by free neuromasts. No trunk canal, no canal in extension of the trunk canal on the caudal fin, and no accessory canals on the trunk and on the caudal fin are developed in P. glenii. These canals, present in the more-basal Gobioidei Rhyacichthys, Protogobius and Terateleotris, are replaced by free neuromasts. The pattern of the lateral-line system on the caudal fin is T-shaped. It consists of free neuro- masts arranged in two rows on each side of the fin, one row transversal (Ict) on the base of the fin and the second longitudinal (lem) in its midline (Fig. 1B). The transversal row is discontinuous. In adults, it extends as three distinct sections of neuro- masts (short rows or accumulations) transversely between the fourth and twelfth branched caudal- fin rays. The median section extends anteriorly to the single longitudinal neuromast row. In juve- niles, these three sections of the row Iet are each represented by a single neuromast. The number of these neuromasts increases with size: in specimens > 70 mm SL, each section is represented by a short row, whereas in specimens >100 mm SL, they are multiple rows or aggregations of neuro- masts. Iet is distinctly separated from the last row of the lateral median trunk series (Im) and extends over the small scales which cover the caudal fin base. Only a single longitudinal row of neuromasts is developed, the lateral caudal median row (lem). The dorsal and the ventral lateral caudal rows (Ied and lev), the first generally present in Gobioidei, the second in many species lacking, are absent. lem extends along the interradial mem- brane between the eighth and ninth (adult) or seventh and eighth (juveniles) branched caudal-fin rays. This row of neuromasts is long but does not reach the rear margin of the caudal fin. It extends over about three quarters of the fin in adults. The length of this row is not known for juveniles | because their caudal fins were damaged in their rear parts. lem begins immediately behind the last row of small scales that cover the base of the caudal fin, opposite the median section of let. At the AHNELT AND GOSCHL: CAUDAL FIN LATERAL-LINE SYSTEM IN GOBIOIDEI 365 origin of lem, the neuromasts are closely set, but they are more dis- tant from each other towards its end. In specimens >100 mm SL, this row may be doubled at its beginning. The pattern of the lateral-line system on the caudal fin of P. glenii differs distinctly from the general pattern found in gobioid fishes and seemingly represents a derived condition. With only one longitudi- nal row of neuromasts developed, large parts of this fin are not cov- ered by the lateral-line system. Such an arrangement of neuro- masts is not known in other basal gobioid fishes including Eleotridae. LATERAL-LINE SYSTEM ON THE TRUNK AND THE CAUDAL FIN OF RHYACICHTHYIDAE.— Within the Rhyacichthyidae, sensu Shibukawa Ficure 1. Stylized pattern of the lateral-line system on the caudal fin of et al. (2001 ), the lateral-line system Gobioidei. A-B: Three longitudinal rows of free neuromasts (A: Gobius of Protogobius attiti is obviously niger, B: Micropercops swinhonis). C_-D: Two longitudinal rows of free i. neuromasts (C: Odontobutis aurarmus, D: Quietula guaymasiae). E: One more specialized than those of longitudinal row of free neuromasts (Perccottus glenii). F: Eight longitudi- Rhyacichthys aSspro. Compared nal rows of free neuromasts (Aphia minuta) (neuromasts of Iced, lem and Icv with the sensory system of the lat- are shown enlarged). The neuromasts are not drawn to scale, and their num- ter, it shows reductions in the later- ber is larger in each species than shown. In most Gobioidei, the neuromasts decrease in size rearwards, as shown in A only. Iced = dorsal longitudinal al-line canals on the head and on row, lem = medial longitudinal row, Ict = transversal row, Icv = ventral lon- the caudal fin (Watson and gitudinal row. Pédllabauer 1998; Shibukawa et al. 2001). Such differences are also found in the trunk lateral-line canal. In R. aspro, this canal is formed by two parts: an anterior continuous and a posterior discon- tinuous part. The anterior part of the trunk canal is continuous with the cephalic canal system (Akihito et al. 2000b:1272, fig. 4-1) and extends uninterrupted along about the first 12 scales of the trunk. Two canaliculi (short side branches) extend dorsally and ventrally from the canal along the posterior margin of every second lateral-line scale. Between the dorsal and ventral end of these canaliculi, the posterior margin of the scales is convex and free of ctenii. This gives the anterior lateral-line scales a characteristic shape: the center of the posterior rim of the scales is emarginat- ed and separates the ctenii into a dorsal and ventral series. The continuous part of the trunk later- al-line canal ends with a single terminal pore. In its further course, the trunk canal is discontinuous and consists of a consecutive series of short canals, each on a single scale. The discontinuous part of the trunk canal starts somewhat dorsal to the end of the continuous trunk canal, which is bent downwards [“lateral-line interrupted at midpoint of body” of Shibukawa et al. (2001)]. These canals extend over nearly the entire surface of the lateral-line scales. Each canal ends at the poste- rior edge of the scale and starts below the margin of the scale in front of it. Where the canals end E 366 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16 in a terminal pore, the ctenii of these scales are separated by a narrow gap and/or a “slight median notch” (Miller 1973). A few free neuromasts accompany the anterior continuous part of the lateral-line canal (Akihito et al. 2000b:1272, fig. 4-1). Their number increases with its end. Short transversal rows of free neuromasts are associated with most canals of the discontinuous trunk canal. They run immediately before the anterior pore. Additionally, these canals are accompanied by short longitu- dinal rows of free neuromasts, one row dorsal and one ventral to the canal. Such longitudinal rows occur also at the end of the continuous part of the trunk lateral-line canal. Dorsal and ventral to the accessory trunk canal of R. aspro, which is confined to a single scale ventral to the first dorsal fin base, occur transversal accessory neuromast rows. These rows are short, do not extend onto neigh- bouring scales, and seemingly correspond to the anterior rows of the lateral dorsal series (Id) of Sanzo (1911). Three lateral-line canals extend over about half of the caudal fin in R. aspro, followed by short longitudinal rows of neuromasts. These three canals (one in extension of the trunk lateral-line canal plus two accessory canals) are restricted to three single scales on the base of the caudal fin in P attiti, followed by long longitudinal rows of neuromasts (Shibukawa et al. 2001; Ahnelt unpub- lished). [Like Shibukawa et al. (2001) these canals in the smaller of the two specimens investigat- ed have not been found. Data of more individuals are needed for a better knowledge of the vari- ability of this feature of Protogobius. Watson and Péllabauer (1998:148) mention: “not present on Protogobius are two short accessory lateral lines on the caudal fin base reported in Rhyacichthys. ... Therefore, we conclude that these authors found the lateral-line canal in the midline of the cau- dal fin base.] In both Protogobius and Rhyacichthys, the longitudinal lateral lines on the caudal fin are asymmetrically arranged: the dorsal and the median lateral lines are separated by three caudal- fin rays, the median and the ventral lateral lines by two rays. This asymmetrical pattern is ple- siomorphic for the Gobioidei. Short transversal rows occur on the first canal carrying scales on the caudal fin of R. aspro immediately before the anterior pore of each lateral-line canal, together forming a transversal row (Iet) consisting of three parts. Additionally, as single neuromasts, longitudinal neuromast rows occur on each first scale of the three lateral-line canals. We did not find neuromasts on the corre- sponding scales on the caudal fin of the two investigated specimens of P. attiti, but, possibly, they also occur in this gobioid species. Obviously, Iet is present in the basal gobioid family, the Rhyacichthyidae, at least in the basal Rhyacichthys, a feature which had been overlooked by Ahnelt and Géschl (2003). The occurrence of a transversal row on the base of the caudal fin is plesiomor- phic for the Gobioidei. Also plesiomorphic is an Iet consisting of three short transversal rows. In the following, we list differences in features of the trunk and caudal fin lateral-line system between P. attiti and R. aspro not mentioned by Watson and Péllabauer (1998) and Shibukawa et al. (2001) (features of R. aspro in parentheses): (i) trunk lateral-line canal not continuous with the cephalic canal system (continuous with the cephalic canal system); (11) trunk lateral-line canal discontinu- ous, consisting of a consecutive series of short canals along the lateral midline except for a short part over the first two or three scales (anterior third of the trunk canal continuous); (i11) trunk canals with no side branches (= canaliculi) (continuous part of the trunk lateral-line canal with canaliculi); (iv) trunk canals on lateral-line scales short, not interrupting the continuous series of ctenii on the posterior margin of each scale (canals terminate at the posterior margin of the lateral-line scales separating the ctenii into dorsal and ventral series); (v) no longitudinal rows of accessory neuro- masts dorsal and ventral to the trunk canals (short longitudinal rows of accessory neuromasts pres- ent dorsal and ventral to the canals of the discontinuous part of the trunk canal); (vi) no short trans- versal rows (present) on each scale of the three lateral-line canals on the caudal fin (three short AHNELT AND GOSCHL: CAUDAL FIN LATERAL-LINE SYSTEM IN GOBIOIDEI 367 transversal rows present, together forming a discontinuous lateral caudal transversal row Ict); (vii) no longitudinal rows of accessory neuromasts dorsal and ventral to the lateral-line canals on the caudal fin (longitudinal rows of accessory neuromasts present dorsal and ventral to the lateral-line canals on each first scale of the three lateral-line canals); (viii) no transversal accessory neuromast rows at the accessory trunk lateral-line canal (transversal accessory neuromast rows present dorsal and ventral to the accessory trunk lateral-line canal). GENERAL PATTERN OF THE LATERAL-LINE SYSTEM ON THE CAUDAL FIN IN GOBIOIDEI EXCLUSIVE OF RHYACICHTHYIDAE The following two patterns of the lateral-line system seemingly occur in the majority of gob- ioid fishes: PATTERN |: Plesiomorphic for Gobioidei, exclusive of Rhyacichthyidae, are four lateral lines on the caudal fin, which are all formed by free (superficial) neuromasts. A discontinuous transver- sal row (Iet) extending over the caudal fin base and three longitudinal rows: one in extension of the deleted trunk canal in the middle of the caudal fin (lem) and two accessory rows, one on the dor- sal part (Ied) and on the ventral part (lev) of this fin (Fig. 1A—B). No lateral-line canals are devel- oped. Obviously, plesiomorphic for Gobioidei is a discontinuous Ict, which consists of at least three short parts, each anterior to one of the three longitudinal neuromast rows. A continuous transversal row at the base of the caudal fin or its loss is seemingly apomorphic. The three longitudinal rows are generally long and extend onto the interradial membranes between two branched caudal-fin rays in the same asymmetrical pattern as in the Rhyacichthyidae. Typically, the gap between Ied and Iem is larger than the gap between Iev and Iem. Generally, these neuromast rows do not run in the midline between two fin rays: Ied and Iem follow the ventral side of a fin ray, lev follows the dorsal side of a fin ray. This pattern of the longitudinal lateral lines on the caudal fin seemingly is shared by the majority of Gobioidei. In species with rounded or point- ed caudal fins, lem generally is the longest row, and the two accessory rows (Iced, lev) are more or less of similar length. PATTERN 2: This pattern is derived from Pattern | and formed by three neuromast rows with one longitudinal row (Iev) lost: one transversal and two longitudinal rows (Fig. 1C). The transver- sal row is often continuous and in its length not affected by the loss of Iev. It still extends onto the ventral part of the caudal fin base. As in species with three longitudinal lateral lines, led and Iem are separated by three fin rays. Generally, lem is somewhat longer than Ied (rounded caudal fin) or distinctly longer (pointed caudal fin). In several species with a rounded caudal fin, a seemingly derived condition occurs with Iem shortened and Icd the longest row. A deviation of Pattern 2 towards a pattern with only one longitudinal row occurs in the California bay gobies of the Chasmichthys group, sensu Birdsong et al. (1988). In some species, led is shortened to only a few neuromasts (Ahnelt and Géschl 2003; Ahnelt unpublished) (Fig. 1D). DEVIATIONS FROM THE GENERAL PATTERN.— We found three distinct deviations from the two general patterns of the lateral-line system on the caudal fin. These deviations occur in the group of Gobioidei exclusive of the Rhyacichthyidae, the unassigned Zerateleotris, and the Eleotridae, but including the Odontobutidae. All investigated species of the Rhyacichthyidae and Eleotridae (and Terateleotris) are characterized by three lateral lines on the caudal fin except for two eleotrine species, Kribia nana and K. kribensis. (i) Longitudinal neuromast rows on the caudal fin arranged symmetrically: This is a deviation from Pattern | with the three longitudinal lateral lines arranged asymmet- rically. In this deviation both accessory neuromast rows (led, lev) are separated from Iem by three 368 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16 fin rays. Such a pattern occurs in two species, Synechogobius ommaturus and Acanthogobius flay- imanus. Seemingly, the symmetric arrangement of the neuromasts is a derived character that devel- oped from a pattern with Iev separated from Iem by two fin rays. An indication for this hypothesis is the occurrence of a second Iev between the second and third fin ray below lem in 4. flavimanus. Such doubled Iev occurs in 46% of the investigated specimens, but only in 10% on both sides of the fin. Except for one specimen with a second Ied, this row and Iem were never doubled. Possibly because of the low number of specimens investigated, no fourth row has been found in S. ommatu- rus. The question of whether three longitudinal rows arranged symmetrically on the caudal fin are a characteristic feature for the Acanthogobius-group, sensu Pezold (1993), is currently under inves- tigation by the senior author. (ii) Longitudinal neuromast rows decreased in number, loss of Ied and lev: This deviation, only a single longitudinal neuromast row (Iem) developed, obviously occurs independently twice within the Gobioidei: in the Odontobutidae, and in the derived Gobiidae. Perccottus glenii is the only known representative of an assemblage of basal Gobioidei (Rhyacichthyidae, Odontobutidae and Eleotridae) with a single neuromast row on the caudal fin. The pattern of the lateral-line system of this species is described above. It is not known whether this arrangement derived from a pattern with two or with three longitudinal neuromast rows. Two species of Gobiidae, Lebetus guilleti and Speleogobius trigloides, are also peculiar with only one longitudinal row (lem) on the caudal fin developed (but concluding from Miller (1963), L. guilleti can also have two longitudinal rows developed). The arrangement of the lateral-line sys- tem on this fin is very similar for both species and represents a derived condition, a further indica- tion of a close relationship as recently proposed by Herler and Kovacic (2002). L. guilleti and S. trigloides are tiny gobies of about 20 mm SL generally characterized by a reduced neuromast pat- tern on the body like, as known for many small gobies (Ahnelt and Bohacek 2004, and authors summarized therein). Some neuromast rows on the head are completely absent and many reduced to one or only a few neuromasts (Miller 1963; Zander and Jelinek 1976). The loss of two longitu- dinal rows on the caudal fin is seemingly not necessarily the result of miniaturization. Other tiny gobies, such as Corcyrogobius liechtensteini (the smallest Mediterranean gobiid fish) and Gobiopterus chuno, have three or more longitudinal rows on this fin developed. (iii) Longitudinal rows increased in number: Sporadically, gobioids can be found with a supernumerary longitudinal neuromast row on the caudal fin. These specimens display four longitudinal rows but, generally, only on one side of the fin (Ahnelt unpublished). Such supernumerary rows occur rarely and are not typical for the species. Gobioids with distinctly more than three longitudinal neuromast rows have these rows gener- ally separated by one fin ray only (Mortara 1918; Scattolin and Ahnelt, unpublished). This highly specialized pattern is so far only known within the Gobiidae and seemingly evolved independent- ly several times within this gobioid family. Three species are known with eight rows of neuromasts on the caudal fin: the north-eastern Atlantic species Aphia minuta, and the Pacific gobies Gobiopterus chuno and Leucopsarion peter- sii. In these species, Iet is continuous and long, extending nearly over the entire caudal fin (Fig. IF). The eight longitudinal rows extend onto the interradial membranes between the second and the tenth segmented and branched fin rays, each separated from the other by one fin ray. These three species can be considered as derived forms and are characterized by paedomorphic features such as completely absent head canals, transparent bodies and pelagic life style. For these evolved pelagic Gobiidae, a convergent evolution of free neuromast pattern on the caudal fin in adaptation | to similar habitats is assumed (Scattolin and Ahnelt, unpublished). Two Atlantic-Mediterranean species of the genus De/tentosteus also have an increased num- ~ AHNELT AND GOSCHL: CAUDAL FIN LATERAL-LINE SYSTEM IN GOBIOIDEI 369 ber of neuromast rows on the caudal fin, but with four (D. guadrimaculatus) and six (D. colloni- anus) rows distinctly less then the above-mentioned species. A detailed description of the neuro- mast pattern on the caudal fin of these two species and a discussion if the increase in neuromast rows possibly following a certain order are under study by the senior author. DISCUSSION Three longitudinal lateral lines on each side of the caudal fin are characteristic for the basal gobioid genera Rhyacichthys Boulenger, 1901, Protogobius Watson and Péllabauer, 1998 and Terateleotris Shibukawa et al., 2001 (Springer 1983; Shibukawa et al. 2001). This pattern is also known from basal Perciformes (Jakubowski 1966, 1967). In all Gobioidei except the Rhyac- ichthyidae, sensu Shibukawa et al. (2001), these lateral lines are formed only by free neuromasts. Further, a discontinuous transversal lateral line on the base of the caudal fin formed by three short sections and plesiomorphic for Gobioidei is a feature seemingly also occurring in basal Perci- formes (Jakubowski 1966, 1967). Three or two longitudinal rows of neuromasts occur on the caudal fin of most Gobioidei with a fourth or third, transversal row more or less distinctly developed on the fin base (Sanzo 1911; Miller and El—Talwil 1974; Ahnelt and Duchkowitsch 2001; Shibukawa et al. 2001) (Fig. 1!A—B). In species with two longitudinal rows on the caudal fin, the ventral row lev is absent (Ahnelt et al. 2000; Ahnelt and Géschl 2003), a feature that occurs in the Odontobutidae, Eleotridae (Eleotrinae) and Gobiidae (Gobiinae and Gobionellinae) (Fig. 1C—D). [From a figure of a generalized sicydiine gobiid shown by Parenti and Maciolek (1993, fig. 3), Ahnelt and Géschl (2003) concluded that Sicydiinae have only two longitudinal rows of neuromasts on the caudal fin. However, all species investigated for this study have three longitudinal rows with the neuromasts invaginated and diffi- cult to detect]. Three longitudinal neuromast rows occur in the Rhyacichthyidae (preceded by lateral-line canals), 7erateleotris (unassigned genus), Odontobutidae, Eleotridae, Gobiidae (Amblyopinae, Gobiinae, Gobionellinae, Oxudercinae, Sicydiinae), Kraemeriidae, Microdesmidae and Ptereleotr- idae. Each of these longitudinal neuromast rows extends onto the interradial membrane between two branched caudal-fin rays in a characteristic pattern. Typically for Gobioidei with three longi- tudinal lateral lines is a larger distance between Iced and Icm than between lev and Iem (Fig. 1 A—B). This is the case because Ied is separated from Iem by three fin rays and Iem from lev by two fin rays. In species with two longitudinal lateral lines (led and Iem), these are also separated by three fin rays. Additionally, the neuromast rows do not run in the midline between two fin rays: Ied and lem follow the ventral side of a fin ray, lev follows the dorsal side of a fin ray. Seemingly, this pat- tern of the longitudinal lateral lines on the caudal fin is shared by most gobioids. A transversal row (ct) at the base of the caudal fin seems to be developed regularly and has been found in all above-mentioned families and subfamilies of Gobioidei including the Kraemeriidae, Microdesmidae and Ptereleotridae (Fig. 1). In Gobiinae, Iet often consists of only a few neuromasts, and in some species it is difficult to distinguish from the longitudinal rows. In gob- iine species this row consists often only of a single neuromast immediately anterior to each of the three longitudinal rows. Often these three neuromasts are somewhat larger then those of the longi- tudinal rows and separated from them by a more or less distinct gap. Additionally, a single neuro- mast or a short neuromast row below the origin of the most ventral longitudinal row (generally lev) forms the ventral end of Ict (Fig.1 A). Nevertheless, in such species the pattern of the transversal- ly arranged neuromasts follows the asymmetric pattern of the longitudinal neuromasts. A deviation from this arrangement occurs in L. guilleti. In this species, three neuromasts are arranged transver- 370 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16 sally (Ict) at the base of the caudal fin, distinctly separated from the single longitudinal row (lem). The dorsal neuromast of Ict is separated from the medial neuromast by only two caudal-fin rays, and the ventral neuromast by only one ray. This peculiar pattern is possibly the result of a reduced number of caudal-fin rays and a specialization. The caudal-fin rays of the related and similar tiny S. trigloides are more numerous, and the three neuromasts of Ict are separated from each other in the characteristic pattern described above. A transversal neuromast row occurs independently in Gobioidei, generally at the base of the caudal fin if two or three longitudinal rows are present (Ahnelt and Scattolin 2003; Ahnelt and Bohacek 2004); it also occurs in the basal Gobioidei Rhyacichthyidae and in 7erateleotris. T. aspro belongs to an assemblage of basal Gobioidei with a trunk lateral-line canal developed, but is derived in the absence of lateral-line canals on the caudal fin. The pattern with four neuromast rows on this fin (one transversal and three longitudinal rows) is plesiomorphic for all Gobioidei with lat- eral lines consisting only of free neuromasts. The transversal row (Iet) on the caudal fin base of T. aspro, figured by Shibukawa et al. (2001:fig. 6A), is discontinuous, similar to that of the Rhyac- ichthyidae. A discontinuous Iet, consisting of three parts, is likely plesiomorphic for the Gobioidei. The Odontobutidae are treated by most authors as the sister group of all other non-rhyaci- chthyid Gobioidei (e. g., Hoese and Gill 1993; Akihito et al. 2000; Shibukawa et al. 2001; Wang et al. 2001; Thacker 2003). It is, therefore, interesting that in the five species we investigated (Micropercops swinhonis, Odontobutis aurarmus, O. obscura, O. potamophila and Perccottus glenii) the lateral-line system on the caudal fin is developed in three different patterns: (1) M. swin- honis, with three longitudinal rows (Ied, lem, Iev) and a transversal row (Iet) reduced to three (sometimes four) neuromasts each close to the origin of the longitudinal rows (Fig. 1B), (ii) O. aurarmus and O. obscura with two longitudinal rows (led, lem) and a transversal row (Fig. 1C), and (iii) Perccottus glenii with one longitudinal row (lem) and a transversal row (Fig. 1E) [Wongrat and Miller (1991) and Miller (2003) mention three longitudinal rows on the caudal fin of P. glenii]. All five odontobutid species are peculiar in having a derived pattern of free neuromasts on the cau- dal fin, with P. g/enii having the most specialized one. The intra-familial relationships of the Odontobutidae (sensu Hoese and Gill 1993) are unre- solved (Akihito et al. 2000; Shibukawa et al. 2001; Thacker 2002). The arrangement of the later- al-line system on the caudal fin is possibly a further indication that the Odontobutidae are not monophyletic. The presence of three longitudinal lateral lines in M. swinhonis is plesiomorphic, but a transversal neuromast row reduced to three single neuromasts is apomorphic for Gobioidei. Two longitudinal lateral lines in O. auwrarmus and O. obscura and one in P. glenii are also derived char- acters. This neuromast pattern on the caudal fin of P g/enii is the most specialized within the Odontobutidae. The pattern of the lateral-line system on the caudal fin of odontobutid species is obviously more diverse than those of other gobioid species. In the group consisting of all gobioids except the Odontobutidae and Rhyacichthyidae, the Eleotridae are seemingly conservative in this character. We investigated species of 20 of the 34 or 35 eleotrid genera (Thacker 2003). All, except for one genus (Kribia), have the plesiomorphic three longitudinal neuromast rows developed. The lateral-line system on the trunk and caudal fin is less variable than on the head of Gobioidei. This explains why the lateral-line system of the head is widely used as an important tax- onomic tool for the classification of gobioid fishes. On the other hand, the more conservative char- acter of the neuromast pattern on the caudal fin allows us to hypothesize that the group of gobioids classified as Odontobutidae (sensu Hoese and Gill 1993) appeared early in the evolution of the Gobioidei and possibly evolved from a sister group within the Eleotridae. This hypothesis is sup- ported by a series of plesiomorphic features that place the Odontobutidae closer to the origin of the AHNELT AND GOSCHL: CAUDAL FIN LATERAL-LINE SYSTEM IN GOBIOIDEI 371 Gobioidei than to the Eleotridae (Hoese and Gill 1993). The Eleotridae have a more conservative pattern of neuromasts on the caudal fin as the investigated odontobutid taxa of Micropercops, Odontobutis and Perccottus. Finally, it can be concluded from the above results that the decrease and the increase of the number of longitudinal neuromast rows on the caudal fin in Gobioidei follow a certain order. A reduction in the number of neuromast rows obviously first affects the ventral row (lev) (Fig. 1A — E). By far, the majority of Gobioidei has three or two longitudinal rows developed on the caudal fin. Only a few species are known with the number of longitudinal rows increased or further decreased. Obviously, the dorsal row (Ied) is the next affected by reduction. In many species with only two longitudinal rows developed, this dorsal row is as a first step often distinctly shortened (e.g., Ahnelt and Géschl 2003) (Fig. 1D). In species with a single longitudinal row, the median row (lem) is always present (Fig. 1E). As with a decrease in neuromast rows, so too an increase of longitudinal neuromast rows on the caudal fin follows a scheme, a phenomenon currently under study by the senior author. In the first step (four longitudinal rows), the additional row is added ventrally, and with the addition of more rows, the gaps between Iev, Iem and Iced are closed. Finally, all longitudinal neuromast rows are separated by only a single fin ray. ACKNOWLEDGMENTS We thank D.J Siebert, O.A. Crimmen (BMNH), W.N. Eschmeyer, T. Iwamoto, D. Catania (CAS), E. Mikschi, H. Wellendorf (NMW), P.A. Hastings, R.H. Rosenblatt, C. Klepadlo, H.J. Walker (SIO) for their support and hospitality during our visits and for the loan of material. We also thank B.W. Coad, S. Laframboise (CMN), I. Nakamura, S. Harada (FAKU), F. Schiemer (IECB), J. Dorda-Dorda, A. Garvia (MNCN), P. Pruvost (MNHN), A. Sevesic (NMH), S.O. Kullander, E. Ahlander (NRM), GR. Smith, D.W. Nelson (UMMZ) and R.P. Vari, S.L. Jewett (USNM), for mak- ing material available. LITERATURE CITED AHNELT, H., ABDOLI, A., NADERI, M. AND B.W. Coap. 2000. Anatirostrum profundorum: a rare deep-water gobiid species from the Caspian Sea. Cybium 24:139-159. AHNELT, H., AND V. BOHACEK. 2004. The lateral-line system of two sympatric eastern Pacific gobiid fishes of the genus Lythrypnus (Teleostei: Gobiidae). Bulletin of Marine Science 73:31—S1. AHNELT, H., AND M. DUCHKOWITSCH. 2001. The lateral-line system of two Ponto-Caspian gobiid species (Gobiidae, Teleostei): a comparison. Folia Zoologica 50:217—230. AHNELT. H., AND J. GOsCHL. 2003. Morphological differences between the eastern Pacific gobiid fishes QOuietula guaymasiae and Quietula y-cauda (Teleostei: Gobiidae) with emphasis on the topography of the lateral-line system. Cybium 27:185-197. AHNELT, H., AND G. SCATTOLIN. 2003. The lateral-line system of a blind goby, Zyphlogobius californiensis, Steindachner 1879 (Teleostei: Gobiidae). Annalen des Naturhistorischen Museums in Wien 104 B:11—25. AKIHITO, A. IwaTA, T. KOBAYASHI, K. IKEO, T. IMANISHI, H. ONO, Y. UMEHARA, C. HAMAMATSU, K. SUGIYAMA, Y. IkKEAD, K. SAKAMOTO, A. FUMIHITO, S. OHNO, AND T. GosOBORO. 2000a. Evolutionary aspects of gob- ioid fishes based upon a phylogenetic analysis of mitochondrial b genes. Gene 259:5—15. AKIHITO, K. SAKAMOTO, Y. IKEDA, AND A. Iwata. 2000b. Suborder Gobioidei. Pages 1139-1310 in T. Nakabo, ed., Fishes of Japan with Pictorial Keys to the Species, second edition. Tokai University Press, Tokyo. BIRDSONG R.S., E.O. MurDY, AND F.L. PEzZoLD. 1988. A study of the vertebral column and median fin osteol- ogy in gobiid fishes with comments on gobioid relationships. Bulletin of Marine Science 42:174-214. Herver, J., AND M. Kovacic. 2002. Lebetus guilleti (Teleostei: Gobiidae) in the northern Adriatic Sea: first record and details on the species’ morphology. Annales 12:177—188. 37) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 16 Hoese, D.F., AND A.C. GILL. 1993. Phylogenetic relationships of eleotridid fishes (Perciformes: Gobioidei). Bulletin of Marine Science 52:415—-440. ILmN, B.S. 1930. Le systeme des Gobiidés. Zrabajos del Instituto Espanol de Oceanografia, Madrid 2:\—63. JAKUBOWSKI, M. 1966. Cutaneous sense organs of fishes. IV. The lateral-line organs in the perch-pike (Lucioperca lucioperca L.) and perch (Perca fluviatilis L.), their topography, innervation, vascularization, and structure. Acta Biologica Cracoviensia, Series: Zoologia 9:137—149. JAKUBOWSKI, M. 1967. Cutaneous sense organs of fishes. Part VII. The structure of the system of lateral-line canal organs in the Percidae. Acta Biologica Cracoviensia, Series: Zoologia 10:71-81. LARSON, H.K. 2001. A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its system- atic placement. Records of the Australian Museum, Supplement 62: \-vi, 1-233. Leviton, A.E., R.H. Gipps, H. HEAL, AND C.E. DAwson. 1985. Standards in herpetology and ichthyology: Part 1. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832. MILLER, P.J. 1963. Taxonomy and biology of the genus Leberus (Teleostei — Gobioidea). Bulletin of the British Museum (Natural History), Zoology 10:207-256. MILLER, P.J. 1973. The osteology and adaptive features of Rhvacichthys aspro (Teleostei: Gobioidei) and the classification of gobioid fishes. Journal of Zoology, London 171:397-434. MILLER, P.J.. 1986. Gobiidae. Pages 1019-1085 in P.J.P Whitehead, M.-L. Bauchot, J.-C. Hureau, J. Nielsen, and E. Tortonese, eds., Fishes of the North-eastern Atlantic and the Mediterranean, vol. 3. UNESCO: Paris. MILLER, P.J. 2003. Family Odontobutidae Hoese and Gill, 1993. Pages 131-132 in P.J Miller ed., The fresh- water fishes of Europe, vol. 8, pt. 1. AULA Verlag, Wiebelsheim. MILLER, P.J., AND M.Y. EL-TALWIL. 1974. A multidisciplinary approach to a new species of Gobius (Teleostei: Gobiidae) from southern Cornwall. Journal of Zoology, London 174:539-574. Mortara, S. 1918. La disposizione degli organi ciatiformi del genere Aphya e suoi rapporti con quella del genere Gobius. Revista Comitato Talassografico Italiano, Memoria 65:5—23. PARENTI, L.R., AND J.A. MACIOLEK. 1993. New sicydiine gobies from Ponape and Palau, Micronesia, with comments on systematics of the subfamily Sicydiinae (Teleostei: Gobiidae). Bulletin of Marine Science 53:945-972. PEZOLD, F. 1993. Evidence for a monophyletic Gobiinae. Copeia, 1993:634—643. SANZO, L. 1911. Distribuzione delle papille cutanee (organi ciatiformi) e suo valore sistematico nei Gobi. Mitteilungen aus der Zoologischen Station zu Neapel, Berlin 20:249-328. SHIBUKAWA, K., A. IWATA, AND S. VIRAVONG. 2001. Terateleotris, a new gobioid fish genus from the Laos (Teleostei, Perciformes), with comments on its relationships. Bulletin of the National Scicence Museum, Tokyo, Ser A. 27:229-257. SPRINGER, V.G. 1983. 7vson belos, new genus and species of western Pacific fish (Gobiidae, Xenisthminae), with discussions of gobioid osteology and classification. Smithsonian Contributions to Zoology 390:1—40. TAKAGI, K. 1988. Cephalic sensory canal system of the gobioid fishes of Japan: comparative morphology with special reference to phylogenetic significance. Journal of the Tokyo University of Fisheries 75:499-568. THACKER, C.E. 2003. Molecular phylogeny of the gobioid fishes (Teleostei: Perciformes: Gobioidei). Molecular Phylogenetics and Evolution 26:354—-368. WarTSON, R.E. AND C. POLLABAUER. 1998. A new genus and species of freshwater goby from New Caledonia with a complete lateral line. Senckenbergiana biologica 77:147-153. Wonarat, P. AND P.J. MILLER. 1991. The innervation of the head neuromast rows in Eleotridine gobies (Teleostei: Gobioidei). Journal of Zoology, London 225:27-42. WANG H.-Y., M.-P. Tsal, J. DEAN AND LEE S.-C. 2001. Molecular phylogeny of gobioid fishes (Perciformes: Gobioidei) based on mitochondrial 12S rRNA sequences. Molecular Phylogenetics and Evolution 20: 390-408. ZANDER, Z.D., AND H. JELINEK. 1976. Zur demersen Fischfauna im Bereich der Grotte von Banjole (Rovinj, YU) mit Beschreibung von Speleogobius trigloides n. gen., n. sp. (Gobiidae, Perciformes). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 73:265—280. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 17, pp. 373-376, 5 figs. September 30, 2004 Gorgasia thamani, a New Species of Garden Eel from Fiji (Teleostei: Congridae: Heterocongrinae) David W. Greenfield! and Sean Niesz? ! California Academy of Sciences, 875 Howard Street, San Francisco, California 94103; 2 Department of Zoology, University of Hawaii, Honolulu, Hawaii 96822. A very elongate garden eel, Gorgasia thamani, is described from Koro Island, Fiji. The new species is most similar to G barnesi, known from the Philippines, Papua New Guinea, and the Solomon Islands, but differs by being more slender with a shorter head and mouth and a different head pore pattern. While conducting a survey of the marine fishes of Fiji, a colony of very elongate garden eels was discovered at the west side of Koro Island, Fiji, off Kade Village on a fine sand bottom in 14 to 15.2 m of water. The species, although similar in appearance and length to G. barnesi, differs from that species in a number of characters and is described here. Gorgasia barnesi is known from Indonesia, the Philippines, Papua New Guinea, and the Solomon Islands. The new species is placed in the genus Gorgasia because the upper lip is not confluent medi- ally across the face of the snout, the anterior nostrils and ethmoid sensory pores are separate from the upper lip, and the mouth is relatively large and slightly oblique, its angle reaching to below anterior half of eye, characters not present in the only other genus in the subfamily, Heteroconger. MATERIALS AND METHODS Information for the holotype is presented first, followed by the range and mean or mode for all type material. All measurements are presented first as thousandths of total length, and then again as either percentage of snout-anus length or percentage of head length for comparison with data presented in Castle and Randall (1999). Methods of counting and measuring follow Castle and Randall (1999), who followed Boéhlke (1989), Smith (1989) and McCosker et al. (1989). Institu- tional abbreviations are as listed in Leviton et al. (1985). SPECIES DESCRIPTION Gorgasia thamani Greenfield and Niesz, sp. nov. (Figs. 1—5) MATERIAL EXAMINED.— HOLotTyPeE: CAS 219365, 1101.4 mm TL, Fiji, Koro Island, west side off Kade Village, 17°21.261'S, 179°22.435’E, sand, 14-15.2 m, 10 November 2002, field number G02-164, collected by D.W. Greenfield, K.R. Longenecker, K.S. Cole and R.C. Langston. PARATYPES: all collected with holotype: BPBM 39441, 1186 mm TL; FMNH 113585, 1180 mm TL; 3 Research Associate, Department of Ichthyology, California Academy of Sciences and Emeritus Professor, Univer- sity of Hawaii. Mailing address: Moss Landing Marine Laboratory, 8272 Moss Landing Road, Moss Landing, CA 95039. 3s) 374 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 17 USNM 376279, 1185 mm TL; BM(NH) 2004.4.8.1, 1053 mm TL; AMS 1I.43260-001, 913 mm TL; NSMT-P68327, 831 mm TL; SAIAB 74101, 776 mm TL; CAS 219366 (2), 705-752 mm TL. OTHER MATERIAL COLLECTED WITH THE TYPE SERIES — CAS 219367 (29), 680-1141 mm TL. ADDITIONAL MATERIAL EXAMINED — Gorgasia barnesi: Indonesia, Ambon: BPBM 36482(7) 422-1209 mm TL. DraGnosis.— An elongate, slender species of Gorgasia (depth at gills 0.69—0.91 percent total length) having 41—44 preanal pores, 69-75 preanal vertebrae, 213-225 total vertebrae, head pores POM 6+2 and ST 1+2, head 9.2—10.9 percent of snout-anus length, and mouth length 2.7—3.2 per- cent of snout-anus length. DESCRIPTION.— Vertebral formula 7/69/213, 6—8 (usually 7), 69-75, 213-225. Lateral-line pores before anus 41, 41-44 (usually 42-44). Total lateral-line pores 120, 118-123 (usually 122). Pectoral-fin rays 13, 11-13 (usually 11). Dorsal rays before anus 167 (holotype only). Total dorsal- fin rays 571 and total anal-fin rays 381 in holotype. Head pores in holotype SO 1 + 4, IO 2 + 4, POM 6 + 2, ST 1 + 2. Proportions as thousandths of total length: snout-anus length 262.2 (262.2— 318.0; 295.1); head length 28 (26.6—34.6; 29.3); snout length 5.3 (4.8-6.3; 5.3); eye diameter 5.2 (4.5—7.9; 5.7); gill opening 2.7 (2.5—3.3; 2.9); pectoral-fin length 4.1 (3.9-6.0; 4.6); depth at gill opening 7.2 (6.9-9.1; 7.9); depth at anus 4.8 (4.88.8; 6.8); front of upper jaw to rictus 8.4 (7.79.9; 8.8); front of upper jaw to end of maxilla 10.3 (9.5—14.7; 10.7). Proportions as percent of snout-anus length: head length 10.7 (9.2—10.9; 9.9); predorsal length 11.7 (9.1—11.7; 10.6). Propor- tions as percent of head length: snout length 18.9 (17.5—20.3; 18.6); eye diameter 18.6 (17.0—23.0; 19.8); front of upper jaw to rictus 3.2 (2.7—3.2; 3.0); gill opening 9.7 (9.0—11.8; 9.9); pectoral-fin length 14.7 (12.6—18.7; 16.0); depth at gills 25.6 (25.2—31.9; 27.4); depth at anus 17.0 (17.0-28.8; 23.4). Color of fresh specimen: Background coloration gray, paler anteriorly and ventrally on head. ~ «.. 1 ie . et ae -_ FIGURE 1. Holotype of Gorgasia thamani, CAS 219365. SS GREENFIELD AND NIESZ: NEW SPECIES OF GARDEN EEL FROM FIJI 375 Body covered with dense, irregular, brown Vice ee spots; tip of tail and snout almost solid brown. fd tas a uN Brown spots less dense ventrally on head and rs ’ 1 on pectoral fins. Pores on head and anterior Is 0 1 portion of lateral line within distinct white ig Rea i ) spots. Pupil black; iris light yellow. Color in alcohol: Background coloration [& ? 4 I A mottled brown-gray; body covered with dense, BS \ Ie irregular brown spots. Tip of tail light brown. [a3 : 2\ A Jaws and underside of head cream with scat- _/s ‘ tered brown spots. Pores on head and anterior if portion of lateral line within distinct white 2 spots. Pupil of eye white; iris black. FIGURE 2. Dentition of holotype of Gorgasia thamani, EtymoLocy.— Named in honor of Dr, Po aS Randolph R. Thaman, Professor of Pacific Islands Biogeography at the University of the South Pacific in Fiji, who has provided unending assistance to us in arranging for the Survey of Marine Fishes Project funded by the National Science Foundation. Without Dr. Thaman’s assistance this project literally would not have been possible. In addition, Dr. Thaman is a leader in promoting conservation measures in Fiji, both terrestrial and marine. COMPARISONS.— Gorgasia thamani is most similar to G barnesi Robinson and Lancraft, and keys to that species using Castle and Randall (1999) because of its high total vertebral and lateral- line pore counts and coloration. It also is a large species, our largest specimen is 1185.8 mm, whereas the largest specimen of G barnesi examined by Castle and Randall (1999) was 1212 mm, the largest known for a garden eel. It differs from G barnesi, however, by being a more slender, elongate species, with the depth at the gills 0.69—0.91 percent of total length versus 1.1—-1.6 in G barnesi (Fig. 3). It also has a shorter head length, 9.2—10.9 versus 11.2—14.1 percent of snout-anus length (Fig. 4). The mouth, as measured by front of upper jaw to rictus, is shorter in G thamani, 2.7—3.2 percent of snout-anus length, versus greater than 3.6 in G barnesi (Fig. 5). Head pores are POM 6+2 and ST 1+2 in G thamani versus POM 5+4 and ST 2+1 in G barnesi. ———— } i i” i8 4 * fe ae ACKNOWLEDGMENTS | /®@ - > 6 ? 14 ‘ B | @ ee ' @ e1.4- ee mn 13 + 4 o | a i.2 4 ® 12 e 0 Py e ig ‘3 $1.04 fa AY, As A Sage a ° e 4 7 9.8 + Gore q 10 ee A °o ® F] 9° Z B 1 AEA ge RS a eee 9 —————————— 400 900 1400 400 900 1400 TOTAL LENGTH mm TOTAL LENGTH mm FIGURE 3. Body depth at gills as percentage of total length, open circles = Gorgasia thamani, closed circles = Gorgasia barnesi. FicureE 4. Head length as percentage of snout-anus length, open circles = Gorgasia thamani, closed circles = Gorgasia barnesi. 376 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 17 5.070 a m 4-0 ‘ e e oe r r a neta ts 5 g fo) e <0 3.074 Be és ° } DFG lo ee 400 900 1400 TOTAL LENGTH mm FicuRE 5. Mouth length as percentage of snout-anus length, open circles = Gorgasia thamani, closed circles = Gorgasia barnesi. We would like to thank K.R. Longenecker, K.S. Cole, and R.C. Langston for assistance in col- lecting specimens, and Captain B. Vasconcellos and the crew of the Moku Mokua Hine for assis- tance in the field. We are grateful to J. Seeto, GR. South, R.R. Thaman , and R.W. Tuxton of the University of the South Pacific, Fiji for facilitating our collecting in Fiji. We also thank the Fijian Government and local village chiefs for permission to collect fishes. Special thanks are due Susan Monden for drawing the figures. This research was supported by National Science Foundation grants INT97-29666 and DEBO-1027545, and Sea Grant Project R/FM-6PD. LITERATURE CITED BOHLKE, E.B. 1989. Methods and terminology. Pages 1—7 in E.B. Bohlke, ed., Fishes of the Western North Atlantic, Part Nine, Volume One. Orders Anguilliformes and Saccopharyngiformes. Sears Foundation for Marine Research, New Haven, Conn. CASTLE, P.H.J., AND J.E. RANDALL. 1999. Revision of Indo-Pacific garden eels (Congridae: Heteroceongrinae), with descriptions of five new species. /ndo-Pacific Fishes (30):1—52. LeviTON, A.E., R.H. Gress, 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. McCoskeR, J.E., E.B. BOHLKE, AND J.E. BOHLKE. 1989. Famity Ophichthidae. Pages 254-412 in E.B. Bohlke, ed., Fishes of the Western North Atlantic, Part Nine, Volume One. Orders Anguilliformes and Saccopharyngiformes. Sears Foundation for Marine Research, New Haven, Conn. SMITH, D.G. 1989. Family Congridae. Pages 460-567 in E.B. Bohlke, ed., Fishes of the Western North Atlantic, Part Nine, Volume One. Orders Anguilliformes and Saccopharyngiformes. Sears Foundation for Marine Research, New Haven, Conn. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 18, pp. 377-383, 6 figs. September 30, 2004 Notes on Drypta longicollis MacLeay and the Status of the Genus-group Name Desera Dejean, 1825 (Coleoptera: Carabidae: Dryptini) Hongbin Liang!, David H. Kavanaugh2, and Mingyi Tian? ! Institute of Zoology, Beijing 100080, China; ? California Academy of Sciences, 875 Howard Street, San Francisco, California 94103, USA; > South China Agricultural University, Guangzhou 51064, China The taxonomic status of Drypta longicollis MacLeay, 1825, mistreated as Desera longicollis (MacLeay) for over a century, is clarified as a species of Drypta Latreille. With Drypta longicollis as type species, genus Desera Dejean is a subjective junior synonym of Drypta; and Dendrocellus Schmidt-Gobel is the valid generic name for species with pectinate tarsal claws previously included in Desera. Specimens identi- fied as Drypta longicollis MacLeay in Andrewes’ collection (The Natural History Museum, London) are found instead to represent Dendrocellus unidentatus (MacLeay, 1825). Enhanced descriptions of Drypta longicollis and Dendrocellus unidentatus and photographs of their type specimens are provided. Desera Dejean (1825) is one of the two Old World genera currently recognized in tribe Dryptini of family Carabidae. Its members differ from those of the other genus, Drypta Latreille (1796), in having pectinate tarsal claw (Andrewes 1936). To date, 18 Desera species have been described (Lorenz 1998a), mainly distributed in southeastern Asia. While sorting specimens of Carabidae in the collection of the Institute of Zoology (IOZ), Beijing, the first author (Liang) discovered several interesting Desera specimens collected from Yunnan Province, China. These specimens with brown femora, black tibia, and acute outer angles of their elytra keyed out as Desera longicollis (MacLeay 1825), based on the works of Heller (1923:303), Andrewes (1936:136) and Hansen (1976:403). However, Jedlicka’s monograph (1963:486) indicated that, in D. longicollis (MacLeay), the “Aussenwinkel der Flugeldecken stumpf’ [outer angles of elytra acute]. In an effort to resolve this contradiction, Liang wrote to Dr. Thierry Deuve at the Muséum National d’ Histoire Naturelle (MNHN) in Paris to request informa- tion about the type specimens of this species. Deuve kindly examined the types and informed Liang that the outer angle of each elytron in the types of Drypta longicollis MacLeay were rounded (Fig. 1), not acute. Liang then sent his specimens to the third author (Tian) in MNHN for comparison. Subsequent morphological comparison revealed that these specimens were not identical to those of D. longi- collis. Vian also noticed that the types of Drypta longicollis, with simple claws, were actually mem- bers of the genus Drypta, the genus in which this species was originally described, and not of the genus Desera as presently conceived. Subsequently, Liang borrowed specimens of Desera uniden- tata (MacLeay 1825) and Drypta longicollis MacLeay from the Natural History Museum (NHM) in London and MNHN and determined that the specimens from Yunnan were identical with Desera unidentata (MacLeay) (Fig. 2) specimens. ST 378 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 18 Habitus and Labels of Type Specimens (scale lines = 2.0 mm) FIGURE | (above). Lectotype of Drypta longicollis MacLeay, 1825 (from India). FIGURE 2 (below). Holotype of Drvpta unidentata MacLeay, 1825 (from Java). LIANG ET AL.: ON DRYPTA LONGICOLLIS AND GENUS-GROUP NAME DESERA 3)/3) The fact that Drypta longicollis MacLeay is actually a member of genus Drypta Latreille, and not of Desera Dejean, is also problematic because D. longicollis is the type species of Desera Dejean. The valid generic name for the species with pectinate tarsal claws, currently included in genus Desera, needs to be established. The purposes of this paper are: (1) to establish the valid generic name for the species current- ly included in genus Desera Dejean; (2) to remove Drypta longicollis MacLeay from Desera and return it to genus Drypra Latreille, and (3) to provide additional descriptive information that will aid in the identification of specimens of both Drypta longicollis and “Desera” unidentata. The abbreviations used in this paper are as follows: Ant 1 = antennomere | (scape); Ant 3 = antennomere 3; PL = length of pronotum measured along median line; PW = Widest width of pronotum; EL = length of elytra; and EW = widest width across both elytra. STATUS OF THE GENUS NAME DESERA DEJEAN Latreille (1796:75) described genus Drypta but did not list any included species with his orig- inal description. He later designated Carabus emarginatus Gmelin (= Carabus dentatus P. Rossi) as type species (Latreille 1802). The status of Drypta Latreille as a valid genus is unquestioned. In his description of Drypta longicollis, Dejean (1825:28) listed “Desera Bonelliana. Leach” as a synonym of that species, and, at the end of his description, he mentioned that Leach had estab- lished a distinct genus, Desera, for this species. Our review of Leach’s pertinent published works (Leach 1815 and 1817) and of his classification as reported by Samouelle (1819) failed to turn up any mention of a genus Desera or of a species, Desera bonelliana; so Dejean’s assignments must have been based on determinations in Leach’s collection and not on a published account. Hope (1831:21) briefly described Desera nepalensis, but failed to note the presence of pectinate tarsal claws in this species. Later (Hope, 1838), he cited Leach as the author of Desera and designated Cicindela cylindricollis Fabricius (= Carabus distinctus P. Rossi) as the type species of the genus. In a subsequent brief description of Desera, Hope (1838:105) once again omitted any mention of pectinate tarsal claws. His apparent failure to appreciate this distinguishing feature of true “Desera” species probably accounts for his selection of a true Drypta, D. distincta (Rossi), as his type species for Desera. The authorship of genus Desera has remained a point of contention; some authors (e.g., Bousquet 2002; L6bl and Smetana 2002) cite Dejean (1825) as the author, others (e.g., Csiki 1932; Andrewes 1939; Lorenz,1998b) credit Hope (1831). If Hope is the author, based on his use of the name in 1831, then Desera nepalensis is the type species of Desera by monotypy. If his claim of authorship were based on his 1838 paper, with his designation of Cicindela cylindricollis Fabricius as type species, then Desera would be a subjective junior synonym of Drypta Latreille, as was sug- gested by Andrewes (1939). In fact, Bousquet (2002) has settled the argument to our satisfaction, citing Dejean as author, with Article 11.6 of the 1999 International Code of Zoological Nomencla- ture as the justification, and with Drypta longicollis MacLeay as type species by monotypy. This means that Desera Dejean is a junior subjective synonym of Drypta Latreille. In 1846, Schmidt-Gébel described the genus Dendrocellus, listed distinguishing features, including the presence of pectinate tarsal claws, and included three species. Two of these were described as new. The first, Dendrocellus discolor, is now recognized as a junior synonym of Desera nepalensis Hope. The second, Dendrocellus flavipes, which Schmidt-Gébel had misidenti- fied as Drypta flavipes Wiedemann and transferred to his new genus, is conspecific with and a jun- ior synonym of Drypta geniculata Klug. The third species was D. geniculata Klug itself. Schmidt- Gobel did not designate one of these as type species. 380 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 18 Chaudoir (1861:545) first included Drypta longicoillis MacLeay in Dendrocellus, and several other workers followed (Bates 1892; Andrewes 1919 and 1936; Winkler 1924; Jedlicka 1963; and Hansen 1967). Dupuis (1912) cited Dendrocellus as a subgenus of Desera in his description of Desera gilsoni from Taiwan. Andrewes (1939:133) synonymized Desera with Drypta, based on Hope’s (1938) type species designation, recognized Dendrocellus as a valid genus name, and selected D. discolor (= Desera nepalensis Hope) as type species. However, Dendrocellus has been treated as a junior synonym of Desera Dejean (or Hope) by most recent workers (e.g., see Lorenz, 1998b; and L6bl and Smetana, 2003). Our conclusion that the type species of Desera Dejean (i.e., Drypta longicollis MacLeay) should be included in Drypta leads us to join Andrewes (1939) in treating Desera Dejean (not Hope) as a synonym of Drypta Latreille and Dendrocellus Schmidt-Gébel as the valid name for the species with pectinate tarsal claws formerly included in Desera Dejean (or Hope [see above]). SPECIES REDESCRIPTIONS Drypta longicollis MacLeay, 1825 (Figs. 1, 3-4) Drypta longicollis MacLeay, 1825:28 (India) Drypta longicollis Dejean, 1825:185 (India). Dendrocellus longicollis Dejean; Bates, 1892:385 (Burma). Desera longicollis MacLeay; Jedlicka, 1963:486 (Burma, India). SPECIMENS EXAMINED.— Lectotype (Fig. 1), a male (MNHN), “Type, Coll. Dejean”, “Ex Museo, Chaudoir”;‘‘/ongicollis Dej., Indes Orient.”; paralectotype, 1 female (MNHN), “Ex Museo, Chaudoir’, “/ongicollis Dej., Indes Orient.” Both lectototype and paralectotype are designated here. DESCRIPTION.— Tarsal claws in type specimens sim- ple (Fig. 3) and outer angle of each elytron round (Fig. 4). 3 4 Ant | with basal third brown, apical two-thirds black. Ratio Ant | : Ant 3 = 4.20 (paralectotype); ratio PL : PW = 1.65 (lectotype), 1.64 (paralectotype); ratio EL : EW = ana, aa 1.74 (lectotype), 1.78 (paralectotype); ratio EW: PW= 5 6 2.30 (lectotype), 2.29 (paralectotype). For additional mor- phological characteristics, refer to Dejean’s description (1825: 185-186). sewer = a DISTRIBUTION.— India and Myanmar (= Burma). FIGURES 3-4. Drypta longicollis Macleay, REMARKS.— Drypta longicollis was described by 1825 (lectotype, from India): (3) fore-tarsal both MacLeay and Dejean in 1825, based on the same claws; (4) outer angle and sutural angle of left specimens (in Dejean’s collection) from India (MacLeay Rue eis $6. Diplo 1825:28; Dejean 1825:185). MacLeay’s work was several 1925 (nolo aoe ap ©) ie months earlier than Dejean’s, and therefore this species claws; (6) outer angle and sutural angle of left name is attributed to MacLeay (Andrewes 1919:134). elytron. Considering his generally excellent work on the cara- a ar ee peice int ies ae bids of Middle and Southeast Asia, it is strange that ~ Fit Andrewes, after examination of the type of Drypta longicollis MacLeay, still mistreated it as rep- resenting a Desera species. He also misidentified specimens of Dendrocellus unidentatus (MacLeay) as Drypta longicollis MacLeay (see “Specimens examined” below). LIANG ET AL.: ON DRYPTA LONGICOLLIS AND GENUS-GROUP NAME DESERA 381 Dendrocellus unidentatus (MacLeay), 1825, new combination (Figs. 2, 5—6) Drypta unidentata, MacLeay, 1825:28 (Java). Desera unidentata (MacLeay), Andrewes, 1919:167 (Java) Dendrocellus longicollis Dejean; Chaudoir, 1861: 45 (India). Desera longicollis MacLeay; Andrewes, 1936:136 (India). Desera longicollis Dejean; Hansen, 1967:403 (Laos). SPECIMENS EXAMINED.— Total 15 specimens. Holotype, | female (NHM), “wnidentata Mac.”, “Java, Horsfield, 60-15”, “60-15, E.I.C.”, “Type, H.T.”, “58”, “54” ; 1 specimen (NHM, sex status undeterminable), “Bonvouloir collection”, “Desera unidentata MacLeay, compared with type H.E.A.”, “wnidentata Macleay (Dendrocellus) Schmidt-Goebel, Java”, “H.E. Andrewes coll., B. M. 1945-97”; 1 female (NHM), “Dammerman, N.W. Seomba, Laora 100M, 104 IV 1925”, “Ex Mus., Buitenzurg”, “Desera unidentata Macl., H.E. Andrewes det.”; 1 male (NHM), “Andaman Is., 1915-38”, “Ex Coll., Brit. Mus.”, “Desera longicollis De}. (See back), compared with type, H.E.A., [on the back of the label] in type the angle of truncture is not dentate”; 1 male (NHM), “Annam, Keng trap, May 1917, R.V. de Salvaza’”, “Desera longicollis Dejean, H.E. Andrewes det.”, “H.E. Andrewes Coll., B.M. 1945-97”; 1 female (NHM), “47252”, “Captn Wimberley”, “Andaman Islands”, “Fry Coll., 1905.100, “Desera longicollis Dej., H.E. Andrewes det.”, “Dendrocellus longicollis Dej., Andaman Is.”; 1 female (MNHN), “Malacca, H. Deyrolle”, “Ex Museo, Chaudoir”, “wnidentata, Java, C. Gary, Reiche”; | male (MNHN), “Java”, “Ex Museo, Chaudoir”, “unidentata, Java, C. Gary, Reiche”; 2 females (MNHN), “Ex Museo, Chaudoir’’, “wnidentata, Java, C. Gary, Reiche”; | male (IOZ), “China, Yunnan Province, Nujiang Prefecture, Lushui County, Liuku Township, Liuku, 800m”, N25.86010°, E98.85155°, 25-26 June 2000, Stop #00-7, D.H. Kavanaugh & H.-B. Liang collectors”; | male and 2 females (IOZ), “Yunnan, Mengla’, “20 April 1982, Peiyu Yu collector, by light trap”; 1 female (IOZ), “Yunnan, Xishuangbanna, Xiaomengyang, 850m”, 25 April 1987, Shuyong Wang collector”. DESCRIPTION.— Because Andrewes (1919:167—168) gave a quite detailed redescription, we simply add the following: Ratio Ant 1 : Ant 3 = 3.67 (type), 3.26—-3.56 (others); ratio PL : PW = 1.62 (type), 1.50—1.76 (others); ratio EL : EW = 1.67 (type), 1.70—1.82 (others); ratio EW : PW = 2.47 (type), 2.24—2.52 (others). DISTRIBUTION.— Indonesia (Java), India (Andaman Is.), Vietnam (central), and China (Yunnan). REMARKS.— Specimens of Dendrocellus unidentatus (MacLeay) can be distinguished from those of all other known Dendrocellus species by the following combination of character states: femora brown except for apices (black), tibia black, outer angles of elytra strongly dentate (Fig. 6), elytra widened posteriad, elytral intervals densely punctate, tarsal claws markedly pectinate with length of individual pectinations (“teeth”) subequal to width of base of tarsal claw (Fig. 5). Andrewes (1919:167) gave a redescription of this species; but there are several errors included, namely: 1) this species was reported as restricted to Java (but present materials indicate that its dis- tribution extends from Java north to India, Laos, Vietnam, and China); 2) antenomere 3 was described as having its base and apex red and separated by a black subapical ring (but in all spec- imens we examined, the basal half of this antennomere was blackish brown and its apical half brown); 3) the prothorax was described as nearly twice as long as wide (but it is only 1.50—1.75 times as long as wide among specimens we have seen); and 4) the tarsal claws were described as finely pectinate (but they are markedly pectinate in all specimens that we examined). 382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 18 ACKNOWLEDGMENTS The authors thank Dr. Thierry Deuve (MNHN) and Ms. Christine Taylor (NHM) for loans of specimens. Thanks are also due to Dr. Yves Bousquet (Eastern Cereal and Oilseed Research Centre, Canada) and Mr. Paul Marek (California Academy of Sciences), for providing essential literature, to Boris Kataev (Institute of Zoology, St. Petersburg) for alerting us to a potential problem with the generic name, Desera, and to Professor Peiyu Yu (IOZ) for reviewing a draft of this paper. Photographs of habitus were taken by senior photographer Guoging Mai (IOZ). This work was sup- ported by National Natural Science Foundation of China (Grant No. 30000026) and U.S. National Science Foundation (Grant No. 0103795). This paper represents contribution no. 23 of the China Natural History Project of the California Academy of Sciences. LITERATURE CITED ANDREWES, H.E. 1919. On the types of Oriental Carabidae in the British Museum, and in the Hope Department of the Oxford University Museum. Transactions of the Entomological Society of London 1919:19-217. ANDREWES, H.E. 1936. Keys to some Indian genera of Carabidae (Col.) VI. The genera Drypta and Desera. Proceedings of the Royal Entomological Society of London, ser. B, 5: 134-136. ANDREWES, H.E. 1939. Papers on the Oriental Carabidae. XX XV. On the types of Indian genera. Annals and Magazine of Natural History, ser. 11, 3:128—139. Bates, H.W. 1892. Viaggio di Leonardo Fea in Birmania e Regioni Vicini XLIV. List of the Carabidae. Annali del Museo Civico di Storia Naturale di Genova, ser. 2, X11:267-428. Bousquet, Y. 2002. Additions and corrections to the world catalogue of genus-group names of Geadephaga (Coleoptera) published by Wolfgang Lorenz (1998). Folia Hevrovskyvana, Supplementum 9:1—78. CuHAuporR, M. 1861. Matériaux pour servir a |’ édude des cincindéletes et des carabiques. Bulletin de la Société Impériale des Naturalistes de Moscou 34:545-—551. Csiki, E. 1932. Pars 124. Carabidae: Harpalinae VII. Pages 1279-1598 in W. Junk and S. Schenkling, eds., Coleopterorum catalogus, Volumen III. Carabidae III. W. Junk, Berlin. DEJEAN, P.PF.M. A. 1825. Species général des coléoptéres de la collection de M. de Baron Dejean. Tome I. Crevot, Paris. xxx + 463 pp. Dupuis, P. 1912. H. Sauter’s Formosa-Ausbeute Carabidae (2™* contribution). Annales de la Société Ento- mologique de Belgique 56:308-338. HANSEN, W. 1967. Contribution a la connaissance des coléopteres carabiques du sud-est asiatique, sur le genre Desera Hope (Coleoptera Carabidae Dryptinae). Bulletin et Annales de la Société Royal Entomologique de Belgique 103:397-413. HELLER, K.M. 1923. Some new Malayan Carabidae, especially Philippine. Zhe Philippine Journal of Science 23:295-305. Hope, F.W. 1831. Synopsis of the new species of Nepaul insects in the collection of Major General Hardwicke. Zoological Miscellany \:21—32. Hope, F.W. 1838. The Coleopterist'’s Manual, part the second, containing the predaceous land and water bee- tles of Linneus [sic] and Fabricius. Henry Bohn, London. xvi + 168 pp. JEDLICKA, A. 1963. Monographie der Truncatipennen aus Ostasien, Lebiinae — Odacanthinae — Brachyninae (Coleoptera, Carabidae). Entomologische Abhandlungen Berichte aus dem Staatliches Museum fiir Tierkunde in Dresden 28(7):269—-568. LATREILLE, P.A. 1796. Précis des caractéres génériques des insectes, disposs dans un order naturel. A. Brive, Bourdeaux. xiv + 208 pp. LATREILLE, P.A. 1802. Histoire naturelle, générale et particuliére des crustacés et des insects. Ouvrage faisant suite a l’histoire naturelle générale et particuliére, composée par Leclerc de Buffon, et rédigée par C. S. Sonnini, membre de plusieurs sociétés savantes. Familles naturelles des genres. Tome troisieme. Dufart, Paris. xii + pp. 13-467. Le LIANG ET AL.: ON DRYPTA LONGICOLLIS AND GENUS-GROUP NAME DESERA 383 LEACH, W.E. 1815. Entomology. Pages 57-172 in D. Brewster, ed., The Edinburgh Encyclopaedia, vol. 9, pt. 1. Blackwood, Edinburgh. LEACH, W.E. 1817. The Zoological Miscellany; being Descriptions of New, or Interesting Animals, vol. 3. E. Nodder and Son, London. 151 pp. Losi, I., and A.-Smetana. 2003. Catalogue of Palaearctic Coleoptera. Volume 1. Archostemata — Myxophaga — Adephaga. Apollo Books, Stenstrup. 819 pp. LORENZ, W. 1998a. Systematic List of Extant Ground Beetles of the World (Insecta Coleoptera “Geadephaga’” : Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhyzodinae). W. Lorenz, Tutzing. 502 pp. LoRENZ, W. 1998b. Nomina carabidarum. A Directory of the Scientific Names of Ground Beetles (Insecta, Coleoptera “Geadephaga”: Trachyvpachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). W. Lorenz, Tutzing. 937 pp. MacLeay, W.S. 1825. Annulosa Javanica, or an Attempt to Illustrate the Natural Affinities and Analogies of the Insects Collected in Java by Thomas Horsfield, M.D. F-L. & GS. and Deposited by Him in the Museum of the Honorable East-India Company. No. |. Kingsbury, Parbury & Allen, London. xii + 50 pp. SAMOUELLE, G. 1819. The Entomologist's Useful Compendium; or an Introduction to the Knowledge of British Insects. Thomas Boys, London. 496 pp. SCHMIDT-GOBEL, H.M. 1846. Faunula Coleopterorum Birmaniae, adjectis nonnullis Bengaliae indigenis. Med. Dr. Joh. Wilh. Helfer ’s hinterlassene Sammlungen aus Vorder- und Hinter-Indien. Nach seinem Tode im Auftrage des béhm. National-Museums unter Mitwirkung Mehrerer. Gottlieb Haase Sdhne, Prague. viii + 94 pp + 3 pl. WINKLER, A. 1924. Catalogus Coleopterorum regionis palaearcticae, Pars 2, pp. 113-240. Winkler and Wagner, Vienna. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 19, pp. 384-394, 3 figs., 2 tables. September 30, 2004 Two new Scorpionfishes (Scorpaenidae) from the South Pacific John E. Randall!:3 and David W. Greenfield? ! Bishop Museum 1525 Bernice St., Honolulu, Hawaii 96817-2704; Department of Ichthyology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103. Scorpaenopsis eschmeyeri, formerly tentatively identified as S. possi, is described as new from 17 specimens from Fiji, Chesterfield Islands in the Coral Sea, New Caledonia, and the southern Great Barrier Reef. It is separated from S. possi by the lack of the pretympanic spine that is unique to S. possi, by its smaller size (largest specimen, 142 mm SL, compared to 194 mm for S. possi), and by a longer head, snout, upper jaw, and predorsal length, compared to specimens of possi of the same size range. Scorpaena lacrimatus is described from a single specimen, 198 mm SL, taken in 400 m off Tahiti, Society Islands. It is distinct from other species of the genus by the combination of the following characters: deep body (depth 2.5 in SL), 17 pectoral rays, and 61 scales in longitudinal series. Scorpionfishes, aptly named for their venomous fin spines, are not well represented at islands of the South Pacific, compared to the rest of the Pacific. Only 60 species of scorpaenids are known from the South Pacific from Lord Howe Island and New Caledonia to Easter Island (Solomon Islands and Vanuatu not included). By contrast, Poss in Carpenter and Niem (1999) listed 131 scor- paenids (12 with a question mark) for the central and western Pacific (Hawaiian Islands not includ- ed). The Indo-Pacific scorpionfish genus Scorpaenopsis was revised by Randall and Eschmeyer (2001); 24 species were recognized, of which eight were described as new. One of these, the wide- ranging S. possi, included ten specimens from the southwest Pacific from Fiji to the southern Great Barrier Reef that lack the characteristic small spine (called the pretympanic spine) dorsally on the head before the tympanic spine. No typical S. possi were found at any of these localities. These ten specimens were not listed as paratypes of S. possi. Recent collections of shore fishes in Fiji by the authors and associates resulted in 12 additional specimens of the same form. Our study of these specimens, plus seven of the nontype “possi” (one CAS lot of three specimens from the Great Barrier Reef has been lost) has enabled us to determine that they represent a new species. The Bishop Museum received a specimen of Scorpaena from Tahiti, caught at the unusual depth of 400 m. Four species of Scorpaena are reported from South Pacific islands of Oceania west to Australia: S. thomsoni Giinther from Juan Fernandez Island, S. orgila Eschmeyer and Allen from Easter Island, S. cookii Giinther from the Kermadec Islands to New South Wales, and S. cardinalis Richardson and S. papillosa (Forster) from New Zealand and southeastern Australia. In addition, S. thomsoni Giinther was described from Juan Fernandez Island in the eastern Pacific. Comparison of the Tahitian specimen with these species, as well as others from the Indo-Pacific region, revealed that it represents an undescribed species. The objective of the present paper is the description of these two new scorpionfishes. 3 Research Associate, Department of Ichthyology, California Academy of Sciences. 384 a RANDALL AND GREENFIELD: TWO NEW SOUTH PACIFIC SCORPIONFISH 385 MATERIALS AND METHODS Type specimens of the new Scorpaenopsis have been deposited at the following institutions: Australian Museum, Sydney (AMS); Bernice P. Bishop Museum, Honolulu (BPBM); California Academy of Sciences, San Francisco (CAS); Museum National d’Histoire Naturelle, Paris (MNHN); National Science Museum, Tokyo (NSMT); and the U.S. National Museum of Natural History, Washington, D.C. (USNM). Lengths of specimens are given as standard length (SL), measured from the front of the upper lip to the base of the caudal fin (posterior end of the hypural plate); head length is measured from the same anterior point to the posterior end of the opercular membrane; body depth is the maximum depth from the base of the dorsal spines (as they emerge from the body), and body width the greatest width just posterior to the gill opening; orbit diameter is the greatest bony diameter, and interorbital width the least bony width; upper-jaw length is taken from the front of the upper lip to the posterior end of the maxilla; caudal-peduncle depth is the least depth, and caudal-pedun- cle length the horizontal distance between verticals at the rear base of the anal fin and the caudal- fin base; lengths of fin spines and rays are measured to their extreme bases; length of base of dor- sal fin is measured from the front of the base of the first dorsal spine to the rear base of the last dor- sal ray (not to the end of the membrane connecting ray to caudal peduncle); pectoral-fin length is the length of the longest ray. Morphometric data presented in Tables | and 2 are given as percent- ages of the standard length. Proportional measurements in the text are rounded to the nearest 0.05. Pectoral-ray counts include the slender uppermost unbranched ray; counts were made of the rays on both sides. The scales in longitudinal series on the body were counted as oblique rows from the upper end of the gill opening to the base of the caudal fin. Counts of gill rakers were made on the first gill arch, the raker at the angle is contained in the lower-limb count. Scorpaenopsis eschmeyeri Randall and Greenfield, sp. nov. Figs. 1—2; Table 1 MATERIAL EXAMINED.— HOLortyPe: CAS 218804, female, 101.5 mm, Fiji, Vanua Levu, Rabi Island, northwest shore, 16°26.701'S, 179°56.261'W, 25 m from rocky shore, fringing reef, 1—2 m, rotenone, D.W. Greenfield, T.A. Greenfield, R.C. Langston, and J. Pilippoff, 20 May 2003. PARATYPES: CAS 214577, 115 mm SL, Australia, Queensland, Great Barrier Reef, Capricorn Group, One Tree Island, off reef crest on west side just above dropoff, rotenone, V.G. Springer, 7 December 1966; USNM 243332, 108.0 mm, Fiji, Malolo Island, south side of channel at northwest end of island barrier reef, sand channels and rock walls, 0-18 m, rotenone, V.G. Springer et al., 25 May 1982; USNM 259405, 84.0 mm, Fiji, Viwa Island, small islet on Viwa Reef (17°11’S, 176°54’E), rotenone, V.G. Springer et al., 27 May 1982; BPBM 33775, 75.5 mm, Coral Sea, Chesterfield Islands, lagoon, south end, patch reef, 2-4 m, rotenone, M. Kulbicki, J.E. Randall, P.J. Doherty, and C. Goiran, 29 August 1988; BPBM 34264, 2: 64.7-65.5 mm, New Caledonia, lagoon near southeast end of St. Vincent Pass, 22°2.1'S, 165°57.8’E, reef in 2-4 m, rotenone, M. Kulbicki, J.E. Randall et al., 21 March 1990; NSMT-P 67891, 101.0 mm SL, Fiji, Viti Levu, barrier reef off Suva, main channel between wreck and end of reef, 18°8.90'S, 178°23.91'E, spur and groove, 4.5-8 m, rotenone, D.W. Greenfield, K.R. Longnecker, and K.S. Cole, 31 May 1999; BPBM 39423, 63.0 mm SL, same locality as preceding, spur and groove, 12—18.5 m, rotenone, D.W. Greenfield, K.R. Longenecker, K.S. Cole, and R.C. Langston, 27 January 2002; CAS 219133, 2: 27.0-45.5 mm, Fiji, Viti Levu, barrier reef off Suva, fish patch, backside of dropoff towards reef, 18°9.59'S, 178°23.96’E, 7.5—-10.5 m, rotenone, D.W. Greenfield, K.R. Longenecker, K.S. Cole, and 386 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 19 R.C. Langston, 2 February 2002; USNM 375892, 94.0 mm, Fiji, nearly the same locality as pre- ceding, 18°9.545'S, 178°23.974'E, 8.5-15 m, same collectors, rotenone, 3 February 2002; AMS 1.42980-001, 79.0 mm, Fiji, Viti Levu, off Suva, Makuluva Island, west side, 18°11.227'S, 178°31.040'E, edge of cut in reef, very silty, 3-5 m, same collectors, rotenone, 4 February 2002; BPBM 39424, 2: 38.5—95.5 mm, Fiji, Viti Levu, off Suva, outside barrier reef about | mile west of sand bank cut, 18°11.161'S, 178°26.757'E, spur and groove, but mostly coral, 12 m, rotenone, D.W. Greenfield and K.S. Cole, 9 February 2002; CAS 219135, 107 mm, same locality as BPBM 39424, 14-15 m, rotenone, K.R. Longenecker and R.C. Langston, 9 February 2002; CAS 219134, 28 mm, Fiji, Viti Levu, off Suva, outside barrier reef, about 1 mile west of sand bank cut, 18°11.163'S, 178°26.28'E, sandy hole in reef surrounded by coral (mostly dead), 23-24 m, rotenone, D.W. Greenfield, K.R. Longenecker, K.S. Cole, and R.C. Langston, 10 February 2002; MNHN 2004-01171, 97.0 mm, Fiji, Northern Lau Group, Vanua Balavu Island, north side of island at Balavu Harbor, 17°11.212'S, 179°0.095'W, outside barrier reef, sand under an overhang, 12—15 m, rotenone, D.W. Greenfield, K.R. Longenecker, R.C. Langston, and Bio Koroi Mataitini, 8 January 2003. DraGnosis.— Dorsal rays XII,9; anal rays III,5; pectoral rays 17—18 (rarely 18); longitudinal scale series 44-48 (modally 46); body depth 2.8—3.05 in SL; head length 2.2—2.3 in SL; snout length 3.0—3.15 in head length; orbit diameter 4.65-4.95 in head length; nearly one-half of orbit extending above dorsal profile of head; interorbital width 6.75—7.2 in head length; pair of interor- bital ridges flaring posteriorly, then curving medially to join slight incurved ridge at front of occip- ital pit; median interorbital ridge extending half way back in interorbital space; occipital pit shal- low and not flat; no coronal spines; no pretympanic spines; suborbital pit not well developed; sub- orbital ridge with four spines, the first on lacrimal; two ventral spines on lacrimal, one directed mainly anteriorly and the other curving posteriorly; first dorsal spine |.9—2.4 in second spine; third dorsal spine longest, 2.2—2.7 in head length; eleventh dorsal spine 1.55—2.0 in twelfth spine; sec- ond anal spine 1.65—2.0 in head length; no supraoccipital tentacle; color variable, the darkest blotches usually above tip of upper opercular spine, two on lateral line, two obliquely above and forward of these, two in soft portion of dorsal fin, and one in anal fin. Largest type specimen, 115 mm. DESCRIPTION.— Dorsal rays XII,9; anal rays III,5; all dorsal and anal soft rays branched, the last to base; pectoral rays 17 (17, two of 28 counts are 18), the second to fourth branched (second to fourth, fifth, or sixth branched in paratypes); pelvic rays I,5, all soft rays branched; principal cau- dal rays 13, the middle 11 branched; upper and lower procurrent caudal rays 7, the most posterior segmented; lateral-line scales 22 (22—23), plus two on base of caudal fin, the first at upper end of gill opening spinous, the next few progressively less so; longitudinal scale series 46 (44-48; one of 14 with 44, three with 45, six with 46, three with 47, and one with 48); scales above lateral line to middle of dorsal fin 8; scales below lateral line to origin of anal fin about 16; median predorsal scales about 9; circumpeduncular scales about 25; gill rakers 5 + 9; pseudobranchial filaments of holotype 31 (26 for 79-mm paratype); vertebrae 24. Body depth 2.8 (2.55—3.05) in SL; body width 1.4 (1.25—1.6) in body depth; head length 2.2 2.2—2.3) in SL; snout length 3.15 (3.0—3.05) in head length; orbit diameter 4.8 (4.65—5.2) in head length; nearly one-half of orbit extending above dorsal profile of head; interorbital space deeply concave, the least width 6.75 (6.8—7.35) in head length; caudal-peduncle depth 3.65 (3.7—3.95) in head length; caudal-peduncle length 3.1(2.95—3.6) in head length. Mouth moderately large, the maxilla extending to a vertical at posterior edge of pupil, the upper-jaw length 2.0 (1.9—1.95) in head length; lower jaw projecting; mouth oblique, forming an angle of about 25° to horizontal axis of body; upper jaw with a dense band of small, slender, con- RANDALL AND GREENFIELD: TWO NEW SOUTH PACIFIC SCORPIONFISH 387 FIGURE 2. Paratype of Scorpaenopsis eschmeyeri, BPBM 33775, male, 75.5 mm, Chesterfield Islands, Coral Sea (J.E. Randall). ical, incurved, and inwardly depressible teeth in about six to seven rows, narrowing to two or three rows posteriorly, the symphyseal gap at front of jaw about one-third pupil diameter in width; lower jaw with a similar band of teeth in about five or six rows anteriorly, narrowing to one or two pos- teriorly; three irregular rows of small, slender, incurved, conical teeth forming a V on vomer; no teeth on palatines. Tongue short, very thick and broadly rounded, with a slender rod-like median anterior projection. Gill rakers short, with small spinules, the longest at angle, equal in length to longest gill filaments. Head spination typical of the genus; no coronal spines; pair of interorbital ridges flaring pos- teriorly, then curving medially to join slight incurved ridge at front of occipital pit; median interor- bital ridge extending half way back in interorbital space; occipital pit shallow and not flat; tympan- 388 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. XX ic spines about equal in length to postocular spines; parietal, and nuchal spines strongly recurved, sharing a broad base; three very small sphenotic spines, followed by the pterotic, lower posttem- poral and supracleithral spines; upper posttemporal spine very small; no developed postocular spines; nasal spines small; suborbital pit between anterior part of orbit and large bony knob on lacrimal not deep; suborbital ridge with four spines, the first on lacrimal more erect; lacrimal with a dorsoanterior ridge, an anterior ridge (without a sharp anterior tip), and two ventral spines, one strongly directed anteriorly and the other curving posteriorly; two opercular spines, the upper flat, without a ridge, the lower preceded by a slightly curved ridge with a short ridge above and adja- cent to its base; four preopercular spines, the uppermost with an anterior supplemental spine, the lowermost very short; clavicular spine strong, projecting dorsoposteriorly, without a median ridge. No supraocular tentacle; a small tentacle associated with anterior ventral spine of lacrimal and two large flat fimbriate tentacles following posterior spine; a series of four small cutaneous flaps on lower jaw just below lower lip; a few small flaps on lower edge of preopercle and three slender ones posteriorly on upper part of maxilla; body with very small scattered cutaneous flaps; lateral line with flaps on fourth, sixth, ninth (largest), thirteenth, sixteenth (small), and twentieth scales. Anterior nostril below base of nasal spine, with a posterior fimbriate flap about twice as long as spine; posterior nostril at edge of orbit below preocular spine, with a low fleshy rim. Scales on body ctenoid, becoming cycloid ventrally on abdomen, chest, and prepectoral area, extending onto base of pectoral fins in a broad V-shape; cycloid scales basally on opercle, includ- ing a few embedded scales above base of ridge of lower spine; scales on opercular flap between opercular spines; no scales on fins except three vertical rows on base of caudal fin. Origin of dorsal fin above supracleithral spine, the predorsal length 2.45 (2.3—2.5) in SL; first dorsal spine 2.2 (1.92.4) in second spine, 7.1 (6.2—7.35) in head length; third dorsal spine longest, 2.7 (2.2—2.65) in head length; eleventh dorsal spine 1.75 (1.55—2.0) in twelfth spine; last dorsal soft ray joined by membrane to dorsal edge of caudal peduncle above base of penultimate lateral-line scale; origin of anal fin below base of first dorsal soft ray, the preanal length 1.4 (1.35—1.4) in SL; first anal spine 4.1 (3.5-4.2) in head length; second anal spine much longer than third, 2.0 (1.65—1.9) in head length; first anal soft ray longest, 1.9 (1.5—2.0) in head length; caudal fin slight- ly rounded, 1.6 (1.55—1.65) in head length; sixth and seventh pectoral rays longest, 1.45 (1.35—1.6) in head length; origin of pelvic fins below base of third dorsal spine (hence distinctly posterior to base of pectoral fins), the prepelvic length 2.5 (2.5—2.6) in SL; pelvic spine 2.7 (2.4—2.8) in head length; pelvic fins approaching or extending slightly posterior to anus, the second ray longest, 1.6 (1.5—1.7) in head length. Color of holotype in alcohol: body brownish gray, becoming white on abdomen and chest, with six irregular blackish blotches along back, progressively smaller posteriorly, a black spot above clavicular spine, and less distinct blackish blotches along or near lateral line; ventral part or body from above anus to base of caudal with irregular near-white blotches, the largest lightbulb- shape on lower half of caudal peduncle; head brownish gray, pale ventrally, with light gray-brown blotches; a black spot on opercular membrane above upper opercular spine, and another on lower part of opercle below level of upper preopercular spine; a large dark blotch above and posterior to upper corner of maxilla; maxilla blotched with white; lips crossed with dark gray-brown spots; spinous portion of dorsal fin mottled gray and whitish, the soft portion gray along about basal third, whitish on outer two-thirds, with two slightly oblique blackish cross bands, the more anterior the most distinct; anal fin white with four irregular oblique dark bands, the third darkest; caudal fin pale with four or five brownish gray spots forming irregular bars; pectoral fins pale gray-brown, whitish distally, with irregular dark brownish gray cross bands; pelvic fins whitish with a few faint gray blotches on outer part of first four soft rays. RANDALL AND GREENFIELD: TWO NEW SOUTH PACIFIC SCORPIONFISH 389 TABLE |. Proportional measurements of type specimens of Scorpaenopsis eschmeyeri expressed as percentages of the standard length. CAS CAS BPBM BPBM AMSI USNM BPBM NSMT-P CAS 218804 219133 34264 33775 42980 375892 39198 67891 214577 Sex female immature male male female female female female female Standard Length (mm) 101.5 45.5 65.5 TSeS 79 94 95.5 101 115 Body depth Som 32.8 BSES B52) 337) 337, 34.5 34.2 39.4 Body width 259 22 22.6 24.1 25.2 26.7 24.6 23.8 24.7 Head length 45.3 43.8 4h 44.6 43.9 44.7 44.8 45.5 45.5 Snout length 14.3 14.5 14.6 S22 14.3 14.6 14.6 14.8 15 Orbit diameter 3 9.2 9.2 9:3, 8.9 9.6 9.6 NS) 8.8 Interorbital width 6.7 6.6 6.3 6.1 6.2 6.6 6.5 6.6 6.2 Caudal-peduncle depth 12.4 HES ilites 11.9 Die 11.7 Al eS 12.3 Caudal-peduncle length 14.7 14.5 13.5 13.9 14.9 14.3 12.6 12.9 12.7 Upper-jaw length 22.9 22D) 223, 2B 22.8 22.9 235 D3), 23.4 Predorsal length 4] 40 39.9 40.4 40.2 40.5 41.5 41.7 40.5 Preanal length TZ 122 71 70.8 71.3 72.6 72.8 73 Vall Prepelvic length 40.1 39.2 38.3 39.7 38.8 39 39.3 Sey 41.1 Dorsal-fin base 57.2 56.4 Sp 57.6 58.6 i Spzi D2 57.9 First dorsal spine 6.4 ell i 6.8 6.8 6.9 U2 6.2 7.1 Second dorsal spine 133 IN 7/aI 14.9 14.8 14 1319 13.8 14 14.9 Longest dorsal spine 16.7 19.8 18.7 19.6 18.3 18.1 18.3 17a 18.4 Eleventh dorsal spine 7.9 Gell esl 8 7.6 8.5 8.4 9 8.2 Twelfth dorsal spine 13.8 14.5 14.3 15.2 Syl 129. 14.6 13.9 14.1 Longest dorsal soft ray 18.9 19.8 19.9 20.2 18 18.4 19.9 18.5 19.6 Anal-fin base 14.5 15.9 14.8 16.4 16.8 15.6 14.7 15 14.5 First anal spine 10.8 11.4 10.5 Wal 12.6 10.7 12 10.9 12.4 Second anal spine 22.6 24.8 26.7 PUM 259) 24.2 25D. 23.8 25.3 Third anal spine 19.3 20.4 De]. Me? 21.4 20.1 Die2 19.4 19.4 Longest anal soft ray 24.1 24.5 26.8 24.9 24.8 Died 24 23 23%3 Caudal-fin length 27.8 28.7 29.3 29.2 27.8 27.7 27.6 27.6 28.4 Pectoral-fin length 31 32.6 30.7 31.8 31.4 30.9 30.7 28.7 29.6 Pelvic-spine length 16.7 73) ES) 18.7 16.7 15.6 16.9 16.4 17 Pelvic-fin length 27.8 28.3 28.1 Di 29 28.2 27.2 26.5 27 Color of holotype when fresh: body above pectoral fin to below ninth dorsal spine mottled red- dish brown, becoming mottled reddish gray posteriorly to caudal peduncle where irregularly marked with pale red and dark reddish brown; dark brown blotches more conspicuous than on pre- served specimen; cutaneous flaps dorsally on body and on lateral line pale bluish gray, those on lower side pale tan with white edges; abdomen pale salmon; head mottled brownish red, pale orange-red ventrally and on chest and lower part of prepectoral area; head, body, median fins, and upper part of pectoral fins with numerous white dots, many grouping to form irregular markings; 390 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 19 areas of dorsal and anal fins indicated as pale in preservative were light red when fresh; caudal-fin membranes transparent, the rays banded with pale red and dark reddish brown; pectoral fins simi- lar in color to caudal fin, the lower ten rays more banded in light red; pelvic fins with a large cir- cular light red area covering most of outer three-fifths of fin, the margin white, the base pale salmon. ETyMoOLoGy.— Named in honor of William N. Eschmeyer in recognition of his research on the Scorpaenidae. REMARKS.— In their revision of Scorpaenopsis, Randall and Eschmeyer (2001:59) listed seven lots of the genus from Fiji, New Caledonia, Chesterfield Islands in the Coral Sea, and One Tree Island of the Capricorn Group, southern Great Barrier Reef as nontype specimens of S. possi. These specimens lack the pretympanic spine that is unique to S. possi. No typical specimens of S. possi were found at any of these localities. Randall and Eschmeyer suggested that these specimens represent a southwestern population of S. possi in the Pacific. Our additional 12 specimens from Fiji, 27-108 mm SL, provided for more comparison with S. possi and the conclusion that this pop- ulation represents a new species. Our first suspicion that the specimens from the southwest Pacific represent a new species was noting their small size compared to S. possi, with mature females as small as 79 mm SL. Twenty specimens of S. eschmeyeri are 142 mm SL or less (these include the lot of three uncataloged CAS nontype specimens of S. possi listed by Randall and Eschmeyer, which have been lost). Nineteen of these are 117 mm SL or less. Seventy type specimens of S. possi are larger than 117 mm SL, and 18 range from 142-194 mm SL, the largest from Rarotonga, Cook Islands. The 142-mm specimen of S. eschmeyeri is one of two specimens from the Capricorn Group of the southern Great Barrier Reef listed as nontypes of S. possi by Randall and Eschmeyer (then as CAS uncat.). The smaller specimen, 115 mm SL, is now CAS 214577. The larger specimen is ten- tatively identified as S. eschmeyeri, but it is not listed as a paratype because of its unusual large size compared to the other type specimens and because it was preserved with the mouth rigidly and fully open and the body curved, hence making accurate measurements difficult. It is cataloged sep- arately as CAS 219194. Although we found no meristic differences to separate S. eschmeyeri from S. possi, four meas- urements serve to differentiate the two species when comparing specimens of the same size range: head length, snout length, upper-jaw length, and predorsal length. The head length of 10 Bishop Museum paratypes of S. possi from 59-121 mm SL is 42.5—43.4% SL, compared to 43.9-45.5% for 17 S. eschmeyeri. The snout length of S. possi is 13.4-14.2% SL (except BPBM 29332, 86 mm SL, from the Ryukyu Islands with 14.9%), compared to 14.3—15.9% for S. eschmeyeri. The upper-jaw length of S. possi is 20.6—22.5% SL, compared to 22.2—24.5% SL for S. eschmeyeri. The predorsal length of S. possi is 37.0-39.8% SL, compared to 39.9-42.1% SL for S. eschmeyeri. The best character to separate the two species is the presence of pretympanic spines in S. possi (larger than about 60 mm SL) and their absence in S. eschmeyeri. The two species are not known to coexist. Scorpaenopsis possi ranges from the Red Sea and east coast of Africa to the islands of French Polynesia except Rapa. The distribution in the western Pacific is from Taiwan to the Solomon Islands. Scorpaenopsis eschmeyeri is currently represented by specimens from Fiji, New Caledonia, the Chesterfield Islands in the Coral Sea, and the south- ern Great Barrier Reef. RANDALL AND GREENFIELD: TWO NEW SOUTH PACIFIC SCORPIONFISH 39] Scorpaena lacrimata Randall and Greenfield, n. sp. Fig. 3; Table 2 MATERIAL EXAMINED.— Holotype: BPBM 31706, female, 198.0 mm, Society Islands, Tahiti, 400 m, hook and line, Michel Kung (via Louise Wrobel), 24 October 1990. DiaGcnosis.— Dorsal rays XII,9; anal rays III,5; pectoral rays 17; longitudinal scale series 61; body depth 2.5 in SL; head length 2.15 in SL; snout length 3.2 in head length; orbit diameter 4.6 in head length; about one-third of orbit extending above dorsal profile of head; interorbital width 7.35 in head length; pair of interorbital ridges curving at front of occipital pit to join tympanic spines; median interorbital ridge extending half way back in interorbital space; occipital pit quad- rangular and moderately deep; no coronal spines; no suborbital pit; suborbital ridge with four spines, the first on lacrimal; first dorsal spine 1.8 in second spine; third and fourth dorsal spines longest, equal to second anal spine, 2.35 in head length; eleventh dorsal spine 1.65 in twelfth spine; supraoccipital tentacle present; small tentacles associated with ventral spines of lacrimal; no tenta- cles or cutaneous flaps on chin or on body; a row of small teeth on palatines; pale yellowish in alco- hol, with scattered small dark brown spots on head, body, and dorsal fin; a short, narrow, dark brown bar extending ventrally from middle of edge of orbit. DESCRIPTION.— Dorsal rays XII,9; anal rays III,5; all dorsal and anal soft rays branched, the last to base; pectoral rays 17, the second to eighth branched; pelvic rays I,5, all soft rays branched; principal caudal rays 14, the median 12 branched, the lower simple rays thickened; upper and lower procurrent caudal rays 7, the posterior two segmented lateral-line scales 22, the first two spinous; longitudinal scale series 61; scales above lateral line to middle of dorsal fin 10; scales below later- al line to origin of anal fin 19; median predorsal scales about 9; circumpeduncular scales about 25; gill rakers 4 + 10; pseudobranchial filaments 52; vertebrae 24. Body deep for the genus, the depth 2.5 in SL; body width 1.6 in body depth; head length 2.15 in SL; snout length 3.2 in head length; orbit diameter 4.6 in head length; about one -third of orbit extending above dorsal profile of head; interorbital space deeply concave, the least width 7.35 in head length; caudal-peduncle depth 3.9 in head length; caudal-peduncle length 3.1 in head length. Mouth terminal and large, the maxilla extending to below posterior edge of orbit, the upper-jaw length 2.0 in head length; mouth oblique, forming an angle of about 25° to horizontal axis of body; upper jaw with a dense band of small, slender, conical, incurved, and inwardly depressible teeth in about eight rows, narrowing to two or three rows posteriorly, the symphyseal gap at front of jaw one-half pupil diameter in width; lower jaw with a similar band of teeth in about six or seven rows anteriorly, narrowing to two rows posteriorly; a single row of small, slender, incurved, conical teeth forming a “V’ on vomer; similar teeth in a single row on palatines. Tongue thick and triangular with a fleshy rounded tip, the upper surface coarsely plicate. Gill rakers short, with spinules, the longest at angle nearly as long as longest gill filaments. Head with the usual complement of head spines, most prominent and retrorse; no coronal spines; pair of interorbital ridges flaring posteriorly, each curving laterally at anterior edge of occipical pit to join tympanic spine; median interorbital ridge extending half way back in interor- bital space; occipital pit quadrangular, flat, and moderately deep; tympanic, parietal, and nuchal spines broad-based and of about equal size; two sphenotic spines, followed by the pterotic, lower posttemporal (upper posttemporal directly above), and supracleithral spines; two very small post- ocular spines (only as small bumps on right side); no suborbital pit; suborbital ridge with four spines, the first on lacrimal more erect; lacrimal with four other spines, the first mainly a forward- directed ridge with a very short free tip; second and third spines directed dorsoventrally (the third clearly shortest), the last spine curving ventroposteriorly; two opercular spines, slender and not 392 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 19 divided, each preceded by a prominent ridge, the upper TaBLe 2. Proportional measurements of holo- curved, and the lower slightly curved; five preopercular type of Scorpaena lacrimata as percentages spines, the uppermost with a strong anterior supple- of the standard length mental spine, the lower two very short; clavicular spine BPBM strong, projecting dorsoposteriorly, with a slight medi- 31706 an ridge. A fleshy supraorbital tentacle about three-fourths ae Go) ee orbit diameter in length between supraocular and post- Body width 153 ocular spines; a small slender tentacle ventrally at front Head length 46.4 of lacrimal, another at base of second ventral spine of Snout length 14.4 lacrimal, and one with last lacrimal spine, broad and as Orbit diameter Oi. long as spine; a few small tentacles on lower edge of _Interorbital width 6.3 preopercle, the largest with the fifth preopercular spine; Caudal-peduncle depth 11.6 no flaps or tentacles detected on chin or elsewhere on Caudal-peduncle length iS head and body. Upper-jaw length DPS) , ; ; ; Predorsal length Sia Anterior nostril below base of nasal spine, with a Prosmniienets 738 posterior fimbriate flap longer than spine; posterior Prepelvic length 43.5 nostril at edge of orbit below preocular spine, with a Dorsal-fin base (2D slight membranous rim, the posterior edge with a few First dorsal spine 9.1 cirri. Second dorsal spine 16.3 Scales on body ctenoid, becoming cycloid ventral- Longest dorsal spine 19.7 ly on abdomen and on chest; prepectoral scales embed- Eleventh dorsal spine 9.6 ded: cycloid scales dorsally on opercle above upper __Welfth dorsal spine UL see j 2 aN Longest dorsal ray 16.4 spine; no scales on opercular flap above upper spine or ee eas 161 below lower spine, but many on flap between spines; First anal spine 9.9 no scales on fins except three vertical rows on base of Second anal spine 19.7 caudal fin. Third anal spine 18.2 Origin of dorsal fin above upper posttemporal Longest anal ray 28.7 spine, the predorsal length 2.7 in SL; first dorsal spine Caudal-fin length DD 1.8 in second spine, 5.1 in head length; third and fourth Pectoral-fin length 27.8 dorsal spines longest, 2.35 in head length; eleventh dor- Pelvic-spine length 19.0 Pelvic-fin length 26.5 sal spine 1.65 in twelfth spine; second and third dorsal soft rays longest, 2.8 in head length; last dorsal soft ray joined by membrane to dorsal edge of caudal peduncle above base of penultimate lateral-line scale; origin of anal fin below base of twelfth dorsal spine, the preanal length 1.35 in SL; first anal spine 4.7 in head length; second anal spine slightly longer than third, 2.35 in head length; first and sec- ond anal soft rays longest, 1.6 in head length; caudal fin rounded, 2.2 in head length; ninth pectoral ray longest, 1.65 in head length; origin of pelvic fins posterior to base of pectoral fins, the prepelvic length 2.3 in SL; pelvic spine 2.45 in head length; pelvic fins extending posterior to anus, the sec- ond ray longest, 1.8 in head length. Color in alcohol: pale yellowish with scattered dark brown spots, most on body no larger than scales, the most prominent on body above clavicular spine, above lower posttemporal and supra- cleithral spines, and as a curved transverse band across scaled part of nape; a short, narrow, dark brown bar extending ventrally from middle of orbit in line with a spot below second suborbital spine; a dark brown spot at convergence of opercular ridges; postorbital, sphenotic, and pterotic spines each in a dark brown blotch; occipital pit dusky with two small dark brown spots posterior- ly; fourth interspinous membrane with a dusky blotch containing a vertically elongate dark brown Ee RANDALL AND GREENFIELD: TWO NEW SOUTH PACIFIC SCORPIONFISH 393 FIGURE 3. Holotype of Scorpaena lacrimata, BPBM 31706, 198 mm, Tahiti, 400 m (J.E. Randall). spot; two dark blotches on fifth membrane and one on sixth, eighth, and ninth membranes; six small dark blotches on soft portion of dorsal fin, mainly on outer half; anal fin with two small dusky spots; caudal fin with dusky pigment on membranes forming a broad band across middle of fin; pectoral fins with two such transverse bands, one at base and one two-thirds out on fin; pelvic fin pale (one was removed). EtyMoLoGcy.— Named /acrimata from the Latin meaning to shed tears or weep, in reference to the short dark brown bar below the eye, reminiscent of a tear. REMARKS.— No information was given on the color in life, but the fish was probably mainly red, in view of its capture in 400 m. The fins were pinned in erect position, and one pelvic fin was removed, suggesting that a photograph may have been taken, but none was provided. Four other species of Scorpaena are known from islands of the South Pacific from Lord Howe Island to Easter Island. Scorpaena lacrimata differs from all in its greater body depth (2.5 in SL, compared to 2.7—3.1 for the mean depth of the four other species; data from Eschmeyer and Allen 1971 and Paulin 1982) and its count of 61 scales in longitudinal series. Paulin (1982) reported 48-50 scales for S. cardinalis, 64-67 for S. cookii, and 43-47 for S. papillosa. Eschmeyer and Allen (1971) counted 50-55 scales for S. orgila. The pectoral-ray count of 17 differentiates lacrimata further from S. orgila and S. papillosa, which have 15—16 pectoral rays. The chest of S. lacrimata is fully scaled, whereas that of S. cookii is naked and S. cardinalis has embedded scales. Elsewhere in the Pacific, the genus Scorpaena is represented by two species in the Hawaiian Islands (Eschmeyer and Randall 1975), three in Japan (Nakabo 2002), two of which range to the south, and one in the Philippines (S. hemilepidotus Fowler). All are readily separated from S. lacrimatus by pectoral-ray counts except the two Hawaiian species,which differ in having 45 scales in longitudinal series, and S. onaria Jordan and Snyder from Japan, which has 17—19 (usually 17) pectoral rays (Motomura and Iwatsuki 1997); it is distinct in its low scale count and sharply descending anterior part of the lateral line. Giinther (1880:24, pl. 12) described Scorpaena thomsoni from the Juan Fernandez Island in the eastern Pacific off Chile. It seems to be the closest species to S. Jacrimata from its color pat- 394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 19 tern and general morphology. It differs in its lower count of scales in longitudinal series (Giinther gave a count of 42, but his very good illustration shows 52), scaled prepectoral area (scales so embedded on S. /acrimata that scale outlines not readily seen), cutaneous flaps on body, a near- truncate caudal fin, and nine instead of ten lower pectoral rays unbranched. ACKNOWLEDGMENTS We thank Mme Louise Wrobel of Service de Ressources Marines of Papeete, Tahiti for pro- viding the specimen of Scorpaena lacrimata, Loreen R. O’ Hara of the Bishop Museum for X-rays, and the following persons for assistance in collecting fishes in Fiji: Kathleen S. Cole, Teresa A. Greenfield, Ross C. Langston, Kenneth R. Longenecker, Bio Koroi Mataitini, and Joanna Pilippoff. Captain Bruce Vasconcellos and the crew of Moku Mokua Hine are gratefully acknowledged for their logistic support, as are Johnson Seeto, G. Robin South, Randolph R. Thaman, and Robert W. Tuxton of the University of the South Pacific for facilitating our collections in Fiji. We also thank the Fijian Government and local village chiefs for permission to collect fishes. This research was supported by NSF grants INT97-29666 and DEB01-02745 and Sea Grant Project R/FM-6PD. LITERATURE CITED CARPENTER, K.E., AND V.H. NIEM, eds. 1999. The Living Marine Resources of the Western Central Pacific. Volume 3. Batoid Fishes, Chimaeras and Bony Fishes, Part 1 (Elopidae to Linophrynidae). Food and Agriculture Organization of the United Nations, Rome. vi + 1398-2068 pp. ESCHMEYER, W.N., AND G.R. ALLEN. 1971. Three new species of scorpionfishes (family Scorpaenidae) from Easter Island. Proceedings of the California Academy of Sciences 37(19):515—527. ESCHMEYER, W.N., AND J.E. RANDALL. 1975. The scorpaenid fishes of the Hawaiian Islands, including new species and new records (Pisces: Scorpaenidae). Proceedings of the California Academy of Sciences 40(11):265-333. GUNTHER, A. 1880. Report on the Shore Fishes procured during the voyage of H.M.S Challenger in the years 1873-1876. Pages 1—82 in Report on the Scientific Results of the Voyage of H.M.S. Challenger during the Years 1873—76. Zoology. Vol. 1, pt. 6. (Reprint edition, 1963, J. Cramer, Weinheim). MorTomura, H., AND Y. IWATSUKI. 1997. A preliminary report of scorpaenid, synancelid, tetrarogid and aploac- tinid fishes in Miyazaki waters, southern Japan. Bulletin of the Faculty of Agriculture, Miyazaki University 44(1—2):127-138. NAKABO. 2002. Fishes of Japan with Pictorial Keys to the Species, English edition, Vol. 1. Tokai University Press, Tokai, Japan. lxi + 866 pp. PAULIN, C.D. 1982. Scorpionfishes of New Zealand (Pisces: Scorpaenidae). New Zealand Journal of Zoology 9:437-450. RANDALL, J.E., AND W.N. ESCHMEYER. 2001. Revision of the Indo-Pacific scorpionfish genus Scorpaenopsis, with descriptions of eight new species. /ndo-Pacific Fishes (34):1—79. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 20, pp. 395-407, 4 figs., 2 tables September 30, 2004 The California Academy of Sciences Gulf of Guinea Expedition (2001) I. The Taxonomic Status of the Genus Nesionixalus Perret, 1976 (Anura: Hyperoliidae), Treefrogs of Sao Tomé and Principe, with Comments on the Genus Hyperolius Robert C. Drewes and Jeffery A. Wilkinson Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103, USA; Email: rdrewes@calacademy.org; jwilkinson@calacademy.org The endemic hyperoliid treefrogs of Sao Tomé and Principe currently recognized as Nesionixalus thomensis (Bocage, 1986) and N. molleri (Bedriaga, 1892) are re-exam- ined. The results of two molecular analyses indicate that these taxa are closely relat- ed to each other (monophyletic) but nest within the genus Hyperolius. A comparison of the morphological character states used by Perret (1976; 1988) to erect the genus Nesionixalus from within Hyperolius Rapp with a broad range of Hyperolius species reveals that most of these are not unique to the island endemics; all except the char- acters of size and digital tip shape are distributed among a number of other species within the latter genus. Nesionixalus Perret, 1976 is returned to the synonymy of Hyperolius Rapp and Nesionixalus thomensis and N. molleri to Hyperolius thomensis Bocage and H. molleri Bedriaga, respectively. The islands of SAo Tomé and Principe (which together comprise the tiny African Republic of Sao Tomé e Principe) lie along a magmatic geological feature known as the Guinea Line (or Cameroon Volcanic Line), which is a flaw (or hotline — Meyers et al. 1998) in the African tecton- ic plate between 1000-1500 km long (Simkin et al. 1994; Burke 2001) that has served as a chan- nel for magmas for millions of years. The Guinea Line extends across the ocean-continent transi- tion, and magmatic extrusions up through it have given rise to major oceanic and continental topo- graphic relief extending from southwest to northeast including the oceanic islands of Annobon (Pagalu; 4.9 my), Sdo Tomé (13+ my), Principe (31+ my), the recent continental island of Bioko (known in colonial times as Fernando Poo and now part of the Republic of Equatorial Guinea), and the mainland features of Mount Cameroon, the various ranges that comprise the Cameroon Highlands, and the Jos Plateau of Nigeria (Lee et al. 1994). The islands of Annobon, Principe and S40 Tomé are oceanic islands in the sense of Darlington (1957), Carlquist, (1965) and MacArthur and Wilson (1967) and long known for the high levels of endemism in their biota; some taxa are shared between the latter two. The three islands are separated from each other and from the West African coastlines of Equatorial Guinea and Gabon by ocean depths in excess of 3000 m. Principe is geologically the oldest and only about 130 km? in area, but it is well-watered and has significant relief provided by a central volcanic peak of just under 1000 m. It is about 220 km southwest of Bioko, the only continental and most northerly island of the chain, about 220 km from the African B95 396 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 20 mainland, and approximately 146 km northeast of Sao Tomé. The latter is geologically younger than Principe and considerably larger, at 836 km?: with the central Pico do Sao Tomé rising to above 2000 m. Sao Tomé is situated 280 km off the mainland and 180 km northeast of the southern-most of the oceanics, tiny (17 km?) Annobon or Pagalu, which is part of the Republic of Equatorial Guinea. (Juste et al. 1994). From late March to early June 2001, the California Academy of Sciences conducted a large, multi-disciplinary research expedition to both islands of the Republic of SAo Tomé e Principe (see Drewes 2002 for a popular account). Voucher specimens and tissues were collected of all of the amphibian and reptile species. This material allows us to address questions pertinent to both taxonomy and bio- geography such as the status of the genus Nesionixalus relative to other genera in the family Hyperoliidae as well as to add natural history information based on our field observa- tions. In this paper, we address the first issue. The large, flamboyant treefrog, hitherto known as Nesionixalus thomensis (Bocage, 1886), is endemic to the island of Sao Tomé (Fig. 1A); a second, recently elevated species, N. molleri (Bedriaga 1892), is a smaller tree- frog (Fig. 1B) present and widespread on both Sao Tomé and Principe. Nesionixalus thomen- sis was originally described as a member of the & : y Figure 1A—B. (A) Nesionixalus (~Hyperolius) thomensis genus fyperolius by Bocage (1886), the type 2; CAS 218934, Sao Tomé Id: Macambrara. Photo by. D. material, five specimens from Roca Saudade Lin. (B) Nesionixalus (=Hyperolius) molleri 3; CAS (ca. 800 m), Rio Quiza (probably Rio Quija in 2!9203-07, Principe Id: Baie das Agulhas. Photo by R.C. SW part of the island; exact locality unknown), P™*: and Ile Sao Tomé was housed in the Museu Bocage in Lisbon. There the series was re-examined in the mid 1970s by Perret, who placed the species in a new genus, Nesionixalus, based on large snout-vent length (males: 30-35 mm; females: 42-47 mm), shape of digital tips (oval, broader than long), dorsal skin of males covered by fine spinosities (smooth in females), absence of vocal sac in males (m. interhyoideus undifferentiated), absence of buccal vocal sac openings in males, and large size but small number of eggs (Perret 1976). In 1978, the Museu Bocage was destroyed by fire and all herpetological material was lost (E.G. Crespo, pers. commun.). In the course of his phylogenetic analysis of the Hyperoliidae which included 63 species of 15 genera (including Semnodactylus as Kassina wealei), Drewes (1984) located what appeared to be the only four museum specimens of N. thomensis still in existence, a male and female in London (BM 1951.1.1.91 and 98.3.30.39) and a pair in Vienna (NHMW 3695-6). The male specimens DREWES AND WILKINSON: TAXONOMIC STATUS OF GENUS NESIONIXALUS 397) exhibited vocal sacs consistent with Drewes’ character 21, state 1 (e.g. sac simple; fibers of m. interhyoideus parallel) and buccal vocal sac openings consistent with character 20, state 2 (anteri- or slits). The four specimens were x-rayed but no additional character states were observed to sup- port Nesionixalus as a separate genus within the context of Drewes’ largely myo-skeletal data set, and he returned Nesionixalus thomensis to the synonymy of Hyperolius thomensis Bocage. Four years later, Perret (1988) resurrected the genus Nesionixalus; he re-examined the Vienna material and while conceding the presence of the vocal sac and vocal sac openings as observed by Drewes (1984), he reiterated large snout-vent length, male integumental spinosities, transversely enlarged toe pads, and large size but small number of eggs (which he considered indicative of a specialized reproductive niche) as diagnostic. He added that the testes were large in size, the gular gland narrow and not conspicuous, and terminal phalanx obtuse or weakly notched (“status 2 de Drewes, non hyperolien”). He did not indicate which terminal phalanx nor how this latter observa- tion was made. In the same work, Perret (1988) described three additional genera to accommodate Cameroonian species formerly accepted as members of the genus Hyperolius Rapp 1842: Alexteroon to include Hyperolius obstetricans Ahl, 1931, Arlequinus for H. krebsi Mertens, 1938, and Chlorolius for H. koehleri Mertens, 1940. In 1992, Loumont reported on the results of field work undertaken in July 1988 and January 1990 on Sao Tomé and Principe; she revised the known amphibian fauna of both islands and pro- vided the first chromosome data and advertisement calls for the endemic anurans. She recognized three hyperoliid tree frogs: Leptopelis palmatus (Peters 1868) known only from Principe; Nesionixalus thomensis (Bocage, 1886), endemic to Sao Tomé, and Hyperolius molleri Bedriaga, 1892, a green, moderate-sized treefrog which is numerous, widespread and endemic to both islands. Loumont (1992:51) placed H. molleri in the genus Nesionixalus along with N. thomensis, citing Perret’s 1988 diagnostic characters. She also listed a number of character states in N. mol- leri that were more similar to typical Hyperolius than to N. thomensis. Loumont evidently collect- ed no tissue samples and did not deal with anuran larvae — the tadpoles of the anuran endemics remain undescribed. In the same year, Loumont joined Schatti in a publication on the herpetofau- na of Sao Tomé Island (Schatti and Loumont 1992). In his additions and corrections to Frost’s (1985) Amphibian Species of the World, Duellman (1993) recognized the genus Nesionixalus (N. thomensis and N. molleri) as well as the genera Alexteroon, Arlequinus and Chlorolius of Perret. Schigtz (1999) included these taxa in his book on African treefrogs as well. In his generic description Schietz (1999, p.309) reiterates Perret’s absence of vocal sac and vocal sac openings character states as diagnostic of the genus in spite of Drewes’ (1984) demonstration of their presence in Nesionixalus thomensis, and Perret’s (1988) subsequent acknowledgment that such is the case. Following his treatment of N. thomensis, Nesionixalus molleri is described as with a large gular sac (Schietz 1999:311). To date, none of the species of Nesionixalus, Alexteroon, Arlequinus or Chlorolius has been tested or confirmed in a published molecular and/or morphological phylogenetic analysis. MATERIALS AND METHODS Five species of Hyperolius and the two species of Nesionixalus (N. thomensis and N. molleri from both SAo Tomé and Principe) were used as the in-group in an initial analysis. Leptopelis palmatus was designated as the outgroup based on results from previous studies indicating that Leptopelis is at least basal to all other hyperoliid genera (Liem 1970; Drewes 1984; Channing 1929; Richards and Moore 1996; Emerson et al. 2000; Vences et al. 2003). An approximately 2 kb sequence that includes part of the 12S and 16S ribosomal genes and 398 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 20 the t-RNA for valine was amplified and sequenced with the primers and protocols in Wilkinson et al. (1996) and Wilkinson (1997). All sequences produced for this study were deposited in GenBank (see Appendix, Table | for a list of sequences, their sources, and accession numbers). The sequences were aligned following the procedure of Kjer (1993) using a putative second- ary structure for Rana catesbeiana as a standard (Nagae 1988). Because of insertion and deletion events in variable regions, those bases considered to have ambiguous alignments were removed from the analyses. We performed a maximum unweighted parsimony analysis using an exhaustive search. We also performed a bootstrap analysis (Felsenstein 1985) under maximum parsimony with 1000 repli- cates using the branch-and-bound search, and calculated Bremer’s decay indices (Bremer 1994) to evaluate the amount of support for the resulting clades. The decay index analysis was facilitated by the program Autodecay (Eriksson 1998). We performed a second analysis on approximately 500 b of only the 16S ribosomal gene in which, in addition to the species used in the first analysis, included DNA sequences retrieved from GenBank and Richards and Moore (1996) for two species each from the genera Afrixalus, Heterixalus, Kassina, the species Tachycnemis seychellensis, two more species and one more sub- species of the genus Hyperolius, and one more species from the genus Leptopelis (see Appendix, Table 1). Because of the additional taxa in this analysis, we used a heuristic search with 100 ran- dom stepwise additions followed by TBR branch swapping. We also performed a bootstrap analy- sis under maximum parsimony with 1000 replicates using the heuristic search with 10 random step- wise additions followed by TBR branch swapping and calculated Bremers decay indices on this second data set. Museum symbolic codes used herein follow Leviton et al. (1985). RESULTS AND DISCUSSION The first parsimony analysis resulted in one cladogram of 1167 steps with a consistency index (excluding uninformative characters) of 0.691, a retention index of 0.662, and a gl statistics (Huelsenbeck and Hillis 1993) of -0.648 for the distribution of all possible trees in the exhaustive search. The second analysis resulted in four most parsimo- nious cladograms of 865 steps long with a consistency index (excluding uninformative char- acters) of 0.494 and a retention index of 0.613. In both analyses Nesion- ixalus thomensis and N. molleri formed a well-supported clade nested within the genus FIGURE 2. The most parsimonious cladogram resulting from an exhaustive x search in a maximum parsimony analysis on approximately 2000 bases for i 7 five of the species of Hyperolius and two species of Nesionixalus in Table 1. meoventris as the sister taxon tO Numbers above branches represent bootstrap support for the respective clade, the Nesionixalus clade (Figs. 2 while numbers below branches are decay indices. Leptopelis palmatus Hyperolius viridiflavus ferniquei Hyperolius frontalis Hyperolius alticola Hyperolius aff. ocellatus Hyperolius cinnamomeoventris Nesionixalus thomensis Nesionixalus molleri Sao Tome Id. Nesionixalus molleri Principe |d. Hyperolius, with H. cinnamo- DREWES AND WILKINSON: TAXONOMIC STATUS OF GENUS NESIONIXALUS 399 and 3). In the second analysis, the genus Hyperolius formed a well-supported clade separate Leptopelis palmatus Leptopelis concolor from all other genera in the 100 Reseina cc neg ale Tsis analysis except Nesionixalus 20 Kassina maculata (Fig. 3). Also, iN molleri (CAS 65 HOTTA OIE 218848) from Sdo Tomé formed oats | a clade with NV. thomensis instead = ; NEM TUE UNS (ole (ese of NV. molleri (CAS 219125) from 90 Tachycnemis seychellensis Principe in two of the four most 4 ES een, Bae parsimonious cladograms, indi- : cating that the reduced number of 2 pixels Diachy ene iis characters in this analysis could 100 Nesionixalus molleri Principe td. N. ri fi , 99 pol pebarate calles aoe IN ey Nesionixalus molleri Sao Tome Id. thomensis. 64 4 6 Our molecular results indi- mE Nesionixalus thomensis cate two possible conclusions: Hyperolius cinnamomeoventris either (1) Nesionixalus is not a 8 fypatolivs burtienanis valid genus (Drewes 1984), and 61 both NV. thomensis and N. molleri 3 79 Hyperolius frontalis are members of the genus 3 Hyperolius alticola Hyperolius, or (2) the species of 5 & : (2) P o Hyperolius aff. ocellatus Nesionixalus are a clade deserv- F ing of generic recognition, and 65 Hyperolius viridiflavus ferniquei their inclusion in the genus 97 3 Hyperolius v. mariae HAyperolius as currently recog- 8 nized renders the latter para- Hyperolius argus FIGURE 3. Strict consensus of four most parsimonious cladograms result- ing from an heuristic search in a maximum parsimony analysis on approxi- mately 500 bases for all species used in the analysis (Table 1). Numbers above and below branches are as in Fig. 2. phyletic. In light of the results of the analyses above, we reexamined the characters used by Perret (1976, 1988) to define the genus Nesionixalus. We relied heavily on an unpublished M.A. disser- tation by L.G. Wilson (2000). This work is important to a discussion of the genus Hyperolius for at least three reasons: (1) it is the first attempt at a phylogenetic analysis of the genus (Wieczorek et al. [2001] focused on a subset — the H. viridiflavus complex), (2) it includes the largest sample size (31) of Hyperolius species studied to date (fifty-one morphological characters were employed, and outgroup species included four of the genus Afrixalus, three of the Malagasy Heterixalus, and Cryptothylax greshoffi); and (3) it is the first cladistic analysis that includes three out of four of Perret’s (1988) new genera, Alexteroon, Chlorolius and Nesionixalus. Specimens of Arlequinus have been unavailable for study thus far. Perret’s Diagnostic Characters Most of the characters used by Perret (1976, 1988) to define Nesionixalus thomensis are equiv- ocal with the exception of the transverse oval shape of the finger and toe tips: Size Snout-vent lengths in the lost type series were given as 30.0 and 35.0 mm for the two adult 400 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 20 males, and 42.0 and 47.0 mm for the females; the fifth specimen of the series was evidently a juve- nile at 18.0 mm (Perret, 1976). The 18 specimens in Loumont’s (1992) collection were of snout- vent lengths 27.0-35.0 mm in males and 36.0-41.0 mm in females. CAS material includes two females at 47.5 and 49.2 mm and 11 adult males ranging from 35.4-41.1 mm; one male, likely subadult, of 31.0 mm was collected as a singleton (see Appendix, Table 2). Snout-vent lengths of Nesionixalus thomensis females do indeed exceed published records for members of the genus Hyperolius, but the females of a number of African mainland species attain or exceed 40.0 mm snout-vent lengths, including Hyperolius torrentis, H. balfouri, H. sankuruensis, H. major and H. horstocki with H. bobirensis of Ghana attaining snout-vent lengths of 47.0 mm (Schigtz 1999). The overall large size of N. thomensis may be attributed to the phenomenon of island gigantism, espe- cially in light of the fact that there are many taxa endemic to S4o Tomé and/or Principe that exhib- it the same phenomenon. For instance, the distantly related hyperoliid frog (sensu Drewes 1984), Leptopelis palmatus, endemic to the tiny island of Principe is the largest member of that genus and the largest African treefrog, with females reaching 110 mm snout-vent length. Another frog endem- ic to Sao Tomé, the ranine Ptychadena newtoni 1s the largest member of its genus (Loumont 1999). The gekkonid lizard, Hemidactylus greeffii, endemic to both islands, is the largest African species in the genus (Loveridge 1947) and the giant sunbird of Sao Tomé, Dreptes thomensis, is substan- tially larger than all other species in the Afro-Asian family Nectariniidae (Cheke et al. 2001). Among the angiosperms endemic to Sao Tomé is the giant, 2-m tall, Begonia baccata, a paleoen- demic considered to be at least 3 million years old (Plana et al. 2004). Dorsal epidermal asperities Males of both Nesionixalus thomensis and N. molleri (sensu Loumont, 1992) possess fine asperities (“spinosités cornées” of Perret 1988) which are relatively evenly spaced on dorsally exposed surfaces of the body. These structures are visible with little or no magnification; some, but not all of these asperities are black in Nesionixalus and the dorsal skin of females of both species lack them. This character is not unique to the island species; it is also shared by a number of main- land species of Hyperolius. Males of H. spinigularis have dorsal asperities, although these are not always pigmented (Schietz 1975; 1999). All males in a large series of H. alticola (see Appendix, Table 2) from southwestern Uganda exhibit uniformly distributed spinosities (non-pigmented) which are absent in the females. Wilson (2000) coded the character state of dorsal spinules of N. thomensis and N. molleri as the same in Hyperolius bobirensis, H. endjami, H. laurenti, H. spinigu- laris, H. sylvaticus, H. tannerorum, and H. viridigulosus, as well as in Chlorolius koehleri and Alexteroon obstetricans. Gular gland Perret (1988) described the gular gland of NV. thomensis as narrow (“mince”), not obvious (“peu apparente”) and not very projecting [posteriorly?] (“ne faint pas saillie”). Wilson (2000) coded states of three characters of the male gular gland: size (four states based upon percent of gular area covered by the gland), thickness (four states based on gland thickness relative to that of surrounding, non-gland gular skin) and overlapping or free margins of the gland. The latter char- acter, modified from Liem (1970), is a measurement of how much of the lateral and posterior bor- ders of the gland cover distinct folds of distensible skin of the vocal pouch (Drewes 1984). The male gular glands of NV. thomensis and N. molleri occupy one-half or less of the total gular area, a character state shared by Hyperolius laurenti, H. montanus, H. occidentalis, H. sylvaticus and Alexteroon obstetricans; the thickness of the glands is the same as that of the glands of 24 of the DREWES AND WILKINSON: TAXONOMIC STATUS OF GENUS NESIONIXALUS 401] 31 species of Hyperolius examined by Wilson (2000), and the extent of the glands free margins is the same as in Hyperolius guttulatus, H. lamottei, H. laurenti, H. montanus, H. ocellatus and H. v. viridiflavus. Terminal Phalanx In the second description, Perret (1988) describes the terminal phalanx of N. thomensis as obtuse or weakly notched, as in Drewes (1984) state 2 [of character 14] and thus “non hyperolien.” Perret does not indicate which terminal phalanx nor the method of preparation, but this description is in error. All terminal phalanges in our cleared and stained material of both taxa are slender, and peniform with basal constriction, consistent with Drewes character 14, state 1, found in nine of the 12 species of Hyperolius he examined (Drewes, 1984, Fig. 10C; H. balfouri — not coded, H. argus and H. phantasticus had claw-shaped terminal phalanges —14(0)). This character is of doubtful intergeneric utility as Wilson (2000) found three states of this character among species of Hyperolius. The condition of the terminal phalanges of both NV. thomensis and N. molleri (MHNG specimens utilized by Loumont) was comparable to that in 22 of the 31 Hyperolius species cited in her study. Size of testes Perret (1976, 1988) was impressed with the large size of the testes of the two males in the type series of NV. thomensis (7 and 8 mm) and presented these data as a defining character in both the original and second description. Loumont (1992) reiterated this trait, noting that testes of H. mol- leri were also large (to 6.7 mm) and suggested this character together with the dorsal asperities jus- tified placement of H. molleri in Nesionixalus Perret. In both descriptions, Perret (1976, 1988) added large size of eggs and small clutch size as further distinguishing NV. thomensis. Wilson (2000) examined male testis size in Hyperolius and relatives and coded four states of this character based on testis length expressed as a percentage of total snout-vent length. Nesionixalus thomensis and N. molleri grouped with the eastern Hyperolius tuberilinguis at 19-21% of SVL and H. bobirensis and H. montanus had even larger testes at 25% and 34% SVL, respectively. All remaining Hyperolius fell within the 6-17% SVL range. Obviously the validity of this character depends upon the male frogs being at the peak of reproductive activity with testes fully enlarged; the fact that male Hyperolius are almost always located and collected while sexually active, together with Wilson’s large sample size of males of 31 species of Hyperolius, suggest that these observations are reasonably accurate and that the testis size in Nesionixalus does not serve to differentiate the genus from Hyperolius. Eggs and clutch size Comparative data regarding size of eggs and clutches among hyperoliid species are scarce. Members of the genus Hyperolius are generally described as having small eggs (0.8—1.5 mm) which in a few species are deposited in water or typically on vegetation above the water surface in clutches of 100 to 500 (Channing, 2001). Perret (1976) found 25 large (2.0-2.5 mm) presumably ovarian eggs in each of two of the three females in type series of N. thomensis,; in erecting the genus he suggested that the small number and large size of the eggs might indicate a specialized repro- ductive mode “wn cycle biologique particulier.” Loumont (1992) added new data for H. thomensis (50 to 60 ovarian [?] eggs of less than 0.5 mm diameter in two females) and H. mo/lleri (70+ mature eggs per clutch, 1.5 mm diameter). However, Fahr (1993) presented a figure of an H. molleri egg mass, which included at least 110 eggs. Egg and clutch size are of questionable taxonomic value 402 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 20 for hyperoliid treefrogs; there are almost no data available on the former except perhaps scattered references in the experimental literature and relative to the number of species recognized, data are very sparse for the latter. In his comprehensive book, Schiotz (1999) treated 85 species and species groups within the genus Hyperolius but was able to provide clutch size data for only 14 of them. Of these, five are known to deposit fewer than 50 eggs per clutch: H. frontalis, H. mitchelli, H. mosaicus, H. pusillus and H. semidiscus. Channing (2001) added clutch size data for four addition- al species including H. pickersgilli with 50 eggs. Finger and toe tips The size and nature of expanded digital finger and toe tips was examined by Liem (1970); Drewes (1984) did not employ the character in his hyperoliid work. With respect to this character, Channing (1989) re-coded and used Liem’s data on rhacophorids only. All of our specimens of Nesionixalus thomensis exhibit toe tips that are as previously described: enlarged, oval and wider in the horizontal plane than they are long — consistent with Liem’s character 33, state 4 which he found common in a number of rhacophorid genera but not in any hyperoliid species. As Perret cor- rectly observed (1976), the condition of N. thomensis digital expansions is very similar to that in Acanthixalus spinosus (see Appendix, Table 2), a West African hyperoliid not examined by Liem (1970). In addition, nearly identically-shaped digital tips are found in Kassina maculifer (K. park- eri of Scortecci 1932; Drewes 1984, 1985; Tandy and Drewes 1985; Appendix, Table 2 herein) of the arid Somali Horn. On the other hand, the expanded digital tips of all of our NV. molleri from both Sao Tomé and Principe Islands are disc-shaped, consistent with Liem’s (1970) character 33, state 2, which he found in all Hyperolius and the majority of other hyperoliid genera he examined. We did not make a broad survey of hyperoliid digital pads and concede that there are no species of Hyperolius known with digital tips similar to those of NV. thomensis. However, this sin- gle character state is not sufficient to support recognition of a distinct genus; moreover, this state is not shared by the purported congener, NV. moll/eri. Other than toe pad shape and body size, all of the morphological characteristics used by Perret (1976, 1988) to define Nesionixalus and Loumont (1992) to subsequently reassign H. molleri are shared by other species of Hyperolius or attributa- ble to island effects. This coupled with the results of our molecular analysis indicate that the genus Nesionixalus Perret is not a lineage separable from Hyperolius and that N. thomensis and N mol- leri should be relegated to the synonymies of Hyperolius thomensis Bocage, 1886 and Hyperolius molleri Bedriaga, 1892 respectively. In all molecular studies to date, Hyperolius has been shown to be monophyletic with respect to other hyperoliid genera; however, sample sizes have been extremely limited (Richards and Moore 1996, three species; Emerson et al. 2000, three species; Wieczorek 1999, 14 species and 16 subspecies of the 1. viridiflavus complex, but with Afrixalus as lone outgroup; Vences et al. 2003, six species). Similarly, all morphology-based phylogenetic studies support the monophyly of Ayperolius (Liem 1970, 14 Hyperolius species examined; Drewes 1984, 12 examined; Channing 1989, data from Drewes, op. cit. and Liem, op .cit) and widely-cited, regional works have assumed it (e.g., Schiotz 1967, 1975, 1999; Rédel 1996; Channing 2001). Hyperolius species can be extremely difficult to determine, especially after preservation; however, the genus has, until the recent descriptions of Nesionixalus, Chlorolius, Arlequinus and Alexteroon (Perret 1988; Amiet 2000) been rather easy to discern by initial determination of pupil shape (Drewes 1984:45—46). For instance, if the specimen in hand has horizontally-oval pupils (Fig. 4 B), is obviously arboreal with expanded digital tips (as opposed to Chrysobatrachus, which is terrestrial [Laurent, 1964]) and has a single gular gland in males (as opposed to Acanthixalus, in which gular glands are paired), it is a member of the genus Hyperolius. DREWES AND WILKINSON: TAXONOMIC STATUS OF GENUS NESIONIXALUS 403 At this time, we offer no taxonomic judgments on the validity of Perret’s (1988) Alexteroon, Chlorolius and Arlequinus. We lack sequence data for these taxa, and morphological work suggest- ing the first two be returned to the genus Hyperolius is unpublished (Wilson 2000). The morpho- logical definitions for Alexteroon and Chlorolius do not seem to include any synapomorphies that would serve to distinguish them unequivocally from Hyperolius, and in all cladograms presented by Wilson (2000), Alexteroon, Chlorolius and Nesionixalus nested within Hyperolius regardless of outgroup (4frixalus, Cryptothylax). Diagnostic morphological character states used in the past (Liem 1970; Drewes 1984) would suggest that Are/quinus is most likely to have a separate evolu- tionary history from Hyperolius. Arlequinus shares the quadratic pupil shape with Acanthixalus, Afrixalus, Heterixalus, Opisthothylax and Cryptothylax within the Hyperoliidae (Fig. 4). Monophyly in the genus Hyperolius has been either assumed by researchers or demonstrated (either morphologically or molecularly) on the basis of very small sample sizes relative to the num- ber of currently recognized species. We contend that to make taxonomic decisions on member taxa in the absence of robust knowledge of relationships within the largest hyperoliid genus runs the risk of adding to an already confused situation. A modern revision of the genus Hyperolius based on both molecular and morphological data is a daunting but long overdue task. We do recognize that H molleri and H. thomensis is a molecularly highly supported clade (as shown in Figs. 2 and 3). Moreover, both species share a chromosome number of 2n= 26 (Loumont 1992; Schatti and Loumont 1992; Fahr 1993), a condition rare among known Hyperolius species but not unique (Morescalchi 1968). However, to designate this clade a genus separate of Hyperolius would render Hyperolius paraphyletic, a result not supported by morphological analysis in this study. An intrigu- B C Callixalus* Alexteroon Acanthixalus Kassina Chlorolius Afrixalus Kassinula Chrysobatrachus* Arlequinus Leptopelis Hyperolius Cryptothylax Paracassina Heterixalus Phlyctimantis Opisthothylax Tachycnemis Ficure 4. Pupil shapes within the Hyperoliidae. Liem (1970) and Drewes (1984) both recognized three states of pupil- shape among hyperoliid frogs, but both combined two of these shapes, the vertical ellipse and rhomboidal/quadrangular, into a single character state; Channing (1989) followed suit, utilizing the same data. Schiotz (1999:43) provided a key that attempts to clarify pupil-shape in both living and preserved specimens: “la. Pupil horizontal, square or (after preservation) round”, and “2b.Pupil vertically elliptical or rhomboidal.” We feel that these treatments mask states which are probably not homologous. Here we present the range of hyperoliid pupil shapes based on photographs of living specimens. A. vertical (vertical ellipse). Kassina maculata (CAS 184054; Photo by J. Vindum); B. horizontal (horizontal oval, round). Hyperolius parkeri (CAS 154572-72; Photo by J. Vindum); C. rhomboidal (quadrangular, square). Cryptothylax greshoffii (MHNG specimen; Photo by J.-L. Perret). *To our knowledge, members of these genera have never been photographed in life. 404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 20 ing result of this study is the fact that this clade is more closely related to H. cinnamomeoventris than any other member of the genus studied; in fact, the support for this relationship is higher than within the molleri/thomensis clade itself (a Bremer’s decay index of 44 as opposed to 35; Fig. 2), and to our Hyperolius aff. ocellatus sample from Bioko, the northern-most member of the island chain, which might be expected due to geographic proximity. Hyperolius cinnamomeoventris, usu- ally readily identifiable, is among the most widespread member of the genus and one of the few species that inhabits both savannah and forest (Schiotz 1999). Such questions as the possible com- mon ancestry of H. cinnamomeoventris and the island endemics can only be determined by a much more broadly representative sample of Hyperolius and other hyperoliid genera. So far, GenBank has sequence data for only 15 identified species of Hyperolius — along with 26 subspecies of the H. viridiflavus complex and four unidentified entities. Not counting Nesionixalus, Arlequinus, Alexteroon and Chlorolius, this represents about 12% of the species currently recognized in Hyperolius that are distributed throughout sub-Saharan Africa. There are undoubtedly tissues in museums that have not yet been sequenced, or sequences not yet entered into GenBank. At CAS, we have unsequenced tissues and vouchers for six such species and with the increase in fieldwork during the past decade especially in West Africa, there are no doubt many more at other institu- tions. ACKNOWLEDGMENTS We are grateful to Christina Richards of Wayne State University for loan of samples, Robert Jachens of USGS, Palo Alto, California for reviewing portions of the introduction and Lindsay G Wilson for use of some of her data. The California Academy of Sciences Gulf of Guinea Expedition owes its success to the assistance of Ned Seligman, Director of STePUP, S40 Tomé, Angus Gascoigne of Voice of America, S40 Tomé, and Quintino Quade and our other field companions on both islands. We received necessary permits and much cooperation from Dr. Theresa D’Espiney, then-Director of ECOFAC and through the kind offices of the Hon. Maria das Neves Batista de Sousa, then-Minister of the Economy of the Republic of S40 Tomé and Principe. Additional thanks go to all of the members of the CAS expedition: Ricka Stoelting, Jens Vindum (Herpetology), Tomio Iwamoto (Ichthyology), Norman Penny, Charles Griswold, Joel Ledford (Entomology), Sarah Spaulding (Invertebrate Zoology), Douglas Long (Mammalogy), Dong Lin (Photography) and Fabio Penny (Education). The Expedition was funded by the Curators Research Fund and the G. Lindsay Field Research Fund, California Academy of Sciences and generous contributions to CAS Department of Herpetology by the late Richard A. Dumke, the late Malcolm and Jeanne Miller, David L. Jameson, and Emily Date. Early reviewers include R. Altig, and L.G. Wilson. This paper represents contribution no. 35 of the Center for Biodiversity Research and Infor- mation of the California Academy of Sciences. LITERATURE CITED. AMIET, J.-L. 2000. Les Alexteroon du Cameroun (Amphibia, Anura, Hyperoliidae). A/ytes 17:125—164. BocaGE, J.V.B. 1886. 2. Reptiles et batraciens nouveaux de l’ile de Thomé. 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Evolutionary relationships of a polymorphic African reed frog complex, Hyperolius spp. Ph.D Dissertation. University of the Western Cape, Bellville, South Africa. ix + 213pp. Wieczorek, A.M., R.C. DREWES, AND A. CHANNING. 2001. Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia 22:155—166. WILKINSON, J.A. 1997. Historical biogeography of rhacophorid frogs from Japan, the Ryukyu Archipelago and Taiwan inferred from a phylogenetic perspective. Ph.D. Dissertation, Texas Tech University, Lubbock Texas. 149 pp. WILKINSON, J.A., M. MATSUIE, AND T. TERACHI. 1996. Geographic variation in a Japanese tree frog (Rhacophorus arboreas) revealed by PCR aided restriction site analaysis of mtDNA. Journal of Herpetology 26:6—16 WILSON, L.G. 2000. Comparative morphology and evolutionary relationships of the genus Hyperolius (Anura: Hyperoliidae). M.A. Thesis. San Francisco State University, San Francisco. x + 106 pp. DREWES AND WILKINSON: TAXONOMIC STATUS OF GENUS NESIONIXALUS APPENDIX TABLE 1. Sources for DNA sequences used in this study. Species Source GenBank Accession Number Afrixalus brachycnemis Afrixalus fornasinii Heterixalus boettgeri Heterixalus tricolor Hyperolius argus Hyperolius alticola Hyperolius cinnamomeoventris Hyperolius frontalis Hyperolius aff. ocellatus Hyperolius puncticulatus Hyperolius viridiflavus ferniquei Hyperolius v. mariae Kassina maculata Kassina senegalensis Nesionixalus molleri Nesionixalus molleri Nesionixalus thomensis Leptopelis concolor Leptopelis palmatus Tachycnemis seychellensis Species Richards and Moore (1996) GenBank AFU22071 GenBank HBU24672 Richards and Moore (1996) Richards and Moore (1996) CAS 207322 AY 603984 CAS 202493 AY603985 CAS 201986 AY 603986 CAS 207321 AY 603988 Richards and Moore (1996) CAS 191296 AY603987 GenBank HVU22064 GenBank KMU22072 Richards and Moore (1996) CAS 219125 AY 603990 CAS 218848 AY 603989 CAS 218925 AY603991 GenBank LCU22079 CAS 219177 AY603992 GenBank TSU22080 TABLE 2. Preserved specimens examined. Museum Number Locality Acanthixalus spinosus Hyperolius alticola Nesionixalus (=Hyperolius) thomensis CAS 218925937; 219404 Sao Tomé e Principe: Sao Tomé Id: Kassina maculifer CAS 153799-800 CAS 180449-481 Cameroon: Sangmelima, Foulassi, Ngam Uganda: Rukungiri Dist: Bwindi Impenetrable NP: Munyaga Rv. Macambrara. CAS 140351; 140353-354; Kenya: Wajir Dist: Wajir 140356-361 Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. 407 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 21, pp. 408-413, 3 figs. September 30, 2004 A New Species of Dendrodoris (Mollusca: Nudibranchia: Dendrodorididae) from the Pacific Coast of North America David W. Behrens!3 and Angel Valdés23 'Department of Invertebrate Zoology, California Academy of Sciences 875 Howard Street, San Francisco, California 94103; *Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007 A new species of the genus Dendrodoris, D. azineae, is described based on a specimen collected from La Jolla Canyon, San Diego County, and photographs of animals from Sycamore Banks, Malibu and Carmel Bay, California. Dendrodoris azineae is characterized externally by its unique color, dissimilar to any described species worldwide, having purple gills and rhinophores. Internally, D. azineae differs from other eastern Pacific species in the size of the prostate and bursa copulatrix, and shape and size of the ampulla and the genital atrium. Thus far D. azineae is only known from southern to central California. The status of species assigned to the family Dendrodorididae along the eastern Pacific coast has changed several times in recent years. Behrens (1991) reported seven nominal species — three yellow species: Doriopsilla albopunctata (Cooper, 1863), Dendrodoris fulva (MacFarland, 1905), and Dendrodoris sp. 1; one red to black species: Dendrodoris krebsii (Moérch, 1863); and three grayish pale or white species: Dendrodoris nigromaculata, Dendrodoris sp. 2, and Dendrodoris sp. 3. Later, Valdés and Behrens (1998) added Doriopsilla spaldingi Valdés and Behrens, 1998, to this fauna. One additional eastern Pacific dendrodoridid species, Doriopsilla janaina Marcus and Marcus, 1967, is found to the south within the Panamic Province (Marcus and Marcus 1967). Gosliner, Schaefer, and Millen (1999) described Dendrodoris sp. 1 as a new species, Doriopsilla gemela Gosliner et al., 1999, and at the same time regarded D. fulva as a junior synonym of D. albopunctata. Behrens (2004) noted the name change of D. krebsii on the Pacific coast of the Americas to Dendrodoris fumata (Riippell and Leuckart, 1830), after Valdés et al. (1996) reported that the specimens from the Caribbean and Eastern Pacific were distinct. Dendrodoris sp. 3 remains undescribed. The present paper deals with recently collected material of Dendrodoris sp. 2. The first known collection of Dendrodoris sp. 2 was made in 1977 in a benthic trawl sample taken by Dr. Rim Fay and Robert Henderson of the then Pacific Biomarine Laboratories, of Venice, California. The collectors noted that it was distinct form all other porostomes on this coast. Periodic trawls in the same area over the next several years failed to produced any additional specimens. This specimen has subsequently been lost. On 12 July 2000, Clinton Bauder photographed the species in about 54 m, in Carmel Bay, California (Rudman 2000; Behrens 2004). Bauder and his team of deepwater technical divers have searched for another specimen ever since with no success, Additionally, there have been several sightings of this species from 45—54 m, respectively in La | 3 Research Associate, Department of Invertebrate Zoology, California Academy of Sciences. 408 BEHRENS AND VALDES: NEW SPECIES OF DENDRODORIS 409 Jolla Canyon, San Diego, California by George Spalding III (Behrens 2004), but specimens were not collected. The present paper describes this species based on the La Jolla specimen, which has been deposited in the type collections of the Natural History Museum of Los Angeles County. SPECIES DESCRIPTION Dendrodoris azineae Behrens and Valdés, sp. nov. (Figs. 1-3) White Porostome Behrens, 1980:100-101 Dendrodoris sp. 2 Behrens, 1991:72 Dendrodoris sp. 3 Rudman, 2000:http://www.seaslugforum.net/dendsp3.htm Dendrodoris sp. 2 Behrens, 2004:43 TYPE MATERIAL. — Holotype: LACM 3035, La Jolla Canyon, San pUEE Se California, March 14, 2004, 54 m depth, one specimen 35 mm preserved length, collected by George Spalding III. The specimen is dissected. EXTERNAL MORPHOLOGY. — Living ani- mals reach up to 75 mm in length. The general color of the living animals varies from opaque white to a creamy-tan (Fig. 1). The margin of the notum is slightly ruffled and is edged with a white band. The clavus of the rhinophore is deep purple, below which the shaft has dense patches of the same purple on the posterior side. This gill is also purple. The body is oval (Fig. 1), high, lacking spicules. The dorsum is smooth, with no tuber- cles. The posterior end of foot is translucent white. The rhinophores are perfoliate with 17 lamellae. The gill is composed of six bipinnate leaves which are somewhat separated from one another. The anus is central between the Ficure |. Living animals of Dendrodoris azineae sp. Beerehial leaves nov. (A) Holotype from La Jolla Canyon, California, 54 m depth (LACM 3035), photograph by Eric Hanauer. Ventrally there are no oral tentacles. The g) Specimen from Carmel Bay, California, 54 m depth, 30 mouth is reduced to a pore. mm long, not collected, photograph by Clinton Bauder. INTERNAL ANATOMY. — The digestive sys- tem has an oval buccal bulb (Figs. 2A—B), with two strong retractor muscles attached. From the posterior end of the buccal bulb emerge the pharynx and a narrow duct that splits into two and con- nects to the ptyaline glands. The paired ptyaline glands are large and situated ventral to the anteri- or portion of the digestive system. The pharynx is long and connects directly to the esophagus through a narrow duct. The esophagus is also long and has two esophageal glands situated next to the connection of the pharynx. The esophagus widens progressively towards the posterior end and becomes wide and muscular proximally. The intestine is long and straight and bears a small, oval pyloric gland near its proximal end. The hermaphroditic gland is separated from the digestive gland. The heart is large and connects to the blood gland through a conspicuous aorta. The blood 410 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 21 1mm pr FIGURE 2. Anatomy of Dendrodoris azineae sp. nov. (LACM 3035). A. Dorsal view of anatomy. B. Anterior portion of the digestive system. C. General view of the reproductive system. D. Dissected view of ampulla and prostate. E. © Dissected reproductive organs. Abbreviations: a = aorta, am = ampulla, bb = buccal bulb, be = bursa compulatrix, bg = | blood gland, cns = central nervous system, dg = digestive gland, dd = deferent duct, eg = esophageal gland, es = esopha- © gus, fg = female gland, ga = genital atrium, hg = hermaphrodite gland, ht = heart, i = intestine, m = retractor muscle, ph= 7 pharynx, pr = prostate, ptg = ptyaline gland, pyg = pyloric gland, rs = renal sac, sr = seminal receptacle, vg = vagina. | BEHRENS AND VALDES: NEW SPECIES OF DENDRODORIS 411 gland is situated posterior to the central nervous system and covers portions of the reproductive system. There is an oval renal sac situated on the right hand side of the heart and connected to the pericardium. The reproductive system is triaulic (Figs. 2C—E). The ampulla is long and convoluted, and branches into a short oviduct and the prostate. The oviduct enters the female gland in the center of the mass. The prostate is flattened and short, covering most of the bursa copulatrix. The prostate connects with a narrow duct that expands into the long, wide and muscular deferent duct. The vagi- nal duct is very long and wide. Both the vaginal duct and the deferent duct open into a large, mus- cular genital atrium. The proximal end of the vaginal duct joins the rounded bursa copulatrix. From the bursa copulatrix leads another duct that branches into a short uterine duct and the pyriform sem- inal receptacle. The penis is very long and smooth, covered with several rows of penial hooks (Figs. 3A-B). The distal spines seem to include some longer, blunt spines. EtyMoLocy. — The name azineae was chosen at the request of George Spalding HI of Solana Beach, California who collected the holotype. It is meant to honor his daughter Azine, who gives him the inspiration to continue his deep dives in search of new marine species in the La Jolla sub- marine canyon. DISTRIBUTION. — Thus far the species is known from La Jolla Submarine Canyon (present study), Sycamore Banks, offshore of Malibu, California (Behrens 1980, 1991) and Carmel Bay, California (photographs by Clinton Bauder). NATURAL HISTORY. — Little is known about the natural history of this species. It has not been observed with any sponge species. All specimens have been observed on sand or rock substrates. In his usual manner, Mr. Spaldling recorded the water temperature at the time of collection of the holotype as 10.6°C, at the collection depth of 60 m. We have no information on the species’ egg mass. DISCUSSION This species is placed in the genus Dendrodoris Ehrenberg, 1831, because of the presence of the following combination of characters: dorsum lacking a network of spicules, anus located cen- trally between the branchial leaves, digestive system with two ptyaline glands, radula and jaws absent, and esophagus with a pair of esophageal glands. Valdés and Gosliner (1999) found these characters to be diagnostic for Dendrodoris, which is a monophyletic group. A review of the literature shows that there are no other species with a similar external col- oration to Dendrodoris azineae. The Atlantic species of the genus were reviewed by Valdés et al. (1996), who found considerable external variability within the nine taxa examined. However, none of the Atlantic species have a uniform white background color with purple rhinophores and branchial leaves. Brodie et al. (1997) examined the systematics of the tropical Indo-Pacific species Dendrodoris nigra (Stimpson, 1855) and Dendrodoris fumata (Riippell and Leuckart, 1830). These authors also found great external variability in these two species, but none of the color forms are similar to D. azineae. . There are no other described species of the genus Dendrodoris along the coast of California. Dendrodoris fumata is the only other species of Dendrodoris described from the eastern Pacific. This tropical species is found within the Panamic Province, from México to the Galapagos Archi- pelago (Behrens 2004). Valdés et al. (1996) described the external morphology of eastern Pacific specimens of this species under the name Dendrodoris nigra (Stimpson, 1855). It differs from Dendrodoris azineae in several regards, including the presence of a uniformly colored (yellow, pink, gray, orange, red, or black) background (Brodie et al. 1997) and wider mantle margin with conspicuous striations. Brodie et al. (1997) described the anatomy of D. fumata, which has a 412 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 21 | FiGURE 3. Scanning electron micrographs of the penial spines of the holotype of Dendrodoris azineae sp. nov. (LACM 3035). A. Distal portion of the penis, B Proximal portion of the penis. BEHRENS AND VALDES: NEW SPECIES OF DENDRODORIS 413 prostate much smaller than that of D. azineae and the bursa copulatrix and seminal receptacle are about the same size, whereas in D. azineae the bursa copulatrix is much larger. Other differences include the shape and size of the ampulla, which is shorter and wider in D. fumata, and the genital atrium, which is much larger in D. azineae. Dendrodoris sp. 3, sensu Behrens (1991), is also a white species, but it differs from Dendrodoris azineae by having large brown spots on the dorsum and white rhinophores and branchial leaves. This species remains undescribed. ACKNOWLEDGMENTS The authors would like to recognize the generous efforts of all of those who have spent many hours searching over the past 20-plus years for specimens of this new species: Clinton Bauder and his fellow members of the Bay Area Underwater Explorers, George Spalding III, Peter Brueggeman, Jim Lance, Rim Fay and Bob Henderson. Thanks also to Eric Hanauer and Clinton Bauder for permitting us to use their photographs for preparation of the color images. This paper was supported by the National Science Foundation through the PEET grant DEB- 9978155 Phylogenetic systematics of dorid nudibranchs to Terrence M. Gosliner and the junior author. LITERATURE CITED BEHRENS, D.W. 1980. Pacific Coast Nudibranchs, A Guide to the Opisthobranchs of the Northeastern Pacific. Sea Challengers, Los Osos, California. 112pp. BEHRENS, D.W. 1991. Pacific Coast Nudibranchs, A guide to the opisthobranchs Alaska to Baja California. Second edition. Sea Challengers, Monterey, California. 107pp. BEHRENS, D.W. 2004. Pacific Coast nudibranchs — Supplement II: New species to the Pacific Coast and new information on the oldies. Proceedings of the California Academy of Sciences 55:11—54. Bropik, G.D., R.C. WILLAN, AND J.D. CoLLINs. 1997. Taxonomy and occurrence of Dendrodoris nigra and Dendrodoris fumata (Nudibranchia: Dendrodorididae) in the Indo-West Pacific Region. Journal of Molluscan Studies 63:407-423. GOSLINER, T.M., M.C. SCHAEFER, AND S.V. MILLEN. 1999. A new species of Doriopsilla (Nudibranchia: Dendrodorididae) from the Pacific coast of North America, including a comparison with Doriopsilla albopunctata (Cooper, 1863). The Veliger 42:201—210. Marcus, Ev., AND Er. MARcus. 1967. American Opisthobranch Mollusks. Part 2: Opisthobranchs from the Gulf of California. Studies in Tropical Oceanography 6:141—248. RUDMAN, W.B. 2000. Dendrodoris sp. 3. Available via http://www.seaslugforum.net/dendsp3.htm. VALDES, A., AND D.W. BEHRENS. 1998. A new species of Doriopsilla (Mollusca, Nudibranchia, Dendrondorid- idae) from the Pacific coast of North America. Proceedings of the California Academy of Sciences 50:307- 314. VALDES, A., AND T.M. GosLiNer. 1999. Phylogeny of the radula-less dorids (Mollusca, Nudibranchia), with the description of a new genus and a new family. Zoologica Scripta, 28:315-360. VaLpes, A., J. ORTEA, C. AVILA, AND M. BALLESTEROS. 1996. Review of the genus Dendrodoris Ehrenberg, 1831 (Gastropoda: Nudibranchia) in the Atlantic Ocean. Journal of Molluscan Studies 62:1-31. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22, pp. 414-430, 11 figs., 2 tables September 30, 2004 Systematics of Okenia from the Pacific Coast of North America (Nudibranchia: Goniodorididae) with Descriptions of Three New Species Terrence M. Gosliner! and Hans W. Bertsch? Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103; ! E-mail: tgosliner@calacademy.org; ? hansmarvida@sbcglobal.net The systematics of the eastern Pacific species of Okenia are reviewed. The anatomy of Okenia vancouverensis (O’Donoghue, 1921) and O. angelensis Lance, 1966 is described more completely. Three new species of Okenia are described from the trop- ical eastern Pacific. Okenia angelica sp. nov. is characterized by its purplish body color with numerous elongate lateral papillae and short, rounded dorsal tubercles. It is known from Baja California to Jalisco, México. Okenia cochimi sp. nov. has a bright uniformly yellow body with numerous lateral appendages and a single medi- al appendage anterior to the gill. It is known from Cedros Island, the central Gulf of California and Bahia Banderas, México. Okenia mexicorum is white with brick red markings. It has short lateral appendages, and a mid-dorsal crest that terminates in a single elongate papilla. This species is known from the central Gulf of California to Bahia Banderas, México. RESUMEN La posicién sistematica de las especies de Okenia del Pacifico Este es revisada. La anatomia de Okenia vancouverensis (O’Donoghue, 1921), y O. angelensis Lance, 1966, es descrita detalladamente. Tres especies nuevas de Okenia son descritas del Océano Pacifico Este Tropical. Okenia angelica esp. nov. es caracterizada por tener un cuerpo violeta con papilas laterales largas y numerosas y tubérculos dorsales cor- tos y redondeados. Esta especie es conocida de Baja California hasta Jalisco, México. Okenia cochimi esp. nov. tiene un cuerpo amarillo brillante y uniforme, apéndices laterales numerosos y un apéndice medio situado en frente de la branquia. Esta especie es conocida de la Isla Cedros, la region central del Golfo de California y Bahia Banderas, México. Okenia mexicorum esp. nov. es blanca con areas de color rojo teja. Esta especie tiene apéndices laterales cortos y una cresta media que termi- na en una papila larga. Esta especie es conocida de la regién central del Golfo de California hasta Bahia Banderas, México. Recently, the systematics of Okenia has been reviewed based on a preliminary phylogenetic analysis (Gosliner 2004). From this analysis came the conclusion that Hopkinsia, Hopkinsiella and Sakishimaia should be regarded as junior synonyms of Okenia. Four species of Okenia have been recorded from the Pacific coast of North America, Okenia vancouverensis (O’ Donoghue, 1921); O. rosacea (MacFarland, 1905); Okenia angelensis Lance, 1966 and Okenia plana Baba, 1960. In this work, aspects of the anatomy of two northeastern Pacific taxa were presented. The radular teeth 414 GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA 415 and reproductive anatomy of Okenia rosacea (MacFarland, 1905) were described. The anatomy of Indo-Pacific and non-indigenous specimens of Okenia plana Baba, 1960, from the Philippines and San Francisco Bay was described. The description of the anatomy of these species will not be repeated here. O’ Donghue’s (1921) description of Okenia vancouverensis included a description of the external morphology and the radula. In Lance’s (1966) original description of O. angelensis, he described the external anatomy, the radular teeth and the shape of dermal spicules. The description of these species is amplified here. Expeditions to Baja California over the last twenty years have produced collections of two additional species of Okenia. More recent collections from the Pacific coast of México by Orso Angulo Campillo and Alicia Hermosillo have yielded specimens of a third undescribed species. The descriptions of these species are presented here. All specimens examined here are housed in the Department of Invertebrate Zoology and Geology of the California Academy of Sciences (CASIZ). SPECIES DESCRIPTIONS Genus Okenia Menke, 1830 Type species: Okenia elegans (Leuckart, 1828), by monotypy. Okenia vancouverensis (O’ Donoghue, 1921) (Figs. 1—2) Idalia vancouverensis O’ Donoghue, 1921:177, pl., fig. 22. MATERIAL EXAMINED.— CASIZ (035222, one specimen, dissected, Rose Harbor, Queen Charlotte Islands, British Columbia, Canada, 4 July, 1913, Will F. Tompson. DISTRIBUTION.— Known only from the Queen Charlotte Islands (present study), Vancouver Island, British Columbia, Canada to Puget Sound, Washington (Behrens 1991). NATURAL HISTORY.— Found in about 15 meters of water on hard mud substrate. EXTERNAL MORPHOLOGY.— The body is broad and ovoid (30 mm in length, in the preserved specimen) with nine to ten pairs of elongate, acutely pointed papillae situated along the notal mar- gin and with an additional 24 similarly-shaped papillae on the remainder of the notum. The living animal is depicted by Behrens (1991) and Rudman (2003). The body is brownish with opaque white spots and an opaque white medial line along posterior portion of the foot. The rhinophores also contain patches of brown pigment. The rhinophores are elongate with 24 densely arranged lamellae. The gill consists of 12 bipinnate to tripinnate branches. Ventrally the foot is broad. The head (Fig. 1A) surrounding the mouth is broad and contains a pair of rounded oral lobes extending laterally. The reproductive opening is situated on the lateral side of the body, approximately one third of the length of the body posterior to the anterior margin. DIGESTIVE SYSTEM.— The buccal mass (Fig. 1B) is thick and muscular with a an elongate buc- cal pump directed posteriorly. Numerous minute oral glands are present at the opening of the buc- cal mass into the mouth. The radular sac is elongate and extends ventrally and anteriorly from the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass immediately ven- tral to the buccal pump. A large, flat, lobate salivary gland is present on either side of the buccal mass anterior to the junction of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains irregular polygonal plates. The cuticle expands as it enters the buccal pump. The radular formula is 32 X 1.1.0.1.1. in the single specimen examined Fig. 2A). The inner lateral teeth (Fig. 2B) are wide basally with an elongate acutely pointed cusp. The masticatory margin of the inner lateral bears 7—9 triangular denticles that increase in size in 416 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 the direction of the outer margin. The outer laterals are large and hook-shaped (Fig. 2B) with a curved, acutely-pointed cusp. CENTRAL NERVOUS SYS- TEM.— The ganglia of the cen- tral nervous system are highly concentrated and surround the esophagus, at the posterior end of the buccal mass. The cerebral and pleural ganglia are entirely fused. A sessile eye is present at the base of either cerebral gan- glion. The pedal ganglia are smaller than the cerebropleural ganglia and are separated by a short commissure. Paired buccal ganglia are situated ventral to the esophagus. REPRODUCTIVE SYSTEM.— (Fig. 1C) The preampullary duct is thin and elongate and expands into the relatively short, sausage- shaped ampulla. The ampulla FIGURE 1. Okenia vancouverensis (O’ Donoghue, 1921). A. Ventral view divides into a short oviduct that 0f head region, scale = 8.0 mm. B. Buccal mass, bp = buccal pump, e = esoph- agus, r = radular sac, sg = salivary gland, scale = 0.75 mm. C. Reproductive ; system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = a more elongate vas deferens. penis, pr = prostate, rs = receptaculum seminis, v = vagina, vd = vas deferens, The proximal portion of the vas _ scale = 1.0 mm. deferens enters a large spherical prostate that consists of numerous nodular bodies. From the prostate gland the ejaculatory duct emerges as a thin convoluted duct, which widens and forms several coils on top of the female gland mass. It then straightens and continues as an elongate duct that eventually widens slightly into the muscular penial bulb. The vagina is very wide at its exit adjacent to the penis, and consists of sev- eral folds. More distally it narrows and continues as a curved duct that again narrows and consists of a section that is folded and twisted together as convolutions. It then straightens again as it enters the base of the large, rounded bursa copulatrix. From the base of the bursa is a second duct that joins with the equally-sized, more pyriform receptaculum seminis. Near the base of the receptacu- lum, the uterine duct separates from the duct joining the receptaculum and bursa and enters the female gland mass. The female gland mass consists of three portions, the albumen, membrane and mucous glands. The mucous gland is the largest of the nidamental glands. DiscussioN.— Okenia vancouverensis is immediately identifiable by its broad body and brown color with opaque white markings. The present material agrees closely with that originally described by O’Donoghue (1921). O’Donoghue did not describe the anatomy of the reproductive system. Okenia vancouverensis appears to be unique among described Okenia species in having a discrete prostate gland rather than a prostate that merely represents an expansion of the deferent duct. The external anatomy of O. vancouverensis is similar to that described for O. aspersa (Alder & Hancock, 1845) and O. leachi (Alder & Hancock, 1854), both known from the northern Atlantic. enters the female gland mass and GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA 417 _ Ms FIGURE 2. Okenia vancouverensis (O’ Donoghue, 1921). Scanning electron micrographs of radular teeth. A. Entire radula. B. Inner and outer lateral teeth. These two species, like O. vancouverensis, are also found on muddy substrate. Okenia aspersa dif- fers from O. vancouverensis in having unevenly sized lateral appendages, with the anteriormost pair being the most elongate. Okenia leachi is similar to O. vancouverensis in having uniformly- sized lateral and notal papillae except the papillae are much longer in O. /eachi. The radular teeth of O. leachi differ from those of O. vancouverensis in having far more denticles on the inner later- al teeth and in having a proportionately smaller outer lateral tooth with a shorter cusp (Thompson and Brown 1984, fig. 7h). The radular teeth of O. aspersa are very similar in shape and size to those of O. vancouverensis (Thompson and Brown 1984, fig. 7g). The inner lateral teeth of O. aspersa bear up to 17 denticles while those of O. vancouverensis have only 7—9 denticles. 418 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 Okenia angelensis Lance, 1966 (Figs. 3-4) Okenia angelensis Lance, 1966:76, figs. 9-12. MATERIAL EXAMINED.— CASIZ 072351, 4 specimens, one dissected, Morro Bay, California, May-June 1981, David Behrens. CASIZ 170047, 5 specimens, 2 dissected, Mission Bay, San Diego, California, on floating docks, 25 June 1987, Terrence M. Gosliner. CASIZ 070471, 10 spec- imens, 2 dissected, Texaco Marina, Morro Bay, California, 26 October, 1975, Gary McDonald. DISTRIBUTION.— Known from San Francisco Bay to México and then disjunctly to southern Chile (Behrens, 1991, Mufioz et al., 1996). NATURAL HISTORY.— Found from shallow embayments on floating docks or in the shallow subtidal zone. EXTERNAL MORPHOLOGY.— The body is elongate (3—10 mm in length) with seven to eight pairs of elongate, rounded papillae situated along the notal margin and with an additional 8 papil- lae situated on the notum. The body is translucent yellowish white with numerous fine brownish spots scattered uniformly over the surface of the notum. The rhinophores are elongate with two to three well-spaced lamellae. The gill consists of five to seven unipinnate branches. The anterior margin of the head includes two elongate oral tentacles that extend anterolaterally. DIGESTIVE SYSTEM.— The buccal mass (Fig. 3A) is thin and muscular with a rounded buccal pump directed dorsally. Numerous small, elongate, pyriform oral glands are pres- ent at the opening of the buccal mass into the mouth. The radular sac is short and extends ventrally from the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass immediately ventral to the buccal pump. A rounded, lobate salivary gland is present on either side of the buccal mass anterior to the junction of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the open- ing of the mouth. No discrete polygonal plates were evident. The cuticle expands as it enters the buccal pump. The radular formula is 19 X 1.1.0.1.1 and 24 X 1.1.01.1 in two specimens examined. The inner lateral teeth (Fig. 4) are wide basally with an elongate acute bifid cusp. The masticatory margin of the inner lateral bears 15—19 triangular denticles that increase in size in the direction of the outer margin. The outer laterals are small and reduced with two short, acutely pointed cusps along the outer edge. REPRODUCTIVE SYSTEM.— (Fig. 3B) The preampullary duct is thin and elongate and FIGURE 3. Okenia angelensis Lance, 1966. A. Buccal mass, bp = buccal pump, e = esophagus, r = radular sac, sg = salivary gland, scale = 0.15 mm. B. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vd = vas deferens, scale = 0.17 mm. expands into the elongate, sausage-shaped ampulla. The ampulla divides into a short oviduct that enters the female gland mass and a more elongate vas deferens. The proximal portion of the vas GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA 419 FiGure 4. Okenia angelensis Lance, 1966. Scanning electron micrographs of radular teeth. A. Entire radular width. B. Inner and outer lateral teeth. deferens is prostatic but very short. It narrows and continues as an elongate loop that eventually narrows slightly into the highly convoluted muscular, proximal ejaculatory portion. The penial bulb is wide and curved. The vagina is wide at its exit adjacent to the penis. More distally it narrow and continues as a thin duct that enters the base of the large, spherical bursa copulatrix. From the base of the bursa is a second duct that joins with the smaller, more pyriform receptaculum seminis. Near 420 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 the base of the receptaculum, the uterine duct separates from the duct joining the receptaculum and bursa and enters the female gland mass. The female gland mass consists of three portions, the albu- men, membrane and mucous glands. The mucous gland is the largest of the nidamental glands. Discusston.— Okenia angelensis is immediately identifiable by its translucent yellowish white body color with minute brown spots. The present material agrees closely with that original- ly described by Lance (1966). Additional details of the anatomy including the motphelosy of the digestive and reproductive systems are added here. Okenia angelica Gosliner and Bertsch, sp. nov. (Figse5Aci6—7)) TYPE MATERIAL.— Ho.oryee: CASIZ 170088, 1 specimen, Punta Gringa, Bahia de los Ange- les, Baja California, México, Oct. 2, 1984, Terrence M. Gosliner. PARATYPES: CASIZ 166889, 1 specimen, dissected, Baja California, México, Feb. 18, 1999, Sandra Millen. CASIZ 170089, 1 specimen, San Carlos, Sonora, México, Dec. 1970. CASIZ 170090, 2 specimens, | dissected, Punta Gringa, Bahia de los Angeles, Baja California, México, June 30, 1987, Terrence M. Gosliner. CASIZ 116929, 1 specimen, Isla Cedros, Pista, Baja California, México, April, 23, 1998, Hans Bertsch. CASIZ 170085, 1 specimen, Punta Gringa, Bahia de los Angeles, Baja California, México, Sept. 20, 1985, Terrence M. Gosliner. CASIZ 170087, one specimen, Punta Gringa, Bahia de los Angeles, Baja California, México, 6 October, 1984, Terrence M. Gosliner. CASIZ 170086, one specimen, Punta Gringa, Bahia de los Angeles, Baja California, México, 24 October, 1986, Hans Bertsch. CASIZ 167395, 1 specimen, Mismaloya, Bahia de Banderas, Jalisco, México, 28 February 2003, Alicia Hermosillo. ETyMoLoGy.— The specific name, angelica, comes from the fact that this species has an angelic appearance. DISTRIBUTION.— This species is known only from Bahia de los Angeles, Baja California to the Bahia de Banderas region near Puerta Vallarta, Jalisco, México (present study). NATURAL HISTORY.— This species is found on small stones in 1—2 meters of water. It appears to be associated with bryozoans. The white egg mass is a thick cylindrical coil of approximately 1.25-1.5 whorls. This species has been documented over a decade from Bahia de los Angeles where its presence is highly seasonal and variable (Tables 1—2). EXTERNAL MORPHOLOGY .— The living animals (Fig. SA) are small, 10-20 mm in length. The preserved specimens are 3—4 mm long. The body is moderately wide and ovoid and relatively high in lateral profile. There is a well-developed, distinct notal border. The foot extends posteriorly and is devoid of notal papillae. The body color is deep purplish with an extensive opaque white patch on the center part of the notum. The white patch may also have areas of orange pigment. The mar- gins of the notum, foot, bases of the rhinophores, lateral papillae, tubercles and gill are all covered with dark purple pigment. The apices of the lateral papillae and rhinophores are covered with light orange pigment. The tips of the gill branches are covered with dark brown. There is a lateral band of chocolate brown and irregular tubercles of the same color scattered over the notum over the opaque white. Seven to nine elongate and slightly curved papillae are scattered along either side of the notum. The posterior pair of papillae on either side share a common base. No mid-dorsal papil- lae are present on the notum, but rounded irregular tubercles are present. The gill consists of 5-10 unipinnate branches. The rhinophores are elongate with 16-18 congested lamellae. The anterior end of the broad foot (Fig. 6A) contains two elongate triangular velar lobes that are united medial- ly. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA 42] a “a ? he Oe ¥ ys ; cr og a “ Ay) "Nill Se eb FOG bia” ane FIGURE 5. Living animals. A. Okenia angelica sp. nov., Punta Gringa, Bahia de los Angeles, Baja California, México, photo by Terrence M. Gosliner. B. Okenia cochimi sp. nov., Las Ballenas, Espiritu Santo Island, Baja California Sur, México, photo by Terrence M. Gosliner. C. Okenia mexicorum sp. nov., Bahia de Banderas, Jalisco, México, photo by Alicia Hermosillo. 422 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 TABLE |. Summary of annual observations of TABLE 2. Summary of monthly observations of Okenia angelica at Bahia de los Angeles during Okenia angelica at Bahia de los Angeles during the 10 the 10 year period of 1992-2001. Columns indi- year period of 1992-2001. Columns indicate totals for cate year, total hours of search time, number of each month of hours searched, number of specimens, animals seen, and density per hour. and density per hour. Year Hours # Specimens Density/hour Month — Hours # Specimens Density/hour 1992 27.52 2 072 Jan 18.34 0 — (1 egg mass) 1898 Dai, : B59 : aa foot 1 Ne 2 Mar 16.7 7} 12 — ; Apr 2a 0 1995 30.3 5 165 May 71 1 058 1996 24.8 1 04 June 33 0 — (1 egg mass) 1997 36.55 1 027 July 39192 30 .75 (egg masses) 1998 13.7 4 9 Aug 9.65 1 .l (egg masses) Sept 26.47 6 .23 (egg masses 1999 18.15 7 39 : ee Oct 4.55 | JP) 2000 15.7 I 064 Nov 34.35 DD .64 (egg masses) 2001 18.05 13 a Dec 17.08 3 18 DIGESTIVE SYSTEM.— The buccal mass (Fig. 6B) is thick and muscular with a rounded buccal pump directed dorsally. There are 3-4 minute, simple pyriform oral glands on either side of the mouth. The radular sac is short and extends ventrally from the buccal mass. The cuticle expands as it enters the buccal pump. A labial cuticle surrounds the lips at the opening of the mouth. Discrete jaw elements are visible at the apex of the cuticle. The labial elements (Fig. 7A) have a broad quad- rangular shape with 2-4 rounded apical lobes. The radular formula is 25 X 1.1.0.1.1. in one spec- imen examined. The inner lateral teeth (Fig. 7B) are broad basally with an elonage, acutely point- ed cusp and 29-31 elongate fine, acutely pointed denticles along the elongate masticatory margin. The outer laterals are slightly smaller (Fig. 7B) with a quadrangular shape and rounded margin without a distinct cusp. REPRODUCTIVE SYSTEM.— (Fig. 6C) The preampullary duct is short and thin and expands into sausage-shaped ampulla. The ampulla divides into a short, narrow oviduct and the wider prostatic FIGURE 6. Okenia angelica sp. nov., A. ventral view of head, scale = 0.5 mm. B. Buccal mass, bp = buccal pump, ¢ = esophagus, r = radula sac, sg = salivary gland, scale = 0.38 mm. C. Reproductive system, am = ampulla, bc = bursa copu- latrix, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vd = vas deferens, scale = 0.5 mm. GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA B FiGuRE 7. Okenia angelica sp. nov.. Scanning electron micrographs. A. Jaw rodlets, scale = 10 um. B. Radular teeth, scale = 40 um. 424 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is short and wide. The distal end of the prostate narrows into a elongate, curved ejaculatory portion that termi- nates in a slightly wider penis. There is no distinct penial papilla. The vagina exits adjacent to the penis. It is narrow and elongate and connects directly to the large, spherical bursa copulatrix. At the base of the bursa, the receptaculum duct emerges and joins the large, pyriform receptaculum seminis. The uterine duct emerges from near the base of the receptaculum and enters the female gland mass after paralleling the vagina for some distance. Discusston.— The only other described species of Okenia, besides O. angelica, that have bilobed, flat velar tentacles are O. echinata Baba, 1949 and O. opuntia Baba, 1960, both known only from Japan. All three species also have prominent jaw elements and similarly shaped radular teeth. Both of these species differ from O. angelica in having elongate papillae over the entire sur- face of the notum. Okenia echinata is orange or brown with minute opaque white spots. Okenia opuntia which also shares the character of having the posterior pair of notal appendages having a united base, is whitish with papillae that are yellow basally and and white apically. Okenia angel- ica has 25 rows of radular teeth with 29-31 denticles on the inner lateral teeth. There are 40 rows of radular teeth with 30—35 denticles on the inner laterals of O. opuntia while in O. echinata there are 15 rows of radular teeth with 20—25 denticles on the inner laterals. Other details of the anato- my, including the reproductive morphology, were not described for the two Japanese species. Okenia cochimi Gosliner and Bertsch, sp. nov. (Figs. 5B, 8—9) TYPE MATERIAL.— HoLotyPe: CASIZ 170091, one specimen, 13 m depth, Las Ballenas Espiritu Santo Island, Baja California Sur, 26 July, 1985, Terrence M. Gosliner. PARATYPES: CASIZ 170833, one specimen, 13 m depth, Las Ballenas Espiritu Santo Island, Baja California Sur, 26 July, 1985, Terrence M. Gosliner. CASIZ 170092, one specimen, dissected, Isla Cedros, Baja California, México, 29 December, 1985, Hans Bertsch. EtyMoLoGy.— The specific epithet, cochimi, comes from the name for the Cochimi tribe of native Americans that inhabited central Baja California. DISTRIBUTION.— This species is known only from the Isla Cedros on Pacific coast of Baja California, from Isla Ballena near La Paz in the Gulf of California south to Bahia Ballena near Puerto Vallarta, México (present study). NATURAL HISTORY.— This species is has been found in relatively shallow water of 10—15 meters depth, but has not been found in association with any prey species. EXTERNAL MORPHOLOGY.— The living animals are small, 2-4 mm in length. The body is moderately elongate and relatively high. There is a well-developed, distinct notal border. The foot extends posteriorly and is devoid of papillae. The body color is uniformly lemon yellow. There are 5—8 pairs of elongate lateral papillae along the sides of the body. The two anteriormost papillae are situated in front of the rhinophores and are anteriorly directed. A single medial papilla is present mid-dorsally anterior to the gill. The gill consists of four unipinnate branches. The rhinophores are elongate with 5—6 congested lamellae. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. The foot is narrow and is wider anteriorly. The oral region is rounded and lacks distinct tentacles (Fig. 8A). DIGESTIVE SYSTEM.— The buccal mass is thick and muscular with a rounded buccal pump directed dorsally. Numerous, large, elongate, pyriform oral glands are present at the opening of the GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA 425 buccal mass into the mouth. The radular sac is short and extends ventrally from the buccal mass. The esophagus is thin and elon- gate and inserts into the buccal mass immediately ventral to the buccal pump. A rounded, lobate salivary gland is present on either side of the buccal mass anterior to the junction of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains FIGURE 8. Okenia cochimi sp. nov., A. ventral view of head, scale = 0.2 ‘ : mm. B. C. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = irregular polygonal plates. The . tae 2 a vemaete hy oS : female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = enticlesexpands as it enters the —acina scale = 0.17 mm. buccal pump. The radular formu- la is 13 X 1.1.0.1.1. The inner lateral teeth (Fig. 9A—B) are wide basally with an elongate acute cusp. The masticatory margin of the inner lateral bears about 16—19 triangular denticles that increase in size in the direction of the outer margin. The outer laterals are small and reduced (Fig. 9A—B) and are triangular in shape with a short curved cusp at the apex. CENTRAL NERVOUS SYSTEM.— The ganglia of the central nervous system are highly concen- trated and surround the esophagus, at the posterior end of the buccal mass. The cerebral and pleu- ral ganglia are largely fused. A sessile eye is present at the base of either cerebral ganglion. The pedal ganglia are smaller than the cerebropleural ganglia and are separated by a short commissure. Paired buccal ganglia are situated ventral to the esophagus. REPRODUCTIVE SYSTEM.— (Fig. 8B) The preampullary duct is long and thin and expands into an ovoid ampulla. The ampulla divides into a short, narrow oviduct and the wider prostatic portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albu- men gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of a single, folded portion. The distal end of the prostatic segment narrows into a short, straight ejaculatory portion that terminates near the base of the short, wide penis. There is no distinct penial papilla. The vagina exits adjacent to the penis. It is narrow throughout its relatively short length. It connects directly to the irregularly shaped, elongated bursa copulatrix and the smaller, pyriform receptaculum seminis. Near the base of the receptaculum seminis, the uterine duct separates and enters the albumen gland. Discusston.— No other described species of Okenia has a uniformly colored yellow body. Several other species, including O. angelensis and O. cupella have a cream to yellowish color (present study; Valdés and Ortea 1995) but have scattered brownish markings, more widely sepa- rated rhinophoral lamella and a more elongate body. Several other species Okenia have lateral appendages with a single median dorsal papilla. In a study of the Indo-Pacific species (Gosliner 2004), these taxa were found in two primary clades, one that includes the species O. lambat Gosliner, 2004, O. virginiae Gosliner, 2004, and O. kendi Gosliner, 2004. These taxa are united in having a short vagina with the uterine duct emerging directly off the vagina rather than from the base of the receptaculum seminis. This differs from the elongate vagina of O. cochimi. These taxa also have small oral glands, unlike those found in O. cochimi. The second clade that includes species with a single mid-dorsal papilla is the one that includes O. japonica Baba, 1949; O. pur- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 —* FIGURE 9. Okenia cochimi sp. noy. Scanning electron micrographs of radular teeth. A.Radular teeth. B. View of half row of radular teeth. pureolineata Gosliner, 2004 and O. liklik Gosliner, 2004. Members of this group have a more elon- gate body shape and large oral glands, as in O. cochimi. All of these species are clearly distinct from O. cochimi. Okenia japonica is uniformly white with a light gray to opaque white dusting GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA 427 over the entire body surface, while O. cochimi is yellow throughout the body. Both O. purpureo- lineata and O. liklik have a purple body color with darker purple lines in the case of O. purpureo- lineata or with brown and white markings in O. /iklik. Okenia cochimi and O. japonica have a sin- gle elongate mid-dorsal papilla while O. purpureolineata has two elongate mid-dorsal papillae and in O. liklik the papilla has been modified to form a short sail-shaped ridge. The outer lateral teeth of O. japonica and O. purpureolineata are quadrangular while those of O. cochimi and O. liklik are triangular with an elongate, curved cusp. Okenia mexicorum Gosliner and Bertsch, sp. nov. (Figs. 5C, 10-11) TYPE MATERIAL.— HoLoryPe: CASIZ 166888, one specimen, 20 m depth, Mismaloya, Bahia de Banderas, Jalisco, México, 28 February 2003, Alicia Hermosillo. PARATYPES: CASIZ 170044, 5 specimens, 2 dissected, 20 m depth, Mismaloya, Bahia de Banderas, Jalisco, México, 17 November 2003, Alicia Hermosillo. CASIZ 170043, 8 specimens, 20 m depth, Mismaloya, Bahia de Banderas, Jalisco, México, 22 November 2003, Alicia Hermosillo. CASIZ 167394 , 1 specimen, 20 m depth, Majahuitas, Bahia de Banderas, Jalisco, México, 29 May 2003, Alicia Hermosillo. CASIZ 170045, 3 specimens, | dissected, 1 m depth, Ensenada de los Muertos, Baja California Sur, México, 25 October 2001, Orso Angulo Campillo. EtTyMOLoGy.— The specific name, mexi- corum, honors our outstanding young Mexican colleagues, Alicia Hermosillo and Orso Angulo Campillo who have advanced greatly our understanding of the opisthobranch fauna of the tropical Americas. DISTRIBUTION.— This species is known along the Mexican coast from Baja California Sur to Bahia de Banderas, Jalisco. NATURAL HISTORY.— Okenia mexicorum has been found in relatively shallow water from 1-20 m depth. Specimens are found in the open on hard and soft substrate, but this species has not been found in association with prey species. EXTERNAL MORPHOLOGY.— The living animals (Fig. 5C) are small, 5-8 mm in length. The preserved specimens are 2-3 mm. long. The body is moderately wide and ovoid and relatively high in lateral profile. There is a well-developed, distinct notal border. The foot extends posteriorly and is devoid of notal papillae. The body color is translucent white with an extensive opaque white patch on the FicurE 10. Okenia mexicorum sp. nov. A. ventral view of body, scale = 0.3 mm. B. Buccal mass, bp = buccal pump, center part of the notum that extends from the level of the rhinophores to the posterior end of the foot. External to this white patch are brick red pigmented areas that extend along the notal e = esophagus, r = radula sac, scale = 0.2 mm. C. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = recep- taculum seminis, v = vagina, vd = vas deferens, scale = 0.2 mm. 428 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 FiGureE 11. Okenia mexicorum sp. noy. Scanning electron micrograph of radular teeth. A. Inner and outer lateral teeth. B. Half-row of radular teeth. margin and are also present on the eight to ten pairs of lateral papillae and on the single, elongate medial papilla located immediately anterior to the gill. This papilla is found at the posterior end of a long, elevated medial ridge. Red pigment patches are also found on the lateral sides of the body. Two pairs of these papillae are found anterior to the rhinophores while the remaining ones are pos- terior and extend to the level posterior to the gill. The posterior two papillae per side are much longer than the more anterior ones and do not share a common base. The rhinophores are uniform- ly white and have 7-9 simple lamellae. The apices of a few of the more posterior lateral papillae have opaque white or translucent white pigment. The gill consists of five to six unipinnate branch- es which bear mottled brick red pigment on their surfaces. The anterior end of the broad foot (Fig. 10A) is simply rounded without distinct oral tentacles. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. GOSLINER AND BERTSCH: SYSTEMATICS OF PACIFIC COAST OKENIA 429 DIGESTIVE SYSTEM.— The buccal mass (Fig. 10B) is thick and muscular with a rounded buc- cal pump directed dorsally. There are several small oral glands around the mouth. The radular sac is long and curved and extends ventrally from the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth but no discrete jaw elements are visible. The cuticle expands as it enters the buccal pump. The radular formula is 15 X 1.1.0.1.1. in one specimen examined. The inner lat- eral teeth (Fig. 11 A—B) are broad basally with a sharp curvature that places the thickened mastica- tory margin at a 90 degree angle relative to the basal portion. The masticatory portion has an acute- ly pointed cusp with 26-34 short, triangular acutely pointed denticles along its cutting edge. The outer laterals are significantly smaller and reduced (Fig. 11A—B) with a single curved, sharply pointed cusp. REPRODUCTIVE SYSTEM.— (Fig. 10C) The preampullary duct is short and thin and expands into sausage-shaped ampulla. The ampulla divides into a short, narrow oviduct and the wider pro- static portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nida- mental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of two folded segments. The distal end of the prostatic segment narrows into an elongate, curved ejaculatory portion that terminates in a long wider, curved penis. There is no dis- tinct penial papilla. The vagina exits adjacent to the penis. It is narrow and moderately long and connects directly to the large, spherical bursa copulatrix. Near the base of the vagina, the receptac- ulum duct emerges and parallels the vagina for a short distance prior to joining the large, pyriform receptaculum seminis. The uterine duct emerges from near the base of the receptaculum and enters the female gland mass. DiscussioNn.— No other species of Okenia has a whitish body color with red markings. Okenia mexicorum is similar to O. mediterranea (Ihering, 1886), the only other Okenia that has a prominent medial mid-dorsal ridge (Valdés and Ortea 1995). However, O. mediterranea lacks a papilla at the posterior end of the ridge and differs markedly in its radular and reproductive anato- my. In O. mediterranea, the inner lateral radular teeth (Schmekel and Portmann 1982; Valdés and Ortea 1995) have fewer denticles (12—14) than do those of O. mexicorum (26-34) and the curva- ture of the teeth of O. mediterranea is more gradual rather than being sharply angled as in O. mex- icorum. The outer lateral teeth of O. mediterranea have a proportionately wider base and a short- er cusp than do those of O. mexicorum. The vaginal duct of O. mexicorum is very short, whereas it is elongate in O. mediterranea (Valdés and Ortea 1995). Okenia hispanica Valdés and Ortea, 1995, is white with pink and yellow spots, but it lacks a medial ridge and a medial papilla. ACKNOWLEDGMENTS This research has been supported by a grant from the National Science Foundation, PEET Program (DEB-9978155). Fieldwork was supported by the California Academy of Sciences. Specimens were also collected by several colleagues including Alicia Hermosillo, Orso Angulo Campillo, Sandra Millen, Luis Aguilar, Bob Van Syoc, Mike Ghiselin and Mike Miller. Their assis- tance and support are greatly appreciated. LITERATURE CITED BABA, K. 1949. Opisthobranchia of Sagami Bay. \wanami Shoten, Tokyo, Japan. 211 pp. BABA, K. The genera Okenia, Goniodoridella, and Goniodoris from Japan (Nudibranchia — Goniodorididae). Publications of the Seto Marine Biological Laboratory 8(1):79-83. 430 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 22 BEHRENS, D. 1991 Pacific Coast Nudibranchs. Sea Challengers: Monterey. 107 pp., 217 plates. GOSLINER, T.M. 2004. Phylogenetic systematics of Okenia, Sakishimaia, Hopkinsiella and Hopkinsia (Nudi- branchia: Goniodorididae) with descriptions of new species from the tropical Indo-Pacific. Proceedings of the California Academy of Sciences 55(5):125—161. LANCE, J. 1966. New distributional records of some northeastern Pacific Opisthobranchiata (Mollusca: Gastropoda) with descriptions of two species. The Veliger 9(1):69-81. MuNoz, M.A., A. VALDES, AND J. ORTEA. 1996. The genus Okenia Menke 1830 (Nudibranchia: Goniodorid- idae) in Chile. Haliotis 25:101—106. O’ DONOGHUE, C. 1921. Nudibranchiate Mollusca from the Vancouver Island region. Transactions of the Royal Canadian Institute 13(1):147-209. RUDMAN, W.B., 2003 (February 8) Okenia vancouverensis O’ Donoghue, 1921. [In] Sea Slug Forum. http://www.seaslugforum.net/okenvanc.htm SCHMEKEL, L., AND A. PORTMANN. 1982. Opisthobranchia des Mittlemeeres, Nudibranchia und Saccoglossa. Faune flora del Golfo di Napoli 40. Monografia della Stazione Zoologica di Napoli. Springer-Verlag, Heidelberg. 410 pp., 36 pls. THOMPSON, T., AND G. BROWN. 1984. Biology of Opisthobranch Mollusks, vol. Il. Ray Society, London. 229 pp., 41 pls. VALDES, A., AND J. ORTEA. 1995. Revised taxonomy of some species of the genus Okenia, Menke, 1830 (Mollusca: Nudibrancia) from the Atlantic Ocean, with the description of a new species. The Veliger 38(3):223-234. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 23, pp. 431-438, 3 figs., 1 table September 30, 2004 A New Species of Okenia (Gastropoda: Nudibranchia: Goniodorididae) from the Pacific Coast of Costa Rica Yolanda E. Camacho-Garcia ! and Terrence M. Gosliner 2 ! Instituto Nacional de Biodiversidad (INBio), Unidad de Malacologia Apdo. 22-3100 Santo Domingo de Heredia, Costa Rica. Email: ycamacho@inbio.ac.cr; *Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103, USA. Email: tgosliner@calacademy.org A new species of opisthobranch mollusk of the genus Okenia is described from the Pacific Coast of Costa Rica. Okenia academica sp. nov. has a translucent gray-brown body, and short dorsal tubercles, nine pairs of mantle processes which are white in color with orange to brick red apices. The species is compared to other species from the Panamic Province and the West coast of North America. This is the first record of the genus for the Pacific Coast of Costa Rica. Key words: Mollusk, Nudibranchia, Costa Rica, Okenia. RESUMEN Una nueva especie de molusco opistobranquio del género Okenia es descrito de la costa pacifica de Costa Rica. Okenia academica sp. nov. tiene un cuerpo gris cafesuz- co traslucido, nueve procesos en el manto, blancos en color con apices anaranjados o rojo ladrillo y tubérculos cortos presentes en el dorso. La especie es comparada con otras especies de la Provincia Panamica y la costa Oeste de Norte América. Este es el primer registro del género para la costa pacifica de Costa Rica. The genus Okenia Menke, 1830 consists of 10 species found primarily along the Pacific Coast of the Americas: Okenia vancouverensis O'Donoghue, 1921 from Canada, Okenia plana Baba, 1960 from California and Japan, Okenia angelensis Lance, 1966 from California and México, Okenia sp. | and Okenia sp. 2 (in Behrens 2004) from California, Okenia luna Millen, Schrédl, Vargas and Indacochea, 1994 from Pert and Chile, and three other new species from México (Gosliner & Bertsch, 2004). Another species, Hopkinsia rosacea MacFarland, 1905 has recently been placed within the genus Okenia (Gosliner, 2004). The description of the present species is the first report and description of a member of the genus Okenia for the Pacific Coast of Central America. SPECIES DESCRIPTION Genus Okenia Menke, 1830 Type species: Doris elegans Bronn, 1826 Okenia academica Camacho-Garcia and Gosliner, sp. nov (Figs. 1-3) TYPE MATERIAL.— Ho_otyPe. Punta San Francisco, Playa Tamarindo, Parque Nacional Las 43] 432 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 23 Baulas, Area de Conservaci6n Tempisque, Costa Rica (9°03'58"N, 85°51'08"W), January 12, 2001, 1 specimen, dissected, 6 mm preserved length, 0 meters depth, leg. J. Magafia (INB0003118102); SEM stub with radula and jaw (INB0003764988); PARATyPES. Punta San Francisco, Playa Tamarindo, Parque Nacional Las Baulas, Area de Conservacion Tempisque, Costa Rica (9°03'58”"N, 85°51'08"”W), January 12, 2001, 1 specimen, dissected, 6 mm preserved length, 0 meters depth, leg. J. Magana (CASIZ 170030), with the radula mounted on a SEM stub with the same number; San Miguel, Reserva Natural Absoluta de Cabo Blanco, Area de Conservacion Tempisque, Costa Rica (9°34'53”"N, 85°08'26"”W), September 18, 1998, 1 specimen, 3 mm pre- served length, 0 meters depth, leg. F. Alvarado (INB0001496648). GEOGRAPHIC RANGE.— This species is known only from the Pacific Coast of Costa Rica (present study). ETYMOLOGY.— The species is named after the California Academy of Sciences (CAS) in San Francisco, California to honor its contribution to the inventory of opisthobranch mollusks in Costa Rica and to honor the celebration of its sesquicentennial. EXTERNAL MORPHOLOGY.— The body is oval and elongated (Fig 1A—B). The dorsum has as many as 24 small tubercles of different sizes. There are no spicules present on the notum. The head has the form of a bilobed velum (Fig. 2B). The rhinophores are proportionally very long with respect to the length of the animal and curve backwards with about 18 fine lamellae. There are two notal processes above the veil and seven elongated notal papillae on each side of the body that can curl either inward or upward. The last process on each side of the body is much longer and bifid. The branchial leaves are larger anteriorly. The branchial plume is composed of 10 to 13 unipinnate branchial leaves. The anal papilla is located in the cen- ter of the branchial leaves. The viscera are visible through the notum in the anterior part of the body The prolongation of the pointed foot is visible when the animal is in motion. The foot is simply rounded anteriorly. : : Figure 1. A. Drawing of the living animal of Okenia academica sp. nov., The background color of the Costa Rica (INB003118104). B. Photograph of the living animal of Okenia body is translucent gray-brown. academica sp. nov., Costa Rica (INB003118104). CAMACHO-GARCIA AND GOSLINER: NEW OKENIA FROM COSTA RICA 433 The notum and the greater ventral part of the foot are opaque white. The largest tubercles in the center of the back are reddish brown to reddish orange in color (Fig. 1A—B). The rhinophores and gill range from translucent white to reddish brown with small white opaque spots. The apices of the lateral processes are orange to brick red. The anus is reddish brown. The anterolateral corners of the foot are white translucent with some yellowish spots. Ventrally, the foot is translucent gray. ANnaTOMY.— The buccal bulb is large and muscular (Fig. 2A). The buccal pump is large and expands dorsally and posteriorly. A pair of short salivary glands is present adjacent to the entrance of the elongate, thin esophagus. The radular sac descends ventrally from the buccal mass. The radular formula is 36 x (1.1.0.1.1) in a 6 mm preserved length specimen (Fig 3A). The inner lateral teeth have up to 28 strong denticles on the inner surface of the masticatory border (Fig 3C—D). Each outer lateral tooth has one cusp (Fig 3B). The labial disc is composed of several rec- tangular jaw elements which are homogeneously distributed (Fig 3E). Each of the jaw elements bears up to six irregular serrations on the outer edges (Fig 3F). The ampulla is very long and thick. It divides into a short oviduct that enters the female gland mass near the lower right side of the mass and the prostate (Fig. 2C). The prostate is long, tubular A B Ficure. 2. A. Buccal mass from right side of Okenia academica sp. nov., scale bar= 1 mm. Abbreviations: m= mouth; rd = radular sac; bp= buccal pump; sg = salivary gland. B. Head of Okenia academica sp. nov., scale bar= 1 mm. C. Reproductive system of Okenia academica sp. nov., scale bar= 1 mm, Abbreviations: am=ampulla, be=bursa copulatrix, fg=female gland, p= penis, pr=prostate, sr=seminal receptacle, v=vagina. 434 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 23 © Date 19 May 2003 64 Time :11:41 FIGURE 3. Scanning electron micrographs of Okenia academica sp. noy. A. Radula (INB003118102). B. Lateral and marginal teeth (INB0003118104) C. Detail of the lateral teeth (INB0003118104). D. Detail of the denticles of a lateral tooth (INB003118102). E. Jaw (INB003118102). F. Detail of the jaw elements showing the serrated edge (INB003118102). and convoluted near the middle section. It constricts into a thin, tubular and muscular vas deferens that forms a slight loop into itself. The distal part of the vas deferens is inside a pear-shaped penis. No penial spines were observed in the slide preparation of the penis. The vagina is very long. Near its base, the uterine duct branches and enters the female gland mass. More proximally, it enters the oval bursa copulatrix. A shorter duct emerges from the base of the bursa copulatrix and leads to an CAMACHO-GARCIA AND GOSLINER: NEW OKENIA FROM COSTA RICA 435 oval and elongate seminal receptacle. The seminal receptacle is twice as large as the bursa copula- trix. REMARKS.— The seven species of Okenia known to inhabit the Pacific Coast of North America have been reviewed recently (Gosliner and Bertsch 2004), this review included three new species. Of the four species that had already been described, O. rosacea (MacFarland, 1905), and O. plana Baba, 1960, differ from the present species in lacking a distinct lateral side of the body separate from the notum and foot. Okenia vancouverensis (O’Donoghue, 1921) differs from O. academica in its external body color, being brown with opaque white spots and in having shorter lateral papillae. Okenia angelensis Lance, 1966, has an elongate body with a yellowish color orna- mented with brown spots. It has only two to three rhinophoral lamellae in contrast to the 18 found in O. academica. The three new Mexican species also differ from O. academica. Okenia cochimi Gosliner and Bertsch, 2004, has a uniformly yellow body color with a single elongate medial notal papilla anterior to the gill. Okenia mexicorum Gosliner and Bertsch, 2004, is opaque white with red markings and also has a single medial notal papilla anterior to the gill. (See also Table | for sum- mary of diagnostic characters.) The species that is most similar to O. academica is Okenia angelica Gosliner and Bertsch, 2004. Externally this species is similar in that it has a broad body with elongate lateral papillae and low tubercles on the notum. The two posteriormost pairs of papillae also originate from a common base in both species and both have a bilobed velum. However, there are significant differences as well. In O. angelica, the margins of the notum, foot, bases of the rhinophores, lateral papillae, tubercles, and gills are all covered with dark purple pigment. The middle of the notum is covered by opaque white as in O. academica, but the opaque white covers much less of the notal surface in O. angelica. In O. angelica, the lateral papillae are covered with light, opaque orange-white pig- ment and lack the darker orange apices of O. academica. Internally, the two species have similar- ly shaped radular teeth and jaw plates. The radula of O. academica contains 36 rows of radular teeth in a 6 mm specimen, while a 12 mm specimen of O. angelica had only 25 rows of teeth. The inner lateral teeth of O. angelica have a more elongate, acutely pointed cusp and have a few more denticles that are more widely spaced than are those of O. academica. The most significant differ- ences between the two species are found in the morphology of the reproductive system. In O. aca- demica the uterine duct emerges from near the base of the vagina, while in O. angelica, it emerges from the top of the vaginal duct at the base of the bursa copulatrix and receptaculum seminis. In O. academica, the receptaculum seminis is larger than the bursa copulatrix, while in O. angelica, the bursa is larger. In O. academica the prostate is much longer than the ejaculatory portion of the vas deferens, while in O. angelica the prostate is very short and the ejaculatory segment is very long. The penial sac of O. academica is wider than the ejaculatory duct, while in O. angelica they are about the same width. Okenia luna Millen, Schrédl, Vargas and Indacochea, 1994 can be differentiated from O. aca- demica externally by the number and color of the mantle processes, which are white with orange to brick red apexes and the presence of several tubercles or papillae on the notum, and a notched foot. Okenia luna is characterized by a smooth dorsum, a white body with yellow markings, unnotched foot, and flat, triangle-shaped oral tentacles. ACKNOWLEDGMENTS This research was made possible by the Cooperative Agreement between the Ministry of Environment and Energy (MINAE) and Instituto Nacional de Biodiversidad (INBio). The field- work in Costa Rica was funded by the INBio through the Netherlands Government and the project 436 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 23 TABLE |. Comparative morphology of the valid species of the genus Okenia from the Panamic Province and Species Description source Distribution Dorsal color Ventral color Velar appendages per side Okenia rosacea MacFarland 1905; Oregon to Isla San rose pink pink twelve, pink (MacFarland, 1905) Gosliner, 2004 Martin Baja Calif Okenia vancouveren- O'Donoghue, 1921; | Canada Brownish with unknown one, same color as sis O'Donoghue, 1921 Okenia plana Baba, 1960 Okenia angelensis Lance, 1966 Okenia luna Millen, Schrédl, Vargas, and Indacochea 1994 Okenia angelica Gosliner and Bertsch, 2004 Okenia cochimi Gosliner and Bertsch, 2004 Okenia mexicorum Gosliner and Bertsch, 2004 Okenia academica Camacho-Garcia and Gosliner, sp. nov Gosliner and Bertsch, 2004 Baba, 1960; Rudman and Darvell, 1990; Bouchet and Ortea, 1993; Gosliner, 2004 Lance, 1966; Behrens, 1991 and Munoz et.al. 1996; Schrédl, 2003 Millen, Schrédl, Vargas, and Indacochea 1994; Schrédl, 2003 Gosliner and Bertsch, 2004 Gosliner and Bertsch, 2004 Gosliner and Bertsch, 2004; Behrens, 1991 present paper Japan, Philippines, Hong Kong, New Zealand, Australia, introduced into California California to Chile Pert and Chile Bahia de Los Angeles to Bahia Banderas, Mexico Isla Cedros to Bahia Ballena, México Baja California Sur to Bahia de Banderas, Jalisco Costa Rica opaque white spots and opaque white medial line on foot translucent yellowish white, small brown spots translucent white flecked with small yellowish and white granules. Reddish brown and green spots also present hyaline white with a mid-dorsal stripe of yellow spots deep purplish with opaque white on the center of the notum uniformly lemon yellow opaque white patch- with reddish brown patches translucent gray- brown translucent white sparsely speckled w/light yellow & white hyaline white usually without pigment translucent white translucent white translucent white translucent gray the body one, speckled with dark brown one, white one, yellow two, with white apices one, lemon yellow two, reddish brown one, with orange to brick red apices “Development of Biodiversity Knowledge and Sustainable Use in Costa Rica.” This paper is sup- ported in part by the National Science Foundation through the PEET grant DEB-9978155, “Phylogenetic systematics of dorid nudibranchs,” to Terrence M. Gosliner and Angel Valdés and the California Academy of Sciences. Angel Valdés made the drawing of the specimens from Costa Rica and Maribel Zuniga helped with some technical aspects of the paper. David Butvill made con- structive comments on this manuscript. CAMACHO-GARCIA AND GOSLINER: NEW OKENIA FROM COSTA RICA 437 from Western North America. Mantle Rhinophoral — Branchial Dorsal papil- Oral tentacles Foot Cusps of outer Jaws Radula processes per bases leaves lae lateral teeth side : a3 cs numerous, reddish pink _ reddish pink numerous, A N reduced to a small P, short 16-22 x pink pink plate (1.1.0.1.1) 9-10 pairs lat- same coloras same coloras P, 24 papillae P, digitiform U, broad — one single promi- P 32 x erally the body the body on the notum rounded tips nent cusp (1.1.0.1.1) 4, speckled in translucent translucent P, one medial A N two cusps, P, regular 21-29 x dark brown white white with with brown occasionally with rounded (1.1.0.1.1) dark brown specks 2-3 smaller denti- specks cles 6-7, white white w/yel- white with P, five or six P, pointedand U two prominent A, no distinct 21 x low & white yellow and fleshy cusps jaw plates (1.1.0.1.1) specks at brown flecks 19x distal end; (CICIEAOSTESIb)) brown specks near base 8-10, yellow hyaline white hyaline white A P, broad, flat- U one small cusp P, rectangular 23-27 x bases, cream witha yellow tened triangles with six serra- (1.1.0.1.1) leaves w/ streak tions opaque white pigment 6-7 w/white _ purple with dark P, many P, broad U no distinct cusp _—P, rectangular 25 x apices brown apices rounded with 2-4 ser- (1.1.0.1.1) tubercles, rations brown in color 4-7 w/lemon lemon yellow lemon yellow P, 1 medial P, rounded U one single and P, small irreg- 13 x apices lemon yellow curved ular polygo-—_(1.1.0.1.1) nal plates 6-9 reddish _— white translucent P, one medial P, rounded U one single and P, very weak- 15 x brown, some white with red- reddish brown curved ly developed (1.1.0.1.1) w/opaque dish brown with white white apices pigment apex 7 w/orange to translucent translucent P, reddish P, broad U one small cusp P, rectangular 36 x brick red apices with small white opaque spots or very dark reddish brown color w/small white opaque or dark reddish brown spots brown to red- dish orange in color LITERATURE CITED with six serra- tions (1.1.0.1.1) Bapa, K. 1960. The genera Okenia, Goniodoridella and Goniodoris from Japan (Nudibranchia: Goniodorididae). Publications of the Seto Marine Biological Laboratory 8(1):79-83, pls. 7-8. BEHRENS, D.W. 1991. Pacific Coast Nudibranchs, a Guide to the Opisthobranchs of the Northeastern Pacific. Sea Challengers, Los Osos, California. 112 pp., 162 photos. BEHRENS, D.W. 2004. Pacific Coast nudibranchs, Supplement II. New species to the Pacific Coast and new information on oldies. Proceedings of the California Academy of Sciences 55(2):11—S4. BoucHET, P., AND J. OrTEA. 1983. A new Hopkinsia feeding on bryozoa in the South Pacific (Mollusca: Opisthobranchia). Venus 42(3):227-233. 438 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 23 GOSLINER, T. 2004. Phylogenetic systematics of Okenia, Sakishimaia, Hopkinsiella and Hopkinsia (Nudibranchia: Goniodorididae) with description of new species from the Tropical Indo-Pacific, Proceedings of the California Academy of Sciences 55(5):125—161. GOSLINER, T., AND H. BERTSCH. 2004. Systematics of Okenia from the Pacific Coast of North America (Nudibranchia: Goniodorididae) with descriptions of three new species. Proceedings of the California Academy of Sciences 55(21):412—428, figs. 1-11. LANCE, J.R. 1966. New distributional records of some northeastern Pacific opisthobranchiata (Mollusca: Gastropoda) with descriptions of two new species. Veliger 9(1):69-81. MACFARLAND, F.M. 1905. A preliminary account of the Dorididae of Monterey Bay, California. Proceedings of the Biological Society, Washington 18:35—S4. MILLEN, S.V.; M. SCHRODL, N. VAGAS, AND A. INDACOCHEA. 1994. A new species of Okenia (Nudibranchia: Doridacea) from the Peruvian Faunal Province. Veliger 37(3):312-318. Munoz, M.A., A. VALDES, AND J. ORTEA, 1996. The Genus Okenia Menke, 1830 (Nudibranchia: Goniodorid- idae) in Chile. Haliotis 25:101—106. O’DONOGHUE, C.H. 1921. Nudibranchiate Mollusca from the Vancouver Island region. Transactions of the Royal Canadian Institute 13(1):147—209. RUDMAN, W.B., AND B.W. DARVELL. 1990. Opisthobranch molluscs of Hong Kong, Part 1. Goniodorididae, Onchidorididae, Triophidae, Gymnodorididae, Chromodorididae (Nudibranchia). Asian Marine Biology 7:31-80. SCHRODL , M. 2003. Sea Slugs of Southern South America: Systematics, Biogeography and Biology of Chilean and Magellanic Nudipleura (Mollusca: Opisthobranchia). Conch Books, Hackenheim, Germany. 165 pp. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 24, pp. 439-450, 8 figs. September 30, 2004 A New Species of Schizobopyrina Markham, 1985 (Crustacea: Isopoda: Bopyridae: Bopyrinae) Parasitic on a Gnathophyllum Shrimp from Polynesia, with Description of an Associated Hyperparasitic Isopod (Crustacea: Isopoda: Cabiropidae) Jason D. Williams !3 and Christopher B. Boyko 2 ! Department of Biology, Hofstra University, Hempstead, New York 11549, U.S.A.; 2 Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th St., New York, New York 10024, U.S.A. A male and female bopyrid pair was found in the branchial chamber of a bumblebee shrimp, Gnathophyllum americanum Guérin-Méneville, collected from Tonga. Examination of these specimens showed that they belong to the bopyrid genus Schizobopyrina Markham, but cannot be placed in any of the described species. We describe this new species as S. bombyliaster and compare it to the other species in the genus. This is the first described species of branchial bopyrid from the genus Gnathophyllum, and only the second bopyrid described from a member of the Gnathophyllidae. The marsupium of the female bopyrid contained specimens of a new species of the hyperparasitic isopod genus Cabirops Kossmann, which is described based on light and scanning electron microscope investigations and dis- cussed in relation to other species in the genus. Bopyrid isopods of the subfamily Bopyrinae are obligate parasites found in the branchial chambers of caridean shrimp (Decapoda: Caridea). The genus Schizobopyrina Markham, 1985, was erected for species formerly placed in Bopyrina Kossmann, 1881, but which possess a maxil- liped palp, elongate oostegites 2-5, and lateral or complete separation of the pleomeres of the female. A single parasitized specimen of Gnathophyllum shrimp (Fig. 1) was found by one of us (CBB) in the collections of the California Academy of Sciences and was identified as belonging to the widespread species G americanum Guérin-Méneville. Examination of the bopyrid parasites showed they belong to an undescribed species of Schizobopyrina. In addition, the female specimen of Schizobopyrina was hyperparasitized by specimens of a new species of isopod of the genus Cabirops Kossman, 1872. The only other described species of bopyrid isopod on a member of Gnathophyllum, or the family Gnathophyllidae, is the abdominal species Diplophryxus kempi Chopra, 1930 (Hemiarthrinae), from G americanum (originally as G fasciolatum Stimpson) col- lected in the Andaman Islands. A branchial bopyrid on G. modestum Hay in the Gulf of México was reported by Dardeau et al. (1980; repeated by Markham 1985), but was never identified even to the subfamily level. All other hosts of Schizobopyrina species are in either the Palaemonidae, which with the Gnathophyllidae forms part of the Palaemonoidea, or the Hippolytidae (Alpheoidea). 3 Contact author: biojdw@hofstra.edu 439 440 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 24 METHODS Camera lucida sketches made of specimens were scanned into a Macintosh™ computer. Images were then prepared using the programs Adobe Photoshop™ and Adobe IIlustrator™. For Scanning Electron Microscopy (SEM), the male Cabirops specimen was dehydrated in an ascend- ing ethanol series followed by four changes of 100% ethanol. Dehydration was completed with a Samdri 795 Critical Point Drier. The dried specimen was mounted on an aluminum stub, coated with gold (EMS-550 Sputter coater), and viewed with a Hitachi S-2460N SEM. Carapace length (CL), including the rostrum, is provided as an indicator of size for the host shrimp. Isopod size is given as total body length (anterior margin of head to posterior margin of pleotelson). Measurements were made to 0.01 mm using an ocular micrometer. All specimens are deposited in the Division of Invertebrate Zoology, California Academy of Natural Sciences, San Francisco, U.S.A (CASIZ), except for the specimen of Schizobopyrina gra- cilis (Chopra, 1923), which is in the Museum National d’ Histoire Naturelle, Paris (MNHN). DESCRIPTIONS OF TAXA Family Bopyridae Rafinesque-Schmaltz, 1815 Subfamily Bopyrinae Rafinesque-Schmaltz, 1815 Genus Schizobopyrina Markham, 1985 Schizobopyrina bombyliaster Williams and Boyko, new species (Figs. 1-4) MATERIAL EXAMINED.— Holotype: female (4.41 mm), infesting right branchial chamber of female Gnathophyllum americanum (3.71 mm CL; CASIZ 170233), 18°43'S, 174°05’E, on coral reef to coral rubble on sand bottom, leeward side of reef, north side of reef between Langitau and Vaka’eitu Island, Vava’u Island Group, Tonga, 2—10 ft (= 0.6—2.7 m) depth, hyperparasitized by Cabirops bombyliophila sp. nov. (CASIZ 170231—170232), coll. Van Syoc, Ferreira, Campbell, and Cornfield, 30 July 1985 (CASIZ 5707). Allotype: male (1.04 mm), same data as holotype (CASIZ 170234). TYPE LOCALITY.—18°43’S, 174°05’E, leeward side of reef, north side of reef between Langitau and Vaka’eitu Island, Vava’u Island Group, Tonga, Pacific Ocean, 0.6—2.7 m depth. DESCRIPTION.— Female (Figs. 2-3). Body length 4.41 mm, maximal width 2.34 mm, head length 0.94 mm, head width 1.01 mm, pleon length 1.31 mm. Pereon and pleon deflexed dextral- ly. Body outline of pereon broad anteriorly, tapering posteriorly; pleon subequal to posterior pere- on. All body regions and pereomeres distinctly segmented (Fig. 2A—B). Cephalon triangular, tapering posteriorly, median region strongly delineated; well withdrawn into first pereomere, nearly obscuring median of first pereomere; frontal lamina strong, triangular, width approximately one-half of head and distinctly separated from head. Eyes lacking. Antenna and antennule (Fig. 3B) of 3 articles each, distal margin of distal segments with setae. Maxilliped (Fig. 3C) with narrow rounded spur; palp single segmented and non-articulated with 7 thick elon- gate setae at distal margin. Pereon composed of 7 pereomeres, broadest across pereomere 3, tapering anteriorly and pos- teriorly. All pereomeres separated. Coxal plates on sides of pereomeres all similar, indistinct; dor- solateral bosses produced on pleomeres |—-4, strongest on 2-4 and larger on right side of body (Fig. 2A). Oostegites enclosing only approximately one-fourth of brood pouch (Fig. 2B); first oostegit- WILLIAMS AND BOYKO: NEW PARASITIC ISOPODS FROM POLYNESIA 44] es asymmetrical, right one slight- ly smaller, with posterolateral lobe extended, smoothly angled, distal end rounded, not setose (Fig. 3D), left one with postero- lateral lobe short, sharply angled, distal end setose (Fig. 3E). Pereopods (Figs. 2A, 3F—G) of about same size. Propodus of all pereopods with shallow cup-like insertion for dactylus (Fig. 3F); distoventral margin of carpus with numerous scales and few short setae at distal tip. First 2 pereopods surrounding head region; no large gaps between any pereopods. Pleon with 5 distinct pleo- meres. Pleomeres 1-4 increas- ingly concave on median of pos- terior margin, with extended ovate, distally tapering and acute, biramous pleopods and unira- mous, distally produced and rounded lateral plates (Figs. 2B, 3A); pleomere 5 fused with pleo- telson (Figs. 2A), anteriorly sharply convex and _ posteriorly straight, lacking pleopods and with subquadrate, distally direct- Ficure 1. The bumblebee shrimp (Gnathophyllum americanum) harbor- ing Schizobopyrina bombyliaster sp. nov. and Cabirops bombyliophyla sp. 3 : nov. in right branchial chamber, CASIZ 170233 (shrimp), 5707 (female bopy- Ing (Fig. 2A). rid), 170232 (female cabiropid). A, dorsal view; B, lateral view (arrowhead Male (Fig. 4). Length 1.04 _ indicates Cabirops female within brood chamber of Schizobopyrina female). mm, maximum width 0.40 mm, Seale = 1.0 mm. head length 0.15 mm, head width 0.27 mm, pleon length 0.29 mm. Occurring on ventral surface of female’s pleon between pleopods (Fig. 2A); directed anteroposteriorly. Head anterior margin subovate, posterolateral margins slightly concave, posterior margin straight, widest medially, incompletely fused with Ist segment of pereon. Moderate sized eyes located mediolaterally. Antenna and antennule of 2 articles each (Fig. 4C), distally setose; extend- ing posterolaterally from head, not overeaching margins of head. Pereomere 3 broadest, tapering slightly anteriorly and posteriorly. Pereomeres 1-3 directed lat- erally; 4-7 with posterolateral margins directed slightly posterolaterally. Body lacking pigmenta- tion. All pereopods (Fig. 4D—-E) of subequal size, all articles distinctly separated; distoventral sur- faces of propodus and carpus with large flat scales and few short setae. Pleon tapering posteriorly, pleomeres directed laterally and fringed with setae. All segments fused, but with deep lateral indentations and faint dorsal suture lines demarcating segments. No ed, lateral plates; uropods lack- 442 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 24 FiGure 2. Schizobopyrina bombyliaster sp. nov., female, 4.41 mm, CASIZ 5707, holotype (with male in situ). A, dor- sal view; B, ventral view. Numbers indicate pleomere. Scale = 500 um. midventral tubercles; 5 pairs of subquadrate tuberculiform pleopods, last pair markedly smaller and more ovate than other pairs (Fig. 4B). Pleotelson (Fig. 4A—B) slightly produced medially, with sin- uous lateral margins, uropods absent. DISTRIBUTION.— Found on the shrimp, Gnathophyllum americanum from Tonga; 0.6—2.7 m depth. EtyMoLoGy.— The specific name, literally bumblebee-eater, is given to call attention to the occurrence of this parasite on a bumblebee shrimp of the genus Gnuathophyllum. REMARKS.— Of the 12 other species currently placed in Schizobopyrina (see Campos and Campos 1990), this new species most closely resembles S. gracilis (Chopra, 1923). The female of S. gracilis reported by Bourdon (1983, as Bopyrina gracilis) was examined for comparison to the material from Tonga, as the types of B. gracilis are in the Zoological Survey of India and unavail- able for examination. Bourdon’s (1983) specimen (MNHN-Ep 347, female 1.88 mm, Amboine (= Ambon, Indonesia) coll. Rumphius II, 1975, ex Periclimenes tenuis Bruce) corresponds well with the type of S. gracilis as described and figured by Chopra (1923), although the lateral plates and dorsal segmentation of the pleomeres are even less distinctly separated in the Ambon specimen. Unfortunately, the male specimen from Ambon has become lost and could not be examined, but all the characters given for it by Bourdon (1983) match those of the male type of S. gracilis. WILLIAMS AND BOYKO: NEW PARASITIC ISOPODS FROM POLYNESIA 443 FIGURE 3. Schizobopyrina bombyliaster sp. nov., female, 4.41 mm, CASIZ 5707, holotype. A, posterolateral view, left side (P=pleopod; numbers indicate pleomere); B, left antenna and antennule; C, left maxilliped, external view; D, right oost- egite 1, external view; E, left oostegite 1, external view; F, right pereopod 7; G, right pereopod 2. Scale = 250 um (A), 500 um (B), 150 um (C), 250 um (D, E), 50 um (F-G). The female of S. hombyliaster sp. nov. resembles S. gracilis in the overall shape of the head, the number of pleomeres (pleomere 5 fused with pleotelson), and the shape of the posterior mar- gin of the pleomere 5/pleotelson. It differs from S. gracilis in that the frontal lamina of the head and the dorsolateral bosses on the pereomeres are much more developed, the pleomere segments are distinct, the lateral plates on pleomeres 1-4 are much more separated, and the lateral plates of 444 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 24 FIGURE 4. Schizobopyrina bombyliaster sp. nov., male, 1.04 mm, CASIZ 170234, allotype. A, dorsal view; B, ventral view; C, left antenna and antennule; D, right pereopod 1; E, right pereopod 7. Scale = 150 um (A-B), 125 tum (C), 250 pm (D-E). pleomere 5/pleotelson are subquadrate instead of rounded. The shape of the left oostegite | is also quite different from that of S. gracilis or any other Schizobopyrina species. The male of S. bom- byliaster sp. nov. is very much like that of S. gracilis, but the posterior margin of the pleotelson is only slightly produced, while that of S. gracilis is strongly produced. The genus Schizobopyrina now contains 13 species, although we concur with Campos and Campos (1990) that S. /obata Bourdon and Bruce, 1983, does not appear to belong to this genus. Schizobopyrina bombyliaster sp. nov. belongs to the group of Schizobopyrina with females having pleomere 5 partly or wholly fused with the pleotelson that also includes S. amakusaensis (Shiino, 1939), S. andamanica (Chopra, 1923), S. bruscai Campos and Campos, 1990, S. gracilis (Chopra, 1923), and S. mivakei (Shiino, 1942). Both Chopra (1923) and Campos and Campos (1990) described S. andamanica (as andamica |sic| in Campos and Campos [1990]) and S. gracilis as hav- ing 5 pleomeres plus pleotelson, but the illustrations of Chopra (1923) clearly show only slight lat- eral distinction between the segments in some specimens of S. andamanica (Chopra 1923: pl. 20, fig. 1, text-fig. 28) and no discernable distinction in S. gracilis (Chopra 1923: pl. 21, fig. 6). This is in agreement with the specimen of S. gracilis reported by Bourdon (1983) and unlike the clear separation of those segments in all other Schizobopyrina species. WILLIAMS AND BOYKO: NEW PARASITIC ISOPODS FROM POLYNESIA 445 Family Cabiropidae Giard and Bonnier, 1887 Genus Cabirops Kossmann, 1872 Cabirops bombyliophila Williams and Boyko, new species (Figs. 1, 5-8) MATERIAL EXAMINED.— Holotype: male (0.91 mm) infesting female Schizobopyrina bom- byliaster sp. nov. (4.41 mm; CASIZ 5707) ex Gnathophyllum americanum (3.71 mm CL; CASIZ 170233), 18°43'S, 174°05’E, on coral reef to coral rubble on sand bottom, leeward side of reef, north side of reef between Langitau and Vaka’eitu Island, Vava’u Island Group, Tonga, 2—10 ft (= 0.6—2.7 m) depth, coll. Van Syoc, Ferreira, Campbell, and Cornfield, 30 July 1985 (CASIZ 170231). Allotype: immature female (2.36 mm), same data as holotype (CASIZ 170232). TYPE LOCALITY.— 18°43'S, 174°05’E, leeward side of reef, north side of reef between Langitau and Vaka’eitu Island, Vava’u Island Group, Tonga, Pacific Ocean, 0.6—2.7 m depth. DESCRIPTION.— Male (Figs. 5—7). Length 0.91 mm, maximum width 0.41 mm at segment 4, head length 0.11 mm, pleon length 0.41 mm; body tear-drop shaped. Cuticular surface with distinct striations, prominent on coxal plates (Fig. 6C—D). Head anterior margin ovate, posterior margin convex, widest at posterolateral junction with segment | (Figs. 5A, 6A—B). Subovate, moderately large eyes (approximately 60 um in maximal length) located mediolaterally, eyes with irregular pigment most prominent around medially direct- ed edge (Fig. 5A). Antenna of 3 articles (Fig. 5C), first article with 3 setae near anterolateral bor- der, article | slightly overlaps basal article of antennule, article 2 with series of shelves, 4 setae on distal margin of article 2, distal bundles of long setae on antenna extend to segment 1, extending slightly beyond margin of head (Fig. 5B). Antennule of 9 articles (4 peduncular and 5 flagellar) (Fig. 5B), first article with rounded anterior edge, articles | and 2 with apophysis on posteromedi- al edge, articles 3 and 4 lacking apophysis; flagellar articles much narrower than peduncular arti- cles, each with a terminal seta, proximal flagellar article shortest, flagellar articles 2—5 of increas- ing length, article 5 with approximately 2 long and 2 short, distal setae. Pereomere 4 broadest, tapering anteriorly and posteriorly. Body pigmentation lacking. Pereomeres with toothed coxal plates; three denticles in plates 1—5, 1 denticle in plates 6—7 (Figs. 5B; 6C—E), medial tooth largest in all segments, lateral two teeth with rounded distal end, dimin- ishing in size over segments 1—5S (Fig. 6C—E). Pereopods 1 and 2 gnathopodal with bifid dactyli (Fig. 7A—C); distoventral surfaces of propodus and carpus with large flat scales and a few setae, rows of minute setae along distal edge of propodus around joint with dactylus, with setae along edge apposed to dactylus, three stout setae on propodus near tip of dactyl (Fig. 7A—C). Pereopods 3—7 ambulatory, carpi of pereopods 3—7 with one stout terminal seta, proprodi with long axis curved, inner margins with two (pereopods 3-5) or one (pereopods 6 and 7) stout setae toward dis- tal end (Figs. 5B; 7D—F) and comb of minute setae along edge (Fig. 7F), dactyli of pereopods 3—7 with long axis curved, tapering to a sharp point (pereopods 3-5), or blunt tip (pereopods 6 and 7), with comb of minute setae (Fig. 7D—E). Pleon with 5 pleopods composed of basis (sympod), exopod, and endopod (Fig. 6F). Sympods with 1 or 2 medially directed spines, endopods with plumose setae (5 setae on pleopod 1), exopods with plumose setae (4 setae on pleopod 1) and one laterally directed seta (short on pleopod 1, longer on pleopods 2-5); setae on endopods and exopods reduced in number on pleopod 5. Pleotelson quadrangular, endopods approximately twice as long as exopods, basis with 1 or 2 pos- terior setae, endopods and exopods with terminal setae (Fig. SA—B). Immature female (Figs. 1, 8). Maximal body length 2.36 mm (from pereonal lobe 2 to pereon- 446 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 24 FiGuRE 5. Cabirops bombyliophila sp. nov., male, 0.91 mm, CASIZ 170231, holotype. A, dorsal view; B, ventral view; C, first antenna, ventral view (terminal setae only partially drawn). Scale = 150 um (A, B), 25 um (C). al lobe 7), maximal width 1.81 mm, pleon length 0.73 mm. In lateral aspect, body highly curved ventrally, forming U-shape with head region and pleon apposed (Fig. 8). Slight light-brown pig- mentation on rostrum (Fig. 8B). Pereon with 7 pereonal lobes, lobes 4—6 largest, on right side all lobes rounded (Fig. 8A), on left side lobes 1-3 rounded and lobes 4—6 with semi-circular depres- sions (Fig. 8B). Faint indication of segmentation on ventral side of pereon, no appendages. Pleon WILLIAMS AND BOYKO: NEW PARASITIC ISOPODS FROM POLYNESIA 447 FiGURE 6. Cabirops bombyliophila sp. noy., male, 0.91 mm, CASIZ 170231, holotype. A, ventral view; B, lateral view; C, ventral view of segments 5 and 6 showing dentition, left side; D, ventrolateral view of segments 4-6, right side; E, ven- trolateral view of segments 2-7, left side; F, left pleopods 1 and 2. Scale = 200 um (A, B), 25 um (C, F), 50 um (D), 100 um (E). conical in shape, with faint indication of segmentation. Pleon with 4 semi-circular depressions on right side in a trapezoidal configuration (Fig. 8A). DIsTRIBUTION.— Found in marsupium of female Schizobopyrina bombyliaster sp. nov. ex Gnathophyllum americanum from Tonga; 0.6-2.7 m depth. EtTyMOLOGy.— The specific name refers to the finding of this hyperparasite in association with the parasite of a bumblebee shrimp of the genus Gnathophyllum. REMARKS.— The genus Cabirops now contains 17 species (six of these remain unnamed) and this likely represents only a fraction of the species that await description (Sassaman 1985; Boyko and Williams 2004). Sassaman (1985) reviewed the genus and indicated that it could be divided into two groups based on coxal plate dentition. Group one contains species with 2 or 3 denticles on 448 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 24 FIGURE 7. Cabirops bombyliophila sp. nov., male, 0.91 mm, CASIZ 170231, holotype. A, right pereopod 1; B, left pereopod 2; C, left pereopod 2; D, left pereopod 3; E, left pereopod 4; F, left pereopod 6. Scale = 10 tum (A, F), 20 pm (B, D), 25 um (C, E). coxal plates 1—5 and | denticle on coxal plates 6 and 7, group two contains species with 2 or 3 den- ticles on all coxal plates. Members of group one parasitize hosts of the subfamilies Pseudioninae or Orbioninae while members of group two are found on hosts of the subfamilies Ioninae and Bopyrinae. While the morphology of Cabirops bombyliophila sp. nov. places it in group one (exhibiting only one denticle on plates 6 and 7), the species is a parasite of a bopyrine host. Such discrepancy is not surprising considering recent studies on Cabirops species from the Bahamas have indicated two species with dentition patterns unlike those in either group (Boyko and Williams 2004). In addition, other characters such as the dactyli of pereopods | and 2 (bifid vs. simple) and internal apophysis on article 2 of antennae 2 (strong vs. weak) differ between groups. WILLIAMS AND BOYKO: NEW PARASITIC ISOPODS FROM POLYNESIA 449 Cabirops bombyliophila sp. nov. most closely resembles C. codreanui Bourdon, 1966, C. montereyensis Sassaman, 1985, and C. orbionei Bourdon, 1972 but can be distin- guished from these species based on the denti- tion pattern (3 denticles on coxal plate 1 in C. bombyliophila sp. nov. vs. two denticles on coxal plate 1 in C. codreanui, C. montereyen- sis, and C. orbionei). In addition, C. codreanui and C. montereyensis are parasites of pseudionines while C. orbionei is found on orbionines; however, due to the potentially FiGurE 8. Cabirops bombyliophila sp. nov., female, large number of undescribed Cabirops species, 2.36 mm, CASIZ 170232, allotype. A, view of right side, conclusions on the degree of host specificity Se eae dorsal pereonal lobes; B, view of left side. are probably premature. Cabirops bombylio- avs phila sp. nov. is further distinguished from C. montereyensis by exhibiting a blunt tip on the dactyli of pereopods 6 and 7 (bifid dactyli are present in C. montereyensis). Finally, the apophysis on arti- cle 2 of antennae 2 is strong in C. bombyliophila sp. nov. but weak in C. orbionei. Cabirops bom- byliophila sp. nov., C. montereyensis, and C. orbionei are apparently the only species in the genus to exhibit a setal comb on the dactyl of pereopod 3; however, as indicated by Sassaman (1985), this feature may have been overlooked in earlier descriptions. ACKNOWLEDGMENTS Dr. Bob Van Syoc (CASIZ) provided loan of specimens and kindly hosted a brief visit by CBB to CASIZ. Dr. Alain Crosnier (MNHN) is thanked for inviting CBB to examine material in the MNHN collections. We thank Ms. Lauren M. Schuerlein (Hofstra University) for her work on the final plates. Financial support from Hofstra University to JDW is greatly appreciated. LITERATURE CITED BOURDON, R. 1966. Sur quelques nouvelles especes de Cabiropsidae (Isopoda Epicaridea). Bulletin du Muséum National d'Histoire Naturelle, 2e Séries 38:846—868. BOURDON, R. 1972. Epicarides de la Galathea Expedition. Galathea Report 12:101—112. Bourbon, R. 1983. Expédition Rumphius II (1975). Crustacés parasites, commensaux, etc. (Th. Monod éd.). VIII. Crustacés isopodes (3e partie: épicarides Bopyridae). Bulletin du Muséum National d Histoire Naturelle, Paris (4) 5 (Section A, no. 3):845—-869. BOURDON, R., AND A.J. BRUCE. 1983. Six bopyrid shrimp parasites (Isopoda, Epicaridea) new to the Australian fauna. Crustaceana 45(1):96—106. Boyko, C.B., AND J.D. WILLIAMS. 2004. New records of marine isopods (Crustacea: Peracarida) from the Bahamas, with descriptions of two new species of epicarideans. Bulletin of Marine Science 74(2):353-383. Campos, E., AND A. ROSA DE Campos. 1990. Taxonomic remarks on Schizobopyrina Markham, 1985, with the description of S. bruscai (Crustacea: Isopoda: Bopyridae). Proceedings of the Biological Society of Washington \03(3):633-642. CHoprA, B. 1923. Bopyrid isopods parasitic on Indian Decapoda Macrura. Records of the Indian Museum 25(5):411—550, pls. 11-21. CHoprA, B. 1930. Further notes on bopyrid isopods parasitic on Indian Decapoda Macrura. Records of the Indian Museum 32:\13—147, pls. 4-6. DARDEAU, M.R., D.L. ADKISON, J.K. SHAW, AND T.S. HopKINs. 1980. Notes on the distribution of four caridean 450 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 24 shrimps (Crustacea: Decapoda) in the northeastern Gulf of Mexico. Florida Scientist 43(1):54—S7. GIARD, A., AND J. BONNIER. 1887. Contributions a l’étude des bopyriens. Traveaux de | ’'Institut Zoologique de Lille et du Laboratoire de Zoologie Maritime de Wimereux 5:\—272, pls. 1-10. KOSSMANN, R. 1872. Beitrage zur Anatomie der schmarotzenden Rankenfiissler. Arbeiten aus dem Zoologischen-Zootomisches Institut der Universitat Wiirzburg 1:97—137. KOSSMANN, R. 1881. Studien tiber Bopyriden. I. Gigantione moebii und Allgemeines iiber die Mundwerkzeuge der Bopyriden. Hl. Bopyrina virbii, beitrage zur Kenntnis der Anatomie und Metamorphose der Bopyriden. Zeitschrift der Wissenschaftlichen Zoologie 35:652—680, pls. 32-35. MARKHAM, J.C. 1985. A review of the bopyrid isopods infesting caridean shrimps in the northwestern Atlantic Ocean, with special reference to those collected during the Hourglass Cruises in the Gulf of Mexico. Memoirs of the Hourglass Cruises 8(3):1—156. RAFINESQUE-SCHMALTZ, C.S. 1815. Analyse de la nature ou tableau de l'univers et des corps organisés. Palermo. 224 pp. SASSAMAN, C. 1985. Cabirops montereyensis, a new species of hyperparasitic isopod from Monterey Bay, California (Epicaridea: Cabiropsidae). Proceedings of the Biological Society of Washington 98(4): 778-789. SHINO, S.M. 1939. Bopyrids from Kytisyt and Rytkyt. Records of Oceanographic Works in Japan 10(2): 79-99. SHINO, S.M. 1942. Bopyrids from the south sea islands with description of a hyperparasitic cryptoniscid. Palao Tropical Biological Station Studies 2(3):437-458. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 25, pp. 451-457, 3 figs., 1 table September 30, 2004 Taxonomic Status of Rhacophorus taronensis Smith, 1940 Jeffery A. Wilkinson!+ and Dingqi Rao? ! Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103; Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan, China 650223; 3H.T. Harvey & Associates, 3150 Almaden Expressway, Suite 215, San Jose, CA 95118 Polypedates gongshanensis (Yang and Su, 1984), described as endemic to the Gaoligong Mountains of western Yunnan Province, China, is demonstrated to be a junior synonym of Rhacophorus taronensis Smith, 1940, the latter known only from the type specimen collected in northern Myanmar. This poorly known species is revisited, and new data derived from comparisons of the original descriptions, exam- inations of the type specimens of R. faronensis and P. gongshanensis, and newly acquired specimens from Myanmar and China are presented. Smith (1940)! described a new species of Rhacophorus based on a single specimen (BMNH 1947.2.8.17) collected by R. Kaulback in the north of Myanmar (Patsarlamdam, Taron Valley; N 27°43’, E 98°10’) on 01 June 1939. He states that this new species (R. taronensis) is related to R. bimaculatus but differs by having less extensive webbing of the fingers, the absence of a dermal projection on the heel, and different coloration. He gives a snout-vent length of 78 mm and also presents a drawing of the right hand (Smith 1940, fig. 3). Since then, no new specimens of R. taro- nensis have been reported. Yang and Su (1984)? described a new species of Rhacophorus from the southern Gaoligong Mountains of western Yunnan Province, China. They compared their specimens only to Rhacoph- orus feae (now Polypedates feae fide Frost 2002), probably due to the similar brown line extend- ing from the tip of the snout to behind the tympanum that is present in both their specimens and R. feae. Their specimens differed from R. feae by the webbing of the first and second fingers being restricted to the base and only one-third webbing between the remaining fingers; the heels overlap- ping when the legs are at right angles to the body; and by the presence of creamy-colored spots on the flank and inner and outer sides of the thigh. Based on these differences, they concluded that these specimens represented a new species, R. gongshanensis, believed to be restricted to the ! Original description (Smith 1940): “Vomerine teeth in strong, almost transverse, groups, commencing at the inner margin of the choanae. Head broader than long, much depressed; snout rounded, scarcely projecting beyond the mouth, shorter than the eye; canthus rostralis distinct; loreal region oblique, feebly concave; nostril a little nearer the tip of the snout than the eye, nearly as broad as the interorbital space, broader than the upper eyelid; tympanum very distinct, 4/5 the diam- eter of the eye, close to it. Outer three fingers with very large discs, which are as broad as long and as large as the tympa- num; more than half webbed, the membrane reaching the disc of the 4th finger and that of the 2nd on the outer side; toes fully webbed, their discs smaller than those of the fingers; the tibio-tarsal articulation reaches to between the eye and the tip of the snout; subarticular tubercles of both fingers and toes strongly developed; a short oval inner metatarsal tubercle; no outer. Skin smooth above, that of the belly and hinder part of the thighs coarsely granulate. A dermal fringe, feebly dis- tinct, on the outer side of the fore-arm, tarsus and foot. Bluish above, green in life, with small black spots; lower parts whitish, thickly speckled with dark grey; hinder part of thighs black with white spots. From snout to vent 78 mm.” 45] 452 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 25 Gaoligong Mountains. Subsequently, Fei, Ye, and Huang (1990) placed R. gongshanensis in the genus Polypedates without explanation. In the process of identifying specimens of Rhacophorus collected in northern Myanmar in 2002 by the Myanmar Herpetological Survey Project (a collaborative project of the California Academy of Sciences, Smithsonian Institution, and the Nature and Wildlife Conservation Division, Forest Department, Myanmar), the holotype of R. taronensis (BMNH 1947.2.8.17, re-catalogued from BM 1940.6.1.39; Figs. 1A, C, E) was examined. It was immediately evident that the holotype was similar in appearance to specimens of P. gongshanensis from Baoshan Prefecture, Yunnan Province, China, collected by a California Academy of Sciences (CAS) and Kunming Institute of Zoology (KIZ) field expedition in 2003. METHODS AND MATERIALS In order to determine whether R. taronensis and P. gongshanensis are conspecific, the holo- types of R. taronensis and P. gongshanensis (KIZ 810485; Figs. 1B, D, F) were examined and com- pared to each other and to the original descriptions. In addition, they were compared with a paratype of P. gongshanensis (KIZ 810555) and newly collected specimens from Myanmar. Specimens recently collected in Myanmar are housed in the collections of the Department of Herpetology, CAS and the Myanmar Biodiversity Museum (MBM), Hlawga, Myanmar. Newly collected specimens from Baoshan Prefecture, China (GLGS field numbers in materials examined section below), are as yet not available for measurements, but eventually they will be housed in the ? Literal translation of original description in Chinese (Yang and Su 1984) — “The body is medium in size, the largest among the 6 male specimens is only 72.2 mm long, the only female specimen is 81.5 mm long. The head is slightly longer than wide in males, but wider than long in the female; the rostrum sharply angles downward from the nostril forward, the canthus rostralis is obvious, the loreal region slants slightly outward, the surface of the loreal region is concave; the nos- trils are located midway between the tip of the snout and eyes; the internarial distance is shorter than the interorbital dis- tance; the top of head is flat; the tympanum is elliptical and close to the eye, with a vertical diameter '> of the diameter of the eye, and close in size to the disc of the third finger; the back of the tongue has a deep notch; vomerine teeth are not curved into an arch, the lateral end is slightly beyond the anterior angle of the internal nares, slightly inclined caudally but does not pass the posterior edge of the internal nares. “The forearm of the male frog is strong, that of the female is slightly weaker, the length of the forearm and hand is slightly beyond half of the body. Fingers are flattened, and rank according to size from big to small, as 3, 4, 2, 1. The disc is wide and straight at terminal end, with a transverse groove at the tip. The disc of the first finger is small; the second, third and fourth fingers are approximately '/; webbed, the second finger is webbed to the subarticular tubercle, the third and fourth fingers are webbed to the distal end of the subarticular tubercle, the first and second fingers are only webbed at the base; the inner edge of the first finger and outer edges of the second and third fingers with obvious marginal webbing; the tuber- cles at the distal end of the third and fourth fingers are bigger than those at the proximal end; the inner edge of the first fin- ger is enlarged at the base in a transverse direction. The hind limbs are 1.5 times that of the body size, the specimen KIZ 810485 is 1.7 times, tibiotarsal articulation reaches to the middle or anterior edge of eyes, the left and right heels overlap; feet are shorter than the tibia (6 males), but reverse for the female; webbing on the toes are relatively well developed to the base of the disc, only the fourth toe webbed to the distal subarticular tubercle; the subarticular tubercles are small and round, inner metatarsal tubercle is small, flat and elliptical, no outer metatarsal tubercle. “The male frog is dotted with white asperities on the dorsal aspect of the body and hind limbs, but the female frog with fewer asperities than the male; dense flat areolation on the ventral surface, the areolation on the lower jaw, throat, and pectoral region is smaller than other areas, the front of the lower jaw of the female frog is smooth without tubercles, the supratympanic fold is linear but slopes slightly backward. “Tn life, the pupil is a horizontal ellipse, blue-black, iris is greenish yellow. Upper surface of head, body, and the four limbs is grass-green, with sparsely scattered small brown spots. The flank and inner and outer side of thigh with many creamy yellow spots, big or small, with dark reddish brown to purple edges. The canthus rostralis, lateral edge of eyelid, and supratympanic fold are light brown. The anterior half of the fingers and toes are violet on dorsum, the posterior half remains grass-green. The supracloacal region, from the tibiotarsal articulation to the base of the fifth toe, and outer side of forearm to the base of fourth finger with a yellowish fine linear dermal ridge. Ventral surface light brown to purple, with scattered deep dark spots.” WILKINSON AND RAO: STATUS OF RHACOPHORUS TARONENSIS SMITH, 1940 453 FiGURE |. Dorsal, ventral, and lateral views respectively of the holotype (BMNH 1947.2.8.17) of Rhacophorus taro- nensis (A, C, E) and the holotype (KIZ 810485) of Polypedates gongshanensis (B, D, F). 454 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 25 collection of the Department of Herpetology, CAS and the Department of Herpetology, KIZ. They have been included to report locality data. After tissues were removed from the recently collected specimens (Myanmar and China), the specimens were fixed in 10% buffered formalin before being transferred to 70% ethanol. Latitude and longitude were recorded with a Garmin 12 GPS (WGS84),. Sex was determined by the presence or absence of vocal apertures, or the presence or absence of eggs within the abdominal cavity. 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); head width (HW, width of head at its widest point); internarial distance (IND, distance between nares); interorbital distance (IOD, minimum distance between upper eyelids); upper eyelid width (UEW, maximum upper eyelid width); snout length (SL, from anterior border of eye to tip of snout); eye diameter (ED, horizontal length between orbital borders of eyes); distance from nostril to eye (DNE, from nostril to anterior border of eye); tympanum diameter (TD, greatest diameter); forelimb length (FLL, from elbow to tip of third fin- ger); first finger length (FFL, from palmer end of inner metacarpal tubercle to tip of finger); third finger length (TFL, from base point between third and fourth fingers to tip of 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 tuber- cle to tip of fourth toe); fourth toe length (FTL, from distal end of third subarticular tubercle to tip of toe); inner metatarsal length (IML, at point of greatest length); width of disk of third finger (3FDW, greatest horizontal width); and width of disk of fourth toe (4TDW, greatest horizontal width). RESULTS COMPARISON OF TYPES.— The holotype of R. faronensis (female) is similar to the holotype of P. gongshanensis (male) in shape, color, and pattern. However, the holotype of R. taronensis is longer (SVL 76.6) than the holotype of P. gongshanensis (SVL 70.5), the head is wider than long (versus longer than wide in holotype of P. gongshanensis), the nostrils are nearer to the tip of the snout than the eyes (versus midway between eye and tip of snout in holotype of P. gongshanen- sis), and the third finger is longer with a wider disc (versus shorter with a narrower disc in holo- type of P. gongshanensis). The SVL is also longer and the head wider than long for the female paratype of P. gongshanensis (CIB 810723; referred to as the allotype in Yang and Su) and a female specimen (CAS 224382) in this study, indicating that these are sexually dimorphic characters. The length of the third finger and width of the disc of third finger are not mentioned for the female paratype of P. gongshanensis in Yang and Su (1984) but in the aforementioned female specimen in this study are similar to the holotype for R. taronenesis and different from the male specimens (Table 1), indicating that these may also be sexually dimorphic characters. The relative location of the nostrils was also nearer to the tip of the snout in the aforementioned female specimen and most of the male specimens, but closer to the eyes in two male specimens (KIZ 810555, CAS 224371) suggesting that this character is not sexually dimorphic. To detect whether other species of Rhacophorus also possess the aforementioned apparent sex- ually dimorphic characters, specimens of R. owstoni were measured for SVL, HL, HW, TFL, and 3FDW. All female specimens (N = 14) were larger (SVL x = 58.93, o = 4.13) than the male spec- imens (N = 12; SVL x = 46.86, o = 2.21). Also, several but not all female specimens possessed a head slightly wider (HW x = 22.46, o = 1.56) than long (HL x = 22.28, o = 1.13), whereas, all male specimens possessed a head slightly longer (HL x = 17.92, o = 0.80) than wide (HW x= 17.47, o = 0.76). Also, the length of the third finger and the width of the disc of the third finger were on average respectively longer and wider in the female specimens (TFL x = 9.45, o = 0.73, WILKINSON AND RAO: STATUS OF RHACOPHORUS TARONENSIS SMITH, 1940 455 TABLE |. Measurements of the holotype and newly collected specimens of Rhacophorus taronensis and the holotype and a paratype of Polypedates gongshanensis. Top values for holotypes are original measurements in Smith (1940) and Yang and Su (1984). sex SVL HL HW IND 10D UEW SL ED DNE TD FLL FFL TFL HAL THE TIL FL FIL IML 3FDW 4TDW BMNH 1947.2.8.17 78 (holotype) Be 76.6) 244268964271 Sion 07 916 62.) 516 <42/6N 187 1916. 26.3) 341011 39.7) 350). WZ aa sigumnls'6 CASPIAS SR Ee 73U1E 245) 2535 5:9) wi8t4e 72 11 83 64 Sen, 09240 ei lie Seine 3 920) 830.48 153 opel 2 panes 3) 5.9 4 Average AO RDA me? OS lime OY Leet a1 one 0:45 eel OO 8.9 sm O:S pS Siem le ees SeeelO Si N26) | STLA eS Sal S4r4 mile eoy nese S 5.8 3.8 Ratio of SVL SPe OA Om cel LOS ies ton lA Gy UE G AS teens SS ieee 8) 24.4034 2849:0 5050198 245190 23! seaton 7.7 5.1 KIZ 810485 OO 25245781 85) 96:8 ae NS) 49 40 PB eS e2 34 5 (holotype) iO 232) 2321 0:46) 0:45 1259 8265 Sto 515) 4 0A D2 216:8) 25:3ih 35212 34165 3379) Hy -3 3 5 3.8 CAS 2245s Of tin 24-2 9228 9 O:0e 6:8) 457) eS 729) Sofi) S350 SP 142% 20:6) 6 32'69929'S) 127s N13 130 33 4.3 37, CRSA me Vig O22 le 2027) OS OsSenyoro)n OSs 70) 159 AS G42 ONS es S'S) 2087 362926) 2710) il 3a) 22.6 3.7 71st KIZ 810555 (paratype) MERGES ZR6n 21S 655 1619e= 1551 LOR SoS Zero wmlON melts 3.0 21S 329M ese lyan 29MM Seok mS 4.5 3.3 CBES274579 se Mia60:00 213 19:85 5:5) io) 553), 1929 8 SY UT) BIL © CLO) = ISAS UIA Seah MEKOG YE ieee WMIER iT I O)s hg) 3.1 MBM-JBS 11793 Il GUS) 20 GI EES Oy SENOS OP yt lsy | Cn) Silas Rasy NIA MS Ye shO espe Azsalban PAO ai lek b as Page) 3.6 2.4 CAS 224373 M 59.7 20:9 20:3 5.2 5.8 5 OW)" 70 es AS B31 9:9F NS 55 204293 0329) 8a 2:4 il 4 lia De S 3.7 3 CAS 224372 Mi 92.6205). 1 953) 74:8) S64 Vr46 1916) —2659) 5:45 45) | 30S 1919), 122) 188299 ND 7602578) halls: WO 3.4 AY) CAS 224370 M 57.4 20.1 19.7 5 5:9 416 94 16:4) 5:9) CAT SOR S29 Ome S13 iy pS tOINE 2 9 iON 2.6. Sexe 27 lia mS 95:4" 16:6)" Sil LO TA 5351 7457, 33:60 0) 356 2057) SIA 29'S)= 2725) 337) 226 3.9 3 Ratio of SVL S416) 33:3) 357) OM 1853" 6:2) 12) 89476) 54.6) G28 22 1) 733'6) col aos 4a 22.2) eA" 2 6.3 4.9 ratio to SVL 0.20; 3FDW x = 2.59, o = 0.32, ratio to SVL 0.06) than male specimens (TFL x = 12.70, o = 1.16, ratio to SVL 0.22; 3FDW x = 3.88, o = 0.47, ratio to SVL 0.07), though these dif- ferences were more pronounced in the larger females, indicating that they may merely be ontoge- netic. Most of the characters given in the description by Yang and Su (1984) and Smith (1940) are consistent with both holotypes of R. taronensis and P. gongshanensis, with the following excep- tions: (1) the position of the tibiotarsal articulation and the amount of webbing between the fingers for both the holotypes of R. taronensis and P. gongshanensis differ from the description by Yang and Su (1984) and more closely resemble the description in Smith (1940) in that the tibiotarsal articulation reaches to between the eye and end of the snout and the fingers are more than '/ webbed; (2) the relative size of the tympanum and amount of webbing on the toes in both the holo- types of P. gongshanensis and R. taronensis differ from the description by Smith (1940) and more closely resemble the description in Yang and Su (1984) in that the tympanum is only slightly over half the diameter of the eye (Table 1) instead of “th, and the toes are not fully webbed but reach only to the distal subarticular tubercle on the fourth toe; and (3) the holotype of R. taronensis is smaller than the reported size of the female paratype of P. gongshanensis by 3.5 mm (from Smith, 1940) or 4.9 mm (measurements taken in this study). Measurements of all specimens examined are given in Table 1. These measurements differ slightly from those given in Yang and Su (1984) for the holotype of P. gongshanensis (greatest dif- 456 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 25 ference being 1.7 mm for hand length), and given in Smith (1940) for the holotype of R. taronen- sis (a smaller snout-vent length in this study). Measurements of the holotype of R. taronensis and the holotype and a paratype of P. gongshanensis are generally smaller than those reported in Yang and Su (1984) and Smith (1940). This may be due to shrinkage of the type specimens between measurements or different standards used in measuring. The nine specimens newly collected from Myanmar are on average smaller than those collected by Yang and Su (1984). However, the ratios of average measurements of each structure to average snout-vent length for the males are similar between the Yang and Su (1984) study and this study. VARIATION.— Further observations were made on spec- imens newly collected from Myanmar. On several specimens (CAS 224370-224371, 224382, 224397, MBM-JBS _ 11793; Joseph B. Slowinski field num- ber to be deposited in the MBM) the spotting on the flanks is white, more reticulate, and con- tinues onto the ventral surface (Fig. 2). The brown spotting on the back may be very extensive (CAS 224371, 224377, MBM- - JBS 11793) (Fig. 2) or absent BCURe 2. From a color transparency of Rhacophorus taronensis (MBM- (CAS 224369-224370, 224379), 7° ''”””) The discs on the fingers of the females are relatively larger than those of males but the snout lengths are relatively shorter. The rostrum of some specimens (CAS 224371—224372, 224377) is spatulate in form, from the nostril to the tip of the snout. All specimens have two strong brown dor- solateral lines that bifurcate at the tip of the snout and continue along the canthus rostralis, lateral edge of eyelid, supratympanic fold, and terminating behind the insertion of the forelimb (Fig. 2). The preceding data demonstrate that the two types are the same species. Variation between the types and the other specimens in this study can be attributed to sexual dimorphism and normal ranges of differences within a species. Polypedates gongshanensis is, thus, a subjective junior syn- onym of R. taronensis. DISTRIBUTION.— Based on records for the type specimens and recent collections in both China and Myanmar, at present, R. faronensis occurs only in the mountainous regions of the northern por- tion of Kachin State, Myanmar and to the south in Baoshan Prefecture, Yunnan Province, China (Fig. 3). Fei (1999) gives a distribution for P gongshanensis extending from Baoshan Prefecture north along the Chinese side of the China-Myanmar border into Gongshan County, Nujiang Prefecture, indicating that this species occurs throughout the border regions of China and Myanmar. However, this distribution has not been confirmed by voucher specimens or recent sur- veys in these areas. MATERIAL EXAMINED.— BMNH 1947.2.8.17, holotype of R. taronensis, Myanmar, Patsar- lamdam, Taron Valley; 27°43'N, 98°10'E; CAS 224369-224372, MBM-JBS 11793, Myan- mar, Kachin State, Putao District, Nagmung Township, Hkakabo Razi National Park, Nga War Village, 27°49'44.1"N, 97°45'59.3"E; CAS 224377, 224379, 224382, Myanmar, Kachin State, Putao District Nagmung Town-ship, Hkakabo Razi National Park, Nga War Village, 27°49’04.1"'N, 97°46'41.3”E; KIZ 810485, holotype of Polypedetes gongshanensis, and 810555, paratype, China, WILKINSON AND RAO: STATUS OF RHACOPHORUS TARONENSIS SMITH, 1940 457 Yunnan Province. Baoshan Prefecture, Pumanshao: Gaoligongshan (GLGS) field numbers to be deposited in the CAS and KIZ, GLGS 1038-1039, 1047, 1049-1050, 1059-1063, 1065-1068, 1070-1079, China, Yunnan Province, Baoshan Prefecture, Tengchong County, Xiao Di Fang Village, 24°52'12.2"N, 98°45'13.4"E; Rhacoph- orus owstoni, CAS 211426—211447, 211449- 211452, Japan, Okinawa Prefecture, Ryukyu Islands, Iriomote Island. 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 Survey Project. We thank Htun Win, Thin Thin, Kyi Soe Lwin, Awan Khwi Shein, and Hla Tun of the Myanmar Herpetological Survey Field Team for their efforts in collecting, and personnel of the Baoshan Nature Reserve for their help during the joint 2003 CAS/KIZ expedition. We also thank the British Museum for loaning and Edwin N. Arnold for hand We Ee ae carrying the type specimen. Fieldwork was sup- Ficure 3. Map of known localities of Rhacophorus ported by a National Science Foundation Grant ‘aronensis based on specimens used in this study. The (DEB-9971861) to Joseph B. Slowinski and type localities for R. taronensis and P. gongshanensis x : i are indicated by the numbers | and 2 respectively. George R. Zug, a National Science Foundation Grant (DEB-0103795) to Joseph B. Slowinski and Peter W. Fritsch, and a National Geographic Society Grant (7340-02) to Peter W. Fritsch. Dong Lin assisted in creating Figure 1, Hla Tun pro- vided Figure 2, and Michelle S. Koo provided Figure 3. Lihua Zhou provided the translation of the Chinese text. Jens V. Vindum, Guinevere O.U. Wogan, and George R. Zug critically read and pro- vided valuable comments on the manuscript. LITERATURE CITED Fel, L. 1999. Atlas of Amphibians of China. Publishing House for Scientific and Technological Literature, Hunan. 432 pp. Fel, L., C-G YE, AND Y-Z. HUANG. 1990. Key to Chinese Amphibians. Publishing House for Scientific and Technological Literature, Chongqing. 364 pp. Frost, D.H. 2002. Amphibian Species of the World: an online reference V2.21 (July 15, 2002). Electronic database available at: http://research.amnh.org/herpetology/amphibian/index.html. SmitH, M.A. 1940. The amphibians and reptiles obtained by Mr. Ronald Kaulback in upper Burma. Records of the Indian Museum 42:465-486. YANG, D-T., AND C-G. Su. 1984. Rhacophorus gongshanensis, a new species of flying frog from the Hengduan Mountains. Acta Herpetologica Sinica 3:51—53. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. 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The Taxonomic Status of the Genus Nesionixalus Perret, 1976 (Anura: Hyperoliidae), Treefrogs of Sao Tomé and Principe, with Comments on the Genus Hyperolius. DAviID W. BEHRENS AND ANGEL VALDES: A New Species of Dendrodoris (Mollusca: Nudibranchia: Dendrodorididae) from the Pacific Coast of North America TERRENCE M. GOSLINER AND HANS W. BERTSCH: Systematics of Okenia from the Pacific Coast of North America (Nudibranchia: Goniodorididae) with Descriptions of Three New Species YOLANDA E. CAMACHO-GARCIA AND TERRENCE M. GOSLINER: A New Species of Okenia (Gastropoda: Nudibranchia: Goniodorididae) from the Pacific Coast of Costa Rica JASON D. WILLIAMS AND CHRISTOPHER B. BOYKO: A New Species of Schizobopyrina Markham, 1985 (Crustacea: Isopoda: Bopyridae: Bopyrinae) Parasitic on a Gnathophyllum Shrimp from Polynesia, with Description of an Associated Hyperparasitic Isopod (Crustacea: Isopoda: Cabiropidae) JEFFERY A. WILKINSON AND DINGQi RAO: Taxonomic Status of Rhacophorus taronensis Smith, 1940