a } \yY l SEES SEN (8 Gea pes 30, 2004 * VOLUME 55 * NUMBERS 26-35 7 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES CALIFORNIA ACADEMYorSCIENCES 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 Following the devastating earthquake and fire that engulfed San Francisco and the Academy on 18 April 1906, the Academy found temporary quarters to house what little it had rescued from the ruins of its once proud Market Street museum. Its library, once the largest natural history library in the United States west of Saint Louis had, like nearly all of its biological collections, disappeared in the conflagra- tion. Although the significance of the loss of the library can never be fully appreciated, there came such an outpouring of generosity from institutions and individuals worldwide that its library soon regained much of its former luster. The post-earthquake library, which is shown in the cover illustration, was housed in a large Gough Street home, which itself had come through the events of 18 April relatively unscathed. The names of institutions, individuals, and, yes, even book dealers, who opened up their inventories, at no cost to the struggling Academy, were duly recorded in the early recovery-period records of the institution and on bookplates that were affixed to the books received (see also Leviton & Aldrich, Theodore Henry Hittell’s The California Academy of Sciences, 1853-1906. California Academy of Sciences, San Francisco, 1997, pp. 561-563). ISSN 0068-547X ‘The Proceedings of the Califernia rbcademy of Setenced accepts manuscripts for publication in the Natural Sciences that stress systematics and evolutionary biology and paleontology, biogeography, and biodiversity, also relat- ed areas in the Earth Sciences, such as biostratigraphy, regional and global tectonics as they relate to biogeography, and paleoclimatology, and topics in astrobiology, anthropology, as well as the history of science as they relate to insti- tutions of natural history, to individuals, and to activities, such as expeditions and explorations, in the natural sciences. Priority consideration will be given to manuscripts submitted by Academy staff and others who are affiliated with the research programs of the institution and/or manuscripts that include reference in part or in whole to Academy research collections or to Academy-sponsored expeditions. Others who do not meet the above criteria may submit manuscripts for consideration for publication on a space-available basis. Manuscripts accepted for publication are subject to page charges; charges may be waived on a case-by-case basis. Note: All manuscripts submitted for publication in any of the Academy's scientific publication series (Proceedings, Occasional Papers, Memoirs) are subject to peer review. Peer review includes both internal and exter- nal review, internal review by at least one Academy scientist whose interests parallel those of the submission, and external review, ordinarily by two individuals who are recognized experts in the field. Published by the California Academy of Sciences 875 Howard Street, San Francisco, California 94103 U.S.A. Printed in the United States of America by Allen Press Inc., Lawrence, Kansas 66044 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26, pp. 459-497, 12 figs. December 30, 2004 *Cephalaspidean” Heterobranchs (Gastropoda) from the Pacific Coast of Costa Rica Angel Valdés! and Yolanda E. Camacho-Garcia2 ! Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, USA; Email:avaldes@nhm.org; * Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco, California. 94103, USA; Museo de Zoologia, Escuela de Biologia, Universidad de Costa Rica, San Pedro de Montes de Oca, San José, Costa Rica; Email: ycamachogarcia@calacademy.org Based on phylogenetic evidence, Mikkelsen (1996) demonstrated that the traditional taxon “Cephalaspidea” is paraphyletic. As a consequence of this new phylogenetic hypothesis, the Acteonidae, Bullinidae, and Ringiculidae were removed from the traditional Cephalaspidea. Furthermore, the Acteonidae was removed from the Opisthobranchia and included in an unresolved group named “Lower Heterobranchia.” The Bullinidae and Ringiculidae remained in the Opisthobranchia, but were included in another unresolved group, the “Architectibranchia.” After the exclusion of these taxa, the rest of the Cephalaspidea remained as a monophyletic group. Members of the paraphyletic traditional Cephalaspidea are here referred to as “cephalaspidean” heterobranchs. Despite the fact that “cephalaspidean” heterobranchs are paraphyletic, it is still practical to deal with them in monographic treatises due to morphological similarities (presence of large, external shells in most groups), shared habitats and life modes, and consistency with histor- ical treatments. Little is know about the “cephalaspidean” heterobranch biodiversity in Costa Rica. Studies by Houbrick (1968), Robinson and Montoya (1987), Hgiszter, (1998), and more recently by Espinosa and Ortea (2001), provided a few records from the Caribbean and Pacific coast, but included no morphological comparisons for most taxa. The purpose of the present study is to provide an up-to-date catalogue of species of “Lower Heterobranchia,” “Architectibranchia,” and Cephalaspidea found along the Pacific coast of Costa Rica, including descriptions of a new species and a systematic review of the generic placement and synonymy of several other species previously described, based on new anatomical evidence. MATERIAL AND METHODS All specimens studied, including the type material, are deposited at the Instituto Nacional de Biodiversidad, Costa Rica (abbreviated INB), the Natural History Museum of Los Angeles County (abbreviated LACM), the California Academy of Sciences, San Francisco (abbreviated CASIZ), The Natural History Museum, London (abbreviated BMNH), and the National Museum of Natural History, Washington D.C. (abbreviated USNM). The material examined is labeled as “shell” for dried shells (with or without remaining tissue), and “specimen” for complete wet specimens includ- ing shell and soft parts. Taxa have been arranged according to a classification system based on the phylogenetic hypothesis by Mikkelsen (1996). All wet specimens available were dissected and morphological examination was facilitated by 459 460 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 breaking the shell. The internal features were examined and drawn using a dissecting microscope with a camera lucida. Dried shells with remaining tissue were re-hydrated and broken to dissect the radula and other hard parts. The buccal mass was removed and dissolved in 10% sodium hydrox- ide until the radula was isolated from the surrounding tissue. The radula was then rinsed in water, dried, and mounted for examination with a Scanning Electron Microscope (SEM). Gizzard plates were dissected and mounted for examination with a SEM. SPECIES DESCRIPTIONS “Lower Heterobranchia” Family Acteonidae d’Orbigny, 1843 Genus Acteon de Monfort, 1810 Acteon traskii Stearns, 1897 (Figs. 1A—D) Acteon traskii Stearns, 1897:14. TYPE MATERIAL.— Acteon traskii: HOLOTYPE: Spanish Bight, San Diego, California (USNM 2506). MATERIAL EXAMINED.— Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 27 m depth, | shell (CASIZ 170722). 1 mile offshore, between Bahia Elena and Bahia San Juanillo (10°57'20"N, 85°46'08"W), Provincia Guanacaste, 25-53 m depth, 14 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-12.43). From 1.75 miles west of Punta Descarte to 1.25 miles north of start, off Bahia Santa Elena (11°02'23"N, 85°48'05"W), Provincia Guanacaste, 55 m depth, 14 February 1972, | shell, leg. P. LaFollette and D. Cadien (LACM 72-15.15). 5 miles north west of Islas Huevos (10°41'45”N, 85°46'25"W), Golfo de Papagayo, Provincia Guanacaste, 60-64 m depth, 18 February 1972, | juvenile shell, leg. P. LaFollette and D. Cadien (LACM 72-34.25). Middle of Bahia Huevos, north of Bahia Culebra (10°38’41"N, 85°41'55”W), Provincia Guanacaste, 10—23 m depth, 18-19 February 1972, | juvenile shell, leg. P. LaFollette and D. Cadien (LACM 72-35.39). Off beach at Bahia Ballena (9°44'12”N, 84°59'32”W), Provincia Puntarenas, 3-18 m depth, 23 February 1972, 1 juvenile shell, leg. P. LaFollette and D. Cadien (LACM 72-45.37). Off Bahia Herradura (9°38'50"N, 84°40'50” W), Provincia Puntarenas, 37 m depth, 10 March 1972, | juvenile shell, leg. J. McLean (LACM 72-54.50). Isla David (10°57'51.5100N, 85°42'36.9080W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 39 m depth, 2 March 1996, 3 shells, leg. Y. Camacho (INB0003747064). Isla David (10°58'05.5000N, 85°42'38.6000W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 30 m depth, 12 March 1996, 1 shell, leg. Y. Camacho (INB0003747061). 800 meters from Isla Despensa, Murciélago (10°59'45.3350N, 85°44'16.2441W), Provincia Guanacaste, 40 m depth, 9 April 2002, 1 shell, leg. F. Gonzalez (INB0003539758). ADDITIONAL MATERIAL EXAMINED.— MExico: Bahia Chamela, Jalisco, 27-72 m depth, | shell, 16 February 1938, leg. G. Willett (LACM 153410). Bahia Tangola-Tangola and Bahia Santa Cruz (15°45'00"N, 96°06'12"W), Oaxaca, 18-37 m depth, 28 February 1934, 2 shells with body parts (LACM 34-133.26). COLOMBIA: Bahia Choco (3°33'N, 77°38-39'W), southwest of Buenaventura, 80 m depth, 16 September 1966, 1 shell (LACM 66-201.8). SHELL MORPHOLOGY.— Length 12 mm: width 5 mm, in the largest specimen examined from Costa Rica. Shell solid, elongate, with convex sides (Fig. 1A). Body whorl large, from *4 to “s of the shell length. Spire short, conical, with 3-4 short whorls. Suture slightly channeled. Protoconch globose, about 1.5 whorl, 500 um in diameter (Fig. 1B). Umbilicus absent. Aperture long, about */s of the body whorl length, wide anteriorly, narrowing abruptly about its half-length. Anterior end of the aperture expanded into a small lip that is more conspicuous near the middle of the shell. Columellar margin thickened, slightly oblique, with a small, simple fold. Sculpture composed of a VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 461 number of punctured spiral grooves (Fig. 1D). Punctuations oval, fused to each other within each groove, more conspicuous near the anterior end of the shell. Grooves separated by gaps several times as wide as the grooves. Color cream to reddish, with a white band on the posterior end of the whorl. Some specimens with the reddish pigment faded out or completely disappeared. ANATOMY.— Radular formula 121 x 133.0.133 in a specimen from Mexico (LACM 34- 133.26). Radular teeth all similar in shape and size, with an elongate base and a curved cusp bear- ing 15—17 elongate denticles (Fig. 1C). GEOGRAPHIC RANGE.— Recorded from Southern California to Panama (Keen 1971). The present paper includes the first record from Colombia. REMARKS.— Acfeon traskii was described based on Holocene material from southern California (Stearns 1897). Living animals have been reported from southern California to Panama (Keen 1971) and Marcus (1972) provided descriptions of the radula and penial anatomy. This species can be distinguished from Rictaxis punctocaelatus (Carpenter, 1864) by having a more fragile, reddish shell with a single white band on the posterior end of the whorl. In R. punctocae- latus the shell is dark grey to black with three white bands on the anterior end, middle region, and posterior end of the whorl. The radula of R. punctocaelatus described by Marcus (1972) contains a much smaller number of teeth, which are larger and vary in shape along the half-row, from short multidenticulate inner teeth to very elongate outer teeth. Gosliner (1996) studied one specimen from the Santa Barbara Basin (Southern California) with the shell and radular morphology identical to the specimens here examined. It is most likely that Gosliner’s material belongs to the same species. Acteon panamensis Dall, 1908 is a much shorter and fragile species originally described from deep waters in Panama (Dall 1908). Rudman (1971) proposed that Acfeon should be used only for species with a columellar fold and numerous minute hook-shaped teeth. Other genus names are available for species with no col- umellar fold or radulae with a reduced number of teeth. The anatomy of numerous species and gen- era of Acteonidae has not been described, so it is not possible to determine their generic placement or synonymy. Acteon traskii is placed in the genus Acfeon because of the presence of a wide radula with numerous short teeth, similar to that of Acteon tornatilis (Linnaeus, 1758), the type species of the genus. For a description of the radula of A. tornatilis see Thompson (1976). Genus Crenilabium Cossmann, 1889 Crenilabium venustus (d’Orbigny, 1840) (Figs. 1E-G) Acteon venustus d’Orbigny, 1840:399, pl. 56, figs. 4-6. ? Daphnella casta Hinds, 1844:25. TYPE MATERIAL.— Acteon venustus: type material untraceable. Daphnella casta: HOLOTYPE: Golfo de Nicoya, Costa Rica (BMNH 79.2.26.89). MATERIAL EXAMINED.— Five miles north west of Islas Huevos (10°41'45”N, 85°46'25”W), Golfo de Papagayo, Provincia Guanacaste, 60-64 m depth, 18 February 1972, 3 shells, 1 fragment, leg. P. LaFollette and D. Cadien (LACM 72-34.26). South of Punta Uvita to Isla Ballena (9°07'36"N, 83°44'30”W), Provincia Puntarenas, 18 m depth, 13 March 1972, 1 fragment, leg. J. McLean (LACM 72-61.22). ADDITIONAL MATERIAL.— Intertidal drift shells in crab colony, Fort Amador Beach (8°56'N, 79°33'W), Canal Zone, Panama, 14 March 1970, 1 shell, leg. A. Marti (LACM 70-85.1). SHELL MORPHOLOGY.— Length up to 12 mm; width up to 4 mm in the largest specimen exam- 462 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 FIGURE 1. A-D, Acteon traski Stearns, 1897; (A) Shell morphology, specimen from Isla Despensa (INB0003539758), 6 mm long; (B) Protoconch, specimen from Isla David (INB0003747061); (C) Radular teeth, specimen from Bahia Tangola- Tangola, Mexico (LACM 34-133.26); (D) Sculpture, specimen from Isla David (INB0003747061). E-G, Crenilabium venustus (d’Orbigny, 1840); (E) Shell morphology, specimen from Amador Beach, Panama (LACM 70-85.1), 9 mm long; (F) Protoconch, specimen from Punta Uvita (LACM 72-61.22); (G) Sculpture, same specimen. ined. Shell solid, very elongate (Fig. 1E). Body whorl long, between */—% of the shell length; spire conical, elongate, with 2—3 whorls. Suture narrowly channeled. Protoconch elongate, about 1.5 whorl, 600 um in diameter (Fig. 1F). Umbilicus absent. Aperture narrow and short, about *4 of the body whorl. Columellar margin with a conspicuous denticle. Sculpture composed of a number of punctured spiral grooves (Fig. 1G). Punctuations conspicuous, oval, situated next to each other within each groove. Grooves separated by gaps with about the same width as the grooves. Color uniformly whitish. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 463 ANATOMY.— All the specimens examined consisted of empty shell so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— Reported from Costa Rica to Peru (Keen 1971). REMARKS.— Acfeon venustus d’Orbigny, 1840 was originally described from Paita, Peru (d’Orbigny 1840). This species is characterized by having a narrow and elongate shell, with an elongate spire, a relatively short aperture, and a strong sculpture composed of conspicuous punc- tuations. There is no information on the anatomy of this species that was described on the basis of empty shells. The material here examined from Costa Rica and Panama matches the original description and it is clear that it belongs to the same species. Because there are no anatomical data, it is not possible to provide a positive taxonomic place- ment for this species. Bouchet (1975) provided anatomical and conchological evidence for main- taining the fossil genus Crenilabrum as valid, those include a radula with lateral teeth provided with a long and sharp cusp and a rachidian tooth, and a shell with a very elongate spire and com- paratively short aperture. The type material of Acteon venustus, as well as the specimens here examined, resembles the conchological features of Crenilabrum. Thus, this species is here tenta- tively transferred to Crenilabrum until complete specimens become available for study. Daphnella castus (Hinds, 1844) was originally described as a member of the Turridae, and later transferred to Acteon by Keen (1971). The holotype of this species (BMNH 79.2.26.89) is par- tially broken, but the general outline of the shell is very similar to that of Crenilabrum venustus and these two names are most likely synonyms. Genus Rictaxis Dall, 1871 Rictaxis punctocaelatus (Carpenter, 1864) Tornatella punctocaelata Carpenter, 1864:646. Remarks.— This species has been reported in the literature from Alaska to Baja California (Keen 1971; Skoglund 2002). The LACM collection holds one lot containing two shells collected alive from Isla San Miguel, Panama (LACM 157893). No specimens have been collected so far from Costa Rica but it is likely that this species is present in this area. Rictaxis punctocaelatus is characterized by having a solid and elongate shell with a black or dark gray background color and three spiral white bands on the anterior and posterior ends of the spire as well as in the middle region. The radula and external morphology of this species was described by Gosliner (1996). Opisthobranchia Cephalaspidea Family Cylichnidae H. and A. Adams, 1854 Genus Cylichna Lovén, 1846 Cylichna atahualpa (Dall, 1908) (Figs. 2A-C) Cylichnella atahualpa Dall, 1908:243, pl. 11, fig. 2. TYPE MATERIAL.— Cylichnella atahualpa: HOLOTYPE: Gulf of Panama (USNM 123081). MATERIAL EXAMINED.— Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 22 m depth, 1 shell (CASIZ 170721). Bahia Chatham (5°33’20’"N, 86°59'10”W), Cocos Island, 26 m depth, 13 January 1938, 11 464 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 shells (LACM 38-179.4). Golfo Dulce (8°24’20"N, 83°13'40"W), Provincia Puntarenas, 35-88 m depth, 26 March 1939, | shell (LACM 39-44.23). One mile offshore, between Bahia Santa Elena and Bahia San Juanillo (10°57'20"N, 85°46'08”W), Provincia Guanacaste, 25—53 m depth, 14 February 1972, 7 shells and fragments, leg. P. LaFollette and D. Cadien (LACM 72-12.45). Half mile to one mile and half west of Roca Vagares, Bahia de San Juanillo (10°57'28”"N, 85°45'20"W), Provincia Guanacaste, 37 m depth, 14 February 1972, 2 shells, leg. D. Cadien and P. LaFollette (LACM 72-13.33). Five miles north west of Islas Huevos (10°41'45"N, 85°46'25"W), Golfo de Papagayo, Provincia Guanacaste, 60-64 m depth, 18 February 1972, 10 shells, leg. P. LaFollette and D. Cadien (LACM 72-34.28). Off Bahia Herradura (9°38'50”N, 84°40'50”W), Provincia Puntarenas, 37 m depth, 10 March 1972, 4 shells, leg. J. McLean (LACM 72-54.52). Off Isla del Cafio (8°45'N, 83°54’W), Provincia Puntarenas, 55 m depth, 16-17 March 1972, 6 shells, leg. J. McLean (LACM 72-66.40). SHELL MORPHOLOGY.— Length up to 12 mm; width up to 3 mm in the largest specimen exam- ined. Shell solid, elongate, with nearly parallel sides (Fig. 2A). Only one whorl visible, forming nearly the entire shell. Apex rounded, umbilicated, with the aperture lip rising from the right side. Anterior end of the shell rounded, slightly flattened in some specimens. Aperture as long as the shell, wider anteriorly and narrowing gradually at about '/ of its length. Columellar margin con- spicuously thickened. Umbilicus absent. Sculpture with a number of irregular spiral lines more conspicuous near the anterior and posterior ends of the shell. Color dirty white, covered with a pale brown or yellowish periostracum. Spiral lines near the anterior and posterior borders of the shell pale brown. Apex and columella opaque white. ANATOMY.— The digestive system contains three smooth gizzard plates similar in shape and size. All the plates are oval and elongate (Fig. 2C). The radular formula is 12 x 4.1.1.1.4 in a spec- imen from Golfo Dulce (LACM 39-44.23). The rachidian teeth are broad with a number of sharp denticles decreasing in size towards the laterals of each plate. The innermost lateral teeth are hook- shaped, with a long and strong cusp with numerous minute denticles (Fig. 2B). The four outermost teeth of each row are much smaller, but also hook-shaped with a long and thin cusp and several small denticles. REMARKS.— Cylichna atahualpa was described based on shells collected at 590 m depth from the Gulf of Panama (Dall 1908). The original description included an illustration of an elongate shell with no umbilicus and the columellar margin conspicuously thickened. The color of the shell was described as white with a pale yellowish periostracum that shows reddish brown in the incised sculpture. The material here examined is from shallower waters but fits the original description and characteristics of the holotype of Cylichna atahualpa. This species was originally described as a member of Cylichnella, but the anatomy of the spec- imens examined, with three oval and smooth gizzard plates, and a radula with several lateral teeth, are typical of species of Cylichna. Other species assigned to Cylichna, also described or reported from the Panamic Province, are Cylichna luticola (C.B. Adams, 1852), Cylichna inca (Dall, 1908), and Cylichna pizarro (Dall, 1908). According to the illustrations provided by Keen (1971), these species are proportionally shorter and wider than C. atahualpa and lack conspicuous spiral lines on the shell, except for C. pizarro. Cylichna diegensis (Dall, 1919), originally described from San Diego, California, and with a geographic range extending from Santa Monica to Baja California, has a similar shell morphology to C. atahualpa. The anatomy of C. diegensis was described by Gosliner (1996), and it differs from that of C. atahualpa in several regards. The inner lateral teeth of C. diegensis bear 9-11 large den- ticles under the main cusp, whereas in C. atahualpa there are numerous minute denticles. Additionally, the outermost teeth of C. diegensis are smooth, whereas in C. atahualpa they have numerous denticles. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 465 Genus Acteocina Gray, 1847 Acteocina infrequens (C.B. Adams, 1852) (Figs. 2D-H) Bulla (Tornatina) infrequens C.B. Adams, 1852:214, 319. Acteocina magdalenensis Dall, 1919:296. Acteocina angustior Baker and Hanna, 1927:124—125, pl. 4, fig. 5. TYPE MATERIAL.— Bulla (Tornatina) infrequens: LECTOTYPE: Panama (MCZ 186451). Acteocina magdalenensis: HOLOTYPE; Bahia Magdalena, Baja California, Mexico, leg. C. R. Orcutt (USNM 218410). Acteocina angustior: HOLOTYPE: Puerto Escondido, Baja California, Mexico (CASIZ 032116). MATERIAL EXAMINED.— Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 22 m depth, 6 shells (CASIZ 170720). Bahia Cocos (10°33'35"N, 85°42'30"W), south of Puerto Culebra, Provincia Guanacaste, 4 m depth, 13 March 1933, 31 shells (LACM 33-123.32). Puerto Parker (10°57'50’N, 85°48'45”"W), Golfo Santa Elena, Provincia Guanacaste, 55 m depth, 9 February 1935, 11 shells (LACM 35- 113.27). Bahia Salinas (11°03'33"N, 85°43'47"W), 11 February 1935, 4 shells (LACM 35-117.19). Bahia Salinas (11°03’33”"N, 85°44’05”W), 11 February 1935, 1 shell (LACM 35-122.16). Between Punta Isla and 500 m South of Punta Isla (10°56’00”"N, 85°48'55”"W), Bahia Santa Elena, Provincia Guanacaste, 1-11 m depth, 13 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-7.33). Southeast end of Bahia Santa Elena, approximately half mile offshore (10°55'15"N, 85°48'30"W), Provincia Guanacaste, 9-12 m depth, 13 February 1972, >50 shells, leg. P. LaFollette and D. Cadien (LACM 72-9.18). Bahia Potrero Grande (10°50'56"N, 85°48'35"W), Provincia Guanacaste, 9-10 m depth, 17 February 1972, 17 shells, leg. P. LaFollette and A. Ferreira (LACM 72-29.12). Middle of Bahia Huevos, north of Bahia Culebra (10°38'41’N, 85°41'55”"W), Provincia Guanacaste, 10-23 m depth, 18—19 February 1972, 3 shells, leg. P. LaFollette and D. Cadien (LACM 72-35.44). Off beach at Bahia Brasilito (10°25'57”"N, 85°49'18”W), Provincia Guanacaste, 18 m depth, 20 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72-40.35). Off Bahia Herradura (9°38'50”N, 84°40'50” W), Provincia Puntarenas, 37 m depth, 10 March 1972, 3 shells, leg. J. McLean (LACM 72-54.56). Isla del Catto (8°44'00"N, 83°52'30”W), Provincia Puntarenas, 12 m depth, 14-19 March, 1972, 6 shells, leg. J. McLean (LACM 72-64.18). 250 m off Punta Piedra (8°35'24.6720N, 83°11'26.3250W), Provincia Puntarenas, 23 m depth, 14 April 1997, 2 shells, leg. M. Madrigal (INB0001496381). Playa San Miguel (9°34’39.3620N, 85°08’05.5470), Cabo Blanco, Provincia Puntarenas, 0 m depth, 13 November 1995, leg. F. Alvarado (INB0003096615). 1.5 km northwest of Playa Junquillal (10°58'40.7139N, 85°42'27.2867W), Bahia Junguillal, Provincia Guanacaste, 20 m depth, 4 April 2002, | shell, leg. F. Gonzalez (INB0003453953). 200 m northwest of Playa Guaria (10°57'55.1620N, 85°42'23.7922W), Bahia Junquillal, Provincia Guanacaste, 8 m depth, 4 April 2002, 1 shell, leg. F. Gonzalez (INB0003458479). Between Isla Bolafios and Playa Coyotera (11°02'51.0971N, 85°43'21.1040), Bahia Salinas, Provincia Guanacaste, 12 m depth, 10 April 2002, 3 shells, leg. F. Gonzalez (INB0003502789). 1.5 km west of Puerto Soley (11°02'35.2251N, 85°41'47.9633W), Bahia Salinas, 10 m depth, 10 April 2002, 8 shells, leg. F. Gonzalez (INB0003503459).1 km west of Playa Junquillal (10°58'24.5393N, 85°42'04.1590W), Bahia Junquillal, Provincia Guanacaste, 30 m depth, 5 April 2002, 17 shells, leg. F. Gonzalez (INB0003503569). 400 m west of Isla Cabo Blanco (9°32'29.2960N, 85°07'16.0340W), Cabo Blanco, Provincia Puntarenas, 20-50 m depth, 16 May 1998, 14 shells, leg. A. Berrocal (INB0003539964). Between Punta Piedra and Punta Gallardo (8°35’47.5010N, 83°11'58.9990W), Provincia Puntarenas, 30-35 m depth, 13 June 1997, 12 shells, leg. M. Madrigal (INB0003542480). Isla David (10°56'06.0000N, 85°42'53.0000W), Bahia Cuajiniquil, Provincia Guanacaste, 18 m depth, 12 March 1996, 11 shells, leg. S. Avila (INB0003575679). Playa Ballena (9°06'34.2870N, 83°41'46.4390W), Provincia Puntarenas, 1 m depth, 16 April 1996, 1 shell, leg.. Y. Camacho (INB0003722811). Isla David (10°58’05.5000N, 85°42'38.6000W), Bahia Junquillal, Provincia Guanacaste, 30 m depth, 12 March 1996, 2 shells, leg. Y. Camacho (INB0003722813). SHELL MORPHOLOGY.— Length up to 9 mm; width up to 3.5 mm in the largest specimen examined. Shell solid, elongate, fusiform, with almost parallel sides (Fig. 2D—E). Body whorl very 466 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 large; spire long and conical in some specimens (Fig. 2E) or short and compressed in others (Fig. 2D), with 2-3 whorls. Suture canaliculated. Protoconch flattened, about 1.5 whorls, 300 um in diameter (Fig. 2F), attached to the teleoconch by the aperture. Umbilicus absent. Aperture very long, about “io of the shell length, narrow, wider anteriorly, narrowing gradually towards the pos- terior end. Columellar margin thickened, oblique, with a long, thin callus running from its anteri- or end to about ';—'/ of the aperture length. Sculpture with fine spiral lines, crossed by faint axial lines, not visible in all specimens. Color uniformly whitish. ANATOMY.— Digestive system containing three smooth and irregular gizzard plates, all of them similar in shape and size (Fig. 2G). Radular formula 16 x 1.0.1 in a specimen from Bahia Potrero Grande (LACM 72-29.12). Lateral teeth hook-shaped with a number of small denticles on the single, curved cusp (Fig. 2H). GEOGRAPHIC RANGE.— Recorded from Baja California to Panama (Keen 1971). REMARKS.— Marcus (1977) considered that Acteon wetherillii Lea, 1833, the fossil type species of Acteocina, is either a member of the Retusidae or Cylichnidae, but different from the common usage of the name Acteocina by modern authors. Marcus (1977) was unable to determine the position of this species, so she decided to use the next available name for this genus, which is Tornatina A. Adams, 1850. Marcus (1977) also maintained Utriculastra as different from Acteocina because of anatomical differences in the number of denticles on the radular teeth and the position of the esophagus. On the contrary, Mikkelsen and Mikkelsen (1984) commented that Acteon wetherillii 1s a syn- onym of the recent species Acteocina canaliculata (Say, 1826), and therefore the name Acfeocina should be maintained as valid. A comparison of the original descriptions of A. wetherillii (Lea 1833) and the redescription of A. canaliculata by Mikkelsen and Mikkelsen (1984) revealed that their shell morphology is in fact very similar. Mikkelsen and Mikkelsen (1984) also added Utriculastra Thiele, 1925 and Cylichnella Gabb, 1873 to the synonymy of Acteocina, following Gosliner (1979) who regarded Utriculastra and Cylichnella as synonyms. Utriculastra was origi- nally based on A. canaliculata (the type species by original designation), and, therefore, it is an objective synonym of Actfeocina. However, Cylichnella differs considerably from Acteocina in shell morphology. Gosliner (1979) commented that Cylichnella has an involute spire whereas Utriculastra possesses a projecting apex. Despite these differences, he regarded these two genera as synonyms because of a similar anatomy. Due to the remarkable differences in shell morphology between Cylichnella and Acteocina, these two genera are here maintained as distinct. Marcus (1977) and Mikkelsen and Mikkelsen (1984) revised several species of Acfeocina and provided detailed anatomical descriptions. The diagnosis above is based on these two papers. Adams (1852) described Bulla infrequens based on two shells collected from Panama. The shells were not illustrated and described as being cylindrical, with a long aperture having a thick columellar plait, a moderately elevated, deeply channeled spire, and a white and smooth surface. Dall (1919) described Acteocina magdalenensis from Bahia Magdalena, Baja California Sur, Mexico, with similar characteristics to those of Acteocina infrequens. Examination of the holotype (USNM 218410) reveals no substantial differences with the lectotype of 4. magdalenensis and therefore these two names are regarded as synonyms. A second synonym is Acteocina angustior described by Baker and Hanna (1927) from Puerto Escondido, Baja California, Mexico. The description and features of the holotype (CASIZ 032116) are undistinguishable from those of Acteocina infrequens. Baker and Hanna (1927) compared 4A. angustior with Acteocina culcitella Gould, 1852 (a northern species), but not with other Panamic species. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 467 FIGURE 2. A-C, Cylichna atahualpa (Dall, 1908); (A) Shell morphology, specimen from Golfo Dulce (LACM 39- 44.23), 6.5 mm long; (B) Radular teeth, specimen from Golfo Dulce (LACM 39-44.23); (C) Gizzard plate, same specimen. D-H, Acteocina infrequens (C.B. Adams, 1852); (D) Shell morphology, specimen from Isla Cabo Blanco (INB0003539964), 8 mm long; (E) Shell morphology, specimen from Bahia Cuajiniquil (INB0003575679), 4 mm long; (F) Protoconch, specimen from Playa Junquillal (INB0003503569); (G) Gizzard plate, specimen from Bahia Potrero Grande (LACM 72-29.12); (H) Radular teeth, same specimen. 468 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 Acteocina carinata (Carpenter, 1857) (Figs. 3A-C) Tornatina carinata Carpenter, 1857:171. TYPE MATERIAL.— TJornatina carinata: SYNTYPES (6): Mazatlan, Mexico (BMNH). MATERIAL EXAMINED.— Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 27 m depth, 25 shells (CASIZ 170719). Puerto Culebra (10°35’N, 85°40'W), Provincia Guanacaste, 31 m depth, 13 March 1934, 1 shell (LACM 33-153.10). Puerto Culebra, (10°37'N, 85°40'W), Provincia Guanacaste, 27 m depth, 26 February 1934, 2 shells (LACM 34-253.5). Playa Blanca, Bahia Playa Blanca, Provincia Guanacaste (10°56’45”N, 85°53'30"W), 3-5 m depth, 8 February 1935, 3 shells (LACM 35-100.7). Puerto Parker (10°57'50"N, 85°48’45"W), Golfo Santa Elena, 55 m depth, 9 February 1935, 2 shells (LACM 35-113.24). Bahia Salinas (11°03'33"N, 85°43'47”W), 11 February 1935, 3 shells (LACM 35-117.15). One mile offshore, between Bahia Santa Elena and Bahia San Juanillo, (10°57'20”"N, 85°46'08” W), Provincia Guanacaste, 25—53 m depth, 14 February 1972, 15 shells, leg. P. LaFollette and D. Cadien (LACM 72-12.44). Half mile to one mile and half west of Roca Vagares, Bahia de San Juanillo (10°57'28"N, 85°45'20”W), Provincia Guanacaste, 37 m depth, 14 February 1972, | shell, leg. D. Cadien and P. LaFollette (LACM 72-13.32). Southeast corner of Bahia Jobo, off sand beach west of Bahia Salinas (11°02'22”N, 85°45'16”W), Provincia Guanacaste, 1-10 m depth, 14 February 1972, 4 shells, leg. P. LaFollette and D. Cadien (LACM 72-19.43). Bahia Potrero Grande (10°50'56"N, 85°48'35"W), Provincia Guanacaste, 9-10 m depth, 17 February 1972, 7 shells, leg. P. LaFollette and A. Ferreira (LACM 72-29.10). Five miles north west of Islas Huevos (10°41'45’N, 85°46'25"W), Golfo de Papagayo, Provincia Guanacaste, 60-64 m depth, 18 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-34.27). Middle of Bahia Huevos, north of Bahia Culebra (10°38’41’N, 85°41'55”W), Provincia Guanacaste, 10-23 m depth, 18—19 February 1972, 6 shells, leg. P. LaFollette and D. Cadien (LACM 72-35.40). Half mile off mouth of Bahia Huevos (10°38'22"N, 85°42'52”W), Provincia Guanacaste, 36-42 m depth, 19 February 1972, 7 shells, leg. P. LaFollette and D. Cadien (LACM 72-36.24). Off beach at Bahia Brasilito (10°25'57"N, 85°49'18”W), Provincia Guanacaste, 18 m depth, 20 February 1972, 18 shells, leg. P. LaFollette and D. Cadien (LACM 72-40.30). Anchorage at Bahia Ballena off Tambor (10°44'10"N, 84°59'34”"W), Provincia Puntarenas, 10 m depth, 21 February, 1972, 1 shell, leg. P. LaFollette and A. Ferreira (LACM 72-43.17). Off beach at Bahia Ballena (9°44'12”N, 84°59'32”W), Provincia Puntarenas, 3—18 m depth, 23 February 1972, 5 shells, leg. P. LaFollette and D. Cadien (LACM 72-45.38). Islet and rocks 1 km west of Isla Alcatraz (9°47'N, 84°53.5'W), Islas Tortugas, Provincia Puntarenas, 2—8 m depth, 23—24 February 1972, | shell, leg. P. LaFollete, D. Cadien, and A. Ferreira (LACM 72-46.58). Bahia Herradura (9°37'58”"N, 84°40'30"W), Provincia Puntarenas, 21 m depth, 9 March 1972, 13 shells, leg. J. McLean (LACM 72-53.29). Off Bahia Herradura (9°38'50”"N, 84°40'50”W), Provincia Puntarenas, 37 m depth, 10 March 1972, 27 shells, leg. J. McLean (LACM 72-54.51). Anchorage inside of small islet 1.5 km south of Punta Quepos (9°22'43”N, 84°09'41”W), Provincia Puntarenas, 21 m depth, 11 March 1972, | shell, leg. J. McLean (LACM 72-57.38). Small islets of Quepos (9°22'12"N, 84°09'15”W), Provincia Puntarenas, 23 m depth, 12 March 1972, 5 shells, leg. J. McLean (LACM 72-59.30). South of Punta Uvita to Isla Ballena (9°07'36"N, 83°44'30"W), Provincia Puntarenas, 18 m depth, 13 March 1972, 1 fragment, leg. J. McLean (LACM 72-61.23). Off Isla del Cafio (8°45’N, 83°54'W), Provincia Puntarenas, 55 m depth, 16—17 March 1972, 1 shell, leg. J. McLean (LACM 72-66.39). From Punta Piedra to Punta Gallardo (8°35'47.5010N, 83°11'58.9990W), Golfo Dulce, Provincia Puntarenas, 35 m depth, 13 June 1997, 1 shell, leg. M. Madrigal (INB0001496191). Playa San Miguel (9°34'39.3620N, 85°08’05.5470W), Reserva Natural Absoluta de Cabo Blanco, Provincia Puntarenas, 0 m depth, 13 November 1995, 7 shells, leg. F. Alvarado (INB0003096504). 1.5 km northwest of Playa Junquillal (10°58’40.7139N, 85°42'27.2867W), Santa Rosa, Provincia Guanacaste, 20 m depth, 4 April 2002, 6 shells, leg. F. Gonzalez (INB0003501408). Playa Naranjo (10°48'16.0960N, 85°41'12.0680W), Estero Real, Provincia Guanacaste, 0 m depth, 6 March 1995, 1 shell, leg. S. Avila (INB0003539800). Isla David (10°58’05.5000N, 85°42'38.6000W), Bahia Junquillal, Provincia Guanacaste, 30 m depth, 12 March 1996, 5 shells, leg. Y. Camacho (INB0003542874). Isla David (10°58’05.5000N, 85°42'38.6000W), Bahia Junquillal, Provincia Guanacaste, 30 m depth, 12 March 1996, 5 shells, leg. Y. Camacho (INB0003542915). Isla David (10°56'32.2000N, 85°42'51.1000), Santa Rosa, Provincia VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 469 Guanacaste, 18 m depth, 4 shells, 12 March 1996, leg. Y. Camacho (INB0003575624). Isla David (10°56'06.0000N, 85°42'53.0000W), Bahia Cuajiniquil, Provincia Guanacaste, 18 m depth, 12 March 1996, 21 shells, leg. Y. Camacho (INB0003575670). 1 km east of Punta Cirial (9°56'10.2222N, 84°52'43.1248W), Isla San Lucas, Provincia Puntarenas, 22.6 m depth, 9 December 1999, 1 shell, leg. M. Calderon (INB0003722804). 400 m west of Isla Cabo Blanco (9°32'29.2960N, 85°07'16.0340W), Reserva Natural Absoluta de Cabo Blanco, Provincia Puntarenas, 20-50 m depth, 16 May 1998, 6 shells, leg. A. Berrocal (INB0003722806). Golfo de Papagayo (10°22'00.0000N, 86°15’00.0000W), 100 m depth, 10 February 1995, 2 shells, leg. Y. Camacho (INB0003722814). Dock of Playa San Miguel (9°34'36.9190N, 85°08'06.3610W), Reserva Natural Absoluta de Cabo Blanco, Provincia Puntarenas, 0 m depth, 2 May 1995, 4 shells, leg. G. Mena (INB0003722818). Punta Oliva (9°34'39.3370N, 85°08'15.3840W), Cabo Blanco, Provincia Puntarenas, 0 m depth, 16 September 1995, 4 shells, leg. F. Alvarado (INB0003722821). La Viuda Rock (8°36'56.0065N, 83°14’08.0868W), Golfo Dulce, Provincia Puntarenas, 66 m depth, 21 April 1997, 1 shell, leg. M. Lobo (INB0003722824). Golfo Santa Elena (10°58’45.4310N, 85°43'01.8500W), Bahia Junquillal, Provincia Guanacaste, 0-3 m depth, 13 March, 1996, 31 shells, leg. R. Angulo (INB0003722830). Boca de Estero Bocon (8°32'57.0600N, 83°18'51.2380W), Golfo Dulce, Provincia Puntarenas, 0 m depth, 13 June 1995, 1 shell, leg. M. Lobo (INB0003747047). La Viuda Rock (8°36'56.0065N, 83°14'08.0868W), Golfo Dulce, Provincia Puntarenas, 66 m depth, 21 April 1997, 23 shells, leg. M. Lobo (INB0003747054). SHELL MORPHOLOGY.— Length up to 4 mm; width up to 2 mm in the largest specimen exam- ined. Shell fragile, elongate, with almost parallel sides (Fig. 3A—B). Body whorl very large; spire short, with 3 whorls, flattened in some specimens (Fig. 3A) and conical in others (Fig. 3B). Suture not channeled. Posterior end of the whorls with an ornated sharp ridge (carina) with longitudinal bands. Protoconch flattened, about 1.2 whorls, 200 um in diameter (Fig. 3C). It is only attached to the teleoconch by the aperture. Umbilicus absent. Aperture very long, between ‘ho and */o of the shell length, narrow, wider anteriorly, narrowing gradually towards the posterior end. Columellar margin thickened, oblique, with a thin callus running from its anterior end to about '/s—'h of the aperture length. Sculpture with fine spiral lines, crossed by faint axial lines, not visible in all spec- imens. Color uniformly whitish. ANATOMY.— All the specimens examined consisted of empty shells so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— Known from the Gulf of California to Costa Rica (Keen 1971). REMARKS.— Carpenter (1857) described this species as different from Acteocina infrequens by having a more irregular spire, a suture not channeled and a shoulder sharply carinated. Additionally, the shell of A. infrequens is elongate, with the end of the whorls rounded and the pro- toconch is about 300 um in diameter, whereas the shell of Acteocina carinata has the posterior end of the whorls with a sharp ridge ornated with longitudinal bands and the protoconch 1s about 200 um in diameter. Acteocina harpa (Dall, 1871), originally described from California by Dall (1871) has a shell morphology similar to that of A. carinata, but according to Gosliner (1996), the former differs by having more assymetrical gizzard plates and a rachidian tooth with fewer denticles. Acteocina sp. 1 (Figs. 3D—-E) MATERIAL EXAMINED.— 0.9 mile off beach at Bahia Potrero (10°27'42”"N, 85°48'15”W), Provincia Guanacaste, 16-18 m depth, 20 February 1972, | shell, leg. P. LaFollette and D. Cadien (LACM 72-39.13). Off beach at Bahia Brasilito (10°25'57’N, 85°49'18”W), Provincia Guanacaste, 18 m depth, 20 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-40.31). SHELL MORPHOLOGY.— Length up to 4.5 mm; width up to 2.5 mm in the largest specimen examined. Shell fragile, oval, with a convex right side and concave left side (Fig. 3D). Body whorl 470 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 very large; spire short and conical, with 1—2 whorls. Suture canaliculated. Protoconch flattened, about 1.5 whorls, 150 um in diameter (Fig. 3E). Umbilicus absent. Aperture very long, about “/s of the shell length, narrow, wider anteriorly, narrowing gradually towards the posterior end. Columellar margin thickened, oblique, with a long, thick callus. Sculpture with fine spiral lines, crossed by faint axial lines, not visible in all specimens. Color uniformly whitish. ANATOMY.— All the specimens examined consisted of empty shells, so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— This species is only known from Costa Rica (present paper). REMARKS.— This appears to be a distinct species of Acteocina, but it is not described herein because of the absence of anatomical data. Acteocina sp. | is easily distinguishable from both Acteocina infrequens and Acteocina cari- nata by its shell morphology. The shell of Acteocina sp. | is shorter, more oval than that of A. infre- quens and most specimens of A. carinata. In addition, the spire of Acfeocina sp. | is shorter than in the other two species and the posterior end of the whorl lacks the sharp ridge ornated with lon- gitudinal bands of 4. carinata. The protoconch of Acteocina sp. | 1s considerably smaller than in the other two species. Acteocina sp. 2 (Figs. 3F-G) MATERIAL EXAMINED.— Puerto Jiménez (8°32.5'N, 83°19.5'W), Golfo Dulce, Provincia Puntarenas, 20 March, 1972, 1 shell, leg. J. McLean (LACM 72-69.17). Bahia El Hachal (10°56'08.9540N, 85°43'58.7890W), Provincia Guanacaste, 0 m depth, 15 September 1994, 2 shells, leg. C. Cano (INB0001480928). Boca de Estero Caballero (8°40'06.3050N, 83°26'41.8580W), Provincia Puntarenas, 0 m depth, 11 June 1995, 11 shells, leg. M. Lobo (INB0001499798). Quebrada Palma (8°39'06.8780N, 83°26'10.8200W), Playa Blanca, Golfo Dulce, Provincia Puntarenas, 0 m depth, 7 April 1995, >50 shells, leg. G. Mena (INB0003537637). Boca de Estero Caballero (8°40'06.3050N, 83°26'41.8580W), Provincia Puntarenas, 0 m depth, 11 June 1995, 11 shells, leg. M. Lobo (INB0003540727). Punta Palma (8°38'59.55 10N, 83°26'07.5540W), Golfo Dulce, Provincia Puntarenas, 2 m depth, 12 June 1995, 7 shells, leg. M. Lobo (INB0003722822). 1 km south of Puntarenitas (8°36'55.7150N, 83°10'07.7110W), Provincia Puntarenas, 0 m depth, 13 May 1997, 1 shell, leg. S. Avila (INB0003726899). Punta Palma (8°38'59.5510N, 83°26'07.5540W), Golfo Dulce, Provincia Puntarenas, 2 m depth, 12 June 1995, 31 shells, leg. M. Lobo (INB0003747326). Boca de Estero Bocdn (8°32'57.0600N, 83°18'51.2380W), Golfo Dulce, Provincia Puntarenas, 0 m depth, 13 June 1995, 7 shells, leg. M. Lobo (INB0003747047). Sector Playitas (8°44’'19. 1980N, 83°21'57.0660W), Golfo Dulce, Provincia Puntarenas, 0 m depth, 10 February 1996, 2 shells, leg. M. Lobo (INB0003747549). SHELL MORPHOLOGY.— Length up to 3 mm; width up to 1.5 mm in the largest specimen examined. Shell fragile, elongate, posteriorly truncate, with almost parallel sides (Fig. 3F). Body whorl very large; spire short, almost flat. Suture canaliculated. Protoconch flattened, partially embedded in the teleoconch, 200 um in diameter (Fig. 3G). Umbilicus absent. Aperture very long, about */o of the shell length, narrow, wider anteriorly, narrowing gradually towards the posterior end. Columellar margin thickened, oblique. Sculpture with conspicuous spiral lines, more densely arranged near the anterior end of the shell, crossed by faint axial lines. Color uniformly whitish to pale brown. ANATOMY.— AIl the specimens examined consisted of empty shells so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— This species is only known from several localities in Costa Rica (pres- ent paper). VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 47] REMARKS.— This appears to be a distinct species of Acteocina, but it is not described herein because of the absence of anatomical data. This species is superficially similar to Acfeocina carinata in shell morphology, but these two species are easily distinguishable by the presence of a shorter spire in Acteocina sp. 2, with the pro- toconch partially embedded in the teleoconch. Additionally, the sculpture of Acteocina sp. 2 has conspicuous spiral lines, more densely arranged near the anterior end of the shell, which are absent in A. carinata. Genus Cylichnella Gabb, 1873 Cylichnella tabogaensis (Strong and Hertlein, 1939) (Figs. 4A—-C) Retusa tabogaensis Strong and Hertlein, 1939:191—192, pl. 18, fig. 4. TYPE MATERIAL.— Retusa tabogaensis: PARATYPE: off Isla Taboga, Panama (LACM 1341). MATERIAL EXAMINED.— Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 22 m depth, 9 shells (CASIZ 170717). Bahia Cocos (10°33'35”N, 85°42'30”W), south of Puerto Culebra, Provincia Guanacaste, 4 m depth, 13 March 1933, 2 shells (LACM 33-123.27). Puerto Culebra (10°35’N, 85°40’W), Provincia Guanacaste, 31 m depth, 13 March 1934, 4 shells (LACM 33-153.11). Playa Blanca (10°56'45"N, 85°56’W), Provincia Guanacaste, 73 m depth, 8 February 1935, 1 shell (LACM 33-103.14). Bahia Salinas (11°03’33"N, 85°43’47"W), 11 February 1935, 2 shells (LACM 35-117.16). Southeast end of Bahia Santa Elena, approximately 0.5 mile offshore (10°55’15"N, 85°48'30"W), Provincia Guanacaste, 9-12 m depth, 13 February 1972, 10 shells, leg. P. LaFollette and D. Cadien (LACM 72-9.15). One mile offshore, between Bahia Santa Elena and Bahia San Juanillo (10°57’20"N, 85°46'08"W), Provincia Guanacaste, 25—53 m depth, 14 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72-12.46). Half mile to one mile and half west of Roca Vagares, Bahia San Juanillo (10°57'28"N, 85°45'20"W), Provincia Guanacaste, 37 m depth, 14 February 1972, 6 shells, leg. D. Cadien and P. LaFollette (LACM 72-13.34). Southeast corner of Bahia Jobo, off sand beach west of Bahia Salinas (11°02’22"N, 85°45'16”W), Provincia Guanacaste, 1-10 m depth, 14 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72-19.44). Five miles north west of Islas Huevos (10°41'45”"N, 85°46'25"W), Golfo de Papagayo, Provincia Guanacaste, 60—64 m depth, 18 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-34). 250 m off Punta Piedra (8°35'24.6720N, 83°11'26.3250), Provincia Puntarenas, 23 m depth, 14 April 1997, 8 shells, leg. M. Lobo (INB0001496380). Between Isla Bolafios and Playa Coyotera (11°02'51.0972N, 85°43’21.1040W), Provincia Guanacaste, 12 m depth, 10 April 2002, | shell, leg. A. Berrocal (INB0003539773). From 500 m off the beach to 2.5 km south- east (8°34'39.1040N, 83°11'29.6540W), Punta Piedra, Provincia Puntarenas, 14-90 m depth, 10 April 1997, 24 shells, leg. M. Madrigal (INB0003540314). Between Punta Piedra and Punta Gallardo (8°35'47.5010N, 83°11'58.9990W), Golfo Dulce, Provincia Puntarenas, 30-35 m depth, 13 June 1997, 11 shells, leg. M. Madrigal (INB0003540361). 1 Km northwest of Playa Coronado (9°59'54.1430N, 84°58'49.6030W), Isla Caballo, Provincia Puntarenas, 10 m depth, 7 December 1999, 2 shells, M. Calderon (INB0003542209). Isla David (10°56'32.2000N, 85°42'51.1000W), Provincia Guanacaste, 18 m depth, 12 March 1996, 7 shells, leg. Y. Camacho (INB0003575621). Isla David (10°56'06.0000N, 85°42'53.0000W), Bahia Cuajiniquil, Provincia Guanacaste, 18 m depth, 12 March 1996, 3 shells, leg. Y. Camacho (INB0003575666). La Viuda Rock (8°36'56.0065N, 83°14'08.0868W), Golfo Dulce, Provincia Puntarenas, 66 m depth, 21 April 1997, 1 shell, leg. M. Lobo (INB0003722823). Bahia Junquillal (10°59’13.0310N, 85°43'16.7950W), Golfo Santa Elena, Provincia Guanacaste, 3 m depth, 2 shells, 13 March 1996, leg. R. Angulo (INB0003722827). Golfo Santa Elena (10°58’45.4310N, 85°43’01.8500W), Bahia Junquillal, Provincia Guanacaste, 0-3 m depth, 13 March, 1996, 2 shells, leg. R. Angulo (INB0003722831). 250 m off Punta Piedra (8°35'24.6720N, 83°11'26.3250), Provincia Puntarenas, 23 m depth, 14 April 1997, 14 shells, leg. M. Lobo (INB0003747050). Isla David (10°58'05.5000N, 85°42'38.6000W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 30 m depth, 12 March 1996, 11 shells, leg. Y. Camacho (INB0003747059). Playa Mostrencal (10°59'47.3060N, $5°42'53.8890W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 2 m depth, 3 shells, 13 March 472 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 FIGURE 3. A—C, Acteocina carinata (Carpenter, 1857); (A) Shell morphology, specimen from Bahia Junquillal (INB0003722830), 4 mm long; (B) Shell morphology, specimen from Isla Cabo Blanco (INB0003722806), 4 mm long (C) Protoconch, specimen from Isla Cabo Blanco (INB0003722806). D—E, Acteocina sp. 1; (D) Shell morphology, specimen from Bahia Potrero (LACM 72-40.31), 4 mm long; (E) Protoconch, same specimen. F—G, Acteocina sp. 2; (F) Shell mor- phology, specimen from Golfo Dulce (INB0003747549), 2.5 mm long; (E) Protoconch, specimen from Golfo Dulce (INB0003537637). VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 473 1996, leg. R. Angulo (INB0003747411). 1 km east of Punta Cirial (9°56'10.2222N, 84°52’43.1248W), Isla San Lucas, Provincia Puntarenas, 0-22 m depth, 9 December 1999, | shell, leg. M. Calderon (INB0003747485). SHELL MORPHOLOGY.— Length up to 4 mm; width up to 2 mm in the largest specimen exam- ined. Shell fragile, oval, with convex sides (Fig. 4A). Only one whorl visible, forming nearly the entire shell, except for a portion of the top of the spire. Apex rounded, umbilicated, with the aper- ture lip rising from the right side. Anterior end of the shell rounded. Aperture as long as the shell, wider anteriorly and narrowing gradually at about '/ of its length. Columellar margin conspicuous- ly thickened with a small gap that divides the columella into two distinct folds. Umbilicus absent. Sculpture with a number of irregular spiral lines more conspicuous near the anterior and posterior ends of the shell. Color pale brown. ANATOMY.— Digestive system containing three smooth and irregular gizzard plates, com- posed of a thickened central area and a thinner extension (Fig. 4B). Central plate wider than the two lateral plates. Radular formula 16 x 1.1.1 in a specimen from Punta Piedra (INB0003540314). Lateral teeth hook-shaped with a wider area below the elongate cusp, bearing 16 long and strong denticles (Fig. 4C). Rachidian teeth broad, with several short denticles decreasing in size towards the laterals of each plate, and a gap with no denticles on the center of each rachidian tooth. GEOGRAPHIC RANGE.— This species is known from Panama and Costa Rica (Keen 1971). REMARKS.— Retusa tabogaensis was originally described based on several shells collected from Isla Taboga, Panama (Strong and Hertlein 1939). The most distinctive feature of this species was the presence of two denticles on the columella. Examination of the type material and original description revealed that our material from Costa Rica belongs to the same species. Further anatomical examination of Costa Rican specimens revealed the presence of a radula with lateral teeth bearing an extension under the cusp with several long and strong denticles, and three gizzard plates, composed of a thickened central area and a thinner extension, one of the plates wider than the rest. All these features are consistent with those of Cylichnella Gabb, 1873, as defined by Gosliner (1979). Cylichnella goslineri Valdés and Camacho-Garcia, sp. nov. (Figs. 4D-F, 5) TYPE MATERIAL.— Cylichnella goslineri sp. nov.: HOLOTYPE: Golfo Dulce, Provincia Puntarenas, 0 m depth, 10 February 1996, 1 specimen, leg. M. Lobo (INB0001497964). MATERIAL EXAMINED.— Sector Playitas (8°44'19.1980N, 83°21'57.0660W), Golfo Dulce, Provincia Puntarenas, 0 m depth, 10 February 1996, 15 specimens, leg. M. Lobo (INB0003718957). Four hundred and fifty meters southwest of Playa Blanca (10°03'39.5590N, 84°57'23.0410W), Provincia Puntarenas, 0 m depth, 8 December 1999, | shell, leg. M. Calderon (CASIZ 0121109). Shell morphology.— Length up to 4.5 mm; width up to 2 mm in the largest specimen exam- ined. Shell fragile, oval to elongate, with convex sides (Fig. 4D). Only one whorl visible, forming nearly the entire shell, except for a visible portion of the top of the spire. Apex rounded, slightly umbilicated, with the aperture lip rising from the right side. Anterior end of the shell rounded. Aperture as long as the shell, wider anteriorly, narrowing gradually at about ‘4 of its length. Columellar margin slightly thickened, simple, with no folds or denticles. Umbilicus absent. Sculpture with a number of irregular and faint spiral lines. Color uniformly dirty white. ANATOMY.— Buccal bulb oval to elongate, connecting posteriorly to a short esophagus and the small salivary glands (Fig. SA). Two strong retractor muscles attach laterally to the buccal bulb. Esophagus opening into a large gizzard, which contains three smooth and irregular gizzard plates, composed of a thickened central area and a thinner extension (Fig. 4E); central plate wider than the 474 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 FiGuRE 4. A—C, Cylichnella tabogaensis (Strong and Hertlein, 1939); (A) Shell morphology, specimen from Bahia Junquillal (INB0003747411), 4 mm long; (B) Central gizzard plate, specimen from Punta Piedra (INB0003540314); (C) Radular teeth, same specimen. D—F, Cylichnella goslineri sp. nov.; (D) Shell morphology, Holotype (INB0001497964), 4.5 mm long; (E) Lateral gizzard plate, Paratype from Golfo Dulce (INB0003718957); (F) Radular teeth, same specimen. two lateral plates. Radular formula 21 x 1.1.1 in a specimen from Golfo Dulce (INB0003718957). Lateral teeth hook-shaped, with a wider area below the elongate cusp bearing 12 strong denticles (Fig. 4F). Rachidian teeth broad, with several short denticles decreasing in size towards the later- als of each plate, and a gap with no denticles on the center of each rachidian tooth. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 475 FIGURE 5. Cylichnella goslineri sp. nov., anatomy of specimen from Golfo Dulce (INB0003718957); (A) Anterior portion of the digestive system; (B) Male reproductive sys- tem. Abbreviations: bb, buccal bulb; e, esophagus; gp, giz- zard plate; gz, gizzard; pn, penis; pr, prostate; rm, retractor muscule, sg, salivary gland. Reproductive system monoaulic. Penis bulbous, connected to the prostate through a long and convoluted duct (Fig. 5B). GEOGRAPHIC RANGE.— Known _ from Costa Rica (present paper). ETYMOLOGyY.— The species name is dedi- smm cated to our former advisor Terry Gosliner, for his company and support during the field work in Costa Rica. REMARKS.— Cylichnella goslineri sp. nov. is clearly distinct from other Cylichna-like opisthobranchs described from the Panamic region by having a more elongate shell with no con- spicuous spiral lines. The most similar species in shell morphology is Cylichna atahualpa, but these two species clearly differ in the morphology of the radula and gizzard plates (see above). The other member of the genus Cy/ichnella is C. tabogaensis, which has a shorter shell with conspicu- ous spiral lines and radular teeth with stronger denticles. Cylichnella goslineri sp. nov. is assigned to the genus Cylichnella by having a radula with lat- eral teeth bearing an extension under the cusp with several long and strong denticles, and three giz- zard plates, composed of a thickened central area and a thinner extension, one of the plates wider than the rest. All these features are consistent with the definition of this genus by Gosliner (1979). 0.5 mm Family Retusidae Thiele, 1925 Genus Retusa Brown, 1827 Retusa paziana Dall, 1919 (Fig. 6A) Retusa paziana Dall, 1919:297. TYPE MATERIAL.— Retusa paziana: SYNTYPE: La Paz, Mexico (USNM 211418). MATERIAL EXAMINED.— Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 22 m depth, 2 shells (CASIZ 170723). Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 27 m depth, 16 shells (CASIZ 170724). Bahia Cocos (10°33'35"N, 85°42'30”W), south of Puerto Culebra, Provincia Guanacaste, 4 m depth, 13 March 1933, 1 shell (LACM 33-123.28). Puerto Culebra (10°35’N, 85°40'W), Provincia Guanacaste, 31 m depth, 13 March 1934, 1 shell (LACM 33-153.12). Puerto Culebra (10°37’N, 85°40'W), 27 m depth, 26 February 1933, 1 shell (LACM 34-253.6). Puerto Parker (10°55’N, 85°49’W), Golfo Santa Elena, Provincia Guanacaste, 8 February 1935, 2 shells (LACM 35-138.2). Puerto Parker (10°57’50”N, 85°48'45"W), Golfo Santa Elena, Provincia Guanacaste, 55 m depth, 9 February 1935, 3 shells (LACM 35-113.25). Bahia Salinas (11°03'33"N, 85°43’47”W), 11 February 1935, 3 shells (LACM 35.117.17). Bahia Salinas (11°03'33”"N, 85°44’05”W), 11 February 1935, 2 shells (LACM 35-122.15). One mile offshore, between Bahia Santa Elena and Bahia San Juanillo (10°57'20’N, 85°46’/08”W), Provincia Guanacaste, 25-53 m depth, 14 February 1972, 3 shells, leg. P. LaFollette and D. Cadien (LACM 72-12.47). Middle of Bahia Huevos, north of Bahia Culebra (10°38’41”N, 85°41'55”W), Provincia Guanacaste, 10-23 m depth, 18—19 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72-35.41). Half mile off mouth of Bahia Huevos (10°38'22"N, 476 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 85°42'52”W), Provincia Guanacaste, 36-42 m depth, 19 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-36.25). Off beach at Bahia Brasilito (10°25’57"N, 85°49'18” W), Provincia Guanacaste, 18 m depth, 20 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-40.32). Off beach at Bahia Ballena (9°44'12’N, 84°59'32”W), Provincia Puntarenas, 3-18 m depth, 23 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72-45.39). Off Bahia Herradura (9°38'50”N, 84°40'50”W), Provincia Puntarenas, 37 m depth, 10 March 1972, 2 shells, leg. J. McLean (LACM 72-54.53). Isla David (10°58'05.5000N, 85°42'38.6000W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 30 m depth, 12 March 1996, 1 shell, leg. Y. Camacho (INB0003747060). Isla David (10°56'32.2000N, 85°42'51.1000W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 18 m depth, 12 March 1996, 4 shells, leg. Y. Camacho (INB0003700589). Isla David (10°58'45.4310N, 85°43'01.8500W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 0-3 m depth, 13 March 1996, 3 shells, leg. R. Angulo (INB0003747335). Isla David (10°58’05.5000N, 85°42'38.6000W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 30 m depth, 12 March 1996, 3 shells, leg. Y. Camacho (INB0003719254). SHELL MORPHOLOGY.— Length up to 3 mm; width up to | mm in the largest specimen exam- ined. Shell fragile, elongate, wider anteriorly and narrower posteriorly, with nearly parallel or slightly concave sides (Fig. 6A). Only one whorl visible, forming nearly the entire shell, except for a visible portion of the top of the spire and the apex of the protoconch, which is partially embed- ded. Apex umbilicated, with the aperture lip rising from the right side. Aperture lip forming a rounded wing connected to the columellar margin. Anterior end of the shell rounded. Aperture as long as the shell, wider anteriorly and narrowing abruptly at about '/ of its length. Columellar mar- gin slightly thickened. Columella simple, with no folds. Umbilicus absent. Sculpture with a num- ber of conspicuous growth lines crossed by faint spiral lines. Color uniformly whitish. ANATOMY.— All the specimens examined consisted of empty shells so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— Known from the Gulf of California (Keen 1971) and Costa Rica. REMARKS.— Dall (1919) described Retusa paziana based on shells from La Paz, Baja California Sur, Mexico, with a short description and no figures. Baker and Hanna (1927) collected additional specimens from the Gulf of California and illustrated this species for the first time. The shells from Costa Rica here examined match the original description and redescription by Baker and Hanna (1927), and there is no doubt they belong to the same species. Retusa paziana 1s the only member of Refusa from the Panamic region. Its unique shell mor- phology, with the wider area close to the anterior end, and the sculpture, composed of strong growth lines, make this species easily recognizable. Retusa sp. (Fig. 6B) MATERIAL EXAMINED.— 1.5 mile east of Punta Ballena (9°44'15”N, 84°33'45"W), Provincia Guanacaste, 3-15 m depth, 21—22 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72- 42.61). Playa Nancite (10°48’N, 85°42’W), north side of Golfo Papagayo, Provincia Guanacaste, 15 January 1986, | shell, leg. E. Coan and R. Hollywood (LACM 86-26.34). SHELL MORPHOLOGY.— Length up to 2 mm: width up to | mm in the largest specimen exam- ined. Shell fragile, oval, wider near the center, with convex sides (Fig. 6B). Only one whorl visi- ble, forming nearly the entire shell, except for a visible portion of the top of the spire and the apex of the protoconch, which is partially embedded. Apex umbilicated, with the aperture lip rising from the right side. Aperture lip forming a rounded wing connected to the columellar margin. Anterior end of the shell elongate. Aperture as long as the shell, wider anteriorly and narrowing gradually at about '/s of its length. Columellar margin slightly thickened. Columella simple, with no folds. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 477 Umbilicus very narrow. Sculpture with a number of simple spiral lines crossed by conspicuous growth lines. Color uniformly whitish. ANATOMY.— AIl the specimens examined consisted of empty shells so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— Known from Costa Rica (present paper). REMARKS.— This is most likely an undescribed species but it will not be described herein because of the lack of anatomical information. Refusa sp. is similar to Retusa paziana in some respects but clearly distinguishable by the outline of the shell and sculpture. The shell of Retusa sp. has its wider area near the center of the shell, whereas in R. paziana it is near the anterior end. Additionally, Retwsa sp. has a small umbilicus that is absent in R. paziana. The sculpture of R. paziana is composed of conspicuous growth lines crossed by faint spiral lines whereas in Retusa sp. the spiral lines are equally conspicuous. This species is provisionally placed in the genus Retusa because of the morphological similar- ities of the shell with some other species of the genus. Genus Volvulella Newton, 1891 Volvulella cylindrica (Carpenter, 1864) (Figs. 6D-E) Volvula cylindrica Carpenter. 1864:179. Volvulella callicera Dall. 1919:299. Volvulella cooperi Dall. 1919:297-298. Volvulella lowei Strong and Hertlein. 1937:164—165, pl. 35, fig. 2. TYPE MATERIAL.— Volvula cylindrica: HOLOTYPE: Santa Barbara, California (Redpath Museum 2364). Volvulella callicera: HOLOTYPE: Galapagos Islands (USNM 194976B). Volvulella cooperi: HOLOTYPE: Scammons Lagoon [= Laguna Ojo de Liebre], Baja California Sur, Mexico (USNM 105501). Volvulella lowei: HOLOTYPE: Puerto Escondido, Baja California Sur, Mexico (CASIZ 065971). MATERIAL EXAMINED.— Puerto Parker, Golfo Santa Elena, Provincia Guanacaste, 27 m depth, 38 shells (CASIZ 170718). Bahia Cocos (10°33'35"N, 85°42'30”W), south of Puerto Culebra, Provincia Guanacaste, 4 m depth, 13 March 1933, 2 shells (LACM 33-123.29). Puerto Culebra (10°35'N, 85°40'W), Provincia Guanacaste, 31 m depth, 13 March 1934, 4 shells (LACM 33-153.13). Puerto Culebra (10°37’N, 85°40'W), Provincia Guanacaste, 27 m depth, 26 February 1934, 1 shell (LACM 34-253.7). Puerto Parker (10°55'N, 85°49'W), Golfo Santa Elena, Provincia Guanacaste, 8 February 1935, 2 shells (LACM 35-138.3). Puerto Parker (10°57'50”N, 85°48'45”W), Golfo Santa Elena, Provincia Guanacaste, 55 m depth, 9 February 1935, 2 shells (LACM 113.26). Bahia Salinas (11°03'33"N, 85°43'47”W), Provincia Guanacaste, |1 February 1935, 3 shells (LACM 35-117.18). Playa Blanca (10°56'45”N, 85°53'30"W), Bahia Playa Blanca, Provincia Guanacaste, 3-5 m depth, 8 February 1935, 7 shells (LACM 35-100.7). Playa Blanca (10°56’45"N, 85°53'30"W), Bahia Playa Blanca, Provincia Guanacaste, 3—5 m depth, 8 February 1935, 7 shells (LACM 35- 100.7). Southeast end of Bahia Santa Elena, approximately half mile offshore (10°55’15"N, 85°48’30"W), Provincia Guanacaste, 9-12 m depth, 13 February 1972, 8 shells, leg. P. LaFollette and D. Cadien (LACM 72- 9.16). One mile offshore, between Bahia Santa Elena and Bahia San Juanillo (10°57’20"N, 85°46'08”"W), Provincia Guanacaste, 25-53 m depth, 14 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-12.47). Southeast corner of Bahia Jobo, off sand beach west of Bahia Salinas (11°02'22”N, 85°45'16”W), Provincia Guanacaste, 1-10 m depth, 14 February 1972, 3 shells, leg. P. LaFollette and D. Cadien (LACM 72- 19.45). Bahia Potrero Grande (10°50'56”N, 85°48’35”W), Provincia Guanacaste, 9-10 m depth, 17 February 1972, 6 shells, leg. P. LaFollette and A. Ferreira (LACM 72-29.11). Middle of Bahia Huevos, north of Bahia Culebra (10°38'41"N, 85°41'55”W), Provincia Guanacaste, 10-23 m depth, 18-19 February 1972, 18 shells, 478 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 leg. P. LaFollette and D. Cadien (LACM 72-35.42). Off beach at Bahia Brasilito (10°25'57"N, 85°49'18”W), Provincia Guanacaste, 18 m depth, 20 February 1972, 11 shells, leg. P. LaFollette and D. Cadien (LACM 72- 40.33). Anchorage at Bahia Ballena off Tambor (10°44'10”N, 84°59'34”W), Provincia Puntarenas, 10 m depth, 21 February, 1972, 2 shells, leg. P. LaFollette and A. Ferreira (LACM 72-43.18). Off beach at Bahia Ballena (9°44'12""N, 84°59'32”W), Provincia Puntarenas, 3—18 m depth, 23 February 1972, 5 shells, leg. P. LaFollette and D. Cadien (LACM 72-45.40). Off Bahia Herradura (9°38'50”N, 84°40'50”W), Provincia Puntarenas, 37 m depth, 10 March 1972, 11 shells, leg. J. McLean (LACM 72-54.54). Anchorage inside of small islet 1.5 km south of Punta Quepos (9°22'43”N, 84°09'41"W), Provincia Puntarenas, 21 m depth, 11 March 1972, | shell, leg. J. McLean (LACM 72-57.39). Playa Nancite (10°48’N, 85°42'W), north side of Golfo Papagayo, Provincia Guanacaste, 15 January 1986, 2 shells, leg. E. Coan and R. Hollywood (LACM 86-26.35). One km west of Playa Junquillal (10°58'24.5393N, 85°42'04.1590W). Bahia Junquillal, Provincia Guanacaste, 30 m depth, 1 shell, 5 April 2002, leg. F. Gonzalez (INB0003503969). One and half km northwest of Playa Junquillal (10°58'40.7139N, 85°42'27.2867W), Bahia Junquillal, Provincia Guanacaste, 20 m depth, | shell, 4 April 2002, leg. F. Gonzalez (INB0003539752). South of Isla Juanilla (10°58'55.1750N, 85°43'10.1707W), Murciélago, Provincia Guanacaste, 15 m depth, 1 shell, 9 April 2002, leg. F. Gonzalez (INB0003539769). Between Isla Bolafios and Playa Coyotera (11°02'51.0972N, 85°43'21.1040), Bahia Salinas, Provincia Guanacaste, 12 m depth, 2 shells, 10 April 2002, leg. F. Gonzalez (INB0003539771). Four hundred m west of Isla Cabo Blanco (9°32'29.2960N, 85°07'16.0340W), Cabo Blanco, Provincia Puntarenas, 20-50 m depth, 4 shells, 16 May 1998, leg. A. Berrocal (INB0003539975). Isla David (10°58'05.5000N, 85°42'38.6000), Bahia Junquillal, Provincia Guanacaste, 30 m depth, 19 shells, 12 March 1996, leg. Y. Camacho (INB0003542912). Isla David (10°56'32.2000N, 85°42'51.1000), Santa Rosa, Provincia Guanacaste, 18 m depth, 16 shells, 12 March 1996, leg. Y. Camacho (INB0003575622). Playa Matapalo (9°21'48.1193N, 84°05'33.8006W), National Park Manuel Antonio, Provincia Puntarenas, 10-15 m depth, | shell, 18 February 2003, leg. F. Gonzalez (INB0003708499). Playa Ballena (9°06'34.2870N, 83°41'46.4390W), Uvita, Provincia Puntarenas, 1 m depth, | shell, 16 March 1996, leg. Y. Camacho (INB0003722812). Muelle de la Playa San Miguel (9°34'36.9 LION, 85°08'06.3610W), Reserva Natural Absoluta de Cabo Blanco, Provincia Puntarenas, 0 m depth, 14 shells, 2 May 1995, leg. G Mena (INB0003722819). Punta Oliva (9°34'39.3370N, 85°08'15.3840W), Cabo Blanco, Provincia Puntarenas, 0 m depth, 12 shells, 16 September 1995, leg. F. Alvarado (INB0003722820). Playa Mostrencal (10°59'47.3060N, 85°42'53.8890W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 2 m depth, 2 shells, 13 March 1996, leg. R. Angulo (INB0003722825). Bahia Junquillal (10°59'13.03 LON, 85°43'16.7950W), Golfo Santa Elena, Provincia Guanacaste, 3 m depth, 5 shells, 13 March 1996, leg. R. Angulo (INB0003722828). Isla David (10°58'45.4310N, 85°43'01.8500W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 0-3 m depth, 9 shells, 13 March 1996, leg. R. Angulo (INB0003747336). Isla David (10°56'06.0000N, 85°42'53.0000W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 18 m depth, | shell, 12 March 1996, leg. R. Angulo (INB0003747062). SHELL MORPHOLOGY.— Length up to 5.5 mm; width up to 2 mm in the largest specimen examined. Shell fragile, elongate, with nearly parallel sides (Fig. 6D—E). Only one whorl visible, forming the entire shell. Anterior end of the shell rounded. Umbilicus absent. Aperture long, wider anteriorly, narrowing gradually at about 's of its length. Aperture bended apically over the apex of the shell, with the parietal wall forming a conical spine that completely covers the apex. Spine long and more curved in some specimens (Fig. 6D) and shorter and straight in some others (Fig. 6E). Columellar margin thickened, with a small protuberance. Sculpture composed of a number of irreg- ular spiral grooves crossed by numerous fine growth lines all over the shell surface. Spiral grooves situated near the anterior and posterior ends of the shell more conspicuous and separated from each other by wider gaps than those in the center of the shell. Color uniformly shiny whitish. ANATOMY.— AIl the specimens examined consisted of empty shells so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— Known from southern California to Panama and the Galapagos Islands (Keen 1971). REMARKS.— Carpenter (1864) described Volvula cylindrica from Santa Barbara, California, VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 479 based on a single shell. The original description is brief and offers few details. Examination of the holotype, illustrated by Palmer (1958), revealed that the specimens here examined match the orig- inal description of this species. The shell morphology of this species is variable, with some specimens having a more elongate and curved apical spine and other specimens with a short and straight spine. The material exam- ined includes intermediate forms within the entire range of variability. This shell variability is probably the cause of the introduction of several names for this species. Three synonyms have been recognized in the Panamic Province: Volvulella callicera Dall, 1919, from the Galapagos, and Volvulella cooperi Dall, 1919 and Volvulella lowei Strong and Hertlein, 1937 from the Gulf of California. The study of the type material of all these species confirmed that they all fit within the variablility of V. cylindrica. Volvulella californica Dall, 1919, originally described from southern California, was redescribed by Gosliner (1996), who argued that it is a distinct species from V. cylindrica. According to Gosliner (1996), V. californica is consistently devoid of sculpture and pyriform in shape, whereas V. cylindrica has a sculpture and is far more cylindrical. Volvulella catharia Dall, 1919 (Fig. 6C) TYPE MATERIAL.— Volvulella catharia: SYNTYPE: Bahia de Panama (USNM 211784). MATERIAL EXAMINED.— Bahia Chatham (5°33'20”N, 86°59'10"W), Cocos Island, 26 m depth, 13 January 1938, 2 shells (LACM 38-179.5). Ten miles off Punta Guiones (9°43.5’N, 85°44.0’W), Provincia Guanacaste, 320-457 m depth, 13 May 1973, 2 shells (LACM 73-65.4). SHELL MORPHOLOGY.— Length up to 4 mm; width up to 2 mm in the largest specimen exam- ined. Shell fragile, oval, with convex sides (Fig. 6C). Only one whorl visible, forming the entire shell. Anterior end of the shell rounded. Umbilicus absent. Aperture long, wider anteriorly, narrow- ing gradually at about '/; of its length. Aperture bended apically over the apex of the shell, with the parietal wall forming a short, blunt protuberance that covers the apex. Columellar margin thick- ened. Sculpture composed of irregular spiral grooves faded in the specimens examined. Color uni- formly pale brown. ANATOMY.— All the specimens examined consisted of empty shells so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— Known from Costa Rica and Panama (Keen 1971). REMARKS.— Dall (1919) described Volvulella catharia as a small species with a short spine at the apex, subcylindrical and smooth. Examination of the holotype (USNM 211784) revealed that the specimens here studied from Costa Rica belong to the same species. Volvulella catharia is clearly different from other Panamic species of Volvulella. The shell of V. catharia is shorter and proportionally wider, with a small and rounded apical extension. This species is provisionally placed in Volvulella until complete specimens become available for study. Volvulella panamica Dall, 1919 (Fig. 6F) Volvulella panamica Dall. 1919:298. Volvulella tenuissima Willett. 1944:71—72, pl. 14, fig. 1. TYPE MATERIAL.— Volvulella panamica: HOLoTyPE: Bahia de Panama (USNM 212654). Volvulella tenuissima: HOoLoTyPE: LACM 1073 (1 dry shell) 137 m, off Redondo Beach, Los Angeles County, California. 480 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 FIGURE 6. A, Retusa paziana Dall, 1919, shell morphology, specimen from Isla David (INB0003719254), 2.5 mm long. B, Retusa sp., shell morphology, specimen from Punta Ballena (LACM 72-42.61), 1.8 mm long. C, Volvulella catharia Dall, 1919, shell morphology, specimen from Punta Guiones (LACM 73-65.4), 4mm long. D—E, Volvulella cylin- drica (Carpenter, 1864); (D) Shell morphology, specimen from Bahia Junquillal. (INB0003747336), 5 mm long; (E) Shell morphology, specimen from Bahia Brasilito, 4 mm long (LACM 72-40.33). F. Volvulella panamica Dall, 1919, shell mor- phology, specimen from Bahia Herradura (LACM 72-54.5 55), 5.5 mm long. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 48] MATERIAL EXAMINED.— Bahia Cocos (10°33'35"N, 85°42'30” W), south of Puerto Culebra, Provincia Guanacaste , 4 m depth, 13 March 1933, 2 shells (LACM 33-123.30). Off Bahia Herradura (9°38'50’N, 84°40'50” W), Provincia Puntarenas, 37 m depth, 10 March 1972, 7 shells, leg. J. McLean (LACM 72-54.55). SHELL MORPHOLOGY.— Length up to 5.5 mm; width up to 2 mm in the largest specimen examined. Shell fragile, elongate, with nearly parallel sides (Fig. 6F). Only one whorl visible, forming the entire shell. Anterior end of the shell rounded. Umbilicus absent. Aperture long, wider anteriorly, narrowing gradually at about '/s of its length. Aperture bended apically over the apex of the shell, with the parietal wall forming a short spine that completely covers the apex. Spine sharp and narrow because of the presence of a rounded notch on the posterior end of the spire. Columellar margin thickened, with a small protuberance. Sculpture composed of a number of irregular spiral grooves crossed by numerous fine growth lines all over the shell surface. Spiral grooves situated near the anterior and posterior ends of the shell more conspicuous and separated from each other by wider gaps than those in the center of the shell. Color uniformly shiny whitish. ANATOMY.— All the specimens examined consisted of empty shells so anatomical examina- tions were not possible. GEOGRAPHIC RANGE.— This species in known from southern California to Panama (Keen 1971; Gosliner 1996). REMARKS.— Volvulella panamica was described by Dall (1919) on the basis of a single shell from Panama Bay that had a very short and sharp apical spine. Examination of the holotype (USNM 212654) revealed that the spine morphology is caused by the presence of a rounded notch on the posterior end of the spire, not by shell damage or a preservation artifact. Examination of sev- eral shells from Costa Rica of different sizes revealed a similar morphology of the apex, which sus- tains the separation of this species from Volvulella cylindrica. Volvulella tenuissima, originally described from southern California by Willett (1944) has a similar morphology and constitutes a synonym. The study of the holotype (LACM 1073), showed the presence of the sharp apical spine typical of this species. Family Haminoeidae Pilsbry, 1895 Genus Haminoea Turton, 1830 Haminoea ovalis Pease, 1868 (Figs. 7A, 8A-C, 9) Haminoea ovalis Pease, 1868:71, pl. 7, fig. 2, pl. 12, fig. 20. TYPE MATERIAL.— Haminoea ovalis: Untraceable. MATERIAL EXAMINED.— Punta Uvita (9°08'44.4560N, 83°45'42.2750W), Provincia Puntarenas, 0 m depth, 1 shell, 13 April 1996, leg. R. Angulo (INB0001486877). 500 m south of Playa Ventanas (9°05'16.1660N, 83°40'50.7600W), Provincia Puntarenas, 0 m depth, 2 specimens, 17 January 2000, leg. M. Calderon (INB0001495966). 500 m south of Playa Ventanas (9°05'16.1660N, 83°40'50.7600W), Provincia Puntarenas, 0 m depth, | specimen, 17 January 2000, leg. M. Calderon (INB0001496120). San Miguel (9°34'49 0680N, 85°08'28.5260), Cabo Blanco, Provincia Puntarenas, 2 m depth, 16 May 1998, | specimen, leg. A. Berrocal (INB0001496693). San Miguel (9°34'49.0680N, 85°08’28.5260), Cabo Blanco, Provincia Puntarenas, 2 m depth, 16 May 1998, 1 specimen, leg. A. Berrocal (INB0001496694). Puerto Escondido (9°23'02.7920N, 84°08'14.4875W), National Park Manuel Antonio, Provincia Puntarenas, 0 m depth, 3 spec- imens, 19 February 2003, leg. A. Berrocal (INB0003572107). National Park Manuel Antonio (9°23'02.6451N, 84°10'05.9237W), Provincia Puntarenas, 8-11 m depth, 1 specimen, 19 February 2003, leg. A. Berrocal (INB0003572120). Isla Ballena (9°06'18.0003N, 83°43'42.7017W), Provincia Puntarenas, 18 m depth, 18 January 2003, 1 specimen, leg. S. Avila (INB0003572558). Isla Santa Catalina (10°28'37.2000N, 482 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 85°52'06.6000W), Provincia Guana- caste, 15 m depth, 21 July 2000, 2 specimens, leg. M. Calderon (INB0003701269). SHELL | MORPHOLOGY.— Length up to 4 mm; width up to 3 mm in the largest specimen examined. Shell fragile, delicate, oval, with convex sides (Fig. 8A). Only one whorl visible, forming the entire shell. Apex rounded, with the aperture lip ris- ing from the left side. Anterior end of the shell rounded. Umbilicus absent. Aperture long, wide anteriorly, narrowing grad- ually at about '4 of its length. Columellar margin slightly thick- ened. Surface smooth, the sculp- ture being reduced to fine growth lines. Color uniformly pale brown. EXTERNAL MORPHOLOGY.— Body oval, 6 mm long in the largest specimen examined. Cephalic shield elongate, nar- rower posteriorly, comprising about '> of the body length (Fig. 7A). Hancock’s organs com- posed of about 6 simple folds each. Parapodia short and nar- row, covering a small portion of the shell laterally. Posterior half of the body almost entirely cov- ered by the shell. Gill unipinnate, with 15 simple lamellae. Color of living animals pale bluish gray, with darker areas on anterior cen- tral region of the cephalic shield and the posterior lateral sides, as S ; z oN , . well as in some irregular areas on FIGURE 7. Living animals; (A) Haminoea ovalis Pease, 1868; (B) Bulla punctulata A. Adams in Sowerby, 1850; (C) Navanax aenigmaticus (Bergh, 1894). the rest of the body. Entire sur- face of the body covered by small and bright orange dots. Shell translucent, almost transparent. Area covered by the shell clear- ly visible, with the same color as the rest of the body and the orange dots larger in diameter. ANATOMY.— Buccal bulb oval to elongate, connecting posteriorly to a short esophagus and two long salivary glands (Fig. 9A). Esophagus opening into a large gizzard, which contains three VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 483 FiGuRE 8. A-C, Haminoea ovalis Pease, 1868; (A) Shell morphology, specimen from Punta Uvita (INB0001486877), 4 mm long; (B) Radular teeth, specimen from Isla Ballena (INB0003572558); (C) Gizzard plate, same specimen. D—G, Atys exarata Carpenter, 1857; (D) Shell morphology, specimen from Punta Descartes (INB0003747331), 5 mm long; (E) Shell morphology, specimen from between Punta Piedra and Punta Gallardo (INB0003540313), 4 mm long; (F) Gizzard plate, specimen from Punta Piedra (INB0001496350); (G) Radular teeth, same specimen. smooth and irregular gizzard plates. All plates are similar in shape and size, each one having a series of transverse, parallel ridges composed of several rows of small, simple denticles (Fig. 8C). Radular formula 19 x 11.1.11 in a specimen from Isla Ballena (INB0003572558). First inner- 484 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 FIGURE 9. Haminoea ovalis Pease, 1868, anatomy of specimen from Isla Ballena (INB0003572558); (A) Anterior portion of the digestive system; (B) Female reproductive system; (C) Male reproductive system. Abbreviations: am, ampulla; bb, buccal bulb; be, bursa copulatrix; e, esophagus; fg, female glands; gz, gizzard; pn, penis; pr, prostate; sg, salivary gland. most teeth with a wide triangular cusp bearing several small denticles (Fig. 8B). Lateral teeth hook-shaped with a strong cusp lacking denti- cles. Rachidian teeth broad, with a wide base, a single central cusp, and one denticle on each side of the cusp. Reproductive system monoaulic. Penis bulbous, connected to the prostate through a short duct (Fig. 9C). Ampulla long and convo- C luted, connecting to a short and wide post- ampullary duct that opens into the female glands. Bursa copulatrix entering the female glands (Fig. 9B). GEOGRAPHIC RANGE.— This is a widespread Indo-Pacific species. This is the first record of the species for the eastern Pacific. REMARKS.— The specimens here examined resemble the external coloration of Haminoea ovalis Pease, 1868, originally described from Tahiti, French Polynesia. Haminoea angelensis Baker and Hanna, 1927, was described on the basis of shell morphology (Baker and Hanna 1927), so it is impossible to determine its taxonomic status. Two other Panamic species, Haminoea vesic- ula (Gould, 1855) and Haminoea virescens (Sowerby, 1833) have been illustrated by Behrens (1991) and clearly differ from H. ovalis by lacking the bright orange spots characteristic of this species. 1mm Genus Adys de Monfort, 1810 Atys exarata (Carpenter, 1857) (Figs. 8D-G) Bulla exarata Carpenter, 1857:173. Atys casta Carpenter, 1864:314. Atys chimera Baker and Hanna, 1927:126, pl. 4, fig. 4. Cylichna veleronis Strong and Hertlein, 1939:191. Cylichna stephensae Strong and Hertlein, 1939:190. Ans liriope Hertlein and Strong, 1951:71, pl. 8, fig. 2. TYPE MATERIAL.— Bulla exarata: HOLOTYPE: Mazatlan, Sinaloa, Mexico (BMNH). Atys casta: LECTOTYPE and PARALECTOTYPE: Cabo San Lucas, Baja California Sur, Mexico (USNM 4014). Atvs chimera: HOLOTYPE: Puerto Escondido, Baja California Sur, Mexico (CASIZ 032115). Cylichna veleronis: PARATYPE: Bahia Honda, Isla Coiba, Panama (LACM 1340). Cylichna stephen- sae: PARATYPE: Bahia Honda, Isla Coiba, Panama (LACM 1339). Atvs liriope: HOLOTYPE: Arena Bank, Gulf of California (CASIZ 065495). MATERIAL EXAMINED.— Bahia Cocos (10°33'35”N, 85°42'30"W), south of Puerto Culebra, Provincia Guanacaste, 4 m depth, 13 March 1933, 5 shells (LACM 33-123.31). Playa Blanca (10°56’45’N, VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 485 5°53'30"W), Bahia Playa Blanca, Provincia Guanacaste, 3—5 m depth, 8 February 1935, 7 shells (LACM 35- 00.9). Playa Blanca (10°56'45"N, 85°53'30"W), Bahia Playa Blanca, Provincia Guanacaste, 73 m depth, 935, 8 shells (LACM 35-103.15). Southeast end of Bahia Santa Elena, approximately half mile offshore (10°55'15"N, 85°48'30"W), Provincia Guanacaste, 9-12 m depth, 13 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72-9.17). Southeast corner of Bahia Jobo, off sand beach west of Bahia Salinas (11°02'22"N, 85°45'16”W), Provincia Guanacaste, 1-10 m depth, 14 February 1972, 13 shells, leg. P. LaFollette and D. Cadien (LACM 72-19.46). Cove between Isla San José and Isla Cocinero (10°51'50"N, 86°55'30" W), Islas Murciélago, Provincia Guanacaste, 3-11 m depth, 17 February 1972, | shell, leg. P. LaFollette, D. Cadien and A. Ferreira (LACM 72-21.19). Middle of Bahia Huevos, north of Bahia Culebra (10°38’41"N, 85°41'55”W), Provincia Guanacaste, 10-23 m depth, 18-19 February 1972, 17 shells, leg. P. LaFollette and D. Cadien (LACM 72-35.43). Off beach at Bahia Brasilito (10°25'57"N, 85°49'18”"W), Provincia Guanacaste, 18 m depth, 20 February 1972, 16 shells, leg. P. LaFollette and D. Cadien (LACM 72- 40.34). Off beach at Bahia Ballena (9°44'12"N, 84°59'32”W), Provincia Puntarenas, 3-18 m depth, 23 February 1972, 7 shells, leg. P. LaFollette and D. Cadien (LACM 72-45.41). Anchorage inside of small islet 1.5 km south of Punta Quepos (9°22'43”N, 84°09'41"W), Provincia Puntarenas, 21 m depth, 11 March 1972, 1 shell, leg. J. McLean (LACM 72-57.40). North side of Isla del Cafio (8°43'15"N, 83°53'07"W), Provincia Puntarenas, 7-14 m depth, 14-19 March 1972, 6 shells, leg. J. McLean and D. Wheeler (LACM 72-63.80). 250 m off Punta Piedra (8°35'24.6720N, 83°11'26.3250W), Provincia Puntarenas, 23 m depth, 14 April 1997, 12 shells, leg. M. Lobo (INB0001496350). 1 km west of Playa Junquillal (10°58’24.5393N, 85°42'04.1590), Provincia Guanacaste, 30 m depth, 5 April 2002, 1 shell, leg. F. Gonzalez (INB0003503527). 1 km west of Playa Junquillal (10°58’24.5393N, 85°42'04.1590), Provincia Guanacaste, 30 m depth, 5 April 2002, 1 shell, leg. F. Gonzalez (INB0003503528). South of Isla Juanilla (10°58'55.1750N, 85°43'10.1707W), Murciélago, Provincia Guanacaste, 15 m depth, 2 shells, 9 April 2002, leg. F. Gonzalez (INB0003504414). From 500 m off the beach to 2.5 km southeast (8°34'39.1040N, 83°11'29.6540W), Punta Piedra, Provincia Puntarenas, 14-90 m depth, 10 April 1997, 25 shells, leg. M. Madrigal (INB0003540313). Between Punta Piedra and Punta Gallardo (8°35'47.5010N, 83°11'58.9990W), Provincia Puntarenas, 30-35 m depth, 13 June 1997, 8 shells, leg. M. Madrigal (INB0003540366). 1 km west of Playa Junquillal (10°58'24.5393N, 85°42'04.1590W). Bahia Junquillal, Provincia Guanacaste, 30 m depth, 4 shells, 5 April 2002, leg. F. Gonzalez (INB0003542436). Isla David (10°56'32.2000N, 85°42'51.1000W), Bahia Cuajiniquil, Provincia Guanacaste, 18 m depth, 12 March 1996, 2 shells, leg. Y. Camacho (INB0003575625). Isla David (10°56'06.0000N, 85°42'53.0000W), Bahia Cuajiniquil, Provincia Guanacaste, 18 m depth, 12 March 1996, 2 shells, leg. S. Avila (INB0003575665). Playa Blanca (8°38'18.0525N, 83°26'15.7570W), Provincia Puntarenas, 0 m depth, 5 April 1995, 1 shell, leg. G Mena (INB0003722817). Bahia Junquillal (10°58'45.4310N, 85°43'01.8500W), Provincia Guanacaste, 0-3 m depth, | shell, 13 March 1996, leg. R. Angulo (INB0003722829). Boca de Estero Caballero (8°40'06.3050N, 83°26'41.8580W), Provincia Puntarenas, 0 m depth, 11 June 1995, 1 shell, leg. M. Lobo (INB0003747046). Golfo Santa Elena (10°59'13.0310N, 85°43'16.7950W), Bahia Junquillal, Provincia Guanacaste, 3 m depth, 13 March, 1996, 1 Shell. leg. R. Angulo (INB0003747330). Punta Descartes (11°02'21.8310N, 85°43'11.0470W), Playa Coyotera, Provincia Guanacaste, 0 m depth, 14 March 1995, 1 shell, leg. G Bassey (INB0003747331). SHELL MORPHOLOGY.— Length up to 6 mm; width up to 3 mm in the largest specimen exam- ined. Shell fragile, oval, with convex sides (Figs. 8D—E). Shell morphology very variable, some specimens are wider (Fig. 8E) whereas others are more elongate (Fig. 8D). Only one whorl visible, forming the entire shell. Apex rounded, depressed, not umbilicated, with the aperture lip conspic- uously rising from the left side. The aperture lip forms a short wing posteriorly, longer in some specimens (Fig. 8D). Anterior end of the shell rounded. Umbilicus absent. Aperture long, wide anteriorly, narrowing at about '4 of its length continuing in a narrow canal to the end of the shell where it widens again. Columellar margin slightly oblique, with a thickened margin and no folds. Sculpture with a number of simple spiral grooves, less conspicuous in the middle area of the shell, and more densely concentrated near the anterior and posterior ends. Color uniformly whitish to pale gray. ANATOMY.— Digestive system with a gizzard containing three irregular gizzard plates. Each 486 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 plate has a series of transverse, parallel ridges composed of small, simple denticles (Fig. 8F). All plates are similar in shape and size. The radular formula is 23 x 5.1.5 in a specimen from Punta Piedra (INB0001496350). The innermost lateral teeth are hook-shaped with a long and strong cusp bearing numerous denticles (Fig. 8G). The remaining four lateral teeth have a simple cusp, with no denticles. The rachidian teeth are broad, with a triangular base and a wide central cusp lacking den- ticles. GEOGRAPHIC RANGE.— Known from southern California to Panama (Skoglund 2002). REMARKS.— Bulla exarata was introduced based on a shell collected from Mazatlan, Sinaloa, Mexico, with a short description with no illustrations. The holotype of this species (BMNH) is an elongate shell with numerous and conspicuous longitudinal striations and the aperture lip conspic- uously rising from the left side. These and other features of the specimen are very similar to those of the material here examined. A few years later, Carpenter (1864) described Atys casta, with a short text and no illustrations, based on a single shell collected from Cabo San Lucas, Baja California Sur, Mexico. Baker and Hanna (1927) considered the description of A. casta “so vague and indefinite that identifications based thereupon would be entirely untrustworthy.” Consequently these authors introduced a new name, Atvs chimera for similar shells collected from Puerto Escondido, La Paz, and Bahia Concepcion, Baja California Sur, Mexico. A re-examination of a syntype of Atvs casta (USNM 4014) revealed that this specimen is very similar in shell morphology to the specimens here exam- ined, which also agrees with the original description of 4. chimera and the features of the holotype (CASIZ 032115). Three other synonyms of Atvs exarata are Cylichna veleronis Strong and Hertlein, 1939, Cylichna stephensae Strong and Hertlein, 1939, and Atvs liriope Hertlein and Strong, 1951. Both C. veleronis and C. stephensae were described based on shells dredged from Bahia Honda, Panama, and were only compared to species of Cylichna. Examination of the type material of these two taxa shows remarkable similarities with the material here examined and all the shells fit with- in the variability described for this species. Cylichna stephensae was described for wider shells with fewer and more distant spiral lines, whereas C. veleronis was described for narrower shells with more highly compressed spiral lines. Atys liriope was described by Hertlein and Strong (1951) as different from Atys chimera by having a more closely spaced and conspicuous sculpture. These differences are explainable by the normal variability of the species, and there is no doubt that 4. /irope is a synonym of A. exarata. This species is placed in Atys because of the presence of transverse, parallel ridges composed of denticles in the gizzard plates, and the radular morphology, which has broad rachidian teeth, with a wide triangular base and a broad central cusp bearing numerous small denticles as well as hook-shaped lateral teeth with a long and strong cusp bearing numerous denticles on the inner lat- eral teeth. Atys defuncta (Baker and Hanna, 1927) (Figs. 1OA—D) Cylichnella defuncta Baker and Hanna, 1927:127—128, pl. 4, fig. 3. TYPE MATERIAL.— Cylichnella defuncta: HOLOTYPE: Bahia Amortajada, Isla San José, Baja California Sur, Mexico (CASIZ 032118). MATERIAL EXAMINED.— Bahia Cocos (10°33 Guanacaste, 4 m depth, 13 March 1933, 2 shells (L Provincia Guanacaste, 31 m depth, 13 March 1934, '35’N, 85°42'30”W), south of Puerto Culebra, Provincia ACM 33-123.33). Puerto Culebra (10°35'N, 85°40’W), 8 shells (LACM 33-153.14). Bahia Chatham (5°33’N, VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 487 86°59'W), Cocos Island, 31 m depth, 28 February 1933, 7 shells (LACM 33-154.1). Puerto Parker (10°55’N, §5°49'W), Golfo Santa Elena, Provincia Guanacaste, 8 February 1935, 2 shells (LACM 35-138.4). Bahia Chatham (5°33'20"N, 86°59'10"W), Cocos Island, 26 m depth, 13 January 1938, 1 shell (LACM 38-179.6). Playa Blanca (10°56'45”N, 85°53'30"W), Bahia Playa Blanca, Provincia Guanacaste, 3—5 m depth, 8 February 1935, 29 shells (LACM 35.100.10). Puerto Parker (10°57'50”"N, 85°48'45”W), Golfo Santa Elena, Provincia Guanacaste, 55 m depth, 9 February 1935, 9 shells (LACM 35-113.28). Bahia Salinas (11°03'33’N, §5°44'05”",W), Provincia Guanacaste, 11 February 1935, 9 shells (LACM 35-122.17). Between Punta Isla and 500 m south of Punta Isla (10°56'00”N, 85°48'55”"W), Bahia Santa Elena, Provincia Guanacaste, 1-11 m depth, 13 February 1972, 5 shells, leg. P. LaFollette and D. Cadien (LACM 72-7.34). One mile offshore, between Bahia Santa Elena and Bahia San Juanillo (10°57'20"N, 85°46'08”W), Provincia Guanacaste, 25—53 m depth, 14 February 1972, 2 shells, leg. P. LaFollette and D. Cadien (LACM 72-12.49). Southeast corner of Bahia Jobo, off sand beach west of Bahia de Salinas (11°02'22"N, 85°45'16”W), Provincia Guanacaste, 1—10 m depth, 14 February 1972, 24 shells, leg. P. LaFollette and D. Cadien (LACM 72-21.47). Cove between Isla San José and Isla Cocinero (10°51'50”N, 86°55'30"W), Islas Murciélago, Provincia Guanacaste, 3-11 m depth, 17 February 1972, 1 shell, leg. P. LaFollette, D. Cadien, and A. Ferreira (LACM 72-21.20). Bahia Potrero Grande (10°50'56"N, 85°48'35"W), Provincia Guanacaste, 9-10 m depth, 17 February 1972, 5 shells, leg. P. LaFollette and A. Ferreira (LACM 72-29.13). South tip of Punta Santa Elena (10°53'35’N, 85°57'52” W), Provincia Guanacaste, 12-15 m depth, 18 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-30.27). 5 miles north west of Islas Huevos (10°41'45"N, 85°46'25”W), Golfo de Papagayo, Provincia Guanacaste, 60-64 m depth, 18 February 1972, 1 shell, leg. P. LaFollette and D. Cadien (LACM 72-34.30). Middle of Bahia Huevos, north of Bahia Culebra (10°38’41"N, 85°41'55”"W), Provincia Guanacaste, 10-23 m depth, 18-19 February 1972, 11 shells, leg. P. LaFollette and D. Cadien (LACM 72- 35.45). Off beach at Bahia Brasilito (10°25'57”N, 85°49'18”W), Provincia Guanacaste, 18 m depth, 20 February 1972, 46 shells, leg. P. LaFollette and D. Cadien (LACM 72-40.36). 1.5 mile east of Punta Ballena (9°44'15"N, 84°33'45”W), Provincia Guanacaste, 3-15 m depth, 21—22 February 1972, 4 shells, leg. P. LaFollette and D. Cadien (LACM 72-42.62). Anchorage at Bahia Ballena off Tambor (10°44’10’N, 84°59'34”W), Provincia Puntarenas, 10 m depth, 21 February, 1972, 30 shells, leg. P. LaFollette and A. Ferreira (LACM 72-43.19). Off beach at Bahia Ballena (9°44'12"N, 84°59'32”W), Provincia Puntarenas, 3-18 m depth, 23 February 1972, 4 shells, leg. P. LaFollette and D. Cadien (LACM 72-45.42). Bahia Herradura (9°38'45"N, 84°40'55”W), Provincia Puntarenas, 9-17 m depth, 9-10 March 1972, 4 shells, leg. J. McLean (LACM 72-52.63). Bahia Herradura (9°37'58”N, 84°40'30"W), Provincia Puntarenas, 21 m depth, 9 March 1972, 31 shells, leg. J. McLean (LACM 72-53.30). Off Bahia Herradura (9°38'50"N, 84°40’50"W), Provincia Puntarenas, 37 m depth, 10 March 1972, 30 shells, leg. J. McLean (LACM 72-54.57). Anchorage inside of small islet 1.5 km south of Punta Quepos (9°22'43"N, 84°09’41"”W), Provincia Puntarenas, 21 m depth, 11 March 1972, 29 shells, leg. J. McLean (LACM 72-57.41). Small islets off Punta Quepos (9°22'43"N, 84°09'41” W), Provincia Puntarenas, 9-23 m depth, 11-13 March 1972, 3 shells, leg. J. McLean (LACM 72- 58.59). Small islets off Punta Quepos (9°22'12’N, 84°09'15”W), Provincia Puntarenas, 23 m depth, 12 March 1972, 2 shells, leg. J. McLean (LACM 72-59.31). North side of Isla del Cafio (8°43'15"N, 83°53’07"W), Provincia Puntarenas, 7-14 m depth, 14-19 March 1972, 26 shells, leg. J. McLean and D. Wheeler (LACM 72-63.81). Isla del Cafio (8°44’00’N, 83°52'30”W), Provincia Puntarenas, 12 m depth, 14-19 March, 1972, 27 shells, leg. J. McLean (LACM 72-64.19). 1.5 km northwest of Playa Junquillal (10°58’40.7139N, 85°42'27.2867W), Provincia Guanacaste, 20 m depth, 4 April 2002, 1 shell, leg. F. Gonzalez (INB0003501383). Manuel Antonio National Park (9°22'36.6614N, 84°09’23.2807W), Provincia Puntarenas, 15-20 m depth, 1 specimen, 17 February 2003, leg. F. Gonzalez (INB0003575268). Isla David (10°56'06.0000N, 85°42'53.0000W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 18 m depth, 4 shells, 12 March 1996, leg. R. Angulo (INB0003575678). Isla David (10°56'06.0000N, 85°42'53.0000W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 18 m depth, 6 shells, 12 March 1996, leg. R. Angulo (INB0003575681). National Park Manuel Antonio (9°22'36.6614N, 84°09'23.2807W), Provincia Puntarenas, 15-20 m depth, 4 shells, 17 February 2003, leg. F. Gonzalez (INB0003707722). La Viuda Rock (8°36'56.0065N, 83°14'08.0868W), Golfo Dulce, Provincia Puntarenas, 66 m depth, 21 April 1997, 1 shell, leg. M. Lobo (INB0003722832). Boca de Estero Caballero (8°40'06.3050N, 83°26'41.8580W), Provincia Puntarenas, 0 m depth, 11 June 1995, 1 shell, leg. M. Lobo (INB0003747045). Playa Mostrencal 488 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 (10°59’47.3060N, 85°42'53.8890W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 2 m depth, 1 shell, 13 March 1996, leg. R. Angulo (INB0003747048). Playa Mostrencal (10°59'47.3060N, 85°42'53.8890W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 2 m depth, 3 shells, 13 March 1996, leg. R. Angulo (INB0003747334). Playa Mostrencal (10°59'47.3060N, 85°42'53.8890W), Bahia Junquillal, Golfo Santa Elena, Provincia Guanacaste, 2 m depth, 3 shells, 13 March 1996, leg. R. Angulo (INB0003747410). Isla David (10°58’05.5000N, 85°42'38.6000W), Bahia Junquillal, Santa Rosa National Park, Provincia Guanacaste, 30 m depth, 12 March 1996, 3 shells, leg. Y. Camacho (INB0003747596). SHELL MORPHOLOGY.— Length up to 5 mm; width up to 2 mm in the largest specimen exam- ined. Shell fragile, elongate, with convex sides, much wider in the central area (Fig. 1|OA—B). Only one whorl visible, except for a visible portion of the top of the spire. Apex umbilicated with the aperture lip conspicuously rising from the left side, forming a rounded wing connected to the col- umellar margin. Anterior end of the shell elongate. Umbilicus wide and deep. Aperture long, wider anteriorly, narrowing at about ' of its length and widening again near the apex. Columella with a single, conspicuous fold. Sculpture with several spiral grooves near the anterior and posterior ends of the shell, absent from the middle area. Color uniformly dirty white. ANATOMY.— Digestive system with a gizzard containing three irregular gizzard plates. Each plate with a series of transverse, parallel ridges composed of small, simple denticles (Fig. 10C). All plates similar in shape and size. Radular formula 10 x 3.1.3 in a specimen from Manuel Antonio (INB0003575268). Lateral teeth hook-shaped with a long and strong cusp bearing numerous den- ticles (Fig. 10D). Rachidian teeth broad, with a wide triangular base and a broad central cusp bear- ing numerous small denticles. REMARKS.— The shell morphology of the specimens here examined 1s identical to the origi- nal description of Cylichnella defuncta by Baker and Hanna (1927). This species was originally described in the genus Cylichnella and subsequently transferred to Acteocina by Skoglund (2002), based on the synonymization of Cylichnella with Acteocina by Mikkelsen and Mikkelsen (1984). However, the anatomy of Cylichnella defuncta is very different from those of Acteocina and Cylichnella. The gizzard plates have a series of transverse, parallel ridges composed of small, sim- ple denticles, and the radula has broad rachidian teeth, with a wide triangular base and a broad cen- tral cusp bearing numerous small denticles as well as hook-shaped lateral teeth with a long and strong cusp bearing numerous denticles. All these anatomical features as well as the shell morphol- ogy, characterized by the presence of striae near the posterior and anterior ends of the shell and the outer lip clearly rising over the apex, are typical of the genus Afys. Therefore C. defuncta is here regarded as a member of Anys. Family Bullidae Gray, 1827 Genus Bulla Linnaeus, 1758 Bulla punctulata A. Adams in Sowerby, 1850 (Figs. 7B, 10E-G, 11) Bulla punctata A. Adams in Sowerby, 1848—50 [1850]:577, pl. 123, fig. 77 (non Bulla punctata Schroeter, 1804). Bulla punctulata A. Adams in Sowerby, 1848-50 [1850]:604. TYPE MATERIAL.— Bulla punctata: SYNTYPES (2): Panama (BMNH 19760047). MATERIAL EXAMINED.— Playa Ocotal, Provincia Guanacaste, 14 May 1976, 8 shells, leg. A. Hardy and R. Hardy (CASIZ 048833). Puerto Culebra, Provincia Guanacaste, | shell (CASIZ 017943). Cove on northwest side of Isla San Pedrito (10°51'30"N, 86°57'57” W), Islas Murciélago, Provincia Guanacaste, 24 m depth, 17 February 1972, | juvenile shell, leg. P. LaFollette and D. Cadien (LACM 72-22.13). Islas Tortugas, VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 489 FIGURE 10. A-D. Atys defuncta (Baker and Hanna, 1927); (A) Shell morphology, specimen from Playa Junquillal (INB0003501383); 3 mm long; (B) Shell morphology, specimen from Parque Nacional Manuel Antonio, (INB0003575268) 3 mm long; (C) Gizzard plate, same specimen; (D) Radular teeth, same specimen. E—G, Bulla punctulata A. Adams in Sowerby, 1850; (E) Shell morphology, specimen from Golfo Dulce (INB0003540757), 7.5 mm long; (F) Gizzard plate, specimen from Isla Bolafios and Playa Coyotera (INB0003502783); (G) Radular teeth, same specimen. 490 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 Provincia Puntarenas, 2-8 m depth, 23-24 February 1972, 7 shells, leg. P. LaFollete, D. Cadien, and A. Ferreira (LACM 72-46.59). Anchorage inside of small islet 1.5 km south of Punta Quepos (9°22'43’'N, 84°09'41"” W), Provincia Puntarenas, 21 m depth, 11 March 1972, 4 juvenile shells, leg. J. McLean (LACM 72-57.42). Isla del Cario (8°44’00"N, 83°52'30"W), Provincia Puntarenas, 12 m depth, 14-19 March, 1972, | juvenile shell, leg. J. McLean (LACM 72-64.20). Boca del Estero Caballero (8°40'06.3050N, 83°26'41.8580W), Provincia Puntarenas, 0 m depth, 11 June 1995, 1 shell, leg. M. Lobo (INB0001499756). Murciélago (10°58'14.7752N, 85°42'04.1162W), Provincia Guanacaste, 20 m depth, 5 specimens, 6 April 2002, leg. F. Gonzalez (INB0003461244). 200 m northwest of Playa Guaria (10°57'55.1620N, 85°42'23.7922W), Bahia Junquillal, Provincia Guanacaste, 8 m depth, 4 April 2002, 1 shell, leg. F. Gonzalez (INB0003500963). Bahia Tomas (10°55'44.7444N, 85°43'15.9105W), Murciélago, Provincia Guanacaste, | m depth, 3 juvenile shells, 9 April 2002, leg. F. Gonzalez (INB0003501262). Bahia Tomas (10°55'44.7444N, 85°43'15.9105W), Murciélago, Provincia Guanacaste, | m depth, 2 shells, 9 April 2002, leg. F. Gonzalez (INB0003501266). Between Isla Bolanos and Playa Coyotera (11°02'51.0971N, 85°43'21.1040), Bahia Salinas, Provincia Guanacaste, 12 m depth, 10 April 2002, 2 specimens, leg. F. Gonzalez (INB0003502783). 1 km west of Playa Junquillal (10°58'24.5393N, 85°42'04.1590W), Bahia Junquillal, Provincia Guanacaste, 30 m depth, 5 April 2002, 5 shells, leg. F. Gonzalez (INB0003504311). Between Playa Palma and Playa Bejuco (8°43'10.6630N, 83°25'19.9570W), Golfo Dulce, Provincia Puntarenas, 0 m depth, 8 November 1996, 1 shell, leg. M. Lobo (INB0003540757). Bahia Santa Elena (10°54’48.0420N, 85°48'18.5810W), Provincia Guanacaste, 0 m depth, 4 shells, 16 February 1994, leg. G Bassey (INB0003542096). Isla Ballena (9°06'18.0003N, 83°43'42.7017W), Provincia Puntarenas, 18 m depth, 18 January 2003, 1 specimen, leg. S. Avila (INB0003572555). SHELL MORPHOLOGY.— Length up to 7 mm; width up to 5 mm in the largest specimen exam- ined. Shell solid, oval, with convex sides (Fig. 1OE). Only one whorl visible, forming the entire shell. Apex rounded, deeply umbilicated, with the aperture lip rising from the left side. Anterior end of the shell rounded. Umbilicus absent. Aperture long, wide anteriorly, narrowing gradually at about '4 of its length, and continuing posteriorly as a narrow canal. Columellar margin thickened. Surface smooth and shiny. Sculpture reduced to fine growth lines, except for the areas near the anterior and posterior ends of the shell, with A several conspicuous spiral lines. Color reddish brown, scattered with irregular white and black blotches. Columella white. y, EXTERNAL MORPHOLOGY.— Body is oval, S9—€€ 9 mm long in the largest specimen examined. A Cephalic shield short, comprising about '/ of the body length (Fig. 7B). Each side of the head with an involute lateral extension. Hancock’s organs composed of 7 simple folds. Posterior */ of the body almost entirely covered by the shell. Gill simple, composed of 15 unip- innate lamellae. Color of the living animals pale brown, with numerous small white spots. Shell completely opaque. ANATOMY.— Buccal bulb oval to elon- gate, connected posteriorly to the short esopha- gus and the long salivary glands (Fig. 11A). FIGURE 11. Bulla punctulata A. Adams in Sowerby, 1850, anatomy of specimen from Isla Bolafios and Playa ¢ Z : Coyotera (INB0003502783); (A) Anterior portion of the contains three smooth and irregular gizzard digestive system; (B) Male reproductive system. pn pr 1mm Esophagus opening into a large gizzard, which plates (Fig. 10F), all of them similar in shape Abbreviations: bb, buccal bulb; e, esophagus; gp, gizzard and size. Plates conical. with four ridges and _ Pate: gz. gizzard; pn, penis; pr, prostate; sg, salivary gland. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 49] longitudinal striations on the inner side. Radular formula 21 x 2.1.2 in a specimen from Bahia de Salinas (INB0003502783). Lateral teeth wide, with a short base and 5 sharp cusps (Fig. 10G). Rachidian teeth broad, short, with a small, triangular central cusp and four larger denticles on each side of the cusp. Reproductive system monoaulic. Penis small, connecting to the prostate through a long and convoluted duct (Fig. 11B). GEOGRAPHIC RANGE.— Reported in the eastern Pacific from Peru to Baja California, Mexico (Skoglund 2002). REMARKS.— The specimens here examined have similar shell morphology to the two syn- types of Bulla punctulata and there is no doubt they belong to the same species. Family Aglajidae Pilsbry, 1895 Genus Aglaja Renier, 1807 Aglaja regiscorona Bertsch, 1972 (Fig. 12A-B) Aglaja regiscorona Bertsch, 1972:103—104, fig 1. TYPE MATERIAL.— Aglaja regiscorona: HOLOTYPE: Bahia Las Cruces, Baja California del Sur, Mexico (CASIZ 024043); PARATYPES: Bahia Las Cruces, Baja California del Sur, México, three specimens (LACM 1617). MATERIAL EXAMINED.— San Miguel, Reserva Natural Absoluta de Cabo Blanco, Provincia Puntarenas, 2 m depth, 16 May 1998, 1 specimen, leg. A. Berrocal (INB0001498365); SW of Isla Plata, Provincia Guanacaste, 10 m depth, 17 April 2004, 1 specimen, leg. Y. Camacho (INB0003836172). SHELL MORPHOLOGY.— Length | mm; width 0.5 mm in the single specimen examined. Shell calcified with a distinct whorl, a curved apical border, and a broad flat wing (Fig. 12B). Protoconch globose, about 2 whorls, 200 um in diameter. EXTERNAL MORPHOLOGY.— Body elongate, 5 mm long in the specimen examined. Cephalic shield triangular, projecting posteriorly and upwards into a small, three pointed crown (Fig. 12A). Parapodia small, not extending over the dorsal surface. Posterior end of the body with two big lobes lacking a flagellum. Color of the living animal cream white. Dorsum and cephalic shield covered with numerous papillae, speckled with black. Parapodia and posterior lobes with several small black spots. Geographic range Known only from Bahia Las Cruces, Baja California Sur (Skoglund 2002) and Costa Rica (present paper). Remarks.— Aglaja regiscorona was originally described from Baja California based on sev- eral specimens collected from Bahia Las Cruces (Bertsch 1972). An examination of the type mate- rial has confirmed that the specimen from Costa Rica belongs to the same species. The shell mor- phology of this species is described here for the first time. Genus Navanax Pilsbry, 1895 Navanax aenigmaticus (Bergh, 1894) (Fig. 7C) Navarchus aenigmaticus Bergh, 1894:217, pl. 10, figs. 11-12, pl. 11, figs. 6-9, pl. 12, figs. 8-10. TYPE MATERIAL.— Untraceable. MATERIAL EXAMINED.— Bahia Alcyone, Cocos Island, 34 m depth, 25 March 1989, 1 specimen, leg. 492 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 K. Kaiser (CASIZ 073370). North side of Isla del Cafio (8°43'15"N, 83°53'07" W), Provin-cia Puntarenas, 0 m depth, 18—19 March 1972, 1 specimen, leg. J. McLean (LACM 72-68.44). Bahia Junquillal, Sector Sle (Cl0ksvaa ZONE 85°42'07.2810 W), Parque Nacional Santa Rosa, Provincia Guanacaste, | m depth, 9 December 1995, | speci- men, leg. NG Camacho (INB0001486933). Playa Grande GO2OS9RI02N i Sd -5l552 8818 W), 500 m W from Punta Carbon, Provincia Guanacaste, 0 m depth, 11 January 2001, 2 specimens, leg. S. Avila (INB0003118189). Playa San Miguel (9°34'45.8380N, 85°08'8 .6800W), Reserva Natural Absoluta de Cabo Blanco, Provincia Punt are- nas, 2 m depth, 30 April 1995, 1 specimen, leg. G. Mena (INBO00 1498434). Playa Manzanillo (9°56'03.4990N, 84°54'52.7660W), Isla Golfo de San Lucas, Provincia Puntarenas, 2 m depth, 9 December 1999, 2 specimens, leg. M. Calderon (INB0001496008). Estacion San Miguel (9°34'53.9500N, 85°08'28 .5380W), Reserva Natural Absoluta de Cabo Blanco, Provincia Punt are- nas, 0 m depth, 28 January 1999, | FIGURE 12. Aglaja regiscorona Bertsch, 1972, specimen from Cabo Blanco (INB0001498365); (A) Living animal; (B) Shell morphology, | mm long (complete). specimen, leg F. Alvarado (INB0001495820). Punta Uvita (9°08'44.4580N, 83°45'33.2680W), Parque Nacional Marino Ballena, Provincia Puntarenas, 0 m depth, 15 January 2000, 2 specimens, leg. M. Calderon (INB0001496175). Punta Uvita (9°08'50.9650N, 83°45'47.1900W), Parque Nacional Marino Ballena, 0 m depth, 13 April 1996, 6 specimens, leg. S. Avila (INB0001486546). East Side of Isla Ballena (9°06'24.5090N, 83°43'35.8230W), Parque Nacional Marino Ballena, Provincia Puntarenas, 6 m depth, 16 January 2000, 1 specimen, leg. M. Calderon (INB0001495898). 500 meters South of Playa Ventanas (9°05'16.1660N, 83°40'50.7600W), Provincia Puntarenas, 0 m depth, 17 January 2000, 1 specimen, leg. M. Calder6n (INB0001495974). San Pedrillo (8°36'53.6350N, §3°44'18.3791W), Parque Nacional Corcovado, Provincia Puntarenas, 0 m depth, 20 January 2000, 1 speci- men, leg. M. Calderon (INB0001495989). San Pedrillo (8°36'40.6143N, 83°44'16.7412W), Provincia Puntarenas, 0 meters depth, 21 January 2000, 1 specimen, leg. M. Calder6n (INB0001495985). San Pedrillo (8°37'22.9440N, 83°44'18.3830W), Parque Nacional Corcovado, Provincia Puntarenas, 0 m depth, 27 February 1998, 8 specimens, leg. A. Berrocal (INB0001498380). San Pedrillo (8°37'00.1590N, 83°44'11.8370W), Parque Nacional Corcovado, Provincia Puntarenas, 0 m depth, 19, 1 specimen, leg. A. Berrocal (INB0001496543). 100 m SE of Punta Curupacha (8°37'57.8040N, 83°13'14.0600W), Provincia Puntarenas, 7 m depth, 29 January 1998, 3 specimens, leg. A. Berrocal (INB0001498379). Roca Negritas (9°12'58.2630N, 83°50'14.2490W), Provincia Puntarenas, 0 m depth, 17 May 1995, 2 specimens, leg. M. Madrigal (INB0001482914). Rock in front Cabo Matapalo (8°22'17.3370N, 83°17'31.7800W), Peninsula de Osa, Provincia Puntarenas, 18 m depth, 13 February 1997, 1 specimen, leg. S. Avila (INB0001498367). VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 493 Cambutal (9°13'21.0780N, 83°50'22.4180W), Provincia Puntarenas, 0 m depth , 16 May 1995, 3 specimens, leg. Y. Camacho (INB0001485355). Punta Voladera, from the tower to NE (8°37'25.8910N, 83°10'56.7300W), Reserva de Vida Silvestre Golfito, Provincia Puntarenas, 0 m depth, 8 May 1997, 2 speci- mens, leg. M. Madrigal (INB0001498405). Punta Gallardo (8°37'35.0910N, 83°14'16.2310W), Parque Nacional Piedras Blancas, Puntarenas Provincia, 10 m depth, 6 March 1997, 1 specimen, leg. S. Avila (INB0001498361). EXTERNAL MORPHOLOGY.— The body is very elongate, 30 mm long, in the specimens exam- ined. The cephalic shield is also elongate, comprising about '4 of the body length (Fig. 7C). The two Hancock’s organs are composed of about 16 simple folds. There are large sensory processes at each side of the mouth opening. The parapodia are narrow. The posterior shield is rounded poste- riorly, with two broad lobes similar in size and shape. The gill is simple, with 11 groups of lamel- lae. The color of the living animals is pale brown with irregular large brown spots that vary in posi- tion and size. There is a series of bright blue spots on each side of the head and along the edge of the parapodia. GEOGRAPHIC RANGE.— Widespread in the tropical and subtropical eastern Pacific and Atlantic. In the Pacific it ranges from Southern Mexico to Chile (Skoglund 2002). REMARKS.— The systematics of Navanax aenigmaticus has been investigated by Gosliner (1980), who provided a complete list of synonyms. Gosliner (1980) also provided illustrations of the anatomy and shell morphology of the species, so that information is not repeated here. Navanax polyalphos (Gosliner and Williams, 1972) (Figure 13A—C) Chelidonura polyalphos Gosliner and Williams, 1972:424—-436. TYPE MATERIAL.— Chelidonura polvalphos: HOLOTYPE: Bahia San Carlos, 6 km north of Guaymas, Sonora, México (CASIZ 549). MATERIAL EXAMINED.— 500 m W of Punta Carbon, Playa Grande, Provincia Guanacaste, 0 m depth, January 11, 2001, 1 specimen, leg. S. Avila (INB0003118180); South side of Punta Zapotal, Provincia Guanacaste, 8-10 m depth, April 15, 2004, 1 specimen, leg. T.M. Gosliner (INB0003836138). SHELL MORPHOLOGY.— Length 2.5 mm; width 1.25 mm, in the single specimen examined. Shell calcified with a distinct wing-like structure protruding from the body whorl. The inner edge of the wing is thickened (Fig. 13). The protoconch is globose, about 1.5 whorl, 200 um in diame- ter. Color yellowish brown. EXTERNAL MORPHOLOGY.— The body is elongate, 5-20 mm long, 1n the specimens examined. The anterior end of the body has two grooved lobes. The eyes are large and are visible on the dor- sal surface of the cephalic shield. The posterior end of the body terminates in two long acute “tails.” The cephalic shield covers the anterior and pharyngeal region of the animal while the posterior shield covers the mantle and posterior viscera. The color of the living animals is dark brown, almost black, and generally with two rows of bright blue spots near the inner side of the parapo- dia. There are numerous small whitish spots, larger and more numerous in some areas of the dor- sal surface. Often, these spots are aggregated to form patches on the shields. In some specimens there are also some small yellowish spots. GEOGRAPHIC RANGE.— Known from the Channel Islands in southern California to Panama. REMARKS.— Gosliner and Williams (1972) provided a complete description of this species including illustrations of the reproductive and nervous system and shell morphology of the species, so that information is not repeated here. 494 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 FIGURE 13. Navanax polvalphos Gosliner and Williams, 1972, specimen from Guanacaste (A) Living animal, (B) Detail of the ventral side of the shell, 2.5 mm long, (C) Detail of the dorsal side of the shell, 2.5 mm long. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 495 ACKNOWLEDGMENTS The authors would like to recognize the generous help of several individuals and institutions. Most of the material deposited at INBio was collected by F. Alvarado, R. Angulo, S. Avila, A. Berrocal, M. Calderon, C. Cano, J. Espinosa, M. Lobo, M. Madrigal, G. Mena, and F. Gonzalez. Most specimens deposited at LACM were collected by D. Cadien, P. LaFollete, and J. McLean. Maribel Zuniga took photographs of the shells, and Terry M. Gosliner and Leopoldo Moro took photographs of the living animals. Lindsey Groves curated the LACM material and made construc- tive comments on the manuscript. Fieldwork in Costa Rica by INBio was funded by the Instituto Nacional de Biodiversidad (INBio) through the Netherlands’ Government project “Development of biodiversity knowledge and sustainable use in Costa Rica.” The SEM work was conducted at the LACM facility supported by the National Science Foundation MRI grant DBI-0216506. REFERENCES ApbAMS, C.B. 1852. Catalogue of Shells Collected at Panama, with Notes on their Synonymy, Station, and Geographical Distribution. Craighead, New York. viii + [5]—334 pp. BAKER, F., AND G D. HANNA. 1927. Expedition of the California Academy of Sciences to the Gulf of California in 1921. Proceedings of the California Academy of Sciences 16:123-135, pl. 4. BEHRENS, D.W. 1991. Pacific Coast Nudibranchs: a Guide to the Opisthobranchs, Alaska to Baja California. Second Edition. Sea Challengers, Monterey, California. 107 pp. BeERGH, R. 1894. XIII. Die Opisthobranchen. Reports of the dredging operations off the West coast of Central America to the Galapagos, to the West coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission Steamer “Albatross,” during 1891, Lieut. Commander Z.L. Tanner, U.S.N., commanding. Bulletin of the Museum of Comparative Zoélogy 25:125—233, pls. 1-12. Bertscu, H. 1972. Two additions to the opisthobranch fauna of the southern Gulf of California. The Veliger 15:103—106. BertscuH, H. 1978. The Chromodoridinae nudibranchs from the Pacific coast of America. Part II]. The genus Chromodoris. The Veliger 20:307-327. BeRTSCH, H. 1980. A new species of Bornella from tropical West-America. Spixiana 3:33-42. BerTSCH, H., AND A. KERSTITCH. 1984. Distribution and radular morphology of various nudibranchs (Gastro- poda: Opisthobranchia) from the Gulf of California, México. The Veliger 26:264-273. BertTscu, H., A.J. FERREIRA, W.M. FARMER, AND T. L. HAyeEs. 1973. The genera Chromodoris and Felimida (Nudibranchia: Gastropoda) in tropical west America: Distributional data, description of a new species, and scanning electron microscopic studies of radula. The Veliger 15:287-294. BOUCHET, P. 1975. Opisthobranches de profondeur de |’Océan Atlantique. I. Cephaslaspidea. Cahiers de Biologie Marine 16:317—365 + pls 1-4. CARPENTER, P.P. 1857. Catalogue of the collection of Mazatlan shells in the British Museum. Collected by Frederick Reigen, described by Philip P. Carpenter. British Museum, London. iv + [ix]—xvi + 552 pp. CARPENTER, P.P. 1864. Diagnoses of new forms of mollusks collected at Cape St. Lucas by Mr. J. Xantus. Annals and Magazine of Natural History, ser. 3, 13:311—315. DALL, W.H. 1871. Descriptions of sixty new forms of mollusks from the west coast of North America and the North Pacific Ocean, with notes on others already described. American Journal of Conchology 7:93—160, pls. 13-16. DALL, W.H. 1908. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission Steamer “Albatross,” during 1891, Lieut.-Commander Z.L. Tanner, U.S.N., commanding. XX XVIII. Reports on the scientific results of the expedition to the eastern tropical Pacific, 496 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 26 in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer “Albatross,” from October, 1904, to March, 1905, Lieut.-Commander L.M. Garrett, U.S.N., commanding. XIV. The Mollusca and the Brachiopoda. Bulletin of the Museum of Comparative Zoédlogy 43:205-487 + pls 1-22. DALL, W.H. 1919. Descriptions of new species of Mollusca from the north Pacific Ocean in the collection of the United States National Museum. Proceedings of the United States National Museum 56:293-371. Espinosa, J., AND J. ORTEA. 2001. Moluscos del Mar Caribe de Costa Rica: desde Cahuita hasta Gandoca. Avicennia suplemento 4:1—77. FERREIRA , A.J., AND H. BERTSCH. 1975. Anatomical and distributional observations of some opisthobranchs from the Panamic faunal province. The Veliger 17:323-330. GOSLINER, T.M. 1979. A review of the systematics of Cylichnella Gabb (Opisthobranchia: Scaphandridae). The Nautilus 93:85—92. GOSLINER, T.M. 1980. Systematics and phylogeny of the Aglajidae (Opisthobranchia: Mollusca). Zoological Journal of the Linnean Society 68:325-360. GOSLINER, T.M. 1996. Opisthobranchia. Pages 1-52 in F.G. Hochberg and P. Scott, eds., Zaxonomic Atlas of the Santa Maria Basin. Volume 8. Mollusca. Santa Barbara Museum of Natural History, Santa Barbara, California. GOSLINER, T.M., AND G.C. WILLIAMS. 1972. A new species of Chelidonura from Bahia San Carlos, Gulf of California, with a synonymy of the family Aglajidae. The Veliger 14:424-436. HERTLEIN, L.G., AND A.M. STRONG. 1951. Eastern Pacific expeditions of the New York Zoological Society. XLII. Mollusks from the west coast of Mexico and Central America. Part X. Zoologica 36:67—120, pls. 1-11. Hinps, R.B. 1844. The zoology of the voyage of H. M. S. Sulphur, under the command of Captain Sir Edward Belcher during the years 1836-42. Published under the authority of the lords commissioners of the Admiralty. Edited and superintended by Richard Brinsley Hinds. Volume 2. Mollusca. Smith, Elder & co., London. 72 + iv pp. Houprick, J. 1968. A survey of the litoral marine molluscs of the Caribbean coast of Costa Rica. The Veliger 11:4-23. Hois@Ter, T. 1998. Preliminary check-list of the marine, shelled gastropods (Mollusca) of Golfo Dulce, on the Pacific coast of Costa Rica. Revista de Biologia Tropical 46(Suplemento):263—270. KEEN, A.M. 1971. Sea shells of tropical West America. Marine mollusks from Baja California to Peru. Second Edition. Stanford University Press, Stanford, Calfornia. Marcus, Ev. 1972. On some Acteonidae (Gastropoda, Opisthobranchia). Papéis Avulsos de Zoologia 25:167—188, 1 pl. Marcus, Ey. 1977. On the genus Jornatina and related forms. Journal of Molluscan Studies supplement 2:1-35. MIKKELSEN, P.M. 1996. The evolutionary relationships of Cephalaspidea s./. (Gastropoda: Opisthobranchia): a phylogenetic analysis. Malacologia 37:375—-442. MIKKELSEN, P.S., AND P.M. MIKKELSEN. 1984. Comparison of Acteocina canaliculata (Say, 1826), A. candei (d’Orbigny, 1841), and 4. atrata spec. nov. (Gastropoda: Cephalaspidea). The Veliger 27:164—-192. OrBIGNY, A. d’. 1835-1843. Voyage dans 1’Amérique Meéridionale (le Brésil, la République Orientale de l’Uruguay, la République Argentine, La Patagonie, la République du Chili, la République de Bolivia, la République du Pérou), exécuté pendant les années 1826, 1827, 1828, 1829, 1830, 1831, 1532 et 1533. Volume 5, Partie 3: Mollusques. Bertrand, Paris. [Dates of publication: pp. 148 (1835), pp. 49-184 (1836), pp. 185-376 (1837), pp. 377-408 (1840), pp. 409-488 (1841), pp. 489-758 + pls 1-85 (1846)]. OrTEA, J.A., AND E. M. LLERA. 1981. Un nuevo dorido (Mollusca: Nudibranchia) de la Isla Isabel, Nayarit, México. /berus 1:47—5S2. PALMER, K.V. 1958. Type specimens of marine Mollusca described by P.P. Carpenter from the West Coast (San Diego to British Columbia). Memoirs of the Geological Society of America 76:\-376. Pease, W.H. 1868. Descriptions of marine Gasteropodae, inhabiting Polynesia. American Journal of Conchology 4:71—80, pls. 7-10. ROBINSON, D., AND M. Montoya. 1987. Los moluscos marinos de la costa Atlantica de Costa Rica. Revista de Biologia Tropical 35:375-400. VALDES AND CAMACHO-GARCIA: “CEPHALASPIDEAN” HETEROBRANCHS 497 RuDMAN, W.B. 1971. The family Acteonidae (Opisthobranchia, Gastropoda) in New Zealand. Journal of the Malacological Society of Australia 2:205—214. SKOGLUND, C. 2002. Panamic Province molluscan literature. Additions and changes from 1971 through 2001. III Gastropoda. The Festivus 23(supplement):i—x1, 1-286. SowerBy, G.B. 1848-50. Thesaurus Conchyliorum, or Monographs of Genera of Shells. Volume 2. Sowerby, London. [Dates of publication: pp. 493-505, pls 92-108 (1848), pp. 507-552, pls 109-11 8bis (1849), pp. 553-899, pls 119-186 (1850)]. SpHON, G.C., AND D.K. MULLINER. 1972. A preliminary list of known opisthobranchs from the Galapagos Islands collected by the Ameripagos Expedition. The Veliger 15:147—152. STEARNS, R.E.C. 1897. Description of a new species of Actaeon from the Quaternary bluffs at Spanish Bight, San Diego, California. Proceedings of the United States National Museum 21:297-299. STRONG, A.M., AND L.G. HERTLEIN. 1937. The Templeton Crocker Expedition of the California Academy of Sciences, 1932, No. 35. Proceedings of the California Academy of Sciences 22:159-178, pls. 34-35. STRONG, A.M., AND L.G. HERTLEIN. 1939. Marine mollusks from Panama collected by the Allan Hancock Expedition to the Galapagos Islands, 1931-1932. The Allan Hancock Pacific Expeditions 2:177—245. THOMPSON, T.E. 1976. Biology of opisthobranch molluscs. Volume 1. The Ray Society, London. 206 pp. Wittetr, G. 1944. New species of mollusks from Redondo, California. Bulletin of the Southern California Academy of Sciences 43: 71-73. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27, pp. 498-549, 83 figs. December 30, 2004 The Marine Gobies of the Hawaiian Islands David W. Greenfield! and John E. Randall? ' Research Associate, Department of Ichthyology, California Academy of Sciences, and Emeritus Professor, University of Hawaii; Mailing address: Moss Landing Marine Laboratory, 8272 Moss Landing Road, Moss Landing, CA 95039. 2 Research Associate, Department of Ichthyology, California Academy of Sciences and Bishop Museum, Honolulu, Hawaii. A total of 34 species of gobies (Teleostei, Gobiidae) are known from the Hawaiian Islands, four of which are freshwater species. All species are treated in a key, but only marine species are illustrated and treated in detail. Information on their nomencla- ture, counts and measurements, distinctive characters, coloration, distribution and habitat is presented. Two new species, Cabillus caudimacula, and Pleurosicya lar- sonae are described. An unidentified species of Favonigobius from O’ahu, a possible introduction, is discussed. The Gobiidae is the largest family of marine fishes, with many species also occurring in fresh water. J.S. Nelson (pers. commun., September 2003) estimates that there are about 220 genera and 2010 species. Judging from the number of undescribed species of gobies of which we are aware, that number of species will surely be exceeded. Most gobies are small (the shortest goby 1s Trimmatom nanus, with females maturing at as lit- tle as 8 mm standard length; but one species, Glossogobius giuris, attains 500 mm standard length). Gobies generally are recognized by their pelvic-fin structure; the fins usually are fused to form a sucking disc which may have an anterior transverse membrane called a frenum that links the spines. When the pelvic fins are separate, they are close together. Gobies and sleepers (family Eleotridae) have been confused in the past, and some gobies with separate pelvic fins have been misidentified as sleepers. Gobies have five branchiostegal rays, whereas the sleepers have six, and the pelvic fins are more broadly separated than in the gobies with the pelvic fins divided. All Hawaiian gobies normally have six spines in the first dorsal fin, except for Discordipinna griessin- geri With five. Most gobies, and all in Hawaiian waters, also usually have two separate dorsal fins. The Hawaiian Islands have a total of 34 species of gobies, four of which (Awaous guamensis, Lentipes concolor, Sicyopterus stimpsoni, and Stenogobius hawaiiensis) are fresh water. In 1905, Jordan and Evermann recognized eight marine gobies in the Hawaiian Islands. Gosline and Brock (1960) found 13. As a result of collecting since then, the number is now 30. These 30 marine species occur in salt to brackish water from tidepools high in the splash zone to moderate depths. One species was recently taken by trawl in 138-169 m. Although some gobies, such as species of the genus Bathygobius, can readily be seen scurrying around tidepools, most are secretive and not easily observed. They are, however, an important part of the communities in which they are pres- ent. In a survey of the fishes of Kane’ohe Bay, Greenfield (2003) found that the most abundant species taken in collections using an ichthyocide was the goby Eviota epiphanies. In reference to species of Eviota, Greenfield and Randall (1999) wrote “These small gobies may form an impor- tant component of the food of larger piscivorous fishes.” Other gobies, of course, also serve as prey to larger fishes. 498 GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 499 Although all species are treated in our key, only the marine species are included in this paper. Several small individuals of a goby species in the genus Favonigobius were taken at Kahana Bay, O’ahu, but were too small to be identified (see Discussion). MATERIALS AND METHODS All counts and measurements follow Hubbs and Lagler (1964) except that the last two rays of the dorsal and anal fins are not counted as one unless it is clear that they are joined at the base. Measurements were made to the nearest 0.1 mm using dial calipers, and are expressed as percent- ages of standard length (SL). Lengths given in figure captions of specimens are standard length (SL), but for underwater photographs, the length is the estimated total length (TL). In the descrip- tions of the new species, measurement data for the holotype are presented first, followed by the range and mean (in parentheses) for all type material. For other descriptions the count or measure- ment is followed by the mode or mean, respectively, in parentheses. Color descriptions are from 35-mm slides taken under water of either living individuals or ones recently collected, or fresh specimens out of water. When listing type material for various species, often only those at the Bernice P. Bishop Museum in Honolulu (BPBM) are listed. Institutional abbreviations are as list- ed in Leviton et al. (1985). KEY TO THE GOBIES OF HAWAII fasbelvic fins separate, imner rays of both fins not connected (Fig: 1.) -).........-0......5- 2 lb. Pelvic fins fused to form a sucking disc, or at least inner rays of both fins connected together (membrane is easily torn, may only be joined at bases of fins) (Fig. 2)................. 5 2a.(1a) Preopercle with two or more spines (Fig. 3); pelvic fins not fringe-like, lacking many side ‘SPIRCI TSS ig, as ently &, 6 CNG BORE OR ERED Nl OCIA CHO! Ee ea, cee RR Asterropteryx semipunctatus 2b.(1a) Preopercle without spines; pelvic fin with many side branches and fringe-like (Fig. 4) 3a.(2b.) Unbranched 5" pelvic-fin ray not developed (Fig. 4); genital papillae of male and female not ruggose (Fig. 5); subcutaneous dark-brown bar at caudal-fin base (Fig. 6); anteriormost branch of 4!" pelvic-fin ray short, with fewer than 10 segments; IT pore present (Fig. 7) 2 eG 00.8.8 FE FNL SOOT RIE Te te CR EERO OP rime te eee RNG ae nen ae Eviota epiphanes 3b.(2b) Unbranched 5" pelvic-fin ray developed (about '/o length of 4» ray) (Fig. 8); genital papil- lae of male and female ruggose (Figs. 9 and 10); no subcutaneous bar at caudal-fin base; ante- riormost branch of 4" pelvic-fin ray elongate, with 9 or more segments (usually about 15) Cie seSyaridelil)--hT. pore absent Chien). ster ison Uy Wan a Mc Lne BNn lure eiets pes 4 4a(3b) No prominent subcutaneous body bars or spots; POP pores present (Fig. 7); size to 12.8 mm TE TN ah eins te REE Ad, Cousin. Lies hia aol a tie a ete om Eviota rubra 4b(3b) Five-6 prominent subcutaneous body bars, ventral surface with 3 broad subcutaneous bars on belly and 4-6 more spots above anal-fin base and on caudal peduncle (Fig. 12); POP pores ADS ENMU Che AU aTedUceds(EiG2s/)).SIZetOnlS-5 smiMeS eps a2 ayes ene Eviota susanae 5a.(1b) Caudal fin distinctly forked (young of various freshwater gobies) . . . Vitraria clarescens? Smal nealdaiim rouuided.or pomited! =... Sea ad tsen oor ches ae balance Maden se tee 6 3 Vitraria clarescens in literature, a synonym of Sicyopterus stimpsoni (Greenfield et al. 1998). 500 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 6a.(5b) Caudal fin distinctly pointed, more than twice as long as wide (Fig. 13); a well-developed fleshy crest on nape extending forward from first dorsal fin at least past edge of opercle and further im, larger specimens (Fis: 14)C0 cs ee ie ec 7 6b.(5b) Caudal fin rounded, not twice as long as wide; no distinct median crest on nape...... 8 7a.(6a) Body with lateral series of elongate blotches; dark spot on superior margin of eye; promi- nent dark spot on pectoral-fin base (Fig. 14); dark spots on branchiostegal membranes under lower jaw (Fig. 15); 18-21 (usually 20) pectoral-fin rays.......... Oxyurichthys lonchotus 7b.(6a) Body lacking elongate blotches on sides, spots on eye margin, pectoral-fin base, or bran- chiostegal membranes; 21—23 (usually 22—23) pectoral-fin rays ....... Oxyurichthys heisei 8a.(6b) Upper 4-9 pectoral-fin rays terminating in free, silky filaments (Fig.16)............. 9 8b.(6b) Upper pectoral-fin rays not terminating in free, silky filaments ................... 1] 9a.(8a) Pelvic sucking disc about as wide as long (Fig. 17); anterior nostril with a small flap; side of cheek with a deep longitudinal groove, with upper portion of cheek extending down over groove anteriorly, hiding papillae in groove (Fig. 18); predorsal scales extending forward to between eyes; upper 8—10 pectoral-fin rays with free, silky filaments . Bathygobius cotticeps 9b.(8a) Pelvic sucking disc clearly longer than wide (Fig. 19); no flap on anterior nostril; side of cheek with a shallow groove anteriorly, with papillae present in groove visible; predorsal scales not extending forward to between eyes; upper 4-6 pectoral-fin rays with free, silky fil- AMVOMUS Settee. Wiaingtneie ausbevvewetbie Sa thie otimus wesibo btccazs (ie A ciel ebro Reece 10 10a.(9b) Predorsal scales extending forward of a line drawn up from posterior edge of preopercle, almost to eyes in some (in specimens 28 mm SL and larger); mandibular frenum straight with NOmEeCe ODES atstGes (IG. 2 O)\ege nec ry einer en eee get ral ee ee Bathygobius coalitus 10b.(9b) Predorsal scales not extending forward to line drawn up from posterior edge of preoper- cle; mandibular frenum more curved with free lobes at sides (Fig. 21) as ata dis an Me Ae one AA Owe NaS a Berta Nal eet Bathygobius cococensis lla.(8b) Lateral scales on back anterior to second dorsal fin smaller than those under second dor- sal fin and on caudal peduncle (Fig. 23); interorbital area covered with very small cirri (Fig. 2D) ANG POLES OMAP) Ofhiead yo <6. sy Nc ees MA ee Cte ea tere VMugilogobius cavifrons 11b.(8b) Size of scales (if present) not obviously different between anterior and posterior parts of body; interorbital area lacking small cirri; pores on top of head present or absent ....... 12 12a.(11b) About three prominent, knob-like, fleshy projections extending forward from lower por- tion of shoulder girdle inside gill openings (Fig. 24).(freshwater species).............. 13 12b:(1 1b) No knob-like, fleshy projects on shoulder girdle... =... 45. soe 14 13a.(12a) A black blotch extending down and back from eye; caudal fin not crossed by any dark bansm(ineshiewaten ism. part ne fo be Ri Rs Re ee i eee Stenogobius hawatiensis 13b.(12a) No black blotch under eye: caudal fin crossed by about 5 dark bars (fresh water) PE RN RIM hatte Wess ale, wade deel cadynilel gee ase tho pele ee eee eee Awaous guamensis 14a.(12b) Origin of first dorsal fin far forward on body, above posterior end of operculum and with five spines; second dorsal-fin spine greatly elongate, extending well back above second dorsal (NN Qe See a a a en eee mre es. Discordipinna griessingeri 14b.(12b) Origin of first dorsal fin behind origin of pectoral fin and with six spines; second dorsal- linsspine may beiclongate ormotes: es. ooce.am : eee 4A ES ee 15 GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS Ficure 1. Pelvic fins of Asterropteryx FIGURE 2. Pelvic fins of Bathygobius FicureE 3. Head of Asterropteryx semipunctatus. coalitus. semipunctatus. Ficure 4. Pelvic fin of Eviota epi- — Ficure 5. Genital papillae of Eviota — FiGuRE 6. Caudal peduncle and fin of phanes. epiphanes, male on left. Eviota epiphanes. Ficure 7. Cephalic pores of Eviota FiGure 8. Pelvic fin of Eviota rubra. FIGURE 9. Genital papillae of Eviota species, modified from Lachner and rubra, male on right. Karnella (1980). Ficure 10. Genital papillae of Eviota Ficure 11. Pelvic fin of Eviota FiGure 12. Eviota susanae. susanae, male on right. susanae. Figure 13. Caudal fin of Oxyur-- FiGurRE 14. Head of Oxyurichthys lon- FIGURE 15 Ventral view of head of ichthys lonchotus. chotus. Oxyurichthys lonchotus. Ficure 16. Pectoral fin of Bathygob- _- Ficure 17. Pelvic fins of Bathygobius — FiGure 18. Head of Bathygobius cot- ius coalitus. cotticeps. ticeps. 501 502 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 15a.(14b) Pelvic fins with a frenum that is thickened and variously raised, thickened portion with a folded pocket in the center extending back onto disc (probe may be inserted into pocket at anterior pantnor disc) RIG. 25). A. ete sie oe osteo seater 16 15b.(14b) Pelvic fins with or without a frenum, but if present not thickened and raised (usually a thin membrane); no folded pocketanstirenum: 4-5 54--0 5-5 oc ica 22 16a.(15a) Interorbital wide, about one and one-half to two eye diameters; snout broadly rounded from side view (Fig. 26); pelvic-fin frenum lacking pelvic-spine lobes or short papillae . . 17 16b.(15a) Interorbital narrow, less than one eye diameter; snout more pointed from side view (Fig. 27); pelvic-fin frenum with thickened skin around pelvic spines, forming lobes that extend posteriorly over disc (lobes may be less distinct, but if so, are covered with many, short papil- LANCET OS 82 ON ee eee a wid vw eleccrnteaw dt 8S wwe Co etaph ale ec eane ane eas ane 18 17a.(16a) Scales on body extending forward onto head (fresh water) ..... Sicyopterus stimpsoni 17b.(16a) Body naked or with a few scales on posterior part of body near caudal peduncle (fresh NVALCIS) Meare eres eeece es ON ee i, ene een ae ee enn CU Ne Renee ee Lentipes concolor 18a.(16b) Interorbital very narrow, about '4 pupil diameter; a single pore in center of interorbital (CE TSANE))) SRaene ReRae ce te ee ene nara eee Re MPe aoe Ire SS 5.5 60 c 00906 19 18b.(16b) Interorbital wider, slightly narrower than one pupil diameter; two pores in center of interorbital>.onemext toveach eye (Figs 3i)\. ace eas os 213s eo 20 19a.(18a) Pelvic-fin frenum with two distinct lobes of thickened skin around pelvic spines, extend- ing posteriorly over disc; frenum not broad and covered with short papillae (Fig. 28); dark pig- ment on lower portion of caudal peduncle extending posteriorly as stripe onto caudal fin Neches ae be iS me gnc hap ega NeNCRCIaE AINE, cts RAZ CONE Dace Ret MS AO cd Pleurosicya micheli 19b.(18a) Pelvic-fin frenum broader, lobes not as distinct and frenum covered with short papillae (Fig. 32): body peppered with melanophores, no distinct dark stripe on caudal peduncle and eal cl al itn ame er pestis hs athe oo: RHI Spe MOE, 99k 8 ST) eee Pleurosicya larsonae 20a.(18b) Pectoral-fin rays 13 (rarely 12 or 14); a distinct black stripe from eye forward onto snout Bem wet soy sy ay ene ell PoACE LAL ese HETIS (eats elce et Na LIYE oe NRA eee Bryaninops tigris 20b.(18b) Pectoral-fin rays 14-17 (rarely 13); stripe from eye to snout absent or if present indis- WIM Cle osts cere cece Bas ie He «Sib Grew sseiagergee Guanes PRUE ac ee Soke Sas Cee zl 21a.(20b) Scalloped grooves present along lower edge of preoperculum (Fig. 33); head depth 50% or more of head length; body bars golden brown to brown when live, generally distinct in pre- Semvedematenialeyse.s ssw shai che), Moet tae «ota Ny PRR on Seen Bryaninops yongei 21b.(20b) No scalloped grooves present along lower preopercular edge (occasionally slight inden- tations along margin); head depth less than 50% of head length; body bars brownish-orange to red when live; usually indistinct in preserved material ............... Bryaninops amplus 22a.(15b) Body naked, with about 10—11 dark vertical bars separated by narrow white bars; pelvic disc short, space between posterior end of fin and anal-fin origin equal to or greater than Polvic-tinglenGthis se... 3.04 s0.5 5 yee eee en tn) a eee Kelloggella oligolepis 22b.(15b) Body with scales, at least posteriorly; body without 12 dark vertical bars; pelvic disc longer, space between posterior end of pelvic fin and anal-fin origin clearly less than pelvic- CUISCTEN OT eu ce Co han os Gp ete Sie weirs Sed wile 4 ooo aes 23 23a.(22b) Gill membranes broadly fused to isthmus; gill opening not extending anteriorly as far as postenionedce of preopercle:i(s . <<)... ...:20s Sheu bans cee os 24 GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 503 Ficure 19. Pelvic fins of Bathygobius FIGURE 20. Ventral view of head of — FiGure 21. Ventral view of head of coalitus. Bathygobius coalitus. Bathygobius cococensis. FIGURE 22. Back of Mugilogobius FiGuRE 23. Dorsal view of head of | FiGuRE 24. Pectoral-fin base of cavifrons. Mugilogobius cavifrons. Awaous guamensis. FiGureE 25. Pelvic fins of Sicyopterus FIGURE 26. Head of Sicyopterus FIGURE 27. Head of Bryaninops stimpsoni. stimpsoni. amplus. FIGURE 28. Pelvic fins of Pleurosicya FiGuRE 29. Pelvic fins of Bryaninops FiGure 30. Head of Pleurosicya micheli. yongei. micheli. Figure 31. Head of Bryaninops Ficure 32. Pelvic fins of Pleurosicva FIGURE 33. Head of Bryaninops amplus. larsonae. yongel. Ficure 34. Head of Gnatholepis FiGure 35. Pelvic fins of Gnatholepis FIGURE 36. Dorsal view of eyes of anjerensis. anjerensis. Gnatholepis anjerensis. 504 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 23b.(22b) Gill membranes not broadly fused to isthmus, gill openings extending forward to or beyondsposterion edge Olpreopercle: 56 ss a ee 2, 24a.(23a) Narrow black bar extending down from under eye across cheek and under head (Fig. 34); pelvic-fin frenum well-developed and obvious, extending to near ends of pelvic-fin spines (ETCASS) ME yr ee arg cs ales fle ae Me he ayes lols die Saree eC 25 24b.(23a) No narrow black bar under eye; pelvic-fin frenum weak, thin and often not obvious, and nowextending tomearends\or pelvic-finispines +245 4.65--- ae tae eee 26 25a.(24a) Pectoral-fin rays 15—17 (modally 16, rarely 15); a narrow dark bar dorsally on eye, cen- tered over posterior half of pupil, not crossing interorbital space (Fig. 36); small scales usual- ly present on cheek anterior to dark eye bar; size to64mmSL..... Gnatholepis anjerensis 25b.(24a) Pectoral-fin rays 16—18 (modally 17, rarely 16 or 18); narrow dark bar dorsally on eye, centered at pupil, often crossing midinterorbital space (Fig. 37); small scales rarely present on cheek entirely anterior to dark eye bar; size to 42 mm SL 6 9.c1cyio- Ra ho RA ee ra at il i Gnatholepis cauerensis hawaiiensis 26a.(24b) Head about as deep as wide; snout pointed (Fig. 38); small dark spot at center of caudal- fin base (Fig. 39); no scales with enlarged cteni at caudal-fin base . Coryphopterus duospilus 26b.(24b) Head much wider than deep; snout more rounded (Fig. 40); large dark spot covering most of caudal-fin base; two large scales with enlarged cteni at top and bottom of caudal-fin ba Sen Gao eh Seems 8. t crate at tain aw lees nonnee eet meeortohe ae oro. eae Cabillus caudimacula 27a.(23b) Posterior end of jaws extending to or past posterior margin of eye; top of head lacking scales; pelvic-fin frenum very well-developed, extending posteriorly one-third to one-half lene thot (Pig rd D)\ eign ree cernicny teeta th tec nt eel net ce eae Psilogobius mainlandi 27b.(23b) Posterior end of jaws not reaching posterior margin of eye; top of head with at least some scales from front of dorsal fin forward; pelvic-fin frenum absent or less developed, not extend- ing, postenorly one-third-length of fim: 5... 4s. 22.22. sede ee eee 28 28a.(27b) Gill opening extending forward to near posterior edge of preopercle............. 29 28b.(27b) Gill opening extending forward to at least midway between posterior edge of preopercle AMOS ViS ia wigsuen ay age. 4 sole agen io be PRCTIAN GO donrade erste Apne atewol sis a lor see ay 29a.(28a) Head with a series of prominent, dark-bordered light lines radiating out from eye across cheek and top of head (Fig. 43); anal fin with one spine and 7 rays; midline of nape naked ante- LOT CORA ONS alata i. 8: 2s Hoyas cw: Say. cusy hy Spee eae koko al weno aaa ba en Priolepis farcimen 29b.(28a) Head without a series of prominent, dark-bordered lines; anal fin with one spine and 8 or more rays; midline of nape with scales anterior to dorsal fin...................... 30 30a.(29a) Scales present on cheek and opercle (Fig. 44); a deep interorbital trench present between Svesn(hies 45) sins eenerallydark) 2 2c) eae ee ee Priolepis eugenius 30b.(29a) Cheek and opercle lacking scales; interorbital without a trench; fins not dark...... 3)! 31a.(30b) Scales on body with dark borders (Fig. 46); second dorsal fin with 9 rays; anal fin with 8 to 9 rays, usually 8; predorsal scales 5 to 8; longitudinal scale series 24—26, usually 25 BOT Ra yr nae a SS can hey Same Meee Se, ane ot ne a Priolepis limbatosquamis 31b.(30b) Scales on body lacking dark borders; second dorsal fin with 11—12 rays, usually 11; anal fin with 9-10 rays, usually 9; predorsal scales 12—16, usually 15; longitudinal scale series US ean 2 oS. Site sd aS A ae A OA oA Priolepis aureoviridis 32a.(28b) Gill opening extending forward only to posterior margin of eye; first spine in second part GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 505 27 FIGURE 37. Dorsal view of eyes of Figure 38. Head of Coryphopterus FiGurE 39. Caudal peduncle and fin Gnatholepis cauerensis hawaiiensis. duospilus. of Coryphopterus duospilus. Ficure 40. Head of Cabillus caudi- FiGuReE 41. Caudal peduncle and fin — FiGure 42. Pelvic fins of Psilogobius macula. of Cabillus caudimacula. mainlandi. Figure 43. Head of Priolepis farci- FiGure 44. Head of Priolepis euge- FIGURE 45. Dorsal view of head of men. nius. Priolepis eugenius. FIGURE 46. Scales on side of body of | FiGure 47. Dorsal fins of Trimma FiGuRE 48. Caudal-fin base of Priolepis limbatosquamis. unisquamis. Trimma unisquamis. of dorsal fin stout and sharp-tipped; body with a line of dash-like dark marks down side and Seven lsraititctallMes 2 DOVE meat cd.c my act lnie Meno anal he Rie a cates Opua nephodes 32b.(28b) Gill opening extending far forward to under chin and anterior part of eye; first spine in second part of dorsal fin not stout and sharp-tipped; series of dash-like marks not present on SIGENO I D OCS gree te rt a i ne tad Sek ER re nile Pee he ch ae ne tS eR 3)3) 33a.(32b) Both dorsal fins with a black border , more obvious in first (Fig. 47); a dark band at base of caudal fin (Fig. 48); dorsal-fin spines not elongate................ Trimma unisquamis 33b.(32b) Dorsal fins without black borders; no dark band at base of caudal fin; dorsal-fin spines PAV AO INAV INO DesClOM Sater A uy drat Ae Oi eye aes ol tie inten Bobet aa weneme ge WUOR@ tgs pamectce 34 34a.(33b) Bony interorbital narrow, less than half pupil diameter; pectoral-fin rays 17-18; second dorsal spine not prolonged; soft dorsal rays 9, soft anal rays 8.............. Trimma milta 34b.(33b) Bony interorbital wider, about equal to pupil diameter; pectoral-fin rays 13-15; second dorsal spine long and filamentous (may reach caudal-fin base in males); soft dorsal rays 10-11, SOR AMAT AYS 9 —l Ory ne me te Rumer Tae hee nee, onan ee eae ecgtine Meier ee ale Trimma taylori 506 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 Asterropterix semipunctatus Riippell, 1830 (Figs. 3 and 49) Asterropterix semipunctatus Riippell, 1830, Fische Rothen Meeres 1828—30:138, pl. 34, fig. 4. Red Sea. Holotype: SMF 1691. Brachyeleotris cyanostigma Bleeker, 1855, Nat. Tijdschr. Ned. Indié 8:452. Cocos-Keeling Is., Indian Ocean. Syntypes: RMNH 4756(2). DIAGNOSIS.— Counts based on 20 individuals, 24.0-34.6 mm SL. Dorsal-fin elements VI-I,10. Anal-fin elements I,8- 1,9 (1,9). Pectoral-fin rays 17-19 (18). Longitudinal scale series 24—25 (24). Pelvic fins separate, without numerous fringe-like side branches. Lower part of preopercle with two or more spines. Scales ctenoid, covering body; head scaled except interorbital space, snout, and chin. Gill membranes broadly united to isthmus; gill opening ending under aes ee middle of opercle. Mouth oblique, lower wx RE 49. Asterropteryx Saunoonictiainn, 501 mm TL, Kane an se E ts ES Ais jaw protruding: jaws extending past ante- B& 0-2 rior margin of eye but not to pupil. Small conical teeth in bands in jaws, the outer row at the front enlarged. Interorbital narrow, about equal to pupil diameter. Third dorsal spine prolonged to a long filament in large adults. Caudal fin rounded, shorter than head. Body depth 3.0—3.6 in standard length. Reported to 65 mm TL. COLOR IN ALCOHOL.— Male- Background color either light cream or medium brown. Light color morph: body with five longitudinal stripes from head back to caudal-fin base, one at midline, two above, and two below. A dark brown spot at top of pectoral-fin base. Caudal-fin base with dark- brown bar. Top of caudal peduncle dark brown. Head with small dark-brown spot at posteriodor- sal margin of eye. A diffuse light-brown bar from anteroventral margin of eye to posterior end of jaws. A dark bar on chin from mandible to mandible. Pectoral fins clear. Pelvics, caudal, anal, and second part of dorsal fin with scattered melanophores. First part of dorsal fin with a dark-brown spot at posterior portion. Dark-color morph: basic color pattern as for light morph except less dis- tinct because of dark background. Pectoral fins with scattered melanophores. Other fins as in light morph except more melanophores and thus darker. Female- Color the same as dark-color morph of male. LIVE COLOR.— (from photograph taken at Kane’ohe Bay) Background color gray with pattern of black markings, entire body overlaid by small, iridescent blue spots. Body crossed by six, irreg- ular, black saddles that extend down to ventral surface: First from top of pectoral-fin base up to ori- gin of first dorsal fin; second from base of posterior half of first dorsal fin; third from elements three to seven of second dorsal fin; fourth from last few rays of second dorsal fin and onto caudal peduncle; fifth at middle of caudal peduncle; and sixth at caudal-fin base. Ventral surface at anal- fin base black, joining bases of saddles together. A separate black blotch extending up from anal- fin origin to midline between saddles two and three. Nape and top of head mottled black and gray, lower half of side of head black. Pupil of eye black surrounded by a narrow golden ring, remain- der of iris mottled black and gray. Pectoral-fin base black except for a gray patch at center that extends out onto fin-ray bases; remainder of fin dusky. Pelvic, anal, and caudal fins dusky. First GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 507 dorsal fin dusky with alternating black and gray bars on spines. First spine of second dorsal fin with alternating black and gray bars, remainder of fin dusky. DISTRIBUTION.— Throughout most of the Indo-Pacific region from the Red Sea and Indian Ocean to the Society Islands and Hawaiian Islands. REMARKS.— Usually lives in shallow protected waters of bays and lagoons, often on dead reefs (Greenfield, 2003). Classified in the earlier literature as an eleotrid because of the divided pelvic fins. Privitera (2001, 2002) studied its reproductive biology in the Hawaiian Islands. Spawning occurs at various times during the day, year-around, with a peak from May to July. Clutch size varied from 296-1552 (mean 886), independent of length of the female. Eggs are ellip- soidal and varied from 0.67—0.84 mm in length, hence unusually small for a body of this size. Eggs were laid beneath coral; they were tended by the male who periodically fanned them with its pec- toral fins. Eggs hatched in laboratory aquaria shortly after lights were turned off, four to six nights after being deposited in the nest. Newly hatched larvae had a mean notochord length of 1.88 mm. Minimum age at maturity, 17.5—19 mm, estimated as four and one-half to five months after hatch- ing. Also known as the bluespotted goby. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218097 (10), CAS 218089 (10), BPBM 5471 (7), BPBM 22621 (10), BPBM 37315 (9), BPBM 37316 (3); Hawai’i: 28719 (6); Midway Atoll: BPBM 35370 (6). Bathygobius coalitus (Bennett, 1832) (Figs. 2, 16, 19-20, and 50) Gobius coalitus Bennett, 1832, Proc. Zool. Soc. London 1830—31(pt 1):166. Mauritius. Holotype: BMNH 1856.2.15.20. DIAGNOsIS.— Counts based on 26 individuals. measurements on 10 individ- uals 40.2-77.6 mm SL. Dorsal-fin ele- ments V-VI-I,8-9 (VI-I,9). Anal-fin ele- ments I[,7-I,9 (1,8). Pectoral-fin rays 18-20 (19), the upper 4-6 (5) rays fila- mentous, branched to base with mem- brane-free ends. Longitudinal scale series ie ET REG 34-37 (36). Bredorcaleceales1s 20) FIGURE 50. Bathygobius coalitus, mm TL, Maui Ocean Center. Pelvic sucking disc longer than wide, space between end of fin and anal-fin origin less than one fin length; broad frenum with an obvi- ous spine on each side of frenum pointing posteriorly. Body covered with ctenoid scales, becom- ing cycloid on abdomen, chest, and nape; extending forward on head past a line drawn up from posterior edge of preopercle, almost to eyes in some individuals. No scales on opercle or cheek. Gill membranes broadly united to isthmus. Jaws extending posteriorly to mid-pupil. Interorbital narrow, about equal to pupil diameter. No flap on anterior nostril. Side of cheek with a shallow groove anteriorly, with papilla in groove visible. Mandibular frenum straight with no free lobes at sides. Caudal fin rounded, about equal to head length. Head depressed, its width greater than its depth. Body depth 4.34.7 in standard length. Reported to 120 mm TL. COLOR IN ALCOHOL.— Head and body light cream. Upper half of body with four dark brown saddles: First from fourth spine of dorsal fin posterior to just past end of fin base and down to meet mid-side blotches; second saddle from third element of second dorsal fin posterior to base of sev- enth element and down to meet mid-side blotches; third saddle from base of last element posterior 508 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 onto three scales of caudal peduncle and down to meet mid-side blotches; last saddle covering last two scales of caudal peduncle prior to caudal fin and down to meet mid-side blotches; first and third saddles darkest and most prominent. Midside of body with a series of about eight to nine interconnected dark brown blotches that do not reach ventral surface of body. Dark area on top of head immediately in advance of first dorsal-fin spine. Head with four dark brown blotches around eye: one, the most prominent, posterior to orbit at level of top of opercle; second anterior to this; third at posteroventral margin of orbit; fourth at anteroventral margin of orbit extending onto pre- maxilla; two additional dark brown blotches posterior to most prominent spot posterior to orbit, first midway between eye and top of opercle and second at top of opercle. A distinct dark brown spot on middle of opercle adjacent to edge of preopercle. Cheek with mottled pattern. Eye with clear pupil and black iris. Pectoral fin covered with scattered melanophores, a distinct spot present at bases of rays 6-8 and a second more diffuse spot over bases of rays 12—15. Anal fin with scat- tered melanophores, darker on distal *4 of membranes. Caudal fin of female with dark brown blotch at center of base and then crossed by four to five indistinct bars. Caudal fin of male with dark brown blotch at base, proceeded by light area containing two blotches. A circle of dark spots out past '2 of fin length. Fin posterior to this with one dark bar and then generally dusky. First part of dorsal fin covered with melanophores concentrated into dark spots on basal half of membranes between spines four to six. Second part of dorsal fin crossed by six bars angling dorsoventrally across fins. Pelvic fins covered with melanophores in males, fewer in females. COLOR OF FRESH SPECIMEN.— (from photograph of specimen collected at Kane’ohe Bay) Background color golden brown with dark brown to gray mottled pattern scattered over body, dark- er on back, nape, top and sides of head. Body with scattered, small, white spots, some arranged into irregular rows. Side of head with many scattered, small, white spots. Ventral surface of head and body white with scattered peppering of melanophores, isthmus with dark gray blotches. Eye with black pupil and iris dark brown with an orange tinge. Pelvic fins white with a peppering of melanophores. Caudal-fin membranes dusky, rays dark brown with white spots on basal three- quarters. Anal fin dusky with dark gray distal margin. Pectoral-fin membranes dusky, rays brown with white spots on basal one-half. First dorsal fin with rows of white spots along base with a nar- row dark brown bar distal to spots. Another row of white spots above brown bar. Middle third of fin dusky with additional row of white spots. Distal third of fin membrane clear with an orange margin. Second dorsal fin similar to first dorsal fin. DISTRIBUTION.— East coast of Africa to the Hawaiian Islands and Marquesas; Japan to the Great Barrier Reef. REMARKS.— Typically found in the intertidal zone in rock and tidepools. Shafer (1998) report- ed that this species spawns throughout the year, has a larval duration of 29-50 days, settles at 7.02—10.6 mm total length, and reaches its maximum size in 20—30 days. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218099 (13), uncataloged, (G91- 11), CAS 218100- (3), CAS 218099 (13); BPBM 5490 (1), BPBM 34559 (1); Hawai’i: BPBM 34622 (2); Necker: BPBM 4863 (1). Bathygobius cocosensis (Bleeker, 1854) (Figs. 21 and 51) Gobius cocosensis Bleeker, 1854, Nat. Tijdschr. Ned. Indié 7:47. Cocos-Keeling Islands, Indian Ocean. Holotype: lost. Neotype: RMNH 4533, selected by Akihito and Megura (1980); location of neotype unknown. Mapo fuscus: Jordan and Evermann, 1905, Bull. U.S. Fish Comm. 23(1):483, fig. 212 (misidentification). DIAGNOSIS.— Counts based on 25 individuals, measurements on 10 individuals 28.5—56.3 mm GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 509 SL. Dorsal-fin elements VI-I,7-VI-I,10 (VI-I-9). Anal-fin elements 1,7-1,9 (1,8). Pectoral-fin rays 17-20 (18), the upper 4-6 (5) rays filamentous, branched to base with membrane-free ends. Longitudinal scale series 34-38 (35). Predorsal scales 10-12. Pelvic sucking disc longer than wide, space between end of fin and anal- fin origin less than one fin length; broad frenum with an obvious spine on each side — oe. of frenum pointing posteriorly. Body cov- FiGuRE 51. Bathygobius cocosensis, BPBM 33489, 41 mm SL, ered with ctenoid scales, becoming Kane’ohe Bay, O’ahu. cycloid on abdomen, chest, and nape; scales extending forward on head to a line drawn up from posterior edge of opercle. No scales on opercle or cheek. Gill membranes broadly united to isthmus. Jaws extending posteriorly to poste- rior margin of pupil. Interorbital narrow, about equal to pupil diameter. No flap on anterior nostril. Side of cheek with a shallow groove anteriorly, with papillae in groove visible. Mandibular frenum curved with free lobes at sides. Caudal fin rounded, about equal to head length. Head depressed, its width greater than its depth. Body depth about 5 in standard length. Reported to 60 mm TL. COLOR IN ALCOHOL.— Head and body light cream; upper half of body with four dark brown saddles, first from third spine of dorsal fin posterior to just past end of fin base and down to meet midside blotches; second saddle from second element of second part of dorsal fin posterior to base of 6th element and down to meet midside blotches; third saddle from base of 8th element posteri- or onto three scales of caudal peduncle and down to meet midline blotches; last saddle covering last two scales of caudal peduncle prior to caudal fin down to meet midline blotches. Midside of body with a series of about eight to nine interconnected dark brown blotches that do not reach ven- tral surface of body. Dark area on top of head immediately in advance of first dorsal spine, resem- bling a saddle. Head with a prominent dark brown blotch posterior to orbit at level of top of oper- cle: a second spot posterior to this spot midway between eye and top of opercle, third spot at top of opercle. An additional spot on posteroventral margin of orbit. A diffuse dark brown spot on mid- dle of opercle adjacent to edge of preopercle. Pectoral fin covered with scattered melanophores, with no distinct spots. Anal fin evenly covered with melanophores. Caudal fin of female crossed by seven distinct bars. Caudal fin and first and second dorsal fins of males evenly covered with melanophores. First dorsal fin of female crossed by four bars angling dorsoventrally across fin. Second dorsal fin of female crossed by six similar bars. COLOR OF FRESH SPECIMEN.— (from photograph of specimen taken at Kane’ohe Bay) Background color golden brown with dark brown mottled pattern scattered over body, darker on top of back by first dorsal fin, nape, top and sides of head. Body with about five irregular rows of small, white spots from pectoral-fin base to caudal-fin base. Ventral surface of head and body white with scattered peppering of melanophores. Side of head with scattered small, white spots. Eye with black pupil and iris dark brown, lips gray with white spots. Pelvic fins white basally, remaining two-thirds dusky. Caudal fin dark brown with white spots on basal one-third of rays, rays darker than membranes. Anal-fin membranes light gray on basal two-thirds, distal one-third dark brown. Pectoral fin dark brown with small, white spots on basal one-half. First dorsal fin with a peppering of melanophores, crossed by three rows of white spots, distal margin orange. Basal half of second dorsal fin with a peppering of melanophores, distal half dark brown, fin crossed by four to five rows of white spots. 510 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 DISTRIBUTION.— Widespread throughout the tropical Indo-Pacific region. REMARKS.— Typically found in tidepools, shallow reef flats, and sheltered patch reefs. Often misidentified in recent literature (including Gosline and Brock, 1960) as B. fuscus (Rippell). MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218101 (12), CAS 218102 (13), CAS 218103 (28), CAS 218104 (3), CAS 218105 (16), CAS 218106 (19), CAS 218107 (4) BPBM 5483 (49), BPBM 5486 (23), BPBM 5488 (43). Bathygobius cotticeps (Steindachner, 1879) (Figs. 17-18 and 52) Gobius cotticeps Steindachner, 1879, Sitzungsber. Akad. Wiss. Wien. 80(1 Abth.):137. Society Islands. Holotype: NMW 30439. Chlamydes laticeps Jenkins, (1903), Bull. U.S. Fish. Comm. 22(1902):503, Fig. 43, O’ahu , Hawaiian Islands. Holotype: USNM 50716. DIAGNOSIS.— Counts based on 25 individuals, measure- ments on 10 individuals 27.8—57.8 mm SL. Dorsal-fin elements VI-I,8-1,9 (VI-I,9). Anal-fin elements I,7-I,8 (1,8). Pectoral-fin rays 21—25 (24), upper 8—10 (9) rays filamentous, branched to based with membrane-free ends. Longitudinal scale series 35-39 (37). Predorsal scales 21—32. Pelvic sucking disc about as wide as long, moderately long, space between end of fin and s anal-fin origin less than one fin length; broad frenum with an FiGuURE 52. Bathygobius cotticeps, obvious spine on each side of frenum pointing posteriorly. Body BPBM 38464, 57 mm SL, Waikiki, covered with ctenoid scales, becoming small and cycloid on rep head and anterior body, extending forward onto head to between eyes, onto upper portion of oper- cle and onto cheek under eye in larger individuals. Gill membranes broadly united to isthmus. Jaws extending posteriorly to anterior part of pupil. Interorbital narrow, about equal to pupil diameter. Anterior nostril with a small flap. Side of cheek with a deep longitudinal groove, with upper por- tion of cheek extending down over groove anteriorly, hiding papillae in groove. A small fleshy lobe projecting posteriorly from front of chin, without a posterior projection on each side. Caudal fin rounded, shorter than head. Head strongly depressed. Body depth 4.2—5.0 in standard length. Reported to 110 mm TL. COLOR IN ALCOHOL.— Head and body light cream. Body crossed by four brown saddles: first from second spine of first dorsal fin posterior past end of fin base to end of depressed last spine and down to belly; second saddle from first ray of second part of dorsal fin posterior to base of 5‘ ray and down to midline of body where it joins third saddle; third saddle from base of 7" ray pos- terior about two scales past end of fin base and down to midline of body; posterior third of third saddle extends posteroventrally to end of last depressed anal-fin ray on ventral surface of caudal peduncle; fourth saddle encircles caudal peduncle just anterior to caudal-fin base. Head with a dark bar extending posteroventrally from posterior part of ventral surface of eye across cheek * distance to edge of preopercle: second shorter bar extending from anteroventral portion of eye onto premax- illa; distinct round spot at posterior edge of eye, second spot posterior to this spot midway between eye and top of opercle, third spot at top of opercle. Pectoral fin with scattered melanophores on basal */. Anal fin with scattered melanophores on basal */s. Caudal fin crossed by six faint bars com- posed of separate spots on fin rays. First part of dorsal fin with a dark spot covering membrane between spines five and six continuing past sixth spine to end of fin. Second part of dorsal fin crossed by six faint bars angling dorsoventrally across fin. Pelvic fin lacking pigment. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 511 COLOR OF FRESH SPECIMEN.— (from photograph of specimen taken at Waikiki). Background color medium brown with dark brown edges on scales on sides of body and head, entire scales dark brown on dorsal surface of body and head. Ventral surface of head and body white with scattered melanophores. Eye with black pupil, iris brown with gold reflections. Grooves on cheek cream, contrasting with darker background of cheek. Lips dark brown. Pectoral, anal, and caudal fins uni- form dark brown to gray. Pelvic fins tan. First dorsal fin with a clear area on basal one-fifth between spines two and six. Remainder of fin darker, spines orange-brown and membranes dusky brown, an intense black spot between spines five and six and extending slightly posterior to sixth spine. Second dorsal-fin membranes dusky brown, spine and rays orange-brown. DISTRIBUTION.— Found in the Indo-Pacific from East Africa to the Hawaiian Islands and Pitcairn Islands; in the western Pacific from Japan to the Great Barrier Reef. REMARKS.— Usually found in tidepools and rocky shores. This species is more abundant in the low intertidal, in areas that do not get cut off from the ocean during low tide (D.J. Shafer, pers. commun., March 1998) MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: BPBM 4869 (2), BPBM 5492 (1), BPBM 5493 (3), BPBM 5494 (2), BPBM 11659 (3), BPBM 15354 (8), BPBM 15370 (7), CAS 218109 (2), CAS 218165 (1); uncat. Univ. Hawaii (2), uncat. Univ. Hawaii (2), Univ. Hawaii 2018 (4). Bryaninops amplus Larson, 1985 (Figs. 27, 31, and 53) Bryaninops amplus Larson, 1985, The Beagle (Occ. Pap. N. Terr. Mus. Arts Sci.) 2(1):66, figs. 5-6. Lagoon off east tip of Palfrey I., Lizard I., Great Barrier Reef, Australia. Holotype: AMS 1[.22916-001. Paratypes: BPBM 27979 (1), Hawaii, BPBM 29319 (9), Hawaii, other paratypes at various museums. DIAGNOsISs.— Counts based on two individuals 23.9-25.2 mm SL. Dorsal-fin elements VI-I, 8. Anal-fin elements 1,9. Pectoral-fin rays 15. Longitudinal scale series 46-49. Body variably scaled, the scales extending forward to between end of pectorals and below fourth dorsal-fin spine, abdomen naked. Snout pointed and bill-like, longer than eye diameter. A large curved canine tooth at midside of lower jaw. Gill opening reaching ventrally a lit- tle below pectoral-fin base. Pelvic suck- ing disc slightly longer than wide, but short, space between posterior end of fin and anal-fin origin greater than pelvic-fin length. Pelvic-fin frenum with thickened skin around pelvic spines, forming lobes that extend posteriorly over disc. Interorbital slightly narrower than one pupil diameter with two pores in center of interorbital, one next to each eye. No scalloped grooves along lower preopercular edge. Head depth less than 50% of head length. Body depth at anal-fin origin 6.2—9.0 in standard length. Caudal fin slightly emarginated to truncate with round- ed corners. No dark bars on back. Reported to 56 mm TL. COLOR IN ALCOHOL.— Background color light cream. Body with a few scattered melanophores on back with slight concentrations in front of first dorsal fin, between spines five and six, at first three elements of second dorsal fin, at fifth ray of second dorsal fin, behind second dor- FiGuRE 53. Bryaninops amplus, 45 mm TL, on seawhip, Kaua’1. SW PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 sal fin, and at end of caudal peduncle. Lower half of body with heavier concentration of melanophores, becoming more dense towards caudal-fin base and extending onto basal one-third of caudal fin. Remainder of caudal fin without pigmentation. Top of head with scattered, larger dark brown spots. Cheek and opercle with a few scattered melanophores and a heavy concentra- tion of melanophores on anterior end of upper and lower jaws. Pectoral-fin base with scattered melanophores, no pigmentation on fin. Both dorsal fins, anal and pelvic fins lack pigmentation. COLOR IN LIFE.— (from a photograph taken at 21 m at Kaua‘1) Body translucent with most apparent color showing through from internal pigmentation. Internal coloration: A bright white line running along backbone. Area below backbone dark brown with two distinct white bars running across gut area from backbone to ventral surface. A series of about eight bright white spots spaced on ventral portion of body above anal-fin base. Red gills show through side of head, inner surface of eye balls and brain white. External coloration: An orange-red stripe extending the length of body just ventral to backbone. Six evenly spaced, orange-red triangles extending dorsally from stripe to top of back. A small patch of orange-red pigment with a few scattered melanophores on top of head over brain and a similar color combination in front of eyes across snout onto upper jaw. Dorsal sur- face of center of upper jaw with a white spot. Iris of eye black surrounded by a narrow white ring, remainder of iris orange-red. A few larger scattered melanophores on pectoral-fin base. All fins lacking color. DISTRIBUTION— Reported from the Great Barrier Reef, Western Australia, Northern Territory, Philippines, Okinawa, Palau, Guam, Hawaiian Islands, Seychelles, and Madagascar. REMARKS— Usually found on seawhips of the genus Juncella, but also found on mooring lines. Male-female pairs often occur on a single seawhip; eggs are laid in an encircling band on the seawhip. Has been taken in Hawaiian waters from Ni ihau, Kauai, and Kona, Hawai 1. MATERIAL EXAMINED— HAWAIIAN ISLANDS: Nv ihau: BPBM 37292 (1); Kaua’i: BPBM 37904 (1), 10 paratypes listed above. Bryaninops tigris Larson, 1985 (Fig. 54) Bryaninops tigris Larson, 1985, The Beagle (Occ. Pap. N. Terr. Mus. Arts Sci.) 2(1):70, Figs. 7-8. On dropoff halfway between Bird and South Islands, Lizard I., Great Barrier Reef. Holotype: AMS 1.20730-017. Paratypes: BPBM 18073(6), other paratypes at various museums. DIAGNOsIS.— From Larson (1985). Dorsal-fin elements VI-I,7-8. Anal-fin elements 1,8-9. Pectoral-fin rays 12—14 (usually 13), the lower three or four rays unbranched and thickened. Longitudinal scale series 32—59 (mean 47). the scales usually reaching to above pectoral-fin base, occasionally a little anterior to it. Midline of nape usually naked (rarely a few predorsal scales), abdomen naked midventrally, and usually the sides as well. Body slender, the depth at anal-fin v8 origin 6.9-8.6 in standard length. Head FIGURE 54. Bryaninops tigris, 30 mm TL, on black coral, Wetar, width about equal to head depth. Snout — [ndonesia. length about equal to orbit diameter. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAITAN ISLANDS 513 Caudal fin truncate. Interorbital narrower than pupil diameter. Gill opening short, ending at or slightly anterior to lower edge of pectoral-fin base. Pelvic fins short and cup-like. Reported to 55 mm TL. COLOR IN ALCOHOL.— Background color light cream. Sides of body lacking pigment. A slight concentration of brownish pigment immediately anterior to first dorsal-fin spine, and between spines three and four, and at base of spine six. Brownish spots also present along base of second dorsal fin between rays one and two, three and four, and at bases of rays five, six and seven. Scattered melanophores present on top of caudal peduncle. There is a series of seven brownish spots spaced along the anal-fin base and ventral surface of caudal peduncle. Scattered melanophores present on top of head behind eyes. A brownish bar extending from front of eye for- ward onto upper jaw. Caudal-fin base dark brown with less intense pigment extending out to end of fin. First dorsal fin with scattered melanophores, more concentrated anteriorly and distally. Second dorsal fin and anal fin with scattered melanophores. COLOR IN LIFE.— Lower side of body blackish to dusky red or orange. Back transparent except for six or seven narrow dusky orange or red bars that are continuous with color of lower side, and a series of small spots of the same color dorsally on the body, one in each space between the bars. A white line along top of vertebral column. A dusky orange or red stripe from front of snout through eye and across postorbital head. Two transverse dusky orange bands on occiput. A prominent black blotch at caudal-fin base. DISTRIBUTION.— Known from the Chagos Archipelago, Gulf of Thailand, Solomon Islands, Great Barrier Reef, Tahiti, and the Hawaiian Islands. REMARKS— Commensal on black coral (Antipathes). Reported from depths of 15—53 m. The only Hawaiian specimens were collected in 53 m off Kauai on 4. dichotoma. MATERIAL EXAMINED— HAWAIIAN ISLANDS: Kaua’1: PBM 18073 (6) paratypes, specimens small and dried. INDONESIA: BPBM 37376 (7). Bryaninops yongei (Davis and Cohen, 1968) (Figs. 29, 33, and 55) Cottogobius yongei Davis and Cohen, 1969, Bull. Mar. Sci. 18(4):752, Figs. 1, 4-6. Darvel Bay, Borneo. Holotype: USNM 200402. DIAGNOSIS.— From Larson (1985). Dorsal-fin rays VI-I,7-10 (usually 8 or 9). Anal-fin rays 1[,8-9. Pectoral-fin rays 13-17 (usually 15-17, mode at 16). Posterior half of body covered with ctenoid scales, those above midline extending forward to first dorsal fin. Longitudinal scale series 26-58 (mean 40). Body depth at anal-fin origin 5.4-8.1 in standard length. Head width greater than head depth. Snout slightly longer than orbit diameter. Gill opening reaching ventrally to below pectoral-fin base. Shee! : FIGURE 55. Bryaninops yongei, 35 mm TL, on seawhip, Pelvic disc usually cup-like and short, not —olokini. Maui. reaching anus, the spines with fleshy lobes. Recorded to 40 mm TL. 514 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 COLOR IN ALCOHOL.— Background color light cream. Body and fins lacking markings except for a few scattered melanophores on the pectoral-fin base, on the top of the head just behind the eyes, and immediately in front of the first and second dorsal fins. COLOR IN LIFE— (from photograph of fish taken at Molokini, Maui in 27 m). Body translu- cent with backbone showing through as a silver stripe. Sides of body with a series of six dark brown triangular-shaped blotches, their apices pointing dorsally. Apex of first triangle located just anterior to first dorsal fin; second at end of first dorsal fin; third at second dorsal-fin origin; fourth at posterior third of second dorsal fin; fifth on caudal peduncle; and sixth at caudal-fin base. Coloration of head variable, one individual, with snout, mouth, cheeks and lower two-thirds of opercle dark brown to black and top of head white. Head of second fish translucent yellow with scattered melanophores on preorbital, snout, premaxilla and occiput. Pupil of eye black, surround- ed by a narrow lemon yellow ring, remainder of iris orange-red, with a scattering of irregular black pigment on the outer perimeter. First dorsal-fin spines black, membranes clear. Rays of second dor- sal, anal, caudal, and pelvic fins reddish, membranes clear. Pelvic-fin ray reddish and membranes clear except for white distal margin that rests on seawhip. DISTRIBUTION.— Red Sea to the Hawaiian Islands, Marquesas, Society Islands, and Rapa; Japan to Great Barrier Reef. Only Hawaiian specimens taken were three in 1967 (O’ahu, Lahilahi Point: USNM 203238: Moku Manu, O’ahu: BPBM 5571), two in 1968 (Hawaii, Puako: USNM 203237); and four in 2001 at Kane’ohe Bay, O’ahu: CAS 218903. REMARKS.— Lives on the antipatharian seawhip Cirrhipathes anguina at depths of three to at least 45 m, typically with one male and female pair per seawhip, sometimes with a few juveniles as well. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: Oahu: BPBM 5571 (1), Moku Manu; CAS 218903 (4), Kane’ohe Bay. SEYCHELLES: BPBM 35540 (2). Cabillus caudimacula Greenfield and Randall, sp.nov. (Figs. 40-41 and 56-57) MATERIAL EXAMINED.— Holotype: CAS 218110, male, 16 mm SL, Hawaiian Islands, O’ahu, Kane’ohe Bay, sand field at seaward edge of spur and groove habitat outside of barrier reef, adja- cent to rubble patch and pavement, (15.2—16.8 m), collected by R.C. Langston, 29 August 2002, field number L02-28. Paratypes: CAS 218111, 14.8 mm SL, USNM 375433, 16.8 mm SL, FMNH 113491, 14.8 mm SL, NTM S.15733-001, 13.3 mm SL, AMS 1.42940-001, 14.3 mm SL, NSMT- P 67809, 14.2 mm SL, all taken with the holotype. BPBM 37261, 18 mm SL, Hawaiian Islands, O’ahu, Kane’ohe Bay, Sampan Channel near buoy, spur and groove, 12—13.5 m, 30 August, 1991, D.W. Greenfield and G. Cockrell (field number G91-26). BPBM 39246, 16.5 mm SL, Hawaiian Islands, O’ahu, Kahe Point, sand and rubble, (14.5 m), R.R. Holcom, 18 November 1998. BPBM 22628 (3) (former UH 1708) 15.0-17.4 mm SL, Hawaiian Islands, O’ahu, off Waikiki reef, W.A. Gosline, V.E. Brock, J.E. Randall et al., December 1952. DIAGNosiIs.— A small (largest 18.0 mm SL), light colored goby with a depressed head (width 1.4—1.8 into head length and always wider than deep); a bilobed tongue; pelvic-fin frenum greatly reduced, flat and not raised and turned back (outer margins about one-third pelvic spine length, shorter in center and easily torn), and the pelvic spines not thickened; a terminal mouth; chin with a small, slightly curved mental frenum, and no barbels present on underside of head; tips of upper pectoral-fin rays not free; cheeks without prominent vertical fleshy flaps that bear papillae; dorsal- fin origin behind pectoral-fin base; no spines on preopercle: dorsal-fin spines thin and flexible; no dermal crest anterior to first dorsal fin; body scaled forward to pectoral fin; first gill slit open. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAITAN ISLANDS Sls Dorsal-fin elements VI-I,8-9. Anal-fin elements I,8-9. Pectoral-fin rays 16-18. Longitudinal scale series 24-25. Branched caudal-fin rays 13-16, usually 15. Two to three gill rakers on lower arch. Large dark spot covering caudal-fin base and out onto basal portion of fin. Pelvic dise longer than wide, reaching anal-fin origin. Body covered with scales, extend- ing forward to pectoral-fin base. Two large scales with enlarged cteni at top and bottom of caudal-fin base, extending over fin rays. Scales absent from base of first dorsal fin and from head. Gill membranes fused to isthmus, gill openings not extend- ing forward of pectoral-fin base. Caudal fin rounded. Jaws extending posteriorly to anterior margin of pupil. Interorbital very narrow, less than one pupil diameter. Largest specimen 22.3 mm TL. DESCRIPTION.— Data for holotype FIGURE 56. Cabillis caudimacula, out of BPBM 39246 (speci- men lost); about 16 mm SL. 2 ee reS ec ae presented first, followed by range for all FIGURE 57. Cabillus caudimacula, paratype, BPBM 39246, specimens and mean or mode. Measure- 16.5 mm SL, Kahe Point, O’ahu. ments as percentage of standard length. Data from 12 specimens, 13.3-17.9 mm SL. Dorsal-fin elements VI-I,9 (VI-I,8-9, usually 1,9). Anal-fin elements I,8 (1,8-1,9, usually I,8). Pectoral-fin rays 17 (16-18, usually 17). Branched cau- dal-fin rays 15 (13-16, usually 15). Lateral scales 25 (24-25, usually 25). Gill rakers in four paratypes 1 + 2 (2),1+3,2+2. Head length 31.2 (30.3—34.7: 32.1). Head width 20.3 (17.3-24.1: 20.7). Head depth 13.7 (12.2—16.5: 14.4). Eye diameter 10.3 (9.2—11.5: 10.8). Snout length 6.2 (4.6-6.2: 5.6). Body depth 13.1 (12.6—-18.9: 15.6). Caudal-peduncle depth 9.4 (9.3—10.9: 10.0). Caudal-peduncle length 21.6 (15.5—22.3: 19.3). First dorsal-fin base 15.6 (10.4-18.3: 13.9). Second dorsal-fin base 25.9 (21.3—28.5: 25.4). Anal-fin base 20.3 (16.9-23.3: 20.6). Caudal-fin length 26.6 (26.3—31.2: 27.9). Pectoral-fin length 25.6 (25.3-31.1: 28.2). Pelvic-fin length 23.1 (23.1-36.1: 29.3). Anterior nostrils tubular, extending anteriorly to upper lip. Posterior nostril with a low, raised ridge. A band of cardiform teeth present in both jaws, outer row the largest. Cheek with four rows of longitudinal sensory papillae: one directly under the eye, two in the center of the cheek, and the fourth at the bottom of the cheek. The following head pores are present: paired nasal pores, anterior interorbital, posterior interorbital, paired supraotics, paired anterior otics; intertem- poral, a pore between the anterior otic and intertemporal, three preopercular pores. COLOR IN ALCOHOL.— Background color of head and body white, with color pattern of black pigment. A distinctive dark blotch posteriorly on side of caudal peduncle and extending onto cau- dal-fin base. Four small blotches on midline of side of body, evenly spaced between front of sec- ond dorsal fin and blotch at caudal-fin base. A wide band running from ventral base of pectoral-fin axil posteriodorsally to first dorsal-fin base. Small blotches at base of second dorsal fin at elements three, five, eight, nine and 10, with scattered pigment extending onto dorsum to lateral midline from each blotch. A diffuse saddle on top of caudal peduncle joining blotch at caudal-fin base. A band running posteriorly from top of pectoral-fin base to join wide band under first dorsal fin; a 516 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 narrow extension from the band extending up to dorsal surface, crossing anterior to first dorsal fin, joining band on other side. Top of head and snout with scattered melanophores. An intense narrow black spot on midline of head midway between dorsal-fin origin and top of pectoral-fin base. A dis- tinct bar running from anteroventral eye margin forward onto jaws. Cheek under eye and opercle with scattered melanophores. A cluster of melanophores on dorsal quarter of pectoral-fin base, extending out onto fin a short distance, remainder of fin lacks pigment as does the pelvic and anal fins. Caudal fin without pigment except for blotch at its base. Second dorsal fin with blotches at its base as previously described. First dorsal fin of one specimen with scattered blotches, the other almost completly black, an extension of the wide band running up from the body, with only the basal half of the first few spines lacking pigment. (Black pigment fades rapidly in preservation.) COLOR IN LIFE.— (from photograph of live specimen from Kahe Point, O’ahu). Background color of head and body white. A distinctive black triangle on side of caudal peduncle and extend- ing onto caudal-fin base, the apex of the triangle pointing anteriorly and the base extending onto the rays and membranes. A cluster of melanophores on body under pectoral fin. Three small, black spots on body midline spaced under second dorsal fin. A dusky blotch on top of nape and on top of head behind eyes. An oblique black band extending across eye and continuing onto upper and lower jaws. Pupil black, iris white. First dorsal fin clear except for a few black pigment spots at base of membrane behind last spine. Second dorsal fin clear except for small black spots on bases of elements three to five. Caudal, anal, pectoral and pelvic fins clear. EtryMoLocy.— The specific epithet, caudimacula, is a compound adjective from the Latin cauda for tail and macula for spot, in reference to the prominent black blotch posteriorly on the body and caudal-fin base, the most distinctive color marking of the species. DISTRIBUTION.— Known at present only from O’ahu, Hawaiian Islands. COMPARISON.— Cabillus tongarevae (Fowler, 1927) from Tongareva (an atoll also known as Penrhyn Id.) in the northern Cook Islands, the only other described species of the genus in the islands of Oceania, differs in having 13 branched caudal rays, | + 4 gill rakers, not having the two strongly ctenoid scales overlapping the base of the caudal fin, and in color; it lacks the large black spot below the dorsal fin and has prominent black spots on the dorsal fins (still visible on the holo- type in the Bishop Museum). There is an excellent color photograph of C. tongarevae from the Ryukyu Islands in Masuda et al. (1984:274, pl. 354, fig. I). It is also positively known from Kanton Island in the Phoenix Islands (Schultz 1943), Marshall Islands (Randall and Randall 1987), and Great Barrier Reef, Australia (Russell, 1983). Cabillus macrophthalmus (Weber, 1909) was cap- tured in Indonesia at a depth range of 120-400 m. The drawing (Fig. 31) in Koumans (1953), shows that it has a dark spot on the base of the front of the first dorsal fin, has a broad dark mark under the first dorsal fin that runs forward onto the nape, and lacks the large black spot at the caudal-fin base. Cabillus lacertops (Smith, 1959) has been reported from Mozambique, the Ryukyu Islands, and the east coast of northern Australia (Masuda et al. 1984), and Tonga (Randall et al. 2004). In the key to the species, Nakabo (2002) illustrated the coloration of C. /acertops, showing a distinct black mark at the caudal-fin base that turns upward. This species also lacks any black pigment on the body under the first dorsal fin. Hayashi and Shiratori (2003) also have photographs of C. lac- ertops and C. tongarevae (page 116). REMARKS.— Specimens were collected from sand near reefs at depths of 1.5—15 m. MATERIAL EXAMINED.— Cabillus tongarevae: MARSHALL ISLANDS, ENEWETAK ATOLL: BPBM 10980 (1); ANDAMAN SEA, SIMILAN ISLANDS (NW of Phuket): BPBM 22803 (1); CORAL SEA, CHESTERFIELD BANK: BPBM 33709 (1). GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS Sky) Coryphopterus duospilus (Hoese and Reader, 1985). (Figs. 38-39 and 58-59) Fusigobius duospilus Hoese and Reader, 1985, J.L.B. Smith Inst. Ichthyol., Spec. Publ. No. 36:1-9. Escape Reef, Great Barrier Reef, Australia. Holotype: AMS 1.22619-026. Fusigobius neophytes africanus Smith, 1959, Ichthyol. Bull., Dept. Ichthyol., Rhodes Univ. No. 13:208, pl. 11F (in part, Aldabra and Pinda, Mozambique only). DIAGNOSIS.— Counts based on 27 individuals, 16.0-49.7 mm SL. Dorsal-fin elements VI-I,9. Anal-fin elements 1,8. Pectoral-fin rays 18—19 (19). Longitudinal scale series 24—25. Pelvic fins united to form a sucking disc that is longer than wide, reaching to anal-fin origin. Frenum weak and easily torn. Interorbital very narrow, less than '2 pupil diameter. Scales on body ctenoid, becoming cycloid anteri- or to paired fins. No scales on operculum and no median predorsal scales. Jaws extending posteriorly to between anterior eye margin and pupil. Gill opening extending forward nearly to edge of pre- opercle. Head about as deep as wide, snout pointed. Body depth 4.3—5.8 in SL. Caudal fin rounded, slightly shorter than head. A small dark spot at center of caudal fin. First dorsal fin usually with dark markings on outer part, one on membrane between first and second spine, and sec- ond on membrane between second and third spine, angling to base of third spine. Largest specimen 57 mm TL. COLOR IN ALCOHOL.— Background color of head and body pale cream. Body with five irreg- ular lines of light brown spots running length of body, upper two lines extending onto nape and head. A distinct dark brown spot on caudal-fin base, covering bases of five central fin rays. A light brown spot, filling most of the center of a scale, above and slightly posterior to dorsal surface of pectoral-fin base. A second more diffuse spot posteroventral to previous spot, located under pec- toral fin. Top of head with series of small, light brown spots continuing forward from lines on body and a series of spots running forward from insertion of first dorsal fin onto nape, the one closest to dorsal fin the largest and most distinct. Side of head and snout with scattered small, dark brown spots. Eye with black pupil surrounded by silver iris, with small dark brown spots on upper two thirds, lower third under pupil lacking spots. Skin on top of eye with three dark brown lines run- ning towards interorbital area. Pectoral-fin base with two light brown blotches, one on the upper third, the other on the lower third. Upper blotch narrowing and extending onto about four fin rays. A distinct dark brown bar on inside of fin at same location as bar on outside. Remainder of pec- toral fin immaculate. Pelvic and anal fins clear. Caudal fin clear except for a few scattered light brown spots on membranes of basal half. First dorsal fin usually clear except for a dark brown spot at middle of second spine extending onto fin membranes anteriorly to first spine and posteriorly as O; ats 518 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 lighter spots across fin. Distinct dark brown spots at base of third spine and on body between spines five and six. First spine of second dorsal fin crossed by three evenly spaced dark brown bars. Remainder of fin with a few scattered light brown spots on membranes. Distinct dark brown spots at bases of rays two and six of second dorsal fin. COLOR IN LIFE.— (from underwater photograph of fish taken at Kona in 18m) Background color of head and body translucent white. Five subcutaneous black blotches along backbone which has silver reflections: first under origin of first dorsal fin; second between first and second dorsal fins; third under posterior half of second dorsal fin; fourth behind second dorsal fin; fifth at cau- dal-fin base (blotches not visible in fresh specimens when clear tissue turns opaque white). About five irregular lines of small orange-brown spots running length of body, upper two lines extending onto nape and head. A series of silvery spots running along back at fin bases. Another row of sil- very spots running along side from caudal-fin base to pectoral-fin axil about midway between mid- line of side and ventral body surface. Pectoral-fin base with two yellow-orange blotches, one on upper third, other on the lower third. Side of head with scattered yellow-orange spots of varying size. A row of small yellow-orange spots running anteriorly from anteroventral margin of eye across upper lip. Pupil of eye black, iris silver-yellow with a series of evenly spaced brown spots on upper two-thirds; lower third under pupil lacking brown spots. Pectoral, pelvic, and anal fins clear with silvery reflections. Caudal fin clear with scattered small silvery reflections and scattered orange-brown spots. Spinous dorsal fin clear on basal half, upper half with a yellowish tinge. A dark brown spot at middle of second spine extending onto fin membranes anteriorly and posterior- ly. Second dorsal fin clear on basal half, upper half with a yellowish tinge, a few scattered orange- brown spots on membranes. DISTRIBUTION.— East coast of Africa to the Hawaiian Islands and Marquesas; Japan to Great Barrier Reef. REMARKS.— Typically found on sand and rubble next to reefs where it seeks shelter under coral or stones. Collected from 1-46 m. Hawaiian specimens of Coryphopterus were misidentified as Fusigobius neophytes (Gtinther) by Gosline and Brock (1960), a valid non-Hawaiian species. Randall (1995) placed Fusigobius in synonomy of Coryphopterus, a decision followed here; how- ever, Thacker and Cole (2002) have argued that both genera are valid. The second, posterior, dark spot on the first dorsal fin appears to be less developed in specimens from the Hawaiian Islands than from other areas, but we were unable to find any morphological characters to separate the Hawaiian population. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218112(1), CAS 218113 (2), CAS 218114 (1), CAS 218115 (2), CAS 218116 (1), CAS 218117 (1), CAS 218118 (2), BPBM 19657 (3), BPBM 37262 (1), BPBM 37858 (1); Hawai’i: BPBM 15171 (1), BPBM 28733 (2); Maui: BPBM 28710 (5); Midway Atoll: BPBM 34769 (2). OGASAWARA ISLANDS: BPBM 35085 (2), BPBM 35159 (1), BPBM 35205 (3), BPBM 35237 (2). MARQUESAS: BPBM 12097 (1), BPBM 12766 (1). PALAU: BPBM 19714 (1), BPBM 37722 (1). MALDIVES: BPBM 32978 (2). Discordipinna griessingeri Hoese and Fourmanoir, 1978 (Fig. 60) Discordipinna griessingeri Hoese and Fourmanoir, 1978, Japan. Jour. Ichthyol. 25(1):21, figs. 14. Holotype: USNM 214889. Paratypes: BPBM 5884 (1): BPBM 11266 (1). DIAGNosIs.— Count for one Hawaiian specimen presented first, followed by counts from lit- erature. Dorsal-fin elements V-I,8 (I,7-8, rarely 7). Anal-fin elements I,8. Pectoral-fin rays 18 (17-19). Longitudinal scale series 26 (22—25). Scales ctenoid posteriorly, cycloid anteriorly. Large GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 519 cycloid scales on top of head, no scales on cheek or opercle. Body depth 5.2—5.8 in standard length. Head distinctly broader than deep. Origin of first dorsal fin far for- ward on body, above posterior end of opercle, and widely separated from the second dorsal fin. First two dorsal spines greatly prolonged in both sexes, the sec- ond longest, about 1.5 in standard length, extending well back above second dorsal oo S b fin. Pectoral fins large, about 2.3 in stan- . dard length, the rays with free tips. Pelvic is 4 is 7 y sucking disc longer than wide, extending FIGURE 60. Discordipinna griessingeri, CAS 218119, 28 mm to anus. Caudal fin somewhat pointed, TL, Kane’ohe Bay, O’ahu. 2.5-3.0 in standard length. Jaws extend- ing posteriorly to anterior margin of pupil. Interorbital narrower than pupil diameter. Gill openings extending forward to edge of preopercle. Reported to 29 mmTL. COLOR IN ALCOHOL.— A broad dark brown stripe running from pectoral-fin base to caudal-fin base, covering most of lower half of body; body above midline cream with three narrow light brown stripes running posteriorly from above pectoral-fin base; ventralmost stripe joining dark brown stripe on lower body where cycloid scales end and ctenoid scales begin; middle stripe run- ning back to caudal-fin base: dorsalmost stripe running back to top of caudal peduncle. Head cream with distinct dark brown spots on top, sides and ventral surface including spots on snout and upper and lower jaws. Pectoral-fin base with a median dark brown stripe extending out onto about half of fin, fin posterior to and below this stripe cream. Dorsal portion and remainder of fin dark brown. First part of dorsal fin uniform dark brown. Second part of dorsal fin with dark-brown stripe on lower one-third of fin; central third of fin cream with scattered melanophores on rays; distal third dark brown. Central rays of caudal fin cream, dorsal and ventral rays dark brown. Anal fin covered with scattered melanophores. Pelvic fins cream. COLOR OF FRESH SPECIMEN.— (photograph of specimen from Kane’ohe Bay-CAS 218119) Background color of head and body off-white. Upper half of body with three longitudinal light brown stripes running from above pectoral-fin base to caudal-fin base. Lower half of body from behind pectoral-fin base to caudal-fin base black. Sides and top of head covered with distinct black spots, a larger, oblong spot running from top of preopercle ventrally onto opercle. Pupil of eye black with a golden yellow rim, iris off-white with seven, evenly spaced, black spots. Pectoral-fin base black, extending out onto central rays and membranes of fin. Dorsal third of pectoral fin brick red, distal margin black with white edge. Portion of fin between black center and dorsal red part white: ventral portion below central black area white. Anal fin brick red with some dusky black except for white basal line and white distal margin. First spine of first dorsal fin white, remainder of fin brick red with overlay of black except for tips of spines which are white. Basal one-quarter of second dorsal fin black, next distal quarter white, remainder of fin brick red with two black ocel- li, margin of fin white. Caudal fin with three distinct sections: dorsal third brick red with three black ocelli; central third white; ventral third brick red near white center grading into black ven- trally, entire fin with narrow, white margin. DISTRIBUTION.— Known from the Red Sea, St. Brandon’s Shoals, Cocos-Keeling Islands, Papua New Guinea, Great Barrier Reef, Fiji, Tonga, Tahiti, Tuamotu Archipelago, Marquesas, Japan, and the Hawaiian Islands. 520 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 REMARKS.— Collected from depths of 1-37 m. Cryptic in coral, known only from collections made with ichthyocide. The second author collected the first specimen of this species in 1967 in Tahiti from 27 m. The first Hawaiian specimen was collected by W. A. Gosline and his class at Kahe Point, O’ahu in 1968 with rotenone; the label states, “caught emerging from Porites near base.” We both collected the second Hawaiian specimen in Kane’ohe Bay, O’ahu in 3 m. The Tahitian specimen was brilliant red, and some others for which the life color is known have also been bright red. The drab red coloration of the Hawaiian specimen is in sharp contrast; it is not known if this is representative of the population in Hawai'i or if the shallow bay habitat influenced the color. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218119 (1). Eviota epiphanes Jenkins, 1903 (Figs 4-6 and 61) Eviota epiphanes Jenkins, 1903, Bull. U.S. Fish Comm. 22(1902):501. O’ahu , Hawaiian Islands. Holotype: USNM 50720; Paratypes: SU 8707 (6). DIAGNOsSIS.— Counts based on 20 individuals 12.3—-14.7 mm SL. Dorsal-fin elements VI-I,8-VI-I,9 (VI-L9). Anal-fin elements I,7-I,8 (1,8). Pectoral-fin rays 15-17 (17). Longitudinal scale series 23-24 (23). Pelvic fins separate with many side branches and finger-like; small, unbranched fifth pelvic-fin ray absent. Anterormost branch of fourth pelvic-fin ray short, with fewer than 10 segments; Genital papillae not ruggose. IT pore pres- ent. Spines in first dorsal fin not pro- ‘ longedias ilaments, “Ai darko midpeduncu. FIGURE 61. Eviota epiphanes, 17 mm TL, Kane’ohe Bay, lar subcutaneous spot present towards 6-ahy. midbase of caudal fin. Pectoral-fin rays 10-16 may be branched. Body covered with scales, extending anteriorly to a line between top of pectoral-fin base and origin of first dorsal fin. No scales on head. Caudal fin rounded. Interorbital very narrow, less than '4 pupil diameter. Jaws extending posteriorly about to posterior margin of pupil. Gill opening extending forward to a point midway between posterior margin of opercle and preopercle edge. Greatest body depth 4.0—5.3 in SL. Largest specimen 20 mm TL. COLOR IN ALCOHOL.— Background color of head and body cream. Body with six faint subcu- taneous bars, often only the most posterior one on caudal peduncle obvious. Base of scales pig- mented with a line of several small brown spots, often more obvious on dorsal half of body. Four bars made up of small brown spots crossing head and nape. Anteriormost bar extending ventrally onto cheek. Area behind eyes anterior to first bar densely packed with small black spots. A distinct bar from ventral eye margin extending ventrally across cheek. Mouth and snout cream with few or no spots. Eye with black iris, pupil clear. Pectoral-fin base with scattered small brown spots on upper half, remainder of fin immaculate. Caudal and pelvic fins immaculate. First dorsal fin dark- er than second, pigment variable, almost solid black in some, to pigment only present on mem- branes between a few spines. Second dorsal fin with scattered pigment along base. Anal fin vari- able, from immaculate to a few scattered pigment spots. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 521 COLOR OF FRESH SPECIMEN.— (taken from underwater photograph at Kane’ohe Bay) Background color of head and body off-white. All markings on head and body composed of small black spots enclosed in a larger orange-brown area defining a spot or bar. Base of scales on body pigmented as above, forming rows of chevrons along sides. Head and nape crossed by four rows of blotches forming bars, the anterior one extending ventrally onto cheek. Cheeks with small spots and a distinct bar from ventral eye margin down across cheek. Snout and jaws orange-brown. Pectoral-fin base with blotch on upper half, fin immaculate. First dorsal fin orange-brown along base and darker distally, membranes before and after sixth spine black. Second dorsal fin lighter than first, with several orange-brown blotches along base. Pelvic and anal fins immaculate. Caudal fin with red on rays and membranes. A series of six subcutaneous bars on body: first anterior to first dorsal spine, second under first dorsal fin, third at second dorsal fin origin, fourth at center of second dorsal fin, fifth just posterior to second dorsal fin, and sixth on caudal peduncle just ante- rior to caudal fin. Iris and pupil of eye black. DISTRIBUTION.— Hawaiian Islands, Johnston Island, Line Islands, Ogasawara Islands, and southern Japan. REMARKS.— In a survey of the fishes of Kane’ohe Bay, O’ahu, this was the most abundant species taken (Greenfield 2003). Boehlert and Munday (1996) reported that species in the genus Eviota composed the most abundant taxon in their ichthyoplankton samples taken near O’ahu. Because of their small size, these gobies are seldom seen by divers, but because of their numbers they must play an important role in reef ecology. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218096 (276), BPBM 38391 (33). Eviota rubra Greenfield and Randall, 1999 (Figs 8—9 and 62) Eviota rubra Greenfield and Randall, 1999, Copeia 1999: 439-446. Kane’ohe Bay, O’ahu, Hawaiian Islands. Holotype: BPBM 38385. Paratypes: BPBM 35422 (3), BPBM 35424 (11), BPBM 37257 (3), BPBM 38386 (3), BPBM 38387 (14). DIAGNOsIs.— Counts based on 16 individuals 10.4—-14.9 mm SL. Dorsal-fin elements VI-I,8-VI-I,9 (VI-I.8). Anal-fin elements [,7-I,9 (1,8). Pectoral-fin rays 15-17 (16). Longitudinal scale series 22-25 (25). Pelvic fins separate with many side branches and fringe-like; small unbranched fifth pelvic-fin ray present, about one-tenth length of fourth ray (often poorly ossified and difficult to see). Anterormost branch of fourth pelvic-fin ray elongate, with nine or more segments. 7 a ~— Genital papillae of male and female wt FIGURE 62. SUBS, 15 mm TL, off Kane? ohe Bay, O’ahu, gose. IT pore absent, POP pore present. 5 m. No dark ventral-midline spots or midpe- duncular subcutaneous spot towards midbase of caudal fin. Spines in first dorsal fin not prolonged as filaments. Pectoral-fin rays 9-16, may be branched. Body covered with scales, extending ante- riorly to a line between top of pectoral-fin base and origin of first dorsal fin. No scales on head. Caudal fin rounded. Greatest body depth 4.1—5.6 in SL. Interorbital very narrow, less than '/ pupil ae 5h ‘ ‘ PRE is ey = 322 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 diameter. Jaws extending posteriorly about to posterior margin of pupil. Gill opening extending forward to a point midway between posterior margin of opercle and preopercle edge. Largest spec- imen 19 mm TL. COLOR IN ALCOHOL.— Background color of body and head light cream, without pigment. Subcutaneous bars absent in specimens from deeper water, but specimens from shallower stations (12-15 m) with faint subcutaneous bars. Head with a few, scattered, tiny red-brown pigment spots. Elements and membranes of first and second dorsal, anal and caudal fins covered with small, dis- tinct reddish brown spots. No pigment on pectoral or pelvic fins. COLOR OF LIVE AND FRESH SPECIMENS.— Body translucent with bright red markings when alive, but translucent areas turn white at death. Two distinct color forms, male with lemon-yellow on head and a second (females and perhaps immature males) without yellow. Following descrip- tion of female color is from a live specimen photographed by R.R. Holcom at Piptkea, O’ahu: Body translucent, backbone, neurocranium and body cavity showing through as white with silver reflections. Silver reflections also scattered along dorsal and ventral margins of body, most likely ends of neural and haemal spines. Series of nine red, subcutaneous bars extending from dorsal to ventral side of body overlying silver backbone and body cavity. First bar above pectoral-fin base; second at center of first dorsal fin; third between spines five and six of dorsal fin; fourth at rays two-six of second dorsal fin; fifth at rays eight and nine; sixth at last two rays of second dorsal fin; seventh, eight and ninth on caudal peduncle. Additional red pigment on surface of body at loca- tions of subcutaneous bars and also joining bars at midline to form H-like patterns. Pectoral-fin base red, with a silver stripe running dorsoventrally across base separating red into upper and lower portions. Nape to top of head crossed by four red bars. Upper and lower jaws and snout red. Upper half of operculum red, lower half and remainder of cheek translucent except for a narrow red bar from ventral margin of eye down to lower surface of head. Pupil of eye black, iris silver with blotches of bright red. Pectoral-fin rays pinkish, membranes without pigment. Pelvic fins lack pig- ment. Fin elements of first and second dorsal, anal and caudal fins with alternating red and pink bands, membranes with heavy scattering of tiny reddish brown pigment spots. Color of male taken from slide of specimen photographed underwater at Kane’ohe Bay short- ly after capture. Color pattern on body as in female except that transluscent areas now are white due to death. Subcutaneous color pattern still partially visible. Cheek, snout, underside of head and gill membranes bright lemon-yellow. A wash of red over the yellow upper and lower jaws. Bar under eye red. Top of head with scattered larger melanophores. Fin color as in female except that distal two-thirds of both dorsal fins and anal fin with heavy peppering of black pigment. Basal third of second dorsal and anal fins yellow. DISTRIBUTION.— Hawaiian Islands. REMARKS.— Of the three species of Eviota known from Hawaiian waters, this is the deepest dwelling, usually being taken at ledge and deep spur and groove habitats (12.2—-28.7 m) at Kane’ohe Bay, and was the only Eviota species taken at depths greater than 18.3 m (Greenfield, 2003). The species typically is red, hence the name rubra, and can be separated from E. epiphanes, with which it coexists in the deep spur and groove habitat, because it lacks the black midpeduncu- lar subcutaneous spot toward the midbase of the caudal fin that is present in E. epiphanies. Eviota susanae Greenfield and Randall, 1999 (Figs. 10-12 and 63) Eviota susanae Greenfield and Randall, 1999, Copeia 1999:439-442. O’ahu, Hawaiian Islands. Holotype: BPBM 38379. Paratypes: BPBM 38380 (1), BPBM 38381 (7), BPBM 38382 (3),BPBM 38383 (2). GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAITAN ISLANDS 23 DiIAGNosis.— Counts based on 12 individuals 14.7—19.8 mm SL. Dorsal-fin elements VI-I.8-VI-I.9 (VI-L8). Anal-fin elements 1,6—I,8 (1,8). Pectoral-fin rays 15-18 (16). Longitudinal scale series 24-26 (25). Pelvic fins separate with many side branches and finger-like; small, unbranched fifth pelvic-fin ray present, about one-tenth length of fourth ray (often poorly ossified and difficult to see). Anterormost branch of fourth pelvic-fin . be ray elongate, with 10 or more segments. a Br ; ze ° aS Genital papillae of male and female rug- FIGURE 63. Eviota susanae, 19 mm TL, off Kane’ohe Bay, gose. IT pores absent and POP pores O7’ahu, | m. absent or greatly reduced. Spines in first dorsal fin not prolonged as filaments. Four to six (usually six) large dark spots on ventral midline posteriorly from anal-fin origin; subcutaneous bars associated with some ventral-midline spots. No dark midpeduncular subcutaneous spot towards midbase of caudal fin. Pectoral-fin rays 9-17, may be branched. Body covered with scales, extending anteriorly to a line between top of pectoral-fin base and origin of first dorsal fin. No scales on head. Interorbital narrow, less than '/ pupil diame- ter. Caudal fin rounded. Greatest body depth 4.2—6.8 in SL. Jaws extending posteriorly about to posterior margin of pupil. Gill opening extending forward to a point midway between posterior margin of opercle and preopercle edge. Largest specimen 25 mm TL. COLOR IN ALCOHOL.— Background color of head and body cream. Scale pockets from origin of first dorsal fin back to caudal-fin base with crescents of dark brown pigment. A narrow dark brown line running along middle of body. Four-six (usually six) dark brown spots on ventral sur- face of body from anal-fin origin to caudal-fin base. An additional, smaller, less intense spot may be present near insertion of ventral procurrent caudal-fin elements. Corresponding dark brown spots on dorsal surface of body. Some of these external spots are at ends of subcutaneous bars (more obvious in fresh specimens) extending from dorsal to ventral surface of trunk. Pectoral-fin base peppered with melanophores, often concentrated into darker upper and lower blotches. Four dark brown bars crossing back anterior to first dorsal fin and top of head, either distinct or indis- tinct. Ventral side of body with scattered melanophores. Side of head mottled in females with indi- cations of a bar from anterior ventral margin of eye down across cheek; in males the bar is narrow- er and there is less pigment on cheek. Upper and lower jaws with scattered melanophores, pigment under chin. Pectoral fins with light scattering of melanophores on membranes. Pelvic fins lack pig- ment. Caudal fin with bar across base where scales are present, remainder of fin with heavy pep- pering of melanophores. Anal and second dorsal fins with heavy peppering of melanophores. First dorsal fin with heavy peppering of melanophores and black spot on membrane posterior to last spine. COLOR OF FRESH SPECIMEN.— (from photograph of specimen taken at Kane’ohe Bay, male) Lower side of head lemon yellow, including below eye, all of upper and lower jaw, snout onto interorbital, lower half of cheek and opercle, all of gill membranes, and under lower jaw. Markings on head rust-orange over yellow, a distinct bar from ventral margin of eye down across cheek. Pupil of eye black, iris golden-silver with blotches of reddish brown. Top half of head and rest of body white with rust orange markings. Pectoral-fin base rust orange overlaid with scattered melanophores except for a white bar on central portion angling anterodorsally up onto opercular 524 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 membrane. Base of scales on body with rust orange pigment. Genital papilla white. Basal three- fourths of caudal fin peppered with mixed red and black pigment spots giving an overall reddish appearance, distal one-quarter white with fewer black pigment spots. Anal fin with similar mixture of red and black pigment spots, but more black than red; base of fifth ray with a red spot, tips of spine and rays white. First dorsal-fin spines with alternating red and white sections, membranes with same mix of red and black pigment as caudal and anal fins, but more black than in second dor- sal-fin membranes; black spot on membrane behind sixth spine. Second dorsal fin with mixture of red and black spots as in anal fin, but more red than black, basal one-half more red, tips of rays white. Pectoral fins with red pigment on basal two-thirds of rays, scattered black and red spots on membranes of basal three-fourths, distal one-fourth clear, tips of rays white. Females-color same as males except no yellow on head which is white with more rust orange bars radiating out on cheek from eye. DISTRIBUTION.— Hawaiian Islands. REMARKS.— This is a species that typically is found in very sheltered areas, often part of the fouling community (Greenfield, 2003). In fact, the type locality of this species is the floating boat dock at the Hawaii Institute of Marine Biology in Kane’ohe Bay. It is the largest of the Eviota species in the Hawaiian Islands, reaching a length of 18.5 mm SL. Gnatholepis anjerensis (Bleeker, 1851) (Figs. 34-36 and 64) Gobius anjerensis Bleeker, 1851, Nat. Tiydschr. Ned. Indié 1:251, fig. 11. Java. Holotype unknown; Neotype BPBM 26651. Gobius ophthalmotaenia Bleeker, 1854, Nat. Tijdschr. Ned. Indié 7:46 (type locality, New Selma, Cocos- Keeling Islands). Gobius capistratus Peters, 1855. Monats. Akad. Wiss. Berlin 1855:443. (type locality, Ibo, Mozambique). Gobius deltoides Seale, 1901, Occ. Pap. B.P. Bishop Mus. 1(3):125 (type locality, Guam). 1905:487, pl. 58. Gnatholepis corlettei Herre, 1936, Zool. Ser., Field Mus. Nat. Hist. 21:356, fig. 20 (type locality, Bushman Bay, Malekula Island, Vanuatu). DIAGNOsIs.— Counts based on 25 in- dividuals 30.0-47.8 m SL. Dorsal-fin ele- ments VI-I,10-11 (1,11). Anal-fin elements J,11. Pectoral-fin rays 15-17 (70% 16, 13% 17). Longitudinal scale series 30; ctenoid scales on body extending anterior- ly to below origin of spinous dorsal fin (sometimes forward on side of nape to above middle of opercle); predorsal scales extending to posterior interorbital space, cycloid scales ventrally on abdomen, on prepelvic and prepectoral areas, and on opercle and cheek (rarely a few ctenoid FIGURE 64. Gnatholepis Higa aI 58 mm TL, Kane’ohe Bay, scales posteriorly on opercle of large spec- O’ahu, 1.5 m. imens); usually a few small scales on cheek entirely anterior to dark bar below eye. Narrow (less than pupil diameter) subcutaneous black bar extending down from under eye across cheek and under head. Pelvic fins united to form t GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 325 a disk, longer than wide, almost reaching anal-fin origin; frenum broad. Gill membranes broadly united to isthmus, opening ending just below pectoral-fin base. Jaws extending posteriorly to between anterior margin of eye and pupil. Interorbital very narrow, less than one pupil diameter. Body depth 3.9-4.6 in SL Caudal fin rounded, usually slightly longer than head. Largest specimen 105 mm, but rarely exceeds 80 mm TL. COLOR IN ALCOHOL.— Color varies greatly depending on the habitat, with individuals from sand areas being much lighter and those from dark habitats such as mangrove mud, much darker. Color descriptions of specimens from both habitats are given. SAND HABITAT— Background color of head and body light cream. Head with a narrow, subcu- taneous, dark brown to black bar running from ventral margin of eye, down across cheek to its ven- tral margin. Remainder of head covered with very small, scattered melanophores. A light brown blotch above pectoral fin, extending forward to eye as a line. Body with six elongate light-brown blotches along midline. A seventh blotch at caudal-fin base. Lighter bars entending up to dorsal sur- face from each blotch. Two of these blotches are at base of first part of dorsal fin, one on caudal peduncle and remainder at second part of dorsal fin. Scattered dark brown spots on blotches. Pectoral, pelvic and anal fins covered with small, dark brown chromatophores. First dorsal fin with a dark brown stripe running along basal one-third of fin, a narrow less pigmented stripe above this; remainder of fin dark brown. Second dorsal fin dark brown on membranes between rays that lack pigment. Caudal fin either without pigment or a few scattered dark brown spots. MANGROVE HABITAT— Background color of head and body medium cream. Subcutaneous dark bar under eye as in light form. A dark stripe running from posterior margin of eye posteriorly to above pectoral-fin base. A dark brown to black subcutaneous blotch running from margin of cen- tral portion of preoperculum onto operculum. Remainder of head covered with dense, small, dark brown spots, some coalesced into small blotches. Upper lip crossed by five dark brown bars, first at midline, one on each side midway between midline and end of premaxilla and another fainter bar at ends of premaxillas. Body blotches as in lighter color form except pigment darker and denser. Blotches at midline joined together resembling a stripe. A series of dark-brown spots arranged in rows running from pectoral-fin base back to caudal fin. Three rows above midline stripe and two below. Pectoral, pelvic, and anal fins as in lighter color form only darker. First dorsal fin as in lighter form except distal half of fin with an additional clear stripe in its center. Second dorsal fin with distinct dark spots arranged in five bars across fin. Caudal fin crossed by about eight rows of distinct spots. COLOR IN LIFE.— (from color photograph taken of a 50-mm individual in shallow water on sand in Kane’ohe Bay) Body white with silvery reflections on some scales. All markings referred to as “dark” are black-brown. A bright yellow humeral spot, the anterior half of yellow humeral spot surrounded by dark pigment that then runs forward to eye as a line. Pupil black, surrounded by a narrow yellow ring. Iris white, with a small dark spot at center at both the front and back of eye. A dark bar running from ventral edge of pupil down across cheek. A narrow, dark bar dorsal- ly on eye, centered over posterior half of pupil. Snout with reticulated dark markings. Both upper and lower jaws with some dark markings. Opercle with a dark line extending from center of pec- toral-fin base to edge of preopercle where it widens. Six dark blotches running along side of body, just ventral to midline. A series of much smaller, distinct, dark spots scattered along dorsal half of body anteriorly, and extending to lower half of body from second dorsal fin posteriorly to caudal- fin base. Pectoral and pelvic fins immaculate. Dorsal fins with a few, scattered small, dark spots. Caudal fin crossed by a number of rows of small, dark spots. DIsTRIBUTION.— Red Sea and east coast of Africa to the Hawaiian Islands and islands of French Polynesia; southern Japan to Great Barrier Reef and Lord Howe Island. 526 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 REMARKS.— Gobius anjerensis was described by Bleeker (1851) from a simple drawing with- out color markings of a specimen that had been collected in the period of 1821—1823 by Kuhl and van Hasselt from Anjer (now Anyer) in the Sunda Straight, Java. No specimen is extant. Bleeker (1874) selected Gobius anjerensis as the type species of the subgenus Gnatholepis, later elevated to a genus. Although one might guess that the drawing of Gobius anjerensis represents a species of Gnatholepis, it certainly cannot be identified to any known species. Were it not its selection as the type species of Gnatholepis, it would surely have been regarded as a nomen dubium. Randall and Greenfield (2001) have described a neotype of Gnatholepis anjerensis collected in the Java Sea off the southwest end of Sulawesi. It is one of the two most common species of the genus in Indonesia, and the species most often found in shallow water. The Bishop Museum has 95 lots of Gnatholepis anjerensis ranging from the Red Sea and coast of East Africa to the Hawaiian Islands and Society Islands. Of the 62 lots for which there is infor- mation on the depth of capture, the specimens of 36 lots were collected from less than 2 m. Some were taken from tidepools, one from a brackish pool (salinity 12 0/oo). The deepest collection was from 46 m in the Red Sea; the deepest collection in the Hawaiian Islands was 26 m. Typically this species lives on sand very near coral reefs or rocky substrata into which it can seek shelter with the approach of danger. Of the 12 lots listed below that were collected around the island of O’ahu, seven were from Kane’ohe Bay. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: Hawaii: BPBM 28720 (1); Maui: BPBM 38359 (11) Moloka’i-BPBM 15132 (3); O’ahu: BPBM 1846 (1), BPBM 5495 (1), BPBM 7306 (5), BPBM 15040 (5), BPBM 15046 (13), BPBM 15130 (1), BPBM 15042 (21), BPBM 17800 (1), BPBM 19665 (10), BPBM 22649 (49), BPBM 31319 (1), BPBM 31320 (28), CAS 218120 (18), CAS 218121 (121), CAS 218122 (56); Kaua’i: BPBM 15041 (6); Midway Atoll: BPBM 34795 (1), BPBM 34873 (5). JOHNSTON ISLAND: BPBM 15043 (6), BPBM 15134 (3), BPBM 38361 (1). Gnatholepis cauerensis hawaiiensis Randall and Greenfield, 2001 (Figs. 37 and 65) Gnatholepis cauerensis hawaiiensis Randall and Greenfield, 2001, Ichthyol. Bull. J.L.B. Smith Inst. Ichth., no. 69:10, pl. Il C, D. [Described as a subspecies of Gnatholepis cauerensis (Bleeker)]. Gobius cauerensis Bleeker, 1853. Nat. Tijdschr. Ned. Indié 4:269, (type locality, Cauer = Kauer, Sumatra). Gnatholepis scapulostigma Herre, 1953, Philip. Jour. Sci. 82(2):193 (type locality, Enewetak Atoll, Marshall Islands). Gnatholepis inconsequens Whitley, 1958, Proc. Roy. Zool. Soc. N.S.W. 1956—57:44 (type locality, Heron Island, Capricorn Group, Great Barrier Reef). Acentrogobius cauerensis, Bleeker, 1983, Atlas Ichthyologique, unpublished plates for vols. XI—XIV. Smithsonian Inst. Press, Washington, D.C. Pl. 435b, fig. 1 (reproduction of previously unpublished color figure, without final corrections). DIAGNOsIs.— Counts based on 26 individuals 14.0-42.0 mm SL. Dorsal-fin elements VI-I,11. Anal-fin elements I,11. Pectoral-fin rays 16-18 (92% 17, 2% 16). Longitudinal scale series 30; ctenoid scales on body extending at least as forward as below origin of spinous dorsal fin, excep- tionally on side of nape to above posterior margin of preopercle; predorsal scales extending to pos- terior interorbital space, cycloid scales ventrally on abdomen, on prepelvic and prepectoral areas, and on opercle and cheek (rarely a few ctenoid scales posteriorly on opercle); usually no scales on cheek entirely anterior to dark bar below eye. Narrow (less than pupil diameter) subcutaneous black bar extending down from under eye across cheek and under head. Pelvic fins united to form a disk, longer than wide, almost reaching anal-fin origin; frenum broad. Gill membranes broadly united to isthmus; gill opening ending slightly ventral to pectoral-fin base. Interorbital very narrow, GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS S27 less than one pupil diameter. Body depth 4.2-5.1 in SL. Caudal fin rounded, usual- ly slightly longer than head. Largest spec- imen, BPBM 15131, 56 mm TL, from Maui. COLOR IN ALCOHOL.— Body whitish to pale yellowish brown with a row of six large dusky blotches on lower side and brown longitudinal lines following center of scale rows (two rows darker when superimposed on dusky spots); a dark brown to black blotch above pectoral-fin base containing a small pale spot; a nar- FIGURE 65. Gnatholepis cauerensis hawaiiensis, 50 mm TL, row blackish bar dorsally on eye from Kona, Hawai’i, 29 m. above center of pupil, crossing midin- terorbital space; a prominent narrow black bar extending ventrally and slightly posteriorly from below middle of eye: dorsal fins with longitudinal rows of dusky dashes; large specimens develop- ing dark lines, one per membrane, parallel to rays, in soft dorsal fin; caudal fin with a dark brown line in membranes between rays. COLOR IN LIFE.— (from color photograph taken of 50-mm individual at 28.5 m on dark sand in Kona, Hawaii) Body gray with six dark (brownish black) blotches running along side of body, just ventral to midline, with a large, blue area around and below each blotch. A cream to yellowish stripe running along midline just above blotches. A series of about six longitudinal lines along sides, running along center of scale rows. Lines dark brown in this specimen, but may be red or orangish-brown. Numerous small, blue spots along ventral half of sides, extending from caudal peduncle forward to front of eye. A bright yellow humeral spot present, surrounded by a black spot. A narrow, dark brown line running forward from humeral spot to eye, where it widens slightly. Iris of eye cream, pupil black, surrounded by a narrow golden ring. A prominent black bar starting at top of eye, across iris, through center of pupil, and continuing ventrally across cheek as the diag- nostic feature for the genus. The bar across the cheek is as wide as the pupil (narrower in speci- mens living on a light-colored background). A dark brown line running from center of pectoral-fin base to preopercle where it widens. Snout and both jaws with reticulated black markings. Pectoral fins immaculate. Both dorsal fins gray with longitudinal rows of dark brown dashes. Caudal fin with reddish brown lines on membranes. DisTRIBUTION.— Hawaiian Islands and Johnston Island. REMARKS.— Gnatholepis cauerensis was first reported from the Hawaiian Islands by Randall and Greenfield (2001). It ranges from the coast of East Africa to the islands of Oceania; however, it has differentiated into at least four subspecies: wide-ranging from East Africa to the Society Islands; islands of the southeastern Pacific (Rarotonga, Austral Islands, Rapa, and Pitcairn Group), Easter Island, and the Hawaiian Islands. The Hawaiian population is distinct from the Indo-Pacific subspecies in having a slightly longer caudal peduncle (1.5—1.7 in head length, compared to 1.71.9), a dark line on membranes of the caudal fin instead of small dark spots, and numerous small blue spots on the lower side of the body. Also it rarely has small scales on the cheek entire- ly anteriorly to the dark eye bar. The population at Johnston Island differs from the Hawaiian in having a slightly shorter cau- dal peduncle (1.6~1.8 in head) and by consistently having small scales on the cheek anterior to the dark eye bar. Knowledge of the life color is lacking. 528 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 Of the eight lots of Gnatholepis cauerensis taken in Hawaiian waters, one was obtained in 2 m, the others all from 14-29 m. As in G anjerensis, this species is usually found on sand near the shelter of rock or reef. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: Hawai’1: BPBM 28734 (1), BPBM 37860 (1), BPBM 37861 (3); Maui: BPBM 15131 (14); O’ahu: BPBM 7930 (1), BPBM 12284 (1) BPBM 37847 (1), BPBM 37859 (1); Midway Atoll: BPBM 15137 (1), BPBM 34770 (2). Kelloggella oligolepis (Jenkins 1903) (Fig. 66) Enypnias oligolepis Jenkins, 1903, Bull. U.S. Fish Comm. 22 (1902):504, Fig. 45. O’ahu, Hawaiian Islands. Holotype: USNM 50715. DIAGNOsIS.— Counts based on 20 specimens 10.1—24.0 mm SL. Dorsal-fin elements VI-I,10-12 (1,11). Anal-fin ele- ments I,6-8 (1,7). Pectoral-fin rays 12—13 (13). Body naked, with 10—11 dark verti- cal bars separated by narrow white bars. Snout rounded, its end either reaching upper jaw or slightly overhanging mouth. Gill membranes broadly united to isth- mus. Jaws extending to under pupil. Interorbital narrow, about equal to pupil diameter. Pelvic sucking disc longer than : | wide, space between'end of fin and anal; ” ~7rcuen 66, Kelloeeeiia ofcolae =) BEG ie eam fin origin equal to or greater than pelvic- Barber’ Point, O’ahu. fin length; frenum broad. Greatest body depth 5.6—7.2 in SL. Caudal fin rounded. Maximum size 29 mm TL. COLOR IN ALCOHOL.— Mature male color pattern: Background color cream. A distinct dark- brown spot on side of head above operculum, often joined by a light brown bar across top of head. An additional light brown bar across top of head anterior to first at preopercle. Diffuse dark brown spot at dorsoposterior margin of eye. Cheek variously marked with light brown bars reaching from posteroventral margin of eye and under chin. Body crossed by 10 light brown bars from anterior of first dorsal fin to caudal peduncle, eleventh bar on caudal peduncle present or absent: Bars sepa- rated by narrow white bars, narrow bars about '/ width of dark bars; first bar across back 1n advance of first dorsal fin extending to belly; second bar from front of first dorsal fin back to base of fourth spine and down to belly; third bar from bases of fifth to sixth spine down to lower part of side, lighter or absent on ventral surface; fourth bar from posterior end of first dorsal fin to origin of sec- ond dorsal fin, down to lower part of side, lighter or absent on ventral surface; sixth bar from pos- terior base of third dorsal-fin ray to base of sixth ray and down to anal fin; seventh bar extends from posterior of ray six to ray seven and down to anal fin; eighth bar from front of base of eighth ray to base of ninth ray and down to anal fin; ninth bar from bases of rays nine and ten down to ven- tral surface of caudal peduncle; tenth bar from base of last rays to behind end of fin down to ven- tral surface of caudal peduncle; eleventh bar, if present, encircling caudal peduncle. A diffuse spot at caudal-fin base. Ventral surface of body either cream or with a light scattering of melanophores. Pectoral, pelvic, anal, and caudal fins with scattered melanophores. First dorsal fin with scattered melanophores with distal half of fin darker in some specimens. Second dorsal fin with interspersed light and dark areas on membranes between every other fin element. ee GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS Sy) Immature and female color patterns (specimens 13.7 mm SL and less [immature]): Trunk of body without pigment, but about eight subcutaneous pigment bars visible through body. A series of 15 black spots on dorsal surface, two on top of head, two anterior to first dorsal fin, two along base of first dorsal fin, one between first and second dorsal fin, six along base of second dorsal fin and two on dorsal surface of caudal peduncle. Four black spots along anal-fin base and two on ventral surface of caudal peduncle. Distinct black spot on head above opercle. A series of black bars radi- ating out from behind and below eye. Pectoral, pelvic, anal, and caudal fins clear. Distal margin of first dorsal fin black and spots on body extending up onto fin. Second dorsal fin with black bars extending up onto fin from body spots. Specimen 17.8 mm (female): Body bars developing as external black spots along midline, dorsal surface, and adjacent to anal-fin base, at locations of sub- cutaneous pigment bars. A series of distinct black blotches on top, side and ventral surface of head. Pectoral, pelvic, anal, and caudal fins with some scattered melanophores and dark areas on dorsal fins more distinct. COLOR OF FRESH SPECIMENS.— (from color photograph of BPBM 39151, captured from tide- pools at Barber’s Point, O’ahu) Mature male color pattern as in color in alcohol except that back- ground color of body white, bars and spots on body and head a dusky black, and background color of head yellowish-cream. Light areas on fins are clear and dark areas are dusky black, except for spot on posterior portion of first dorsal fin that is intense black. Female color pattern as in color in alcohol except that background color of body is cream and that of head yellowish cream. Pigment spots on body, head and fins intense black. DISTRIBUTION.— Hawaiian Islands and Easter Island. Although previously recorded from the Ryukyu Island, Japan, the Ke/loggella species there is K. quindecimfasciata. REMARKS.— This species occurs in tidepools high in the splash zone. It seems improbable that there would be sufficient gene flow between fishes found in splash zone tidepools at the Hawaiian Islands and Easter Island; however, Hoese (1975) examined specimens from both localities and was unable to find any differences except that the bands on the body are slightly narrower in Easter Island specimens. We also have examined specimens from Easter Island and the Hawaiian Islands, and also were unable to determine any differences other than the color character mentioned by Hoese. Three other fish species are known to have this unusual antitropical distribution, and a num- ber of mollusks found at Easter Island are also only known from the Hawaiian Islands. One can postulate that these are species that were once more widespread when waters were cooler in the past, and now have been isolated to the north and south. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: BPBM 5499 (14), BPBM 26370 (2), BPBM 31259 (1), BPBM 39151 (7), CAS 218123 (4). EASTER ISLAND: BPBM 6741 (2), BPBM 6742 (2). Mugilogobius cavifrons (Weber, 1909) (Figs. 22, 23, and 67) Gobius cavifrons Weber, 1909, Notes Leyden Mus. 31 (note 4). Sula Takomi, Ternate, Moluccas, Indonesia. Syntypes: ZMA 112616 (43). DIAGNOsIs.— Counts based on 12 individuals 21.8-49.0 mm SL. Dorsal-fin elements V-I,7- VI-L.8 (VI-L8). Anal-fin elements I,7-I,8 (1,8). Pectoral-fin rays 15—16 (16). Lateral scales 37-46 (39-40). Interorbital wide, greater than one eye diameter in individuals 37 mm SL or larger, and covered with very small cirri that extend onto snout and under eye on cheek. Pelvic sucking disc longer than wide, but short, space between posterior end of fin and anal-fin origin greater than pelvic-fin length; frenum broad. Body covered with scales, lateral scales anterior to second dorsal 530 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 fin smaller than those under second dorsal fin and on caudal peduncle. Scales extending onto top of head meeting cirri on interorbital and onto operculum. No scales on cheek. Caudal fin rounded. Depth of body about five in SL. Jaws extending to posterior margin of eye. Gill membranes broadly united to isthmus. Attains 58 mm TL. COLOR IN ALCOHOL.— Background color dark cream, entire body and head overlaid by a heavy peppering of small, dark brown spots. A pattern of dark brown marks on upper half of body: the first a line angling posteroventrally from top of head above top of pectoral-fin base down to midline at line down from second dorsal-fin spine; a blotch between this line and front of first dorsal fin, sometimes blotch extends up to nape; second line from second dorsal-fin spine extending posteroventrally to body midline; third line from sixth dorsal-fin spine extending posteroventrally to body midline. Five bars on posterior half of body extending from dorsal surface to body midline: first at second element of second dorsal fin; second at fourth element; third at ele- ments seven and eight; last two across caudal peduncle. A blotch at caudal-fin base. Two dark brown bars from posteroventral eye margin across cheek, upper bar reaching preopercular margin, lower bar midway to margin. Center of opercle near preopercular margin with a dark brown spot. Body patterns may be more diffuse in larger individuals. First dorsal fin with a dark brown stripe angling from front of fin down to base of last spine and following membrane; stripe beginning on upper third quarter of fin, with clear areas on lower half and upper quarter. Larger males may have scattered pigment on lower half of fin and black on distal fin margin. Second dorsal fin with dark brown pigment on lower three quarters being more concentrated on membranes midway up fin. Dark pigment on fin more extensive on larger males. Pectoral, pelvic, and anal fins densely cov- ered with melanophores. Caudal fin crossed by four to five bars. COLOR IN LIFE.— (from photograph of living specimen taken at Kane’ohe Bay) Background color tan, color patterns overlaying tan are black. Head with a series of distinct lines: One from snout to anterodorsal portion of eye; another from middle of side of upper jaw to ventral margin of eye; a line from posteroventral eye margin to edge of preopercle; a long wavy line from end of jaws across ventral portion of cheek onto opercle. Opercle, preopercle and nape with scattered irregular marks. Side of body with a number of irregular, broken narrow bars. Ventral surface of body light tan. Pectoral, pelvic, and anal fins dusky. Caudal fin crossed by four irregular, black bars. First dor- sal fin crossed by two black bars, the first on distal one-quarter of fin and second on second quar- ter up from base, expanded to cover basal one-half posteriorly. Second dorsal fin dusky with some darker areas. DISTRIBUTION.— Known from Indonesia, Papua New Guinea, Philippines, Tarwan, Ryukyu Islands, Kosrae (Caroline Islands), Guam, and O’ahu, Hawaiian Islands. REMARKS.— This species was most likely unintentionally introduced into Hawaiian waters perhaps via ballast water of a ship. It was first recorded from the mangroves at Coconut Island, Kaane’ohe Bay, O’ahu by Randall et al. (1993) as M. parvus, which is a synonym of M. cavifrons (Larson, 2001). This species now is widespread on O’ahu, being most common in shallow man- grove, brackish and freshwater habitats. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: BPBM 32479 (1), BPBM 33492 (1), BPBM 33931 (4), BPBM 34569 (2), BPBM 34997 (5). FIGURE 67. Mugilogobius cavifrons, BPBM 33492, 22 mm SL, Kane’ohe Bay, O’ahu, 0.1 m. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 531 Opua nephodes E.K. Jordan, 1925 (Fig. 68) Opua nephodes E.K. Jordan, 1925, Proc. U.S. Nat’l. Mus. 66(2570):36, pl. 2, fig. 2. O’ahu, Hawaiian Islands. Holotype: USNM 87419. DIAGNOSIS.— Counts based on 11 individuals 20.5—33.3 mm SL. Dorsal-fin elements VI-I,9-VI-I,11 (VI-I,10). Anal-fin elements [,10-I,11 (1,10). Pectoral-fin rays 17-19 (19). Longitudinal scale series 27-29 (28). First spine in second dorsal fin stout and sharp-tipped, first spine of first dorsal fin also stouter than following spines. Body with a line of dash-like dark marks running along midline of side and four fainter lines above. Pelvic sucking disc longer than wide, almost reaching to genital papillae, frenum broad. Body cov- ered with scales extending forward on top of head to eyes. No scales on opercle or cheek. Gill openings extending forward to below posterior margin of eye. Jaws extending posteriorly to between front of eye and pupil. Interorbital very narrow, less than one pupil diameter. Caudal fin rounded, shorter than head. Greatest body depth 4.8—-5.6 in SL. Largest specimen 57 mm TL. COLOR IN ALCOHOL.— Head and body light cream with light brown markings. A series of five dash-like dark marks running along midline of side, first under middle of first dorsal fin; second under first third of second dorsal fin; third under last third of second dorsal fin; fourth at center of caudal peduncle; fifth at caudal-fin base. Four stripes above midline dashes, lower two running from caudal peducle forward to above pectoral-fin base. Third stripe running from top of caudal peduncle forward onto head and top stripe from base of second dorsal fin onto head. Side of head with diffuse bar under eye that extends posteriorly across cheek and onto opercle. Pectoral-fin base with diffuse pigment on upper half, no pigment on pectoral-fin rays or membranes. Pelvics, anal, and both dorsal fins with scattered melanophores. Caudal fin crossed by three light, indistinct bars. COLOR IN LIFE.— (from photograph of live fish taken at Kane’ohe Bay) Background color of head and body white. Body with six dark brown, longitudinal lines composed of separate dashes running the body length. Ventralmost line begins behind pectoral-fin base and joins the line above below second dorsal-fin origin. The latter line continues to caudal-fin base and has five prominent, evenly spaced, dark brown blotches on it, the last blotch at caudal-fin base. Two dorsalmost lines extending posteriorly from top of head onto body. Entire body and head overlaid with iridescent white blotches. Ventral surface of head and body white. Sides and top of head with scattered dark brown blotches. Eye with black pupil surrounded by golden ring, iris dark brown. Dark brown blotch on upper portion of pectoral-fin base. First dorsal fin peppered with melanophores. Second dorsal fin also peppered with melanophores, but with dark brown blotches on basal third of some rays. Anal fin peppered with melanophores. Caudal fin crossed by three irregular rows of dark brown blotches. Pectoral and pelvic fins clear, overlaid with small, scattered, iridescent white spots. DisTRIBUTION.— Hawaiian Islands. REMARKS.— Specimens have been collected from depth of 1—11.5 m on silty sand and also mud bottom. This species is common on the mud bottom of Kane’ohe Bay, where along with FIGURE 68. Opua nephodes, 35 mm, Kane’ohe Bay, O’ahu, 2m. 552 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 Oxyurichthys lonchotus, it is a common food item of hammerhead shark pups. The Honolulu Laboratory of the National Marine Fisheries Service has recently collected this species using a shrimp trawl from deep water (138-169 m) north of Moloka’1. The deep water specimens appear to have larger eyes than those taken in shallow water. Jennifer K. Schultz (pers. commun., April, 2004) compared tissue samples preserved in 70% ethanol of deeper water and shallow specimens using mitochrondial DNA cytochrome b and concluded that they are the same species. More spec- imens are needed to confirm the apparent larger eye of this goby from deeper water. This species has incorrectly been placed in the genus Hazeus. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: Oahu: CAS 218124 (3), CAS 218125 (16), BPBM 4838 (1), BPBM 14648 (6), BPBM 17820 (21), BPBM 22641 (5) cleared and stained, UH 1772 (9); Maui: BPBM 34920 (5); Moloka’1: 39132 (3). Oxyurichthys heisei Pezold, 1998 (Fig. 69) Oxyurichthys heisei Pezold, 1998, Copeia 1998(3):687-689. Holotype: BPBM 15473. Paratypes: BPBM 34513 (4), BPBM 24145 (3), BPBM 24151 (2), BPBM 24140 (1), BPBM 24084 (1), CAS 74809 (2), LIAIP 1968466 (1), LIAIP 1968467 (1), NLU 64915 (2). DIAGNOSIS.— Counts from Pezold (1998) based on type material (49.4—-63.7 mm SL). Dorsal-fin elements VI-I,12. Anal-fin elements 1,13. Pectoral-fin rays 21—23 (22-23). Longitudinal scale series 50-64. Caudal fin distinctly pointed, more than twice as long as wide. Well-devel- oped fleshy crest on nape extending for- ward from first dorsal fin to preopercle. FIGURE 69. Oxyurichthys heisei, 54 mm SL, off NW Moloka’1, 124 m (specimen lost). Pelvic sucking disc longer than wide, not reaching beyond anus: frenum broad. Body covered with small scales extending forward to above midopercle on nape, but with naked median. No scales on cheek and opercle. Gill membranes fused to isthmus at level of middle of opercle. Jaws extending posteriorly to below middle or posterior third of orbit diameter in females, to below posterior margin of orbit in males. Interorbital very narrow, less than one pupil diameter. Greatest body depth 5.5—7.0 in SL. Largest specimen 118 mm TL. COLOR IN ALCOHOL.— From Pezold (1998): Figure 69. Oxyurichthys heisei, 54 mm SL, off NW Moloka’i, 124 m (specimen lost). “No spot on eye; no spots on gular fold beneath preopercle or anterior process of quadrate; anterior nares not darkly pigmented; body uniformly yellowish brown with faint pigment on cheek beneath posterior portion of orbit (in one specimen, five faint midlateral patches of melanophores are discernible with a microscope); D1 lightly pigmented, but with dark streak between first and second spines, variably indicated (weakly represented in holo- type); D2 lightly pigmented; caudal fin with scattered melanophores, dusky distally and between lowermost rays; anal fin lightly pigmented basally and marginally, with clear zone between bands; pelvic fins with few melanophores on connecting membrane, dusky in patches but primarily con- centrated between distal branches of rays and proximally; between rays 2/3 and 4/5 in some spec- imens; pectoral fins unpigmented.” COLOR IN LIFE.— Unknown. DISTRIBUTION.— All known specimens are from deep water (124-143 m) off the coast of Moloka’i, Hawaiian Islands. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIAN ISLANDS 533 ReEMARKS.— This is one of only two deep-dwelling gobies known from Hawaiian waters. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: Moloka’1: CAS 218155 (1). Oxyurichthys lonchotus (Jenkins, 1903) (Figs. 13—15 and 70) Gobionellus lonchotus Jenkins, 1903, Bull. U.S. Fish Comm. 22(1902):503, fig. 44. O’ahu, Hawaiian Islands. Holotype: USNM 50698. DIAGNOsIs.— Counts based on 20 individuals, 28.7—39.1 mm SL. Dorsal-fin elements VI-I,12-VI-I,13 (VI-I,12). Anal- fin elements I,12-I,14 (1,13). Pectoral-fin rays 18-20 (19). Longitudinal scale series 77-88. Caudal fin pointed, more than twice as long as wide. Well-developed fleshy crest on nape extending forward from first dorsal fin at least past edge of FIGURE 70. Oxyurichthys lonchotus, BPBM 37084, 47 mm SL, opercle and farther in larger individuals. Kahana Bay, O’ahu, 0.1 m. Body with a lateral series of elongate blotches. Dark spot on superior margin of eye. Prominent dark spot on pectoral-fin base and dark spots on branchiostegal membranes under lower jaw. Pelvic sucking disc longer than wide, almost reaching anal-fin origin; frenum broad with fringe on edge. Body covered with small scales extend- ing forward on top of head to a line up from edge of preopercle. No scales on opercle or cheek. Gill membranes fused to isthmus at level of midopercle. Jaws extending posteriorly to mid pupil. Interorbital very narrow, less than one pupil diameter. Greatest body depth 5.3—5.7 in SL. Attains 130 mm TL. COLOR IN ALCOHOL.— Background color of head and body cream. Sides of body with a series of light brown marks corresponding to dark gray marks in color description for fresh specimen. Top of head with a narrow dark brown line extending from posterior margin of one eye to the other eye; two faint, light brown bars crossing top of head posterior to this line. A dark brown blotch below posteroventral eye margin. A small dark brown spot on cheek above end of upper jaw. Anterior tubular nostrils black. Distal margin of crest on nape dark brown. Black spot on superior eye mar- gin. Side of head with diffuse brown pigment on opercle. Isthmus with scattered brown chro- matophores. Branchiostegal membranes under lower jaw with one or two dark brown blotches. Upper two-thirds of pectoral-fin base with a dark brown blotch. Pectoral, caudal, dorsal, and anal fins with scattered light brown chromatophores on rays and membranes. Pelvic-fin membranes dark brown, rays cream or with a few light brown chromatophores. COLOR OF FRESH SPECIMEN.— (from photograph of specimen taken at Kahana Bay, O’ahu) Background color of body, top and sides of head grayish brown, ventral surface of head and body light gray. Body and head with small, scattered, iridescent blue spots. Sides of body with a series of dark gray marks, the first as seven narrow bands under the pectoral fin that almost reach dorsal and ventral surfaces. A dark blotch under end of pectoral fin. Four more blotches on side posterior to end of pectoral fin, the first under fourth element of second dorsal fin; second under seventh to ninth; third under 12‘ to 13‘ and past end of fin; and fourth on caudal-fin base. Another series of dark gray blotches along bases of both dorsal fins: first at first dorsal spine; second at third spine; third at fifth; fourth anterior to first spine of second dorsal fin; fifth at second element of second dorsal fin; sixth between elements five and six; seventh between elements six and seven; eighth 534 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 between elements ten and 11; and ninth at base of 12‘ element. Top of head with a black bar extending from posterior margin of one eye to the other; two blotches between this bar and first dorsal-fin origin. Side of head with a narrow dark gray bar running across center of opercle at a slight posterodorsal angle. A second lighter bar on preopercle paralleling opercular bar. A distinct, small, round, black spot on cheek above end of upper jaw. Dusky area between anteroventral mar- gin of eye and upper jaw, black pigment on both nostrils. Eye with black pupil and orange iris, a black triangular-shaped mark on posteroventral margin of eye, narrow portion of triangle pointing anteriorly. Pelvic fins dusky. Pectoral fins with dark gray blotch on upper two-thirds of base, rays and membranes dusky. First dorsal fin with a dusky stripe on basal one-quarter, a white stripe dis- tal to that on next one-quarter, and a narrow white stripe distal to the dark stripe. Ends of spines with an orange tinge. Second dorsal fin crossed by seven dusky stripes alternating with white stripes angling dorsoventrally across fin. Margin of fin with an orange tinge. Dorsalmost leading caudal-fin ray with alternating black and white bars, remainder of fin dusky, with darker pigment on posteroventral margin of fin and orange on margins in mature individuals. Anal-fin rays white basally with dusky blotches on membranes between rays on basal third of fin. Remainder of anal fin dusky with a white distal margin. DISTRIBUTION.— Hawaiian Islands. REMARKS.— This species is common in areas with soft mud bottoms such as that found around mangroves and at the bottom of Kane’ohe Bay, O’ahu. A series of collections using a small seine were made at Kahana Bay, O’ahu, and snapping shrimp (4/pheus malabariacus) were con- sistently taken along with the gobies. It is possible that O. /onchotus uses shrimp burrows for shel- ter MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218126 (26), CAS 218139 (115); BPBM 1884(1), BPBM 1885 (1), BPBM 1886 (1), BPBM 4884 (2), BPBM 5520 (1), BPBM 5521 (1), BPBM 15382 (13), BPBM 15386 (1), BPBM 37084 (2), BPBM 38854 (1). Pleurosicya larsonae Greenfield and Randall, sp. nov. (Figs. 32, 71-72) MATERIAL EXAMINED.— HOLOTYPE: CAS 218127, 11.4 mm SL, female, Hawaiian Islands, O’ahu, Kane’ohe Bay, patch reef N. of Sampan channel, 1.2—3.1 m, 17 Nov. 2000, field no. GOO- 06, D.W. Greenfield, J.E. Randall, R.C. Langston, D.B. Eckert. PARATYPES: CAS 218128 (6) 8.0-14.7 mm SL, taken with holotype. CAS 218129 (2) 12.9-15.9 mm SL, Hawaiian Islands, O’ahu, Kane’ohe Bay, patch reef N. of Sampan Channel, 0—3.1 m, 8 Oct. 1992, field no. G92-03, D.W. Greenfield and N. Burke; BPBM 39152 (5) 12.1—13.5 mm SL, Hawaiian Islands, O’ahu, Kaneohe Bay, patch reef NE of Crashboat channel, 0—3.1 m, 15 Apr. 1993, field no. G93-01, D.W. Greenfield; NTM S.15734-001 (3) 14.9-15.7 mm SL,same data as BPBM 39152; NTM S.15735- 001 (2) 13.0-13.6 mm SL, Hawaiian Islands, O’ahu, Kane’ohe Bay, patch reef N. of Sampan Channel, 2.4—3.1 m, 20 May 1993, D.W. Greenfield and K. Cole; USNM 375434 (1) 15.6 mm SL, Hawaiian Islands, O’ahu, Kane’ohe Bay, patch reef at Crashboat Channel, 0-4.6 m, 4 May 1991, field no. G91-07, D.W. Greenfield, T. A. Greenfield, R.K. Johnson; FMNH 113492 (1) 14.7 mm SL, taken with holotype.; BPBM 39544 (1) 11.3 mm SL, Hawaiian Islands, O’ahu, Haleiwa, 6 m, 21 August 2004, R. R. Holcom. DIAGNosis.— Small Plewrosicyva with a broad head and moderate snout, eyes set dorsolateral- ly. Second dorsal- and anal-fin rays I,7-I.8 (usually I.8). Pectoral-fin rays 17—18 (usually 17). Longitudinal scales 24. TRB 6. Nape naked. Pelvic fin rounded, cup-like, with fleshy, rounded pelvic spine lobes. Tongue round. Lower gill opening margin attached at level of preopercular mar- ee GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 335) gin. Gill membranes broadly joined to isthmus. Live color translucent with orange markings and some melanophores. DESCRIPTION.— Description format follows Larson (1990) for ease of compar- ison. An asterisk indicates counts of the holotype. First dorsal-fin spines VI*. Second dorsal-fin elements I,7 (1), 1,8(12)*. Anal-fin elements 1,7(3), 1,8(10)*. Pectoral-fin rays 17(8)*, 18(2). Lowermost pectoral-fin rays unbranched and thickened distally 4(3),5(7)*. Branched caudal-fin rays 10(7)*, 11(4). Seales ctenoid. Longitudinal scale count 23(1), 24(5)*.25(4). Nape naked. Gill rak- ers on outer face of first arch 2+1+6(1), 2+1+4(3) (holotype not counted). Guill rakers short. Lowermost half of lower limb of first gill arch bounded by mem- brane to opercle. Measurements in percentage of SL based on 10 specimens 11.5—16.1 mm SL. Data for holotype presented first, followed by range and mean of all specimens. Head and anterior half of body roughly triangular in cross- section (apex dorsal) posterior half of body compressed. Body depth at anus 17.5 (13.2—17.5:15.3). Head length 34.1 (31.6—34.7:33.1). Head rather broad, width always greater than depth. Head width 19.2 (16.3—20.6:17.9.) [56 (48.2-60:54.3% of HL)]. Head depth 16.1 (14.7—17.2:15.9) [47 (42.3-51:47 % of HL)]. Snout moderate 10.9 (8.6—11.7:10.3) [32 (24.7-35:27.6% of HL)], round- ed when viewed from above. Mouth subterminal, ending at point below anterior part of pupil of eye, 13.1 (13.1—17.5:14.8) [38 (38-54:44.7% of HL)]. Large upper lip overhanging mouth anteri- orly, 3.9 (2.3-4.4:3.4) [11.5 (7-13:10.4% of HL)]. Lower lip very narrow. Eyes moderate 9.2 (7.9-11.2:9.5) [27 (24-35.3:28.8% of HL)]. Eyes set dorsolaterally, high on head. Interorbital space narrow, 2.2 (1.8—3.3:2.5) [6.4 (5.4-10:7.5% of HL)]. Anterior nostril in a short tube, poste- rior nostril in a larger tube, directly anterior to eye margin. Tongue rounded. Lower margin of gill opening attached at level of preopercular margin. Gill membranes broadly joined to isthmus at line between posterior eye margin and preopercular edge. Fins low, first dorsal shorter than anterior rays of second dorsal. Anal rays unbranched. Caudal fin rounded. Pectoral fins slightly pointed, reaching to gap between dorsals. Pelvic disc round, cup-like, just reaching anus. Frenum and pelvic spine lobes fleshy and fimbriate, lobes rounded. Upper-jaw teeth very small, fine, and pointed; arranged in a band broader anteriorly, narrow- ing at sides. Four or five enlarged, slender, curved teeth present on each side of upper jaw spaced from front to side, mostly hidden by upper lip. Lower jaw with band of very small, fine, pointed teeth across front; band extending over edge of jaw, so that outermost enlarged and curved teeth angle outward. Sides of lower jaw with row of larger slightly posteriorly curved teeth; and one or two large curved canine teeth at each side of jaw symphysis. Lateral-line canals as for genus (Larson 1990: Fig. 5). Nape naked, scales on body extending anteriorly to area between top of pec- toral-fin base and origin of first dorsal fin. Belly midline under pelvic fins naked. Male genital FIGURE 71. Pleurosicva larsonae, holotype, CAS 218127, 16.0 mm SL, Kane’ohe Bay, O’ahu. PS tae we : 2 FIGURE 72. Pleurosicya larsonae, 15 mm TL, Kane’ohe Bay, O’ahu, 2.5 m. 536 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 papilla moderately long, flattened, with several tiny lobes at expanded tip. Female genital papilla short, cylindrical to slightly flattened, with several small lobes on either side of tip. Largest speci- men 19.5 mm TL. COLOR IN ALCOHOL.— Most color lost in alcohol, body straw yellow. A few scattered melanophores on top of head behind eye, cheek and on pectoral-fin base. A fine, short, dark line on middle of nape midway between eyes and first dorsal fin sometimes present (not in holotype). A few scattered melanophores sometimes present on body. Some dark markings as in Figure 71 retained on dorsal fins. Pigmentation shown in Figure 71 taken from underwater photograph show- ing fresh coloration. COLOR OF FRESH SPECIMEN.— (from photograph taken underwater of freshly collected spec- imen at Kane’ohe Bay) Head and body translucent with digestive tract and vertebrae clearly visi- ble. Body overlaid with a pattern of orange markings as well as some melanophores. Head with a distinct orange band running anteriorly from front of eye to end of snout and across jaws, the most obvious color feature. An orange stripe running along length of lower jaw, and a fainter orange bar running up from end of maxilla to ventral eye margin. Scattered orange pigment extending along preopercular edge to eye. A narrow orange stripe extending from posterior eye margin posteriorly along midline, the stripe breaking up past pectoral-fin base into a series of about 10 dashes. Top of head with a distinct pattern of orange markings. A pair of stripes running from the anterodorsal eye margin towards the ascending process of the premaxilla as a V. Interorbital with two dashes cen- tered along its length and a U-shaped marking at posterior portion of interorbital space with closed end posteriorly. A narrow orange stripe running anteriorly from dorsal-fin origin onto nape, split- ting into a V that almost meets the U-shaped mark. Secondary stripes running from V to posterior eye margins. Nape directly posterior to eyes with scattered large melanophores. Pupil of eye black, iris silver-yellow with an orange ring around pupil. Digestive tract with distinct black spots show- ing through body. Spaces between vertebrae orange with orange pigment extending along neural and haemal spines. Body with some faint orange marbling. Pectoral-fin base with distinct large melanophores, base of pectoral-fin rays orange, remainder of fin clear. Pelvic disc clear, caudal fin clear with scattered orange spots. Anal fin clear with four orange spots with some scattered melanophores spaced along its base. First dorsal fin clear with an orange spot at the base of the first three spines, the fin crossed by three narrow black bars angling from the first spine posteriorly to fin base. Second dorsal fin clear, crossed by four orange bars with a few scattered melanophores at the base of last two fin rays. EtyMoLoGy.— Named in honor of Helen K. Larson of the Northern Territory Museum of Arts and Sciences, Darwin, Australia, whose revision of Pleurosicya laid the foundation for all future work on the genus. COMPARISONS.— Separated from the following species with scaled napes because of its naked nape: P. annandalei, P. australis, P. boldinghi, P. elongata, P. labiata, P. micheli, P. mossambica. Separated from P. prognatha by lacking its distinctive elongate, pointed upper lip; from P. spongi- cola by lacking its very short, steep snout. From P. plicata by lacking its elongate pelvic-spine lobes and having a round rather than a trilobed tongue; from P. bilobata by lacking a distinct black blotch at rear of soft dorsal fin, lacking a bilobed tongue and differing in live coloration; from P. Jringilla by lacking a black spot on anal fin anteriorly; from P. carolinensis by lacking large curved teeth at middle of each side of lower jaw and by having more pectoral-fin rays (17—18 versus 14-15, usually 15 in P. carolinensis); from P. muscarum by having the upper lip overhanging the lower jaw, versus a more terminal mouth in PR muscarum, by lacking the scattering of melanophores over body and five pairs of red lines radiating out from eye, and by having four to five lowermost unbranched pectoral-fin rays versus usually three or fewer in P. muscarum. ees GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS S37) Pleurosicyva larsonae is most similar to P. occidentalis and P. coerulea. It differs from P. occiden- talis by lacking the intense elongate black spot on center of nape; however, like P. coerulea, it sometimes has a fine, short, dark line on the center of the nape. It differs from P. coerulea by hav- ing a narrower head (mean width 54.3% HL versus 72% in P. coerulea), shorter snout (mean length 27.6% HL versus 35% in P. coerulea). It also has a narrower upper lip, 7-13, mean 10.4% of HL versus 13.5, 13, and 12 in three paratypes (ANSP 165112) of P. coerulea. Pleurosicya coerulea is “translucent bluish to blue-green with two reddish stripes on head” (Larson, 1990:20), whereas P. larsonae is translucent gray with orange markings. DISTRIBUTION.— Known only from Kane’ohe Bay, and Haleiwa, O’ahu, Hawaiian Islands. REMARKS.— This species is most certainly more widespread in the Hawaiian Islands but appears to be extremely cryptic. It was only collected five times in a total of 75 general ichthyocide collections, with four of those collections from a specific habitat, shallow patch reefs (O-4.6 m) with good circulation. Our sixth collection from that specific habitat only occurred after a number of futile attempts to collect more individuals for photographs. Members of the genus Pleurosicya generally are commensal with invertebrates or found on seagrasses and algae. Despite hours of searching we have never seen this species alive and thus do not know its specific association; how- ever, Ronald R. Holeom (pers. commun., August, 2004) observed and caught a specimen at Haleiwa, O’ahu, that was living on Porites compressa. Pleurosicya micheli Fourmanoir, 1971 (Figs. 28, 30, and 73) Pleurosycia micheli Fourmanoir, 1971, Cah. ORSTOM Ser. Oceanogr. 9(4):499, fig. 8. Caban I., Philippines. Neotype: AMS I[.21918-071. DIAGNOsIs.— Counts based on 10 individuals 14.0—-17.1 mm SL. Dorsal-fin elements VI-I.8. Anal-fin elements I-8-I,9 (1,8). Pectoral-fin rays 16-18 (17). Longitudinal scale series 25-28 (27). Dark pigment on lower portion of caudal peduncle extending posteriorly as stripe onto caudal fin. Pelvic sucking disc longer than wide, almost reaching anus. Pelvic- fin frenum with thickened skin around pelvic spines that extend posteriorly over disc. Interorbital very narrow, about one- half pupil diameter, a single pore in center FIGURE 73. Pleurosicya micheli, 25 mm TL, Kona, Hawai'i, of interorbital space. Snout pointed from 18m. side view, jaws extending posteriorly to below about center of pupil. Body covered with scales, extending forward onto nape. No scales on opercle or cheek. Gill membranes free from isthmus, gill openings extending forward to posterior margin of pupil. Caudal fin rounded. Greatest body depth 5.0—7.1 in SL. Attains 25 mm TL. COLOR IN ALCOHOL.— Background color of head and body light cream. Scattered melano- phores on lower half of body from anal-fin origin extending as a stripe onto lower half of caudal fin, becoming denser posteriorly. A line of melanophores running on dorsal surface of body along sides of both dorsal fins. A line of melanophores extending anteriorly from origin of first dorsal fin, extending anteriorly to cover area over brain. Scattered melanophores extending posteriorly from 538 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 posterior margin of eye onto top of preopercle. A few melanophores at top of pectoral-fin base. First dorsal fin clear except for a few large melanophores on lower half of membranes between spines four and six. Second dorsal, pectoral and pelvic fins clear. Anal fin with scattered melanophores between spine and first ray, remainder of fin clear. Lower three segmented rays of caudal fin clear, next four segmented rays dorsal to those densly pigmented dark brown, remaining rays of fin clear. COLOR IN LIFE.— (from a photograph taken in 18 m at Kona, Hawai’i) Head and body translu- cent with internal coloration along backbone visible. A series of six dusky red blotches along back- bone, with white areas on top of backbone between the blotches. A dusky red stripe running from end of backbone coloration onto lower middle part of caudal fin. A reddish area on side of head behind eye and a red to yellow stripe running from front of eye across snout to upper lip. Scattered black spots overlaid by red on top of head behind eyes. Eye with black pupil, iris silver with con- tinuation of red stripes in front of and behind eye onto iris, top of iris with black edge. Fins clear except for a few scattered black spots on membranes of lower half of first dorsal fin. DISTRIBUTION.— Seychelles and Maldives to Hawaiian Islands and Society Islands; Ryukyu Islands and Taiwan to Great Barrier Reef, New Caledonia, and Loyalty Islands. REMARKS.— This species is commensal on a variety of hard corals, and appears to be the most common species of the genus over most of its range. It has been collected from depths of 15—38 m. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: Hawaii: BPBM 28736 (34). Priolepis aureoviridis (Gosline, 1959) (Figs. 74-75) Quisquilius aureoviridis Gosline, 1959, Pac. Sci. 13(1):68, fig. 1. O’ahu, Hawaiian Islands. Holotype: USNM 175013. Paratypes: BPBM 14203 [ex UH 1703] (8). DIAGNOSIS.— Counts based on 14 individuals 20.5—31.1 mm SL. Dorsal-fin elements VI-I, 11—VI-I,12 (VI-I,11). Anal-fin elements I,9-1,10 (1,9). Pectoral-fin rays 18-20 (19). Longitudinal scale series 28-30 (28). Predorsal scales 12-16. Body covered with scales, extending on top of head to eyes. No scales on opercle or cheek. Interorbital very narrow (less than '2 pupil diameter) but not forming a deep trench between eyes. Pelvic fin longer than wide, reaching past anus; frenum completely absent. Jaws extending posteriorly to anterior margin of pupil. Gill openings extending forward just past edge of preopercle. Greatest body depth 4.0-4.5 in SL. Caudal fin rounded. Largest specimen 47 mm TL. COLOR IN ALCOHOL.— Background color of head and body cream overlaid with a peppering of scattered melano-phores that are more concentrated at scale margins. Body crossed by six faint brown bars extending from dorsal to ventral surface: first under fifth spine of first dorsal fin; sec- ond at origin of second dorsal fin; third at fifth element of second dorsal fin; fourth at tenth ele- ment; fifth about three scales past posterior end of second dorsal fin; sixth bar at caudal-fin base. Side of head with two faint light brown bars; first under center of pupil extending down across cheek; second from posterior eye margin across cheek to ventral margin of preopercle. Dorsal, anal, and caudal fins with scattered melanophores, no pigment on other fins. COLOR OF FRESH ADULT SPECIMEN.— (from underwater photograph of freshly collected spec- imen at Kaneohe Bay) Background color of head and body lemon yellow. Head and body crossed by a series of narrow light gray bars. A bar under center of pupil extending down across cheek. Another bar extending from posterior eye margin down across cheek and also dorsally across top of head behind eyes. Three more bars between eyes across interorbital. Eye with black pupil, sur- rounded by a bright yellow ring, iris lemon yellow. Nape crossed by two bars, anteriormost at pos- ; < GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAITAN ISLANDS Sy) terior edge of preopercle, second one directly anterior to first dorsal fin. Six more bars on body, extending from dorsal to ventral surface: first under fifth spine of first dorsal fin: second at origin of second dorsal fin; third at fifth element of second dorsal fin: fourth at tenth element; fifth about three scales past posterior end of second dorsal fin: sixth bar at caudal-fin base. Scales on body with light brown margins. All fins lemon yellow with a pep- pering of melanophores on basal portions. COLOR OF FRESH JUVENILE SPECI- MEN.— Head and body lemon yellow with 20m. a heavy peppering of melanophores. Light gray bars as in adult. Second dorsal, anal, caudal, and pelvic fins lemon yellow with a heavy peppering of melanophores on membranes between yellow elements. Fin margins with a narrow band of black edged with white distally. First dorsal fin similar to other fins except for a black ocellus ringed with white on anteroventral portion of fin covering first five spines (Fig. 75). Pectoral fins lemon yellow with light peppering of melanophores. DISTRIBUTION.— Hawaiian Islands, Johnston Island, and Caroline Islands (Pohnpei and Ant Atoll). 18 m. REMARKS.— This attractive, yellow species is found on outer reefs, but is not usually seen because it is secretive. In Kane’ohe Bay it was most common at the deep spur and groove and ledge habitats (Greenfield 2003). MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218901 (1), CAS 218902 (3), CAS 218899 (3), CAS 218170 (1), CAS 218169 (3), CAS 218168 (1), CAS 218132 (2), CAS 218898 (8), CAS 218900 (1), BPBM 7902 (6), BPBM 9860 (1), BPBM 12289 (4), BPBM 15461 (1), BPBM 24455 (1), BPBM 31017 (1), BPBM 34570 (1), BPBM 34571 (2), BPBM 34573 (1), BPBM 34535 (2), BPBM 35427 (5), BPBM 35778 (4), BPBM 37267 (4); Hawai’i: BPBM 10901 (2), BPBM 17825 (1), BPBM 17826 (1), BPBM 24799 (1), BPBM 28711 (1). Priolepis eugenius (Jordan and Evermann, 1903) (Figs. 44-45, and 76) Quisquilius eugenius Jordan and Evermann, 1903, Bull. U.S. Fish Comm. 22(1902):203. O’ahu, Hawaiian islands. Holotype: USNM 50674 (missing). Paratypes: CAS-SU 7483 (3). Gobiomorphus eugenius, Jordan and Evermann, 1905, Bull. U.S. Fish Comm. 23(pt. 1)(1903):357. DIAGNOsIs.— Counts based on 20 individuals 21.3—38.5 mm SL. Dorsal-fin elements VI-I,9- VI-I, 11 (VI-I,11). Anal-fin elements I,9. Pectoral-fin rays 18-20 (19). Longitudinal scale series 540 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 27-29 (27). Body covered with scales, extending anteriorly on top of head to eyes and ventrally on head to at least upper half of cheek and opercle. Interorbital very narrow (less than ', pupil diameter), form- ing a deep trench between the eyes. Pelvic fin longer than wide reaching to anus, frenum completely absent. Jaws extending posteriorly to anterior margin of pupil. Gill openings extending forward to edge of preopercle. Caudal fin rounded. Greatest body depth 3.6-4.4 in SL. . : . Largest specimen 54 mm TL. IGURE 76. Priolepis eugenius, 32 mm itl, Kane’ ohe Bay, COLOR IN ALCOHOL.— Background O’ahu,1.5 m. color of head and body light brown. Body crossed by six dark brown, wide bars, positions as in fresh specimen described below. Pattern of dark brown bars on head as in fresh specimen. Eye with gray pupil, surrounded by black iris. First and second dorsal and caudal fins dark brown to black. Anal and pelvic fins dark brown to black with white margins. Pectoral fins light brown. COLOR OF FRESH SPECIMEN.— (from underwater photograph of freshly collected specimen from Kane’ohe Bay) Background color of head and body gray, overlaid with dark brown color pat- tern. Body crossed by six wide bars: first under first four spines of first dorsal fin; second from behind fifth spine to end of membrane of first dorsal fin; third under second to fourth elements of second dorsal fin; fourth under elements seven to nine; fifth behind last ray of fin onto caudal peduncle; sixth at center of caudal peduncle. Caudal-fin base with blotch. A wide bar crossing nape in front of dorsal fin and running down anterior to pectoral-fin base. Two narrow bars across head between posterior margin of eyes and bar on nape. Two narrow bars extending down from ventral eye margin across cheek, one from anteroventral margin of pupil and other from posteroventral pupil margin. Pupil black, surrounded by a red-orange ring, iris gray with bands dark brown radi- ating out from pupil. First and second dorsal fins, anal fin and pelvic fins black with a narrow white margin. Pectoral fins gray. DISTRIBUTION.— Hawaiian Islands and Johnston Island. REMARKS.— This species can be found in shallower water than the other species in the genus in Hawaiian waters. In Kane’ohe Bay it was collected most often in the shallow spur and groove habitat, open patch reefs and even in tidepools, all areas with considerable water movement (Greenfield 2003). MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu, CAS 812130 (2), CAS 218166 (6), CAS 812131 (1), CAS 218167 (15), BPBM 8513 (1), BPBM 15349 (3), BPBM 19662 (6), BPBM 22626 (49), BPBM 22640 (4), BPBM 22671 (5), BPBM 37263 (2); Moloka’i, BPBM 4794 (2), BPBM 30225 (1); Midway Atoll, BPBM 34784 (2), BPBM 34799 (4), BPBM 34822 (1), BPBM 34855 (1), BPBM 34874 (6), BPBM 34891 (1), BPBM 35371 (1). Priolepis farcimen (Jordan and Evermann, 1904) (Figs. 43 and 77) Gobiopterus farcimen Jordan and Evermann, 1904, Bull. U.S. Fish Comm. 22(1902):205. Hawai'i, Hawaiian Islands. Holotype: USNM 50654. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 541 Zonogobius farcimen, Gosline and Brock, 1960, Handbook of Hawaiian fishes. University Press of Hawai’1, Honolulu, p. 270. DiaGnosis.— Based on 7 individuals 10.1-15.5 mm SL. Dorsal-fin elements VI-I,8-VI-1,10 (VI-I.9). Anal-fin elements 1,7-I.8 (almost always I,7). Pectoral-fin rays 17-18 (18). Longitudinal scale series 25-28 (26, often lost). Body covered with scales, extending anteriorly no farther than second spine of first dorsal fin. No scales on head or predorsal area. Interorbital narrow (about '/ pupil diame- ter). Pelvic fins longer than wide, reach- Oe) ite. Be ing past anus, frenum completely absent. Be | in 4 ~¥ . a Jaws extending posteriorly to below mid- FIGURE 77. SS 16 mm, Rane one Bae OPA, dle of pupil. Gill openings extending for- 1 m. ward just past edge of preopercle. Caudal fin rounded. Greatest body depth 3.64.4 in SL. Largest specimen 26 mm TL. COLOR IN ALCOHOL.— Background color of head and body light cream. Body, head and fin membranes covered with a fine peppering of brown chromatophores. Color pattern on head (as described in color in life) fades easily in preservative but represented by faint brown lines. Distal margin of first dorsal fin dusky. COLOR OF FRESH SPECIMEN.— (from underwater photograph of freshly collected specimen at Kaneohe Bay). Background color of head red-orange, overlaid by a series of distinct white lines edged in black: three lines crossing interorbital between eyes; a single line from anterior margin of eye running anteriorly to upper jaw; two lines extending ventrally from ventral margin of eye; a line crossing top of head posterior to eyes and continuing down across side of head to ventral sur- face, this line branching posteriorly to eye and extending as an additional line across top of head; a line crossing top of head at level of top of opercle, splitting above top of opercle, with one branch extending down along preopercular margin to ventral surface, the other branch extending down across pectoral-fin base. A short line crossing nape directly anterior to origin of first dorsal fin. Eye with black pupil, iris brunt orange. A series of about seven very short lines extending from bases of both dorsal fins ventrally to where distinct scale pattern begins. Background color of body gray (red-orange when alive) from pectoral-fin base back to origin of second dorsal fin where back- ground color fades to a yellowish hue. Margins of scales with dark brown (black-orange when alive) edges forming a lattice pattern. Pectoral fin burnt orange at base grading into yellow distal- ly. Spines of first dorsal fin with alternating clear and red-orange bands, membranes clear except for distal margin which is black. Second dorsal fin with alternating clear and burnt orange bands, membranes clear. Anal fin similar to second dorsal fin. Caudal fin with yellow tinge. DIsTRIBUTION.— Hawaiian Islands and Johnston Island. REMARKS.— This is the smallest species of Priolepis in Hawaii, and can easily be separated from the other species in our area by the distinctive dark-bordered light lines that radiate out from the eye across the cheek and top of the head. We took fewer individuals of this species at Kane’ ohe Bay than of the other Priolepis species. It usually was found at the outer reefs in deeper water (Greenfield 2003). MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218133 (2), CAS 218134 (3), CAS 218135 (1), CAS 218135 (1), CAS 218136 (1), CAS 218137 (1), CAS 218138 (1); BPBM 15456 542 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 (4), BPBM 15468 (3), BPBM 19663 (7), BPBM 22625 (2), BPBM 22629 (6), BPBM 35426 (1), BPBM 38477 (1); Midway Atoll: BPBM 22627 (1), BPBM 34783 (1), BPBM 34856 (1), BPBM 34867 (1), BPBM 34942 (1). Priolepis limbatosquamis (Gosline, 1959) (Figs. 46 and 78) Quisquilius limbatosquamis Gosline, 1959, Pac. Sci. 13(1):69, fig. 2. Oahu, Hawaiian Islands. Holotype: USNM 175012. Paratypes: BPBM 14201 [ex UH 1704] (8), additional paratype at USNM. DIAGNOsIs.— Counts based on 10 individuals 15.0-22.5 mm SL. Dorsal-fin elements VI-I,8-VI-I,9 (VI-I,9). Anal-fin elements I,7-I,8 (1,8). Pectoral-fin rays 17-19 (18). Longitudinal scale series 25-27 (26). Body covered with scales, extending anteriorly at most */4 of distance to eyes. No scales on opercle or cheek. Interorbital very narrow (less than ‘> pupil diameter) but not forming a deep trench between eyes. Pelvic fin longer than wide, reaching anus, frenum completely absent. Jaws extending posteriorly to below anteri- or margin of pupil. Gill openings extending forward just past edge of preoprecle. | : ; i Caudal fin rounded. Greatest body depth & 2" 4 3sar ey cee’ s ; 5 SWS Ss ae 4.44.9 in SL. Largest specimen 28 mm FIGURE 78. Priolepis limbatosquamis, 30 mm TL, Kane’ ohe ae. Bay, O’ahu, 10 m. COLOR IN ALCOHOL.— Background color of head and body cream. Scales on body with dark brown margins forming a distinct lattice pattern; scales usually lost leaving only this pattern on scale pockets and no body bars as described in color of fresh adult specimen. Ends of bars often evident on dorsal surface on head and bases of fins. Ventral surface of body cream with peppering of melanophores. Head heavily peppered with melanophores, with a brown bar slightly narrower than pupil diameter under center of eye extend- ing down to posterior end of jaw. A second wider bar extending from posteroventral eye margin across cheek to ventral surface. A third bar running along preopercular margin up and across top of head. A fourth bar extending from pectoral-fin base onto dorsal surface of head. Pectoral and pelvic fins with scattered melanophores on basal membranes, clear distally. Caudal fin with scattered melanophores. Membranes of first and second dorsal fins and anal fin with scattered melanophores. COLOR OF FRESH ADULT SPECIMEN.— (from underwater photograph of freshly captured spec- imen at Kane’ohe Bay) Background color of body clear, with six subcutaneous, dark brown bars running from dorsal to ventral surface showing through body: first bar under first dorsal fin; sec- ond between first and second dorsal fins; third under anterior portion of second dorsal fin; fourth under center of fin; fifth at end of fin; sixth at center of caudal peduncle. Scales on body with dark brown margins forming a distinct lattice pattern. Ventral surface of belly light cream. Background color of head cream with an orange wash. A distinct orange bar under center of eye extending down to posterior end of jaws. A more diffuse orange bar extending from posteroventral eye margin across cheek to ventral surface. A diffuse orange bar on posterior preopercular margin running onto GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 543 top of head. A similar bar extending from pectoral-fin base onto dorsal surface of head. A distinct clear area extending from behind pectoral-fin base onto nape anterior to first dorsal fin. Spines and rays of first and second dorsal fins white with orange spots forming oblique rows, membranes pep- pered with small, white, iridescent white spots. Caudal-fin rays white with orange spots, forming five or six irregular bars. Basal two-thirds of anal fin orange, distal one-third white. Pectoral and pelvic fins clear. COLOR OF FRESH YOUNG SPECIMEN.— (from photograph of specimen collected at Kane’ ohe Bay) Background color of head and body clear. Head and body crossed by nine broad brown bands: anteriormost from preopercle across top of head; second across nape anterior to first dorsal fin: third band at anterior portion of first dorsal fin; fourth at posterior end of first dorsal fin; fifth at anterior portion of second dorsal fin; sixth at center of second dorsal fin; seventh at posterior por- tion of second dorsal fin; eight and ninth bands across caudal peduncle. A red internal band angling posteroventrally from sixth to seventh band (retained from larva). A brown band across head at pos- terior eye margin and two brown bands under eye. Eye with black pupil, iris silver. COLOR OF NEWLY SETTLED LARVA.— (from photograph of specimen collected at Kane’ohe Bay) Body clear with a distinctive red bar running posteroventrally from middle of second dorsal fin. A red blotch at caudal-fin base. A red line along lower jaw. Red chevron marks on body under pectoral fin. Black on posteriodorsal part of gut showing through body. Eye with black pupil, iris silver. DISTRIBUTION.— Hawaiian Islands and Johnston Island. REMARKS.— This is a species that usually is found in the deeper outer part of Kane’ ohe Bay, being most abundant in the deep spur and groove habitat of the bay (Greenfield 2003). It is secre- tive and not usually seen while diving. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218140 (2), CAS 218141 (1), CAS 218142 (1), CAS 218143 (1), CAS 218144 (1), CAS 218145 (1), CAS 218146 (1), CAS 218147 (1), CAS 218148 (2), CAS 218149 (1), CAS 218150 (1), CAS 218151 (2), CAS 218152 (3), CAS 218153 (1), CAS 218154 (1), BPBM 10038 (1), BPBM 17811 (2), BPBM 31016 (1), BPBM 37264 (1); Midway Atoll: BPBM 34802 (1), BPBM 34823 (2), BPBM 34857 (1). Psilogobius mainlandi Baldwin, 1972 (Figs. 42 and 79) Psilogobius mainlandi Baldwin, 1972, Pac. Sci. 26(1):126, fig. 4. O’ahu, Hawaiian Islands. Holotype: USNM 206174. Paratypes: BPBM 5522-25 (1, 2, 17, 11), BPBM 10862 (1), BPBM 10864 (26), BPBM 10865 (4). DIAGNOsIS.— Counts from Watson and Lachner (1985) based on 59 individuals 8.5—37.5 mm SL. Dorsal-fin elements VI-I,9-VI-I,11 (VI-I,10). Anal-fin elements I,8-I,10 (1,9). Pectoral-fin rays 15—19 (16). Longitudinal scale series highly variable, 32—78. Jaws extending posteriorly to or past posterior eye margin. Third (sometimes also second and fourth) dorsal-fin spines elongate, reach- ing past origin of second dorsal fin when depressed. Pelvic sucking disc longer than wide, reach- ing or almost reaching anal-fin origin. Pelvic-fin frenum very well-developed, extending to ends of pelvic spines (about one-third to one-half of fin length). Body covered with small scales that are partially embedded anteriorly near pectoral-fin base. No scales on head or nape. Gill openings extending forward to preopercular margin. Interorbital narrow, less than one pupil diameter. Caudal fin broad and pointed, as long as or a little longer than head length. Greatest body depth 5.0—7.1 in SL. Attains 58 mm TL. COLOR IN ALCOHOL.— Background color of head and body light cream. Body with a series of 10 midlateral brown blotches of varying sizes extending from pectoral-fin base to caudal-fin base. 544 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 Six blotches are larger with smaller blotches in between larger blotches. A sec- ond series of brown blotches along dorsal side of body at fin bases. Six of these blotches larger and extend into fin bases: two under first dorsal fin, three under sec- ond dorsal fin, and sixth on caudal pedun- cle. Five smaller brown spots spaced between larger spots, not crossing back. A brown band, about width of pupil across = = nape above pectoral-fin base. Pectoral-fin fe = Ai oe — base with a light brown blotch on upper Figure 79. Psilogobius mainlandi, 35 mm TL, and Alpheus half. Cheek with a brown bar from top of —“apax, Kane’ohe Bay, O’ahu, 2 m. upper jaw to top of preopercular margin. A second bar from ventral margin of center of eye forward onto upper jaw. Top of head with two brown blotches in advance of band on nape. Pectoral, pelvic, and anal fins with scattered melanophores. First dorsal fin with scattered melanophores on membranes, with extension of large body spots onto basal portion of fin. Dorsal spots also extending onto basal portion of second dor- sal fin. Branchiostegal membranes black in males. COLOR IN LIFE.— (from underwater photograph of living fish at Kane’ohe Bay) Background color of head and body grayish white, ventral surface lighter. Small, iridescent blue spots scattered over head and body. Five distinct, vertical, white lines on side of body: first three under first dor- sal fin; remaining two under anterior portion of second dorsal fin; all except last under pectoral fin. A series of six dark orange-brown blotches along midline of body: first between first two white lines; second between lines four and five, remaining posterior spots evenly spaced along side with last one at caudal-fin base. Corresponding dark brown blotches dorsal to those at midline, running along dorsal-fin bases. An orange-brown blotch at center of pectoral-fin base. Another blotch ante- rior to that blotch on opercle. Side of head with a dark brown line from posterior end of jaws to top of opercle. A dark brown line from upper jaw to anteroventral margin of eye. Pectoral fins clear. Pelvic fins dusky, posterior margin dark brown. First and second dorsal fins dusky with scattered, small, iridescent blue spots on basal third of fins. Second dorsal fin with an iridescent blue margin. Caudal fin dusky with a yellowish tinge, a few small iridescent, blue spots on central rays, and iri- descent blue margin. DISTRIBUTION.— Hawaiian Islands. REMARKS.— This species lives in shallow, protected areas on silty sand. It has a symbiotic relationship with the snapping shrimp, 4/pheus rapax, living in the burrow usually with a pair of shrimp. The shrimp build and maintain the burrow, and the goby, with its superior vision and lat- eralis system, serves as the sentinel. When the shrimp emerge to deposit sediment or repair the bur- row entrance, they usually make contact with the goby with one of their antennae. In return, the goby is provided with a home and perhaps food that is pushed up by the shrimp. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218156 (14), CAS 218157(3), CAS 218158 (2), 62 BPBM paratypes listed above, BPBM 22624 (4), BPBM 22644 (1), BPBM 31323 (43); Maui: BPBM 32844 (1). GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 545 Trimma milta Winterbottom, 2002 (Fig. 80) Trimma milta Winterbottom, 2002, Aqua, Jour. Ichthyology and Aquatic Biol. 5(2):45—52. Moorea, Society Islands. Holotype: ROM 59750. Paratypes: BPBM 8617 (5), BPBM 9410 (2), other paratypes at various museums. DiaGnosis.— Dorsal-fin elements VI-I.9. Anal-fin elements I, 8. Pectoral-fin rays 17-18 (usually 17). Longitudinal scale series 22—24. Predorsal scales 6-8. Ctenoid scales on body extending a little anterior to origin of first dorsal fin; nape, opercle, pectoral-fin base, chest, and mid- line of abdomen with cycloid scales, those on opercle in a single dorsal row (may be absent in juveniles). Pelvic fins united by membrane only basally, the membrane one-tenth length of fifth pelvic rays; fin longer than wide, extending at least to FIGURE 80. Trimma milta, BPBM 38706, 18.7 mm SL, anal-fin origin; fifth pelvic ray un- pfptikea, O’ahu, 23 m. branched and only about half length of longest pelvic ray; no pelvic frenum. Gill opening extending forward to pupil. Jaws extending pos- teriorly to below front of pupil. Interorbital slightly wider than one-half pupil diameter. Caudal fin slightly rounded. Greatest body depth 4.0-4.5 in SL. Largest specimen 28 mm TL. COLOR IN ALCOHOL.— Head and body cream. Scale pockets with dark brown pigment at edges so scales strongly outlined on body. Head and pectoral-fin base peppered with very small dark brown pigment spots. Pupil of eye cream, iris black. Spines and rays of dorsal and anal fins with orange-brown pigment, some similar pigment on fin membranes towards base of fins. Pectoral, pelvic, and caudal fins clear. COLOR IN LIFE.— (from photograph of live specimen from Paiptkea, O’ahu taken by R.R. Holcom-BPBM 38706 also Fig. 4 in Winterbottom, 2002) Head and body orange-red. Scale pock- ets rimmed with dark brown pigment so scales strongly outlined on body, making body darker than head. Pupil of eye black, surrounded by a yellow ring. Iris dusky yellow with neon blue pigment above and below pupil. Fin rays of both dorsal, anal, and pectoral fins orange-red, membranes clear. Caudal fin yellowish. DIsTRIBUTION.— Reported from Society Islands, Hawaiian Islands, Marshall Islands, Caroline Islands, Fiji, Solomon Islands, Great Barrier Reef, Papua New Guinea, Indonesia, Philippines, Taiwan, and western Australia. REMARKS.— Specimens have been collected from depths of 9-29 m from coral reefs and sand-rubble substratum. Our only Hawaiian specimen (BPBM 38706) was collected on O’ahu off Haleiwa (N.W. side of Paptkea reef) in 23 m of water under ledges and in small holes at a dropoff by R.R. Holcom. Holcom reported that the species was sometimes in pairs, side by side. Trimma taylori Lobel, 1979 (Fig. 81) Trimma taylori Lobel, 1979, Breviora (456):3, fig. 1. Oahu, Hawaiian Islands. Holotype: BPBM 19919. Paratypes: BPBM 19920 (1), BPBM 19921 (2), BPBM 19922 (8). 546 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 DiaGNosis.— Dorsal-fin elements VI-I,10-11 (rarely 11). Anal-fin elements 1,9-10 (rarely 9). Pectoral-fin rays 13-15. Longitudinal scale series 23—24 (usually 24). Predorsal scales 6-8. Head naked, scales on body ctenoid, becoming cycloid on pectoral-fin base, chest, and ventrally on abdomen. Second dorsal spine elongate and filamentous, sometimes the third as well. Pelvic fins united by membrane for about one-third length of fins; fin longer Figure 81. Trimma taylori, 23 mm TL, Waikiki Aquarium, than wide, usually extending to base of Yonolulu. third anal soft ray; fifth pelvic ray branched, about four-fifths length of fourth ray; no pelvic frenum. Gill opening extending forward to below pupil. Jaws extending posteriorly to below front of eye. Interorbital about as wide as pupil diameter. Caudal fin truncate. Greatest body depth 4.4—5.6 in SL. Reaches 25 mm TL. COLOR IN ALCOHOL.— Head, body, and fins all cream colored, no dark pigment in preserved material. COLOR OF FRESH SPECIMEN.— (from photographs taken at Kona and Kan’eohe Bay) Body and head translucent pale yellow or pale red. Backbone visible through body with alternating black and white sections on top; these alternating sections extending down below midline on anterior half of body. Head yellow dorsally behind eyes, overlaid by small reddish brown spots; a white bar in this area crossing between eyes. Eye with black pupil, surrounded by golden iris that has reddish blotches on it. Surface of body, head, and fins covered with tiny yellow spots. DISTRIBUTION.— Red Sea, Chagos Archipelago, and Maldive Islands to Hawaiian Islands, and Society Islands. REMARKS.— Generally found in caves or under ledges, mostly in 20-50 m, often in small aggregations above the bottom. Named from O’ahu for Leighton Taylor, former director of the Waikiki Aquarium. MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218159 (1), BPBM 31038 (9), BPBM holotype and three paratypes listed above. Trimma unisquamis (Gosline, 1959) (Figs. 47, 48, and 82) Hazeus unisquamis Gosline, 1959, Pac. Sci. 13(1):70, fig. 3. O’ahu, Hawaiian Islands. Holotype: USNM 175009. DIAGNOSIS.— Counts based on 20 individuals 13.3—17.1 mm SL. Dorsal-fin elements VI-I,6- 7 (VI-I,7). Anal-fin elements I,6-7 (1,7). Pectoral-fin rays 18-19 (19). Longitudinal scale series 24-25. A dark band at caudal-fin base. Both dorsal fins with a black border. Pelvic sucking disc longer than wide, reaching anal-fin origin; frenum well developed. Body covered with scales extending forward onto top of head to eyes. Scales on opercle but not cheek. Gill membranes not fused to isthmus, gill opening extending forward to below anterior margin of eye. Jaws extending posteriorly to mid pupil. Interorbital space very narrow, less than one pupil diameter. Caudal fin truncate. Greatest body depth 4.04.3 in SL. Attains about 26 mm TL. COLOR IN ALCOHOL.— Background color of head and body cream. Dorsal half of body, nape, and sides of head overlaid with scattered melanophores. A distinct dark brown bar at caudal-fin GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 547 base. Eye with black pupil surrounded by silver iris, edged with black. Pectoral, pelvic, and anal fins lacking pigment. Caudal fin clear with dusky area on distal one-third. First and second dorsal fins with scattered melanophores and black distal margins. COLOR OF FRESH SPECIMEN.— (from : photograph of freshly collected specimen FiGuRE 82. Trimma unisquamis, BPBM 37265, 18.5 mm SL, from Kane’ohe Bay) Background color of — Kane’ohe Bay, O’ahu, 12 m. body lemon yellow. Dorsal half of body and nape overlaid by diffuse black pigment, somewhat more concentrated on scale margins. A dis- tinct black bar at caudal-fin base. Background color of head red, overlaid by dusting of melanophores. Eye with black pupil surrounded by golden ring, iris red. Pectoral and pelvic fins yellow. First dorsal fin with reddish orange spines, membranes on basal three-quarters clear, distal one-quarter black. Some black pigment on membranes at bases of spines one through four. Second dorsal fin with same color pattern as first. Anal fin yellow with some black pigment distally. Caudal fin yellow, anterior dorsal and ventral margins red, distal half of fin dusky. DISTRIBUTION.— Hawaiian Islands, Easter Island, Society Islands, Tonga, and Guam. REMARKS.— This species typically is found in more open offshore reefs, being most abundant in the deep spur and groove habitat at Kane’ohe Bay (Greenfield 2003). MATERIAL EXAMINED.— HAWAIIAN ISLANDS: O’ahu: CAS 218160 (18), CAS 218164 (43), CAS 218161 (12), CAS 218163 (46), BPBM 22653 (1), BPBM 37265 (4); Maui: BPBM 14208 (1); Midway Atoll: BPBM 34824 (7). DISCUSSION Several small specimens (about 14 mm SL) of a goby (CAS 218162) were taken at Kahana Bay, O’ahu in 1990 while seining with the University of Hawaii ichthyology class. We could not identify the specimens and thus sent them to Helen Larson in Australia, who placed them in the genus Favonigobius. This genus is not known from Hawaii, and thus is most likely an introduction, perhaps through bilge water. Attempts to collect more specimens were not successful, and it is not known if it has become established. This species keys to 28a in our key (Priolepis), but it clearly is not within that genus. Whereas Priolepis species have horizontal rows of papilla across the cheek and above the upper jaw, they are lacking in Favonigobius. In addition, the mouth is much more oblique in Prio/epis than in Favonigobius. A drawing of one of the Favonigobius specimens is presented (Fig. 83) to aid in its identification if future specimens are collected. FIGURE 83. Favonigobius sp., CAS 218162, 13.5 mm SL, Kahana Bay, O’ahu, less than 0.5 m. ACKNOWLEDGMENTS We would like to thank the following persons for assistance in collecting gobies in Hawai’1: Erin Baumgartner, Nancy C. Burke, Geoffrey J. Cockrell, Kathleen S. Cole, David B. Eckert, 548 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 27 Ronald R. Holcom, Robert K. Johnson, Ross C. Langston, Kenneth R. Longenecker, and Jeffrey L. Mahon. Robert B. Moffitt provided us with deep-water specimens of Opua. Jennifer K. Schultz and Brian R. Bowen compared deep and shallow water Opua populations using DNA. We thank the personnel of the Division of Aquatic Resources of the Hawai’i Department of Land and Natural Resources for granting permission to collect fishes using ichthyocides. Susan G. Monden drew all of the figures, and we thank her for this. We thank Helen A. Randall for her assistance in editing. This research was supported by grant no. R/E1-5PD from Sea Grant. LITERATURE CITED AKIHITO, P., AND K. MEGURO. 1980. On the six species of the genus Bathygobius found in Japan. Japanese Journal of Ichthyology 27(3):215—236. BOEHLERT, G.W., AND B.C. Munpy. 1996. Ichthyoplankton vertical distributions near Oahu, Hawai'1, 1985-1986: data report. NOAA Technical Mem. NMFS-SWSC 235:1-148. FOwLer, H.W. 1927. Fishes of the tropical central Pacific. Bulletin Bernice P. Bishop Museum No. 38:1—32, Plea GOosLINE, W.A., AND V.E. BRockK. 1960. Handbook of Hawaiian Fishes. University Press of Hawai'l, Honolulu. GREENFIELD , D.W. 2003. A survey of the small reef fishes of Kane'ohe Bay, O'ahu, Hawaiian Islands. Pacific Science 57(1):45—76. GREENFIELD , D.W., AND J.E. RANDALL. 1999. Two new Eviota species from the Hawaiian Islands (Teleostei: Gobiidae). Copeia 1999(2):439-446. GREENFIELD , D.W., A.Y. SUZUMOTO, AND C. CHONG. 1998. Vitraria clarescens, a junior synonym of the fresh- water Hawatian goby Sicvopterus stimpsoni (Teleostei: Gobiidae). Copeia 1998(2):501—503. HayAsul, M., AND T. SHIRATORI. 2003. Gobies of Japanese Waters. TBS Buritanica, Tokyo. 223 pp. HoeseE, D.F. 1975. A revision of the gobiid fish genus Kelloggella. Records of the Australilan Museum 29 (17):473-484. Huss, C.L., AND K.F. LAGLER. 1964. Fishes of the Great Lakes Region. University. of Michigan Press, Ann Arbor, Michigan. 213 pp. JORDAN, D.S., AND B.W. EVERMANN 1905. The aquatic resources of the Hawaiian Islands. Part I. The shore fishes. Bulletin U.S. Fish Commission 23. xxvii + 574 pp. KOUMANS, F.P. 1953. Gobioidea. In: Weber and de Beaufort, Fishes of the Indo-Australian Archipelago, Vol. 10, xi + 423. E.J. Brill, Leiden. LARSON, H.K. 1985. A revision of the gobiid genus Brvaninops (Pisces), with a description of six new species. The Beagle 2(1):57-93. LARSON , H.K. 1990. A revision of the commensal gobiid fish genera Pleurosicva and Luposicya (Gobiidae), with descriptions of eight new species of P/eurosicya and discussion of related genera. The Beagle 7(1): ISS" LARSON , H.K. 2001. A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its system- atic placement. Records of the Western Australian Museum, Supplement 6:21—233. LeEvITON, A.E., R.H. Gipss, JR., E. 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. Masuba, H., K. AMAOKA, C. ARAGA, T. UYENO, AND T. YOSHINO, EDS. 1984. The Fishes of the Japanese Archipelago. Vol. 1 (text: xxii + 437 pp.) and vol. 2 (plates). Tokai University Press, Tokyo. NAKABO, T., ED. 2002. Fishes of Japan with Pictorial Keys to the Species, English edition. Vol. 2:867—1749. Tokai University Press, Tokyo. PRIVITERA, L.A. 2001. Reproductive biology of the coral-reef goby, Asterropteryx semipunctata, in Kaneohe Bay, Hawai'i. Environmental Biology of Fishes 65:287-310. PRIVITERA , L.A. 2002. Characteristics of egg and larval production in captive bluespotted gobies. Journal of Fish Biology 58:1211—1220. RANDALL , J.E. 1995. Fusigobius Whitley, a junior synonym of the gobiid fish genus Coryphopterus Gill. GREENFIELD AND RANDALL: MARINE GOBIES OF THE HAWAIIAN ISLANDS 549 Bulletin of Marine Science 56(3):795—798. RANDALL , J.E., J.L. EARLE, T. HAYES, C. PITTMAN, M. SEVERNS, AND R.J.F. SMITH. 1993. Eleven new records and validations of shore fishes from the Hawaiian Islands. Pacific Science 47(3):222-239. RANDALL . J E.. AND D.W. GREENFIELD. 2001. A preliminary review of the IndoPacific gobiid fishes of the genus Gnatholepis. Ichthyological Bulletin , J.L.B. Smith Institute of Ichthyology no. 69:117. RANDALL , J.E., AND H.A. RANDALL. 1987. Annotated checklist of the fishes of Enewetak Atoll and other Marshall Islands. Pages 289-324 in D.M. Devaney, E.S. Reese, B.L. Burch, and P. Helfrich, eds., The Natural History of Enewetak Atoll, Vol. II, Biogeography and Systematics. U.S. Department of Energy, Office of Scientific and Technical Information, Oak Ridge, Tennessee. RANDALL, J-E., J.T. WILLIAMS, D.G. SmitH, M. KULBICKI, G. Mou THAM, P. LABROOSE, M. DRONEN, E. CCLAU, AND B.S. MANN. 2004. Checklist of the shore and epipelagic fishes of Tonga. Atoll Research Bulletin no. 02: 35 pp: RusseLL, B.S. 1983. Checklist of Fishes of Great Barrier Reef Marine Park Capricornia Section. Special Publication, Great Barrier Reef Marine Park Authority no. 1. 184 pp. ScHuLTz, L.P. 1943. Fishes of the Phoenix and Samoan Islands collected in 1939 during the expedition of the U.S.S. “Bushnell.” Bulletin of the U.S. National. Museum 180. x + 316 pp. SHAFER, D.J. 1998. Early Life History Growth and Settlement Dynamics of a Tropical Reef Fish (Gobiidae: Bathygobius coalitus). Ph.D. Dissertation, University of Hawaii. 156 pp. SmitH, J.L.B. 1959. Gobioid fishes of the families Gobiidae, Periophthalmidae, Trypauchenidae, Taenioididae, and Kraemeriidae of the western Indian Ocean. Ichthyological Bulletin, Department of Ichthyology, Rhodes University no 13:185—225. THACKER, C.E., AND K.S. CoLe. 2002. Phylogeny and evolution of the gobiid genus Coryphopterus. Bulletin of Marine Science 70(3):837—850. Watson, R.E., AND E.A. LACHNER. 1985. A new species of Psilogobius from the Indo-Pacific with a redescrip- tion of Psilogobius mainlandi (Pisces: Gobiidae). Proceedings of the Biological Society of Washington 98(3):644-654. WINTERBOTTOM, R. 2002. Two new species of 7rimma (Gobiidae) from the central, western, and south Pacific. Aqua, Journal of Ichthyology and Aquatic Biology 5(2):45—S2. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 28, pp. 550—560, 5 figs. December 30, 2004 Two New Species of Dorid Nudibranchs (Mollusca, Opisthobranchia) from Bahia de Banderas and La Paz, Mexico Alicia Hermosillo! and Angel Valdés2 ! Universidad de Guadalajara, Centro Universitario de Ciencias Biologicas y Agropecuarias, Las Agujas, Zapopan, Jalisco, México; Email: alicia_hg@prodigy.net.mx; * Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, USA; Email: avaldes@nhm.org. Two new species of dorid nudibranchs from the Pacific coast of Mexico are described based on specimens collected from Bahia de Banderas and one specimen from La Paz. Trapania goddardi sp. nov. is most similar to Trapania goslineri (Millen and Bertsch, 2000) and Trapania inbiotica (Camacho and Ortea, 2000), but it is differen- tiated by the absence of yellow markings on the extra-rhinophoral and extra- branchial processes and the presence of marginal, large cusps on the radular teeth. Paradoris lopezi sp. nov. is characterized by a pale grayish-tan background color with reddish tubercles. This species is differentiated from other species of Paradoris from the Pacific Ocean by the presence of the external coloration mentioned above, the presence of two accessory glands and a single accessory sac with a spine, and the number and morphology of radular teeth and jaw elements. KEY WORDS: Goniodorididae, Discodorididae, Tropical eastern-Pacific. RESUMEN Dos especies nuevas de déridos nudibranquios son descritas para la costa Pacifica de México, en base a espécimenes recolectados en Bahia de Banderas y un espécimen en La Paz. Las especies mas similares a Trapania goddardi esp. nov. son Trapania goslineri (Millen y Bertsch, 2000) y Trapania inbiotica (Camacho y Ortea, 2000), pero se diferencian por la ausencia de manchas amarillas en los apéndices extra-bran- quiales y extra-rinoforicos, y por la presencia de cuspides marginales en los dientes de la radula. Paradoris lopezi esp. nov. se caracteriza por tener una coloracién gen- eral grisaceo pardo claro con tubérculos rojizos. Esta especie se diferencia de otras especies de Paradoris del Océano Pacifico por la coloraci6én externa antes menciona- da, la presencia de dos glandulas accesorias y un saco accesorio con una espina, asi como por el nimero y forma de los dientes radulares, y de los elementos de la armadura labial. PALABRAS CLAVE: Goniodorididae, Discodorididae, Pacifico Este tropical. There has been a significant lack of information on the diversity and opisthobranch faunal composition in Bahia de Banderas, Mexico. Only a few papers have been published on the opistho- branchs from the northern coast of the bay, situated in the state of Nayarit (Sphon and Mulliner 1972; Bertsch et al. 1973; Bertsch 1978, 1980; Bertsch and Kerstitch 1984; Ferreira and Bertsch 1975), and Isla Isabel (Ortea and Llera 1981); and there are no published studies on the opistho- branchs from the central and southern sections of the bay, in the state of Jalisco. 550 HERMOSILLO AND VALDES: NEW DORID NUDIBRANCHS FROM MEXICO Spl A recent survey by the senior author has revealed a diverse opisthobranch fauna in Bahia de Banderas, including numerous undescribed species (Hermosillo-Gonzalez 2003). Ninety-six species represented by over 20,000 specimens were identified in the field or collected, including 20 new records for this area, revealing a previously undocumented diverse fauna. In this paper we describe two new species collected during the survey. MATERIALS AND METHODS Most of the material examined was collected in several localities in Bahia de Banderas, Mexico, by the senior author. Bahia de Banderas is located on the west coast of México, in the states of Jalisco and Nayarit. Its large surface, over 1,000 km?, makes it the largest bay along the Pacific coast of mainland México. Punta Mita and Cabo Corrientes, 42 kilometers apart, delimit Bahia de Banderas to the north and south respectively. The specimens are deposited at the Department of Invertebrate Zoology and Geology of the California Academy of Sciences, San Francisco (CASIZ) and the Malacology Section of the Natural History Museum of Los Angeles County (LACM). Specimens were dissected and the inter- nal features were examined and drawn using a dissecting microscope with a camera lucida. A por- tion of the mantle was critical-point dried for the Scanning Electron Microscope (SEM). The buc- cal mass was removed and dissolved in 10% sodium hydroxide until the radula was isolated from the surrounding tissue. The radula was then rinsed in water, dried, and mounted for examination with the SEM. Features of living animals were recorded from field photographs by the senior author. SPECIES DESCRIPTIONS Family Discodorididae Bergh, 1891 Genus Paradoris Bergh, 1884 TyPE SPECIES: Paradoris granulata Bergh, 1884, by monotypy. Paradoris lopezi Hermosillo and Valdés, sp. nov (Figs. 1A-B, 2-3) MATERIAL EXAMINED.— HOLOTYPE: Punta de Pichilingue (24°21.25'N 110°17.001'W), La Paz, Baja California Sur, Mexico, 31 October 2004, 1 specimen 32 mm long, collected under coral rubble at 1 m depth (CASIZ 171661). PARATYPE (1): Majahuitas (20°29.111’N 105°35.057’W), Bahia de Banderas, Jalisco-Nayarit, Mexico, 10 March 2004, 27 mm long, collected under a rock at 12 m depth (LACM 3041). EXTERNAL MORPHOLOGY.— The maximum length of the living animal is 32 mm. The pre- served holotype measures 22 x 12 mm. The body shape is oval with an irregular mantle margin. The notum is larger than the foot, covering it completely (Figs. 1|A—B). The gill is composed of 7 tripinnate leaves and is retractable into an irregular sheath, which is slightly inclined posteriorly. The anus is located in the middle of the branchial plume. The rhinophores have 15 conspicuous lamellae and are retractable into irregular sheaths. The dorsum is porous and lacks caryophyllidia (Fig. 2E), but it is covered with irregular tubercles of different sizes. There are a few larger tuber- cles on the center of the dorsum and smaller and more abundant tubercles towards the mantle mar- gin. Minute tubercles cover the entire surface of the dorsum. Ventrally, the anterior border of the foot is notched and grooved. The oral tentacles are conical and not grooved. The color of the body is pale grayish-tan with the smaller tubercles having the same color as the rest of the body (Fig. 1A). On closer inspection, minute dark ringlets composed of black specks SS PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 28 « & rf ve ee. - ’ a a a a %. q Slee FiGuRE 1. Living animals and egg mass. (A) Paradoris lopezi sp. nov, dorsal view of the paratype (LACM 3041). Same specimen, ventral view. (C) Trapania goddardi sp. nov, holotype (LACM 3039). (D) Trapania goddardi sp. nov, e mass. (E) 7rapania goddardi sp. nov, lighter paratype (CASIZ 171662). ne = HERMOSILLO AND VALDES: NEW DORID NUDIBRANCHS FROM MEXICO SS) and white flecks can be observed. A few irregularly distributed large black spots can be observed on the dorsum. The larger tubercles in the center of the dorsum are of a dark reddish-orange. The mantle margin is encircled by a blotchy tan line. The branchial leaves are clear grayish-tan with darker tips. The rhinophores have a clear stalk with dark lamellae. The ventral side of the body is white with a few tan spots; color varies among individuals from pale to dark grayish-tan. INTERNAL ANATOMY.— The oral tube is small and short, with six strong retractor muscles attached posteriorly (Fig. 3C). The buccal bulb is about twice as large as the oral tube, rounded, with a conspicuous radular sac emerging from the posterior ventral surface and pointing upwards. There is a pair of retractor muscles attached to the sides of the buccal bulb. There are two elongate and folded salivary glands attached to the buccal mass, near the esophageal insertion. The radular formula is 23 x 17.0.17 in a 27 mm long specimen (LACM 3041). There is no rachidian tooth. The innermost lateral teeth are simple hooks with a long, curved cusp and no den- ticles (Fig. 2A). The mid-lateral teeth change abruptly in size, becoming larger at about teeth 3-4; the cusps of the mid-lateral teeth are also shorter and the base longer and more curved than those of the inner teeth (Fig. 2B). The outer teeth are smaller and have a short triangular cusp; the outer- most tooth of each row is reduced to a simple plate (Fig. 2C). The labial cuticle has a pair of jaws composed of numerous, irregular rodlets (Fig. 2D). REPRODUCTIVE SYSTEM.— The reproductive system is triaulic (Figs. 3A—B). The narrow pre- ampullary duct widens into a long and convoluted ampulla that has three folds. The ampulla nar- rows abruptly into the distal portion and connects to the female glands and the prostate. The prostate is broad, glandular and folded, with two regions clearly differentiated. The prostate nar- rows into the deferent duct, which is a long, extremely narrow and irregularly coiled tube that opens in a large common atrium with the vagina. The penis is unarmed. The vagina is a long, thin tube that tapers proximally and runs over the female gland to enter the large, spherical bursa cop- ulatrix. The seminal receptacle is rounded and connects serially to the bursa copulatrix. GEOGRAPHIC RANGE.— This species is only known from Bahia de Banderas, Jalisco-Nayarit, Mexico and La Paz, Baja California Sur, Mexcio. NATURAL History.— This species is found in the shallow subtidal (12 m depth) under rocks. Individuals produce a milky substance when disturbed. EtyMoLoGy.— The specific name is dedicated to the memory of the father of the second author, Angel Valdés Lopez, who passed away in August 2003. REMARKS.— Paradoris lopezi has all the features characteristic of the genus Paradoris (see Valdés 2002) as follows: anterior border of the foot notched and grooved; dorsum covered with simple tubercles, stiffened by integumentary spicules; labial cuticle armature with rodlets; radula composed of simple, strong, hamate teeth, which have a short cusp and lack denticles; reproduc- tive system with a flattened, granular prostate, having two well differentiated regions, and a series of accessory glands and/or accessory sacs containing copulatory spines; penis and vagina devoid of hooks. Four other species of Paradoris are known from the Pacific Ocean, but none has previously been reported from the eastern Pacific. These four species are: Paradoris leuca Miller, 1995 from New Zealand, Paradoris tsurugensis Baba, 1986, from Japan, and Paradoris araneosa Valdés, 2001 and Paradoris imperfecta Valdés, 2001, both from New Caledonian deep waters. Paradoris leuca differs from P. lopezi by lacking accessory glands and copulatory sacs armed with spines (see Miller 1995). Other differences are the shape of the jaw elements, which have a triangular cusp in P. /euca and are rounded or irregular in P. /opezi. Externally, the dorsal tubercles of P. leuca are smaller than those of P. lopezi. Paradoris tsurugensis is the most similar species to P. lopezi in external morphology and col- 554 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 28 FIGURE 2. Paradoris lopezi sp. nov, paratype (LACM 3041), Scanning Electron Micrographs of radula, jaws, and dor- sum. (A) Innermost lateral teeth, scale bar = 100 um. (B) Mid-lateral teeth, scale bar = 100 um. (C) Outermost lateral teeth, scale bar = 50 um. (D) Jaw rodlets, scale bar = 30 um. (E) Dorsum, scale bar = 500 um. oration. Both species share a grayish background color with large yellowish-orange dorsal tuber- cles. However, the tubercles of P. tswrugensis are lighter than those of P. /opezi, and in the former species they are surrounded by a darker line and often have a darker central area. The tubercles of P. lopezi are uniformly dark reddish-orange. Anatomically, P. tsurugensis is distinguishable from P. Ca oa ae ———————— HERMOSILLO AND VALDES: NEW DORID NUDIBRANCHS FROM MEXICO 555 FIGURE 3. Paradoris lopezi sp. nov, paratype (LACM 3041), anatomy. (A) Reproductive system, scale bar = 1 mm. (B) Detail of several reproductive organs, scale bar = 1 mm. (C) Lateral view of the buccal mass, scale bar = 1 mm. (D) Ventral view of the anterior end of the foot, scale bar = 1mm. Abbreviations: am, ampulla; at, genital atrium; ag, accessory gland; be, bursa copulatrix; dd, deferent duct; es, esophagus; fg, female glands; ot, oral tube; pr, prostate; sg, salivary gland; rs, radular sac; sa, accessory sac; sp, copulatory spine; sr, seminal receptacle; tn, oral tentacle; vg, vagina. lopezi in having two accessory glands and two copulatory sacs armed with spines (see Baba 1986), instead of two accessory glands and a single accessory sac. Also, P. tsurugensis has many more radular teeth than P /opezi at comparable size. The radular formula of the former is 90 x 20- 25.0.20-25 whereas it is 23 x 17.0.17 in the latter. . Paradoris araneosa is clearly distinguishable from P. /opezi by its external morphology and coloration. The dorsal tubercles or P. araneosa are smaller and more densely arranged than those of P. lopezi. Additionally, the color of the former is pale brown with a few large, dark brown spots scattered on the dorsal surface, and several, more numerous, small darker dots; the larger tubercles 556 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 28 are opaque white. This contrasts with P. /opezi, which is pale grayish-tan with the smaller tubercles having the same color as the rest of the body and the larger tubercles being dark reddish-orange. Paradoris imperfecta differs from P. lopezi in lacking accessory glands and by having two accessory sacs (see Valdés 2001), instead of one. Externally, P. imperfecta lacks the large dorsal tubercles present in P. /opezi. Also, the color of the living animals is uniformly cream with a few, irregular, pale brown spots, and numerous small dark brown dots on the dorsal surface, which is very different from the pale grayish-tan color with dark reddish-orange tubercles of P. /opezi. Family Goniodorididae H and A. Adams, 1854 Genus Trapania Pruvot-Fol, 1931 TYPE SPECIES: Trapania fusca (Lafont, 1874), by monotypy. Trapania goddardi Hermosillo and Valdés, sp. nov (Figs. 1C-E, 4-5) MATERIAL EXAMINED.— Ho_ortyPe: Islas Marietas (20°42.042'N, 105°33.878'W), Bahia de Banderas, Jalisco-Nayarit, Mexico, 17 April 2003, 7 mm long, collected on a wall at 9 m depth (LACM 3039). PARATYPES (2): Islas Marietas (20°42.042’N, 105°33.878'’W), Bahia de Banderas, Jalisco-Nayarit, Mexico, 17 April 2003, 1 specimen 6 mm long, dissected (CASIZ 171662); Bajo de la Viuda (20°43.973'N, 105°23.544'W), Bahia de Banderas, Jalisco-Nayarit, Mexico, 18 November, 2003, | specimen 3 mm long collected on a wall at 12 m of depth, dissected (LACM 3040). EXTERNAL MORPHOLOGY.— The maximum length of the living animal is 7 mm. The body is smooth, lacking tubercles, and elongate, wider in the middle region (Figs. 1C—E). The border of the mantle is reduced, undifferentiated. Each side of the body bears a digitiform extra-branchial process and a digitiform and posteriorly-curved extra-rhinophoral process. The non-retractile rhinophores are short and stout; they have no sheaths and bear 6—7 lamellae. The three branchial leaves are bipinnate and non-retractile. The anus closes the branchial circlet posteriorly. The digi- tiform oral tentacles are short, situated antero-laterally on the rounded cephalic region. The color of the body is variable from off-white with irregular small brown blotches on the middle and sides of the dorsum to tan with darker blotches. The larger blotches are almost diamond shaped; the two anterior ones join in the middle of the dorsum. The density of the brown blotches can vary between individuals (Figs. 1C—E). The ventral side of the body is off-white. The oral ten- tacles, rhinophores, branchial leaves, posterior end of the foot, and extra-branchial and extra- rhinophoral processes are the same color as the rest of the body, off-white with some brown spots. INTERNAL ANATOMY.— The radular formula is 17 x 1.0.1 in a 6-mm long specimen (LACM 3040). The radula consists of two series of 17 elongated lateral teeth (Fig. 4A). Each tooth has a long, sharp, hamate cusp displaced to the side of the radula, which generally curves inwards. Additionally, each tooth has a series of 4-6 large denticles with 1—3 smaller denticles intercalated between the larger ones. The jaws have a single series of rodlets on the masticatory border (Fig. 4B); they have a sin- gle cusp and a wide base. REPRODUCTIVE SYSTEM.— The reproductive system is triaulic (Fig. SA). The ampulla is pyri- form and connects directly to the female gland and prostate. The prostate is wide and irregular in shape, with a single loop: it connects to the deferent duct, which opens into a common atrium with the vagina. The long, narrow vagina connects directly to the rounded bursa copulatrix (serial) near the insertion point of the smaller, round, seminal receptacle. The penis bears several rows of penial hooks of different sizes and shapes (Fig. 5B). HERMOSILLO AND VALDES: NEW DORID NUDIBRANCHS FROM MEXICO o/ GEOGRAPHIC RANGE.— Trapania goddardi 1s known from the type locality in Bahia de Banderas, Jalisco-Nayarit and from Santa Cruz, Nayarit, Mexico. NaTuRAL History.— This species is found intertidally and in the upper subtidal (maximum depth 12 m) on or under rocks. The cryptic coloration, which blends with the environment, and its small size make this species difficult to find. The egg mass is a clear string of large white eggs with two whorls (Fig. 1D). The size of the egg mass is 6.3 mm x 5.4 mm with an average width of 0.76 > ma AE 2 og & 4 al e di FIGURE 4. Trapania goddardi sp. nov, holotype (LACM 3039), Scanning Electron Micrographs of radula and jaws. (A) Radular teeth, scale bar = 20 um. (B) Jaw rodlets, scale bar = 20 um. Tae : ms 558 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 28 mm. The average diameter of the eggs 1s 0.135 mm. The average number of eggs per square mm is 56. ETYMOLOGY.— The specific name is given in honor of Dr. Jeff Goddard who first found this species in 1985. REMARKS.— The generic placement of Trapania goddardi is based on the shape of the body, which is elongate, wider in the middle, with a pair of extra-branchial and a pair of extra-rhinophoral processes; the radular mor- phology is also diagnostic with a single series of elongate lateral teeth with several pointed denticles and a larger outer cusp, and lacking a rachidian tooth. There are three valid species of Trapania described for the eastern Pacific: Trapania velox (Cockerell, 1901) is known from SSS California to the Pacific side of Baja California FIGURE 5. Trapania goddardi sp. nov, holotype (LACM (Behrens 2004); Trapania goslineri Millen and 3039), anatomy. (A) Reproductive system, scale bar i 0.5 Bertsch, 2000, is known from the Gulf of mm) Feil sinc sae ar 10 um, Athens California, the outer coast of Baja California, — female glands: pr, prostate; sr, seminal receptacle: vg, vagi- and from the Pacific Coast of mainland Mexico © na. in Bahia de Banderas (Hermosillo-Gonzalez 2003); and Trapania inbiotica Camacho and Ortea, 2000 is known only from Costa Rica (Camacho and Ortea 2000). Zrapania goslineri alone has been reported for an area that overlaps the known range of T. goddardi. The three eastern Pacific species have a bright white background color and are easily distin- guishable from 7. goddardi. Both T: velox and T. goslineri have yellow markings on the oral tenta- cles, rhinophores, branchial leaves, extra-rhinophoral and extra-branchial processes, and on the dorsal portion of the posterior part of the foot (Millen and Bertsch 2000). Similar yellow markings can be observed in 7. inbiotica except for the branchial leaves and rhinophores (Camacho and Ortea 2000). Trapania goslineri has black spots irregularly distributed over the entire body; 7: velox has black lines and 7: inbiotica red spots (Millen and Bertsch 2000). On the contrary, T. god- dardi has an off-white background color, with various sizes and densities of brown spots, and it lacks black or red pigmentation. Also, the rhinophores, branchial leaves, extra-branchial and extra- rhinophoral processes, oral tentacles, and posterior end of the foot are the same color as the rest of the body, lacking yellow markings. Internally, Trapania goddardi is easily distinguishable from the other eastern Pacific species of Trapania. The radula of 7. goddardi is remarkably distinct by having an elongate and curved cusp situated on the outermost edge of the lateral teeth. Trapania inbiotica has denticles on the outer side of the cusp, a radular formula with 28 rows (Camacho and Ortea 2000), versus 17 in T. goddardi, and 21—24 inner denticles, many more than the 4-6 inner denticles of 7. goddardi. Trapania goslineri has two outer denticles, a smaller one followed by a more elongate one; its radu- lar formula is 37-41 and each tooth has 7—9 inner denticles. Trapania velox has a radular formula of 22-32 rows, with a long outer denticle and 8-11 inner denticles on each tooth (Millen and Bertsch 2000). pte me ee HERMOSILLO AND VALDES: NEW DORID NUDIBRANCHS FROM MEXICO 55) The reproductive system of 7rapania goddardi is similar to the other species, except for 7. goslineri, which has a semiserial bursa copulatrix (Millen and Bertsch 2000), whereas in 7. god- dardi it is serial. There are two Indo-West Pacific species that have a white and brown coloration with patches: Trapania brunnea Rudman, 1987 and Trapania safracornia Fahey, 2004. The reproductive systems of T. brunnea, T. safracornia and T. goddardi are similar, but their external colorations are very dif- ferent. Trapania brunnea has well-defined bright white blotches on a dark brown background (Rudman 1987). Trapania safracornia has a brown color with symmetrical white patches between the rhinophores, gill and posterior end of dorsum; the white patch on the gill area is saddle shaped (Fahey 2004) as opposed to the diamond shaped tan blotches of 7? goddardi. The rhinophores of T. brunnea are brown in color with a white tip, slender and have a long tip that protrudes from the clavus: the rhinophores of 7: safracornia are translucent with a red clavus and a white tip, where- as T. goddardi has stout rhinophores, off-white in color with brown specks. Trapania brunnea has white processes with a brown band, 7. safracornia has yellow processes while 7: goddardi has off- white processes with tan specks. The gill of 7? brunnea 1s white with brown and red specks, the gill of 7. safracornia is translucent with a pink tinge, and the gill of 7) goddardi is off-white with brown blotches. The radular teeth of 7. brunnea, T. safracornia, and T. goddardi are similar, all having a long curved cusp situated on the outer edge of the lateral teeth. However, the number of primary and secondary denticles and size and shape of the large cusp are different in each species. The outer cusp of 7: goddardi is proportionally longer compared to the width of the tooth than those of 7: safracornia and T. brunnea. Trapania safracornia has 10—14 primary denticles per tooth with a smaller denticle between each pair of larger ones (Fahey 2004). Trapania brunnea has 15 denticles but the secondary denticles are not always present between the primary ones (Rudman 1987). Trapania goddardi has only 4-6 primary denticles which are considerably smaller than the outer cusp, and secondary denticles numbering up to three. The radular formula of the three species are 21 rows of teeth for 7. safracornia; 30 for T: brunnea and 17 for T: goddardi. The jaws of these three species differ as well. Trapania brunnea has two rows of curved rodlets with sharp tips; 7. safracornia has two rows of straight sharp rodlets; and 7. goddardi has one row of rodlets with a single cusp and a wide base. ACKNOWLEDGMENTS We would like to acknowledge Roberto Chavez and Buceo Vallartech in Puerto Vallarta for funding the field work. Also our thanks to Carole and Julius Hertz for a critical review of the man- uscript and the Instituto Técnico del Mar # 6 and the Centro Universitario de la Costa (Universidad de Guadalajara) for their support. This research is part of the National Science Foundation supported project “Phylogenetic sys- tematics of dorid nudibranchs,” through the PEET grant DEB-9978155 to Terrence M. Gosliner and the junior author. The SEM work was conducted at the LACM facility supported by the National Science Foundation under the MRI grant DBI-0216506. REFERENCES Basa, K. 1986. Description of a new species of nudibranchiate Mollusca, Paradoris tsurugensis, Dorididae, from Japan. Boletim de Zoologia 10:1—8. 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. 560 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 28 BertscH, H. 1978. The Chromodoridinae nudibranchs from the Pacific coast of America. — Part II. The genus Chromodoris. The Veliger 20:307-327. Bertscu, H. 1980. A new species of Bornella from tropical West-America. Spixiana 3:33-42. BertscH, H., AND A. KerSTITCH. 1984. Distribution and radular morphology of various nudibranchs (Gastropoda: Opisthobranchia) from the Gulf of California, México. The Veliger 26:264-273. Bertscu, H., A.J. FERREIRA, W.M. FARMER, AND T.L. HAyes. 1973. The genera Chromodoris and Felimida (Nudibranchia: Gastropoda) in tropical west America: Distributional data, description of a new species, and scanning electron microscopic studies of radula. The Veliger 15:287-294. CAMACHO-GARCIA, Y., AND ORTEA, J. 2000. A new species of Trapania (Nudibranchia: Goniodorididae) from the Pacific coast of Central America. Revista de Biologia Tropical 48:317-322. CAMACHO-GARCIA, Y,. AND VALDES, A. 2003. Caryophyllidia bearing dorid nudibranchs (Mollusca, Nudi- branchia, Doridacea), from Costa Rica. Proceedings of the California Academy of Sciences 54:65—79. FAHEY, S. 2004. A new species of Zrapania (Nudibranchia: Goniodorididae) from Western Australia with comparisons to other Indo-West Pacific TZrapania. Zootaxa 514:1—12. FERREIRA, A.J., AND H. BeRTSCH. 1975. Anatomical and distributional observations of some opisthobranchs from the Panamic faunal province. The Veliger 17:323-330. HERMOSILLO-GONZALEZ, A. 2003. New distributional records of opisthobranch mollusks for Bahia de Banderas, México (Tropical Eastern Pacific). The Festivus 35:21—28. MILLEN, S.V., AND BERTSCH, H. 2000. Three new species of dorid nudibranchs from southern California, USA, and the Baja California Peninsula, México. The Veliger 43:354—366. MILLER, M.C. 1995. New species of the dorid nudibranch genus Paradoris Bergh, 1884 (Gastropoda: Opisthobranchia) from New Zealand. Journal of Natural History 29:901—908. OrTEA, J.A., AND E.M. LLERA. 1981. Un nuevo dorido (Mollusca: Nudibranchia) de la Isla Isabel, Nayarit, México. /berus 1:47-52.RUDMAN, W. 1987. The genus Trapania (Nudibranchia: Goniodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53:189-212. SCHRODL, M. 2000. Revision of dorid Nudibranchia collected during the French Cape Horn Expedition in 1882-1883, with discussion of the genus Geifodoris Bergh, 1891. The Veliger 43:197—209. VALDES, A. 2001. Deep-sea dorid nudibranchs (Mollusca, Opisthobranchia) from the tropical West Pacific, with descriptions of two new genera and nineteen new species. Malacologia 43:237-311. VALDES, A. 2002. A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia). Zoological Journal of the Linnean Society 136:353—636. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 29, pp. 561—567, 4 figs. December 30, 2004 Two New Cardinalfish Species of the Genus Apogon from Easter Island David W. Greenfield!? and John E. Randall ! Research Associate, California Academy of Sciences, San Francisco, California and Emeritus Professor, University of Hawaii; 2 Mailing address; Moss Landing Marine Laboratory 8272 Moss Landing Road, Moss Landing, CA95039; 3 Bernice P. Bishop Museum, Honolulu, Hawaii. Three species of Apogon have been reported from Easter Island, two of which are here described as new: Apogon kauteamea (formerly A. coccineus), small, translu- cent red, with VI first-dorsal spines, a free edge of skin near the anterior nasal open- ing, and two rows of scales between the lateral line and the base of the third dorsal spine; and Apogon rubrifuscus (formerly talboti), distinct in having two predorsal bones and 12 pectoral-fin rays. It is similar to A. deetsie and A. caudicinctus, but has a much larger membranous preopercular flap, lacks any dark bars on the body, and attains larger size (101.5 mm SL). Only three cardinalfish species have been recorded from Easter Island, Apogon coccineus Riippell, Apogon talboti Smith, and Apogon chalcius Fraser and Randall (Fraser and Randall 1986; DiSalvo et al. 1988). Two, however, represented misidentifications. When Gon and Randall (2003) restricted A. coccineus to the Indian Ocean, the status of that species at Easter Island needed to be reevaluated. In 1986, Randall et al. collected a large (101.5 mm SL) Apogon specimen from a cave at a depth of 39.9 m that was reported by DiSalvo et al. (1988) as A. tal/boti. A closer examination of that specimen showed that it is not 4. fa/boti. Both of these species were determined to be unde- scribed and are described here. MATERIALS AND METHODS All counts and measurements follow Hubbs and Lagler (1964) except that the last two fin rays of the dorsal and anal fins are not counted as one unless it is clear that they are joined at the base. Measurements were made to the nearest 0.1 mm using dial calipers and are expressed as percent- age of standard length (SL) or in relation to each other. Data for the holotype are presented first, followed by values for all specimens in parentheses. Ranges given in the descriptions are followed by the mean for measurements or mode for counts. Length of dorsal-fin spines was measured by placing one end of the caliper tip at the base of the spine pushed against the posterior base of ante- rior spine and the other caliper tip at the spine tip. Gill-raker counts include rudiments. Because A. rubrifuscus appears to be most similar to A. deetsie Randall (1998), and A. kautamea relates to the two complexes described by Greenfield (2001), the format for the description of each follows the format of each paper respectively to ease comparison. Institutional abbreviations are as listed in Leviton et al. (1985). 56] 562 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 29 SPECIES DESCRIPTIONS Apogon kautamea Greenfield and Randall, sp. nov. (Figs. 1-2; Tables 1-2) Apogon coccineus (non Riippell) Randall and Egafia, 1984. MATERIAL EXAMINED.— HOLotyPE: CAS 219899, 40.3 mm SL. Easter Island, west coast off Tahai, 18-20 m, 13 February 1985, rotenone, J.E. Randall, A. Cea Egafia, and L.H. Di Salvo. PARATYPES: BPBM 39186 (2, 33.3—39.2 mm), taken with holotype; CAS 219900 (3, 38.5-42.4 mm) Easter Island, off Motu Tautara, cave at 39.3 m, 10 February 1986, rotenone, J.E. Randall, A. Cea Egafia, L.H. Di Salvo, J.L. Earle, and M. Garcia; BMNH 2004.6.18.1 (1, 36.9 mm) Easter Island, west shore, wreck about 20 m offshore between Hanga Roa and Hanga Piko, 3.0—-4.6 m, 27 January 1969, Chemfish, J.E. Randall and GR. Allen; USNM 378013 (1, 37.9 mm) Easter Island, between Motu Tautara and Ara O Hara, cave with sand bottom, 16.8 m, 15 February 1985, J.E. Randall and A. Cea Egatia; FMNH 113713 (1, 39.8 mm) Easter Island, west coast, Tahai, large coral head on sand, 10.7 m, 18 February 1986, rotenone, J. E. Randall and M. Garcia ; AMS I.4330- 001 (1, 40.8 mm) same locality as FMNH 113713; NSMT-P 68772 (1, 40.5 mm) same locality as CAS 219900; SAIAB (RUSI) 74299 (1, 39.8 mm) same locality as CAS 219900; BPBM 6634 (1, 25.4 mm), Easter Island, west coast, tidepools between Hanga Roa and Hanga Piko, 0.3—0.9 m, 25 January 1969, rotenone, G.R. Allen and B.A. Baker; BPBM 6635 (1, 41.3 mm), Easter Island, off Ahu akapu, 24.4 m, 5 February 1969 (color photo), J.-E. Randall and G. A. Allen. Apogon kuatamea-non-type material: BPBM 6632 (4, 20.0-44.0 mm) Easter Island, off Ahu Akapu, rock and sand, 10.7 m, 7 February 1969, rotenone, J.E. Randall, GR. Allen and B.A. Baker. Apogon campbelli: Pitcairn, BPBM 16927(2); Oeno Atoll (Pitcairn Islands), BPBM 16499 (2), BPBM 16533 (4); Tuamotu Arch., Mangareva Group, Temoe Atoll, BPBM 13529 (2); Rangiroa Atoll, BPBM 10310 (5); Society Islands, Tahiti, BPBM 10286 (3), Moorea, BPBM 6170 (1); Rapa: BPBM 39423 (3): Cook Islands, Rarotonga, BPBM 13946 (3); Aitultaki, BPBM 5613 (5). Apogon crassiceps. Holotype, Fiji, Viti Levu, MC 28214, BPBM 5795 (25), BPBM 38996 (1), CAS 218904 (12); Tonga, BPBM 38241 (2). DIAGNosis.— A small, usually less than 40 mm SL, semitranslucent, red species lacking stripes or bars or black edges on scales on body sides, with six spines in the first and one spine and 8—9 (almost always 9) rays in the dorsal fin; two spines and eight rays in the anal fin; usually a total of 16 or fewer gillrakers (rudiments included) on the first gill arch; 13—15 (almost always 14) pec- toral-fin rays; two predorsal (supraneural) bones; a free edge of skin near the anterior nasal open- ing (Fig. 1F in Greenfield 2001); and two scales between the lateral line and the base of the third spine of the first dorsal fin. DESCRIPTION.— Dorsal-fin elements VI-I.8 (VI-I,8-9, almost always I,9); anal-fin elements II,8 (11,8); all dorsal and anal soft rays branched, the last to base; pectoral-fin rays 14 (13-15, usu- ally 14), the upper two and lower one unbranched; pelvic rays I,5; lateral line complete, the pored scales 24 (often plus one or two on caudal-fin base); predorsal scales 6; scales above lateral line to origin of first dorsal fin two; transverse scales 10 (9-10, usually 10); circumpeduncular scales 12; gill rakers 3 + 13 (3-4 + 12-14, usually 3 + 12 or 13), in holotype one developed on upper arch and 8 developed on lower arch, the remainder knobs. Measurements based on holotype and 12 paratypes. Measurement for holotype presented first, followed by range for all types and the mean in parentheses all in percentage of SL. Counts based on 13 types plus three non-type specimens. Standard length 33.3-42.4 mm. Greatest body depth 36.7 (30.6—37.9: 35.6). Head length 39.0 (37.6-43.1: 39.5). Eye diameter 15.2 (13.4—15.8: 14.8). i | GREENFIELD AND RANDALL: NEW CARDINAL FISH FROM EASTER ISLAND 563 Snout length 7.3 (5.8—8.7: 7.2). Bony interorbital width 9.3 (8.1-9.4: 8.9). Upper jaw length 20.4 (19.7-21.8: 20.7). Caudal peduncle depth 13.7 (11.5-14.2: 13.1). Caudal peduncle length 31.3 (27.9-31.8: 30.2). First dor- Sal-fm base length 15.9 (15.0-17.5: 15.9). Second dor- Sal-fin base length 15.2 (14.1-15.9: 14.7). Anal-fin base acta lool (13.2—16.3: 15.2): Pectoral-fin length 24.8 (23.7-29.7: 26.6). Pelvic-fin length 26.8 (23.6—28.8: 26.0). First dorsal-fin spine length 5.0 (4.8-7.6: 5.9). Second dorsal-fin spine length 22.9 (21.6—24.1: 22.8). Third dorsal-fin spine feaethe (7-7 (17.5=20:3: 18.9). Fourth dorsal-fin spine length 14.5 (14.2—16.6: 15.3). Fifth dorsal-fin spine length 10.2 (9.4—11.7: 10.6). Sixth dorsal-fin spine length 5.2 (4.7—7.0: 5.9). Length spine of second dorsal fin 14.1 (13.2—14.8: 14.0). Longest dorsal ray (1-3, usually 3") 21.9 (21.4—24.0: 22.5). First anal-fin spine length 4.6 (3.6—4.9: 4.2). Second anal-fin spine length 12.9 (11.8—13.2: 12.8). Longest anal ray (1-3, usually 2"4) 21.1 (18.9—23.9: 22.1). Upper caudal-fin lobe length 32.5 (32.1—35.8: 34.3). Lower caudal-fin lobe length 29.1 Q91-39.2: 32.8). Color of fresh specimen (from 35mm transparency of BPBM 6635). Head and body translu- cent, overlaid with a wash of red that is more concentrated on anterior and dorsal portions of body. Scales on nape strongly outlined in red and black pigment on predorsal scales. Scales on upper half of body outlined in red with a concentration of red along both dorsal-fin bases. Snout and side of head washed in light red, ventral surface of head white. Black area where brain is located showing through body. Back-bone showing through body as a red line, hypural plates also red. Red along anal-fin base. Dorsal and ventral edges of caudal peduncle with a concentration of red. Iris of eye gold, pupil black. Dorsal-fin spines and rays reddish, membranes clear. Rays of other fins with red- dish tinge, membranes clear. Color in alcohol: Head and body straw colored. Area over brain brown. Predorsal scales and scales along base of first dorsal fin with brown pigment. Scales on dorsal surface of caudal pedun- cle with scattered brown pigment spots. A few scattered brown pigment spots on scales at caudal- fin base. Iris of eye black, pupil dark straw. All fins clear. ETYMOLOGy.— The specific epithet is a compound adjective from the Old Rapanui language of Easter Island, combining ‘*ku’ata’ meaning transparent and mea meaning red, referring to the transparent red coloration of the species. Comparisons.— The distinctive nasal flap places 4. kautamea in the Apogon coccineus com- plex; however, it has two scales between the lateral line and the dorsal-fin base that is typical of the Apogon erythrinus complex as defined by Greenfield (2001). The nasal flap is more complex than adding an additional scale row, and thus we consider A. kautamea to belong to the Apogon FIGURE 2. Fresh color of paratype of Apogon kautamea, BPBM 6635. 564 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 29 coccineus complex. Two other mem- TABLE 1. Gill-raker counts from both sides of species of Apogon bers of the 4. coccineus complex are : Upper linb Lower limb Total rakers known ee Pacific Be 3 4 12 13° 12°15 Mice eae belli A. crassi . Both o Wabi e ae A campbeli 24 9 21 2 — 17 eo these species have a single scale row between the lateral line and the dor- sal-fin base. These two species are morphologically very similar, but differ in the number of pec- [apie 2. Pectoral-fin ray counts of toral-fin rays and gill rakers (Tables 1 and 2). Apogon camp- _ species of Apogon. belli appears to be an antitropical species, whereas A. crassi- Dee ize is ceps is more tropical. Apogon kautamea is morphologically A. crassiceps 8 18 7. 16. 3 Sees nies A. kautamea 30 2 [Se 1407 35 Seas eres 1 ein : ane A. campbelli — =3' ~ 29 == similar to both species, and has counts similar to 4. campbel- ; : : : as A. crassiceps C23 ae li, but differs from both by having the unique combination of A. kautamea — 1 28 = a nasal flap and two scales between the lateral line and dorsal- fin base. REMARKS.— With the discovery of A. kautamea, it is necessary to modify the definition of the A. coccineus complex as defined by Greenfield (2001). The single remaining character defining the A. coccineus and A. erythrinus phenetic complexes is the structure of the snout below the anterior nostril. Apogon rubrifuscus Greenfield and Randall, sp. nov. (Figs. 3, 4B) Apogon talboti (non Smith) DiSalvo et al., 1988. MATERIAL EXAMINED.— Holotype: BPBM 39346, 101.5 mm SL. Easter Island off Motu Tautara, cave at 39 m. Taken with rotenone by J.E. Randall, A. Cea Egafia, L.H. Di Salvo, J.L. Earle, and M. Garcia. 10 February 1986. Apogon caudicinctus: BPBM 35192 (4), Ogasawara Archipelago; CAS 219830 (2), Fiji. Apogon deetsie: BPBM 13983 (1), Hawaii. Apogon postero- fasciatus: CAS 219901(3), Fiji; CAS 219902 (1), Fiji. Diagnosis.—Dorsal-fin elements VI-I,9; anal-fin elements II,8; pectoral-fin rays 12; lateral- line scales 24; predorsal scales 6; gill rakers 5 + 15; body depth 2.9 in SL; upper preopercular edge serrate, the corner and lower edge with a protruding, cresentric membranous lobe with a crenulat- ed margin that almost reaches the edge of the sub and interopercles; preopercular ridge smooth; caudal fin forked with rounded lobes; color pinkish red anteriorly with posterior two thirds dusky gray with black scale edges. DESCRIPTION.— Dorsal-fin elements VI-I,9; anal-fin elements II,8; all dorsal and anal soft rays branched, the last to base; pectoral-fin rays 12, the upper two and lower one unbranched; pelvic rays 1,5; principal caudal rays 17, the upper and lower unbranched; procurrent caudal rays three upper and three lower; lateral line complete, the pored scales 24 (plus two smaller pored scales on caudal-fin base); predorsal scales 6; scales above lateral line to origin of first dorsal fin one; scales below lateral line to origin of anal fin 6; circumpedunclular scales 12; gill rakers 5 + 15, all developed: pseudobrancial filaments 14; branchiosteal rays 7; vertebrae 10 + 14; supraneur- al (predorsal) bones two. Body depth 2.9 in SL; body moderately compressed 2.2 in depth; head length 2.4 in SL;; dor- sal profile of head straight (except for rounded front of snout); snout length 5.2 in head; eye diam- eter 2.9 in head; interorbital width 5.5 in head; caudal peduncle depth 4.1 in head; caudal pedun- cle length 3.4 in SL. Mouth slightly inferior and relatively large, maxilla extending to between the GREENFIELD AND RANDALL: NEW CARDINAL FISH FROM EASTER ISLAND 565 posterior pupil margin and poste- rior orbit edge; upper jaw length 2.0 in head; mouth slightly oblique, the gape forming an angle of about 40° to horizontal axis of body; supramaxilla not present; a band of villiform teeth in jaws (maximum of about 12 rows in upper jaw and five in lower jaw); a single row of about 17 very small, conical teeth on palatine; small conical teeth form- ms a V-shaped patch on vomer, FiGuRE 3. Fresh color of holotype of Apogon rubifuscus, BPBM 39346. two rows anteriorly and two rows posteriorly (vomerine teeth larger than those in jaws). Anterior part of tongue slender and spatu- late. Largest gill raker at angle and adjacent to angle on lower limb, its length one-third eye diam- eter. A single, acute, sharp, opercular spine at level of center of eye; upper edge of preopercle ser- rate, with 17 serrae: corner and ventral part of preopercular edge with a large, cresentric, protrud- ing lobe with a crenulated margin that almost reaches edge of sub and interopercles and extends posterior of serrate edge; lateralis system of head with numerous close-set pores having slightly elevated rims; two longitudinal rows of pores from front of snout to occipit, crossed by 24 trans- verse rows; a suborbital row of pores with same transverse pore system. Anterior nostril a short membranous tube at level of bottom of pupil, about a nostril diameter from groove at base of upper lip: posterior nostril a larger triangular aperture, apex towards snout, directly in front of center of eye, the internarial distance about half pupil diameter; nasal chamber cavernous, the skin over the roof thin. Scales weakly ctenoid and thin; scales present on nape, opercle, and preopercle, those on opercle larger than scales on body; no scales on occiput, interorbital, snout, or ventrally on head; no scales on fins except for small scales on about basal fourth of caudal fin. Origin of first dorsal fin over third lateral-line scale; predorsal distance 2.4 in SL; fin spines strong and sharp; first dor- sal spine 45.6 in head; second dorsal spine 2.1 in head; third dorsal spine slightly longest, 2.1 in head: second dorsal fin separated by two median scales (posteromost indented posteriorly) from first dorsal fin, its origin above ninth lateral-line scale; spine of second dorsal fin 2.3 in head; third soft dorsal ray longest, 1.5 in head; origin of anal fin slightly posterior to second dorsal-fin origin; first anal-fin spine short, 16.9 in head; second anal spine 2.5 in head; fourth soft ray longest, 1.8 in head; caudal fin forked with broadly rounded lobes; pectoral fins long and rounded, seventh ray longest, 3.0 in SL; pelvic fins reach to second anal-fin spine, 1.6 in head. FRESH COLOR.— Background color of head and anterior part of body pinkish red; scale edges from first dorsal fin back to caudal fin black, and background color of caudal peduncle gray, mak- ing the posterior two thirds of the body much darker than the anterior third. Lateral-line scales on caudal peduncle paler than other scales, forming a distinct gray line down center of caudal pedun- cle sides. Preopercle and maxilla with silvery reflections. Pupil of eye black, iris silver. Spines and soft rays of dorsals, anal, caudal and pelvic fins pink to dark red, membranes dusky, so that the fins are dark. Pelvic-fin rays pink, membranes clear. COLOR IN ALCOHOL.— Head and body straw yellow, with scale edges outlined in dark brown, particularly in posterior two thirds of body; fins pale except for outer edges of dorsals, anal, pelvics, and caudal fins; pectoral fins clear; iris of eye black with white pupil. 566 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 29 ETYMOLOGY.— The specif- ic epithet is a compound adjec- tive from the Latin ruber (red) and fuscus (dark colored) in ref- erence to the red body color overlaid by black pigment on the posterior two thirds of the body. ComPARISONS.— Within the genus Apogon there are several species that have the same gener- al body shape, one that is typical of Apogon talboti Smith. Because of this similarity, a num- ber of specimens reside on muse- um shelves identified as 4. tal- boti, but they are not. Apogon talboti has three predorsal bones and 13 pectoral-fin rays, whereas another group of specimens we D refer to as “talboti look-alikes” FIGURE 4. Preopercle and membranous flap indicated by arrow: (A) have only two predorsal (supra- 4pegon talboti, USNM 213228, Taiwan; (B) Holotype of Apogon rubrifuscus, BPBM 39346, Easter Island; (C) Apogon deetsie, BPBM 13983, Oahu, Hawaiian Islands; (D) Apogon posterofasciatus, CAS 219901, Fiji; (E) Apogon caudicinctus, BPBM 35192, Ogasawara Archipelago. neural) bones and 12 pectoral-fin rays. Apogon — rubrifuscus belongs to this look-alike group. In addition to the three predorsal bones and 13 pectoral-fin rays, A. ta/boti has a distinctive, small, crenulated membranous flap at the lower corner and along the lower margin of the preoperculum (Fig. 4A). In their description of A. posterofasciatus, Allen and Randall (2002) state that it is most closely related to 4. deetsie Randall (1998) and A. caudicinctus Randall and Smith (1988), and that “This trio is characterized by a membranous flap at the lower corner and along the ventral margin of the preopercle.” Two of these species, 4. deetsie and A. caudicinctus, have two predorsal bones, whereas A. posteriofasciatus has three. Ofer Gon (pers. commun. Oct. 2004) believes that 4. pos- teriofasciatus is a member of the 4. fa/boti group and not associated with A. deetsie or A. cau- dicinctus. Apogon rubrifuscus also has this membranous flap, and it is larger than that found in those three species (Fig. 4B). Apogon rubrifuscus and A. deetsie have a crenulated membranous flap below the suborbital bones, whereas it is smooth in 4. posterofasciatus and A. caudicinctus (Fig. 4C—E). The preopercular margin of A. caudicinctus 1s more weakly serrate than in the other species, or smooth. The posterior end of the maxilla is concave in both A. rubrifuscus and A. deet- sie whereas it is straight in both 4. posterofasciatus and A. caudicinctus. Apogon rubrifuscus dif- fers from A. deetsie by having more gill rakers (5 + 15 versus 3-4 + 12-13) and in coloration. Apogon deetsie has two dusky bars on the body, one below the basal half of the second dorsal fin and a broader bar posteriorly on the caudal peduncle and extending slightly onto the caudal-fin base. Apogon rubrifuscus lacks those bars, but rather has the entire caudal peduncle dusky as is a portion of the body anterior to the caudal peduncle. In preserved material, 4. rwbrifuscus has dis- tinct dark scale margins whereas they are lacking in 4. deetsie. Among the species considered here, A. rubrifuscus and A. deetsie appear to be most similar. | | GREENFIELD AND RANDALL: NEW CARDINAL FISH FROM EASTER ISLAND 567 KEY TO THE EASTER ISLAND CARDINALFISHES la. Seven spines in first dorsal fin; overall life color golden yellow.......... Apogon chalcius Seesiespines im first dorsal fin: overall life:color mainly red)... 5.25255 54205--..524. 57: 2 2a. Corner and lower preopercular edge with protruding, crescentric membranous lobe with a crenulated margin that almost reaches edge of sub and interopercles; posterior two thirds of body dusky gray with black scale edges; five rakers on upper gill arch, all developed 8 Reese Ai Agee soe ae a Sol ar gine Sun Seu hem eeu aaa eae Apogon rubrifuscus 2b. Corner and lower preopercular edge without large membranous lobe; body light without dark scale edges; three to four, usually 3 rakers on upper gill arch, only one developed <= 2c 6 © oy CRY cee eee ER cela mea ih Lar betty ob A oes tla te Me A Apogon kautamea ACKNOWLEDGMENTS The second author thanks foremost the National Geographic Society that funded his 1969 and 1985 fish visits to Easter island, and the Engelhard Foundation for its support in 1986. He is also grateful to Louis H. DiSalvo, Alfredo Cea Egania, John L. Earle, and Henri and Michel Garcia for assistance in the field. LITERATURE CITED ALLEN, G.R. AND J.E. RANDALL. 2002. Four new Indo-Pacific species of cardinalfishes (Apogonidae). Aqua, Journal of Ichthyology and Aquatic Biology 4(3):115-126. DiSAtvo, L.H., J.E. RANDALL, AND A. CEA. 1988. Ecological reconnaissance of the Easter Island sublittoral marine environment. National Geographic Research 4(4):45 1-473. FRASER, T.H., AND J.E. RANDALL. 1986. A new species of the cardinalfish genus Apogon from Easter Island. Copeia 1986(3):641—645. Gon, O., AND J.E. RANDALL. 2003. A review of the cardinalfishes (Perciformes: Apogonidae) of the Red Sea. Smithiana Bulletin \:1-46. GREENFIELD, D.W. 2001. Revision of the Apogon erythrinus complex (Teleoste1: Apogonidae). Copeia 2001 (2): 459-472. Huss, C.L., AND K.F. LAGLER. 1964. Fishes of the Great Lakes Region. University of Michigan Press, Ann Arbor. 213 pp. Leviton, A.E., R.H. Gipss Jr., E. HEAL, AND E.E. DAWSON. 1985. Standards in herpetology and ichthyology. Part 1, Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985(3):802-832. RANDALL, J.E. 1998. Review of the cardinalfishes (Apogonidae) of the Hawaiian Islands, with descriptions of two new species. Agua, Journal of Ichthyology and Aquatic Biology 3(1):25-38. RANDALL, J.E. AND A. CEA EGANA. 1984. Native names of Easter Island fishes, with comments on the origin of the Rapanui people. Occasional Papers of the Bernice P. Bishop Museum 25(12):1—16. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 30, pp. 568-572, 6 figs. December 30, 2004 Allenbatrachus meridionalis, a New Toadfish (Batrachoididae) from Madagascar and Reunion David W. Greenfield! and Wm. Leo Smith2-4 ! California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103; 2 Department of Ichthyology, American Museum of Natural History, New York, NY 10024 The third species in the genus Allenbatrachus, A. meridionalis, is described from Madagascar and Reunion, the southernmost occurrence of the genus. It differs from the other two described species, A. grunniens and A. reticulatus, by having more anal and dorsal-fin rays, differently shaped orbital cirri and maxillary barbels, and being darker in coloration. The genus A/lenbatrachus Greenfield previously included two species, 4. grunniens (Linnaeus), known from India eastward to the Philippines, and A. reticulatus (Steindachner) known from Thailand south to Sumatra (Greenfield 1997, 1999). While examining specimens at the Muséum national d’histoire naturelle, Paris, the first author found several specimens from Mada- gascar and Reunion that represented an undescribed species of Al/enbatrachus. Subsequently, the second author with colleagues from the American Museum of Natural History and Wildlife Conservation Society (WCS) collected three additional specimens of the same undescribed Allenbatrachus species from the Makira region of Madagascar. These specimens included a single specimen collected from a small freshwater stream just north of Ambodivoanongy, Sanavilory, and two specimens purchased from local fishers in the nearby Maroansetra market. The collection of these specimens confirms the previous report of Kiener (1963) who illustrated a toadfish specimen (identified in that publication as Batrachoides grunniens, now A. grunniens) from freshwater streams near the town of Maroansetra. This undescribed species belongs to the genus A/lenbatrachus because it has the following characters: (1) dorsocranium has a foramen on each side of the head behind the eyes, bordering the sphenotic and frontal bones (MNHN A3777 cleared and stained) (this character is shared only with species in the genera Batrachoemoeus and Halophryne); (2) it lacks the pore (foramen) present in the upper part of the pectoral-fin axil that is present in Batrachomoeus species; (3) the lower jaw is terminal, projecting beyond the upper jaw, whereas the upper and lower jaws are about equally terminal in Halophryne species; and (4) the interorbital width is equal to or less than the eye diam- eter and the gill slit is restricted to the upper one-half to two thirds of the pectoral-fin base in Halophrvne species, whereas the interorbital width is greater than the eye diameter and the gill slit extends from the upper three-fourth to four-fifths of the pectoral-fin base in Allenbatrachus, the condition in the new species. 3 Research Associate, Department of Ichthyology, California Academy of Sciences and Emeritus Professor, University of Hawaii. Mailing address: Moss Landing Marine Laboratory, 8272 Moss Landing Road, Moss Landing, CA95039. 4 Columbia University, E3B and CERC, New York, NY. 568 GREENFIELD AND SMITH: NEW TOADFISH FROM MADAGASCAR 569 MATERIALS AND METHODS All counts and measurements follow Hubbs and Lagler (1964) except that the last two fin rays are not counted as one unless it is clear that they are joined at the base. Measurements were made to the nearest 0.1 mm using dial calipers. All measurements are expressed as percentage of stan- dard length (SL). Some counts were made with radiographs. Counts and measurements for the holotype are given first, followed by the range and mean or mode in parentheses for all specimens when variable. Institutional abbreviations are listed in Leviton et al. (1985). SPECIES DESCRIPTION Allenbatrachus meridionalis Greenfield and Smith, sp. nov. (Figs. 1-6) MATERIAL EXAMINIED.— HOLOTYPE: AMNH 233686, 147.2 mm SL, Malagasy Republic, Madagascar, Market at Maroantsetra, October 2003, P. V. Loiselle. PARATYPES: CAS 220508, 169.0 mm SL, same locality as holotype. AMNH 234024, 90.7 mm SL, Malagasy Republic, Madagascar, small tributary of Antainambalana River, just north of Ambodivoanongy, Sanavilory, November 8, 2003, J.S. Sparks, W.L. Smith, K.L. Tang, and party. MNHN 1992-0670, 189.0 mm SL, Madagascar, Toliara, Tuléar, Andavadanova, 23°20'00"S; 43°30'0"E, Mauge. MNHN 1962-0197, 118.7 mm, 144.0 mm, 165.5 mm SL, Madagascar, Maroantsetra, Antainambalana estuary (listed as Antenambalana), Kiener. USNM 379463, 122.2 mm SL, same locality as MNHN 1962-0197. ADDITIONAL MATERIAL EXAMINED.— Allenbatrachus meridionalis (non-types): MNHN A- 3777, 109.6 mm SL, Reunion, 21°7'0"S; 55°35'0”E, cleared and stained. MNHN 1966-0912, about 180 mm and 194 mm SL, Madagascar, 20°0'0"S; 42°30'0"E, specimens bent, Kiener, Therezien. Allenbatrachus grunniens:Thailand: CAS 75217(2), CAS 75218(1). Vietnam: USNM 047986(1). Malaysia: AMS 1.27634008(1). Borneo: CAS-SU 32944(2), CAS-SU 27732(4). Philippines: ANSP 48783(1), ANSP 77373(1), CAS-SU 26909 (1 cleared and stained), CAS-SU 38261(1), CAS-SU 38262(1), USNM 148493(1). India: CAS-SU 41321(1), AMS B.8319(1), ZSI 2099 (1 cleared and stained). Allenbatrachus reticulatus: Singapore: CAS-SU 30658(20, 1 cleared and stained), CAS-SU 33701(1), CAS-SU 35153(2), CAS 82188(1 neotype). Thailand: CAS 66821(1), CAS 75216(1), CAS 17652(1), CAS 88690(7), AMS 1I.21036003(2). Burma: ZSI 10741(1), ZSI 10957(1). Sumatra: USNM 333283(5). DiaGnNosis.— A generally dark colored species of Allenbatrachus with 19 anal-fin rays, 22 dorsal-fin rays, usually a single, simple, pointed orbital cirrus, a simple, pointed barbel at the end of the maxilla, and pointed teeth. DESCRIPTION.— Dorsal-fin elements I]-22. Anal-fin rays 19. Pectoral-fin rays 21 (19-22, usu- ally 19). Vertebrae 9 precaudal plus 19 (18-19) caudal (including terminal centrum). Upper later- al-line pores 27 (22-30, usually 27), lower lateral-line pores 18 (17-23), middle lateral-line pores, if present, indistinct. Head length 33.6 (33.1-39.9; 36.3). Head width 25.2 (25.2-31.6; 27.7). Head depth 16.6 (16.6-25.1; 20.0). Bony interorbital width 5.8 (4.8-7.2; 6.0). Fleshy interorbital width 9.8 (6.7-10.8; 10.1). Orbit diameter 5.4 (4.5-9.8; 6.2). Snout length 8.0 (5.9-9.2; 7.5). Upper jaw length 17.0 (17.0-18.6; 17.9). Width of mouth at rictus 17.9 (17.9-23.1; 20.2). First predorsal-fin distance 38.8 (38.7-41.7; 40.0). Second predorsal-fin distance 50.1 (50.1-54.7; 52.0). Preanal-fin distance 61.6 (58.5-64.2; 61.6). Prepelvic-fin distance 24.0 (21.2-26.8; 23.4). Greatest body depth 18.1 (18.1-28.4; 22.0). Caudal-peduncle depth 7.8 (6.5-8.8; 7.9). Length of second dorsal-fin base 43.5 (43.5-56.0; 49.4). Length of anal-fin base 33.1 (33.1-41.6; 37.7). Caudal-fin length 22.2 (20.4- 24.0; 22.4). Pectoral-fin length 23.9 (22.9-28.7; 25.9). Pelvic-fin length 21.7 (19.7-23.8; 22.3). 570 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 30 ~ FiGuRE |. Holotype of Allenbatrachus meridionalis, AMNH 233686. FIGURE 2. Paratype of A/lenbatrachus meridionalis, AMNH 234024, from fresh water. ie : er Pe222>> Be. Lipp a ee oe CH oP med eS mie MLZ» ee BRS Se : Bjapecrhie sae Oe Sey : ae NN Shy URNS SS FIGURE 3. Drawing of paratype of A//enbatrachus meridionalis, MNHN 1992-0670. Distance between pelvic-fin bases 6.1 (6.1-10.8; 8.6). A sin- gle, simple orbital cirrus, tip pointed (fig. 4e) (one specimen has three simple cirri on one side and two on the other, the central always the largest, and a second specimen had three on one side and one on the other with the central cirrus the largest; usually a single, simple orbital cirrus). Largest bar- bel at end of maxilla simple and pointed (Fig. 5c). All teeth in jaws pointed; vomer and palatines with a single row; den- tary with single row on sides and triple row at symphysis; FiGuRE 4. Orbital cirri. A/lenbatrachus reticulatus: (a) anterior; (b) posterior. premaxilla with a double row on side and triple row at front. Al/enbatrachus grunniens: (c) anterior; (d) Color in alcohol of holotype: Overall coloration of pre- posterior. Allenbatrachus meridionalis: (€) : . single cirrus. served specimens is dark, almost black, much darker than “78° SU™ other species in the genus. Background color dark brown, with black color pattern. Sides of body GREENFIELD AND SMITH: NEW TOADFISH FROM MADAGASCAR wal with irregular black blotches forming a band i = m : = eG Se “2 centered on the middle of the sides, the areas Sy Cy : % above and below this lighter. Ventral surface of < Dp head and body light brown, a distinct white area S 7 around the anus. Top and sides of head dark 2 eat c brown, overlaid with a heavy, irregular mottling Figure 5. Largest barbel at end of maxilla: (a) of black. Pupil of eye white, iris black. Inside of Allenbatrachus grunniens; (b) Allenbatrachus reticulatus; mouth white. Pectoral-fin base dark brown with (©) 4“enPatrachus meridionalis. two black blotches, fin rays and membranes white, crossed by four distinct black bars, the distal margin of the fin white. Pelvic fin light brown, mottled with black blotches. Caudal fin light tan, covered with large, round black spots that form bars across fin, distal margin light tan. Second dorsal fin dark brown, with two black bands running the length of the fin, one along the base and the other along the distal margin, with a few brown tips on some of the mem- branes. Basal half of anal fin dark brown with black mottling, outer half a black band, white tips on the fin membranes. Color of fresh, small, freshwater specimen (AMNH 234024): Background color light tan with straw tinge, head and body overlaid with black markings. Head with black area in front of and under anterior half of eye, a black blotch behind eye with a line from it extending back across side of head. Another black blotch above eye and across interorbital area. Lower side of cheek, lower jaw, and underside of head cream. Area over opercle with black blotches running up across nape. Sides of body with irregular black blotch- es forming a band centered on the middle of the sides, wider on posterior quarter of body. Area below band light tan, with small, black spots. Pectoral fin light tan with black blotches forming indistinct bands, distal margin of fin white. Anal fin cream with some black blotches distally on posterior half. Base of caudal fin cream with black blotches on distal half. Distinct black blotch around base of first dorsal fin. Three black blotches along second dorsal-fin base. Second dorsal fin light tan with black mottling. ETyMOLOGy.— The specific epithet is the Latin adjective meridionalis, meaning southern, referring to the fact that this is the southernmost known species of the genus. COMPARISONS.— Allenbatrachus meridionalis differs from both other species of Allenbatrachus, A. grunniens and A. reticulatus, by having both orbital cirri and maxillary barbels that are simple and pointed rather than with multiple tips that are either rounded or pointed. It also differs from both by having more anal-fin rays, 19 versus 16-17. Allenbatrachus meridionalis has 22 dorsal-fin rays, whereas the other two species usually have 19. The new species is most similar to A. grunniens in having sharp, pointed teeth, rather than the rounded teeth found in A. reticula- tus. Discussion.— The description of this species from Madagascar and Reunion considerably extends the range of the genus Allenbatrachus, previously known from the Ganges River area in India eastward to the Philippines. The capture of a specimen of A. meridionalis from a small fresh- Orbit Diameter as Percentage of Standard Length Standard Length (mm) FIGURE 6. Orbit diameter as percent standard length ver- sus standard length for A//enbatrachus meridionalis type specimens. Sf PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 30 water stream as well as from saltwater habitats, concurs with the capture of the other two species in the genus in both fresh and salt water. The single, small (90.7 mm SL) freshwater specimen has a much larger eye than all other specimens, and at first we considered the possibility that it might be a different species from those taken in salt water; however, when orbit diameter as a percentage of SL was plotted against SL, it became evident that extreme allometric growth in eye size occurs in this species (Fig. 6). It also is lighter in color, with fin markings not as distinct as in larger spec- imens from saltwater habitats. ACKNOWLEDGMENTS We would like to thank Guy Duhamel and Patrice Pruvost for their hospitality while visiting the MNHN and for lending us specimens. We also thank Scott A. Schaefer, Melanie L. J. Stiassny, John S. Sparks (AMNH), and Paul Loiselle (WCS) for making specimens available. Collecting efforts in Madagascar were facilitated by B. Andriamihaja and the MICET (Institute for the Conservation of Tropical Environments, Madagascar) staff, D. Meyers and the WCS staff, Critical Ecosystem Partnership Fund, Conservation International’s Global Conservation Fund, the Government of Madagascar, the Ministry of Environment, Water and Forests of Madagascar, and Antongil Conservation. The AMNH and CAS specimens were collected under permits obtained from the Direction des Eaux et Foréts and the Association National pour la Gestion des Aires Protégées (ANGAP), Antananarivo, Madagascar, in accordance with IACUC guidelines. Funding for W. L. S. and for the AMNH fieldwork in Madagascar was provided by AMNH, Columbia University, WCS, an AMNH Lerner-Gray grant for marine research, and National Science Foundation DEB-0405246. LITERATURE CITED GREENFIELD, D.W. 1997. Allenbatrachus, a new genus of Indo-Pacific toadfish (Batrachoididae). Pacific Science 51(3):306-313. GREENFIELD, D.W. 1999. Batrachoididae. Pages 1999-2003 in K.E. Carpenter and V.H. Niem, eds., F4O species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes part I (Elopidae to Linophrynidae). FAO, Rome. Huss, C.L., AND K.F. LAGLER. 1964. Fishes of the Great Lakes Region. University of Michigan Press, Ann Arbor. 213 pp. KIENER, A. 1963. Poissons, Péche et Pisciculture a Madagascar. Centre Technique Forestier Tropical. Nogen dur Marne (24):1-244. LEVITON, A.E., R.H. Gipss, JR., E. HEAL, AND C.E. DAwson. 1985. Standards in herpetology and ichthyology. Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 31, pp. 573-587, 6 figs. December 30, 2004 The California Academy of Sciences Gulf of Guinea Expedition (2001) II. Additions and Corrections to Our Knowledge of the Endemic Amphibians of Sao Tomé and Principe R. C. Drewes and R. E. Stoelting Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, USA; Email: rdrewes@calacademy.org We present observations on the Sao Tomé Island endemic Hyperolius thomensis which indicate that it is one of the few African treefrogs known to utilize phytotel- mata (tree holes) for breeding. Our field work and observations strongly suggest that this island giant is restricted to primary forest, remnants of which are usually at higher elevations or inaccessible areas of S40 Tomé Island. Our locality data and field observations in a number of circumstances are not congruent with those of Loumont (1992). An examination of the data associated with her collections housed in the Natural History Museum of Geneva reveals that her data are not specific with regard to individual specimens and dates and, as a result, the status and distribution of the amphibian species on both islands may have been misinterpreted. We note that females of the endemic ranine ranid frog, Ptychadena newtonii, attain snout-vent lengths greater than other members of the genus, and that this species should there- fore be considered an island giant. We describe and illustrate for the first time the males of Africa’s largest treefrog, the Principe Island endemic Leptopelis palmatus, provide figures illustrating the range of male and female color pattern polymor- phism and comment on adult size dimorphism and size at metamorphosis. The islands of the Gulf of Guinea off the west coast of Africa are a unique chain that includes one geologically recent continental island, Bioko (formerly Fernando Poo), and three oceanic islands, Principe, Sao Tomé and Annobon (Pagalu). Isolated from the African mainland since their orogeny, the three oceanic islands are noted for their highly endemic but poorly known flora and fauna. In 2001, the California Academy of Sciences conducted a multidisciplinary, two-month research expedition to the two geologically oldest islands, Principe and SAo Tomé, which resulted in the collection of voucher specimens and tissues now available for genetic analysis. In an earlier study, Drewes and Wilkinson (2004) presented a more detailed introduction to the geologic histo- ry of the two islands, and a popular account of the expeditions’ scope and goals is available (Drewes, 2002). Loumont (1992) attempted the first comprehensive analysis of the entire endemic amphibian fauna of both islands. She recognized three hyperoliid species Nesionixalus thomensis endemic to S40 Tomé, Leptopelis palmatus found only on Principe, and Nesionixalus molleri common on both islands, two ranids, Ptychadena newtoni (Sao Tomé) and Phrynobatrachus dispar (common on both), and two species of dermophine caecilians, Schistometopum thomense and S. ephele of Sao Tomé. A subsequent morphological study (Nussbaum and Pfrender 1998) indicated that S. ephele Se 574 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 31 is conspecific with S. thomense, and we confirmed this decision based on mtDNA analysis (Stoelting, in prep.). In a recent systematic work, Drewes and Wilkinson (2004) returned N. thome- nsis and N. molleri to the genus Hyperolius Rapp 1842 from which these species were twice removed by Perret (1976, 1988; Loumont 1992). Although the species that comprise the amphibian fauna of Sao Tomé and Principe have been known to science since the latter half of the 19' Century, these islands have rarely been visited by modern biologists and then usually only briefly. Thus, much of the natural history of these island species including basic data such as the extent of intra-island distributions has remained undocu- mented. We report data suggesting that the endemic treefrog Hyperolius thomensis of Sao Tomé 1s a primary forest, high-elevation, tree-hole breeding frog. We describe the male of the Principe endemic Leptopelis palmatus with comments on adult sexual size dimorphism and metamorph- adult size disparity. We attempt to refine our knowledge of the distributional limits of the amphib- ian species of SAo Tomé and Principe. MATERIALS AND METHODS The CAS Gulf of Guinea expedition afforded the opportunity to examine localities on both islands in considerable detail and from the perspectives of five academic disciplines: herpetology, ichthyology, mammalogy, entomology and invertebrate zoology. Many sites on both islands were visited and sampled on multiple occasions. Three of the 11 CAS staff members involved in all or part of the 10-week expedition were herpetologists who spent roughly 110 person-days in the field. The availability of transportation and accessibility of many sites, especially on SAo Tomé, allowed us to conduct a reasonable amount of sampling and observation at night (until 2300 hr or later). All CAS specimens were hand-collected, euthanized, fixed in 10% buffered formalin and later trans- ferred to 70% ethanol. Anuran larvae were retained in formalin. Tissues were taken from selected individuals of all taxa and preserved in 95% ethanol. Latitude, longitude and elevation were record- ed with a Garmin 12 Global Positioning System receiver (datum WGS 84). All specimens and tis- sues are housed in the Department of Herpetology, California Academy of Sciences; associated data can be accessed on-line at www.calacademy.org/research/herpetology/catalog. Institutional abbreviations follow Leviton et al. (1985). RESULTS AND DISCUSSION I. Reproduction in Hyperolius thomensis In his initial description of the genus Nesionixalus, Perret (1976) found 25 large (2.0—-2.5 mm) presumably ovarian eggs in each of two of the three females in the 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 reproductive mode — “wn cycle biologique particulier!’ Our field data suggest that Perret (1988) was correct. While our observations confirm that H. molleri deposits its egg masses above water in typical Hyperolius mode (Schietz 1999; Fahr 1993; Figure | herein;), there is strong inference that H. thomensis is one of the few African tree-hole breeding frogs. To date, two bufonids, one microhylid, one petropedetine ranid, and five hyperoliids — Acanthixalus spinosus, A. sonjae, Callixalus pictus (possibly), Hyperolius mosaicus and H. acutirostris — are known to breed in phytotelmata (Rédel et al., in press). In 10 weeks on Sao Tomé, we encountered Hyperolius thomensis at only two localities. A sin- gle, rather small male (CAS 219059; 31.0 mm.) was collected on 14 April 2001 among a group of H. molleri males, which were calling from tall grasses along a trail-side ditch on the footpath to Lagoa Amelia above Bom Successo (elevation: 1170+ m.). Our series of 13 adults (CAS i | DREWES AND STOELTING: GULF OF GUINEA AMPHIBIANS SIS 218925—937) was collected on 9 April 2001 within a hole in a buttressed, unidentified tree in pri- mary forest at Macambrara, just west of the site of a large transmission tower at an elevation of 1280 m. This locality is a well-forested ridge top with no open water available. The tree hole was located where two buttresses were joined about 1.7—1.8 m. above the ground. The dimensions of the opening were roughly 7-12 cm wide by 25 cm high. The internal diameter of the hole was about 20 cm, and about 20 cm of stagnant water was in the basin. Two smaller holes were located on the same tree at least 1.7 m above the ground, one of which contained water and egg masses (but no adults): the other was dry. When initially discovered on 9 April 2001, the first referenced tree hole yielded 11 adult H. thomensis — two adult females and nine males, 38 larvae and at least three egg masses adhering to the interior walls of the hole a few centimeters above the water surface. A subsequent visit to the tree on 16 April 2001 resulted in the collection of two more adult males. Of the two females col- lected. CAS 218925 has ovarian eggs whereas CAS 218934 has none. We have not attempted to dismantle the egg masses, but each contains from 20-40 large, pig- mented eggs. In one mass, the eggs are at the 32-cell stage of development (Gosner 1960, stage 7), and in another the eggs are at yolk plug stage (Gosner stage 11). The 38 larvae range from Gosner stages 25—41 with the majority around 25-37. A detailed description of the larvae of Hyperolius thomensis, along with H. molleri, Phrynobatrachus dispar and Ptychadena newtoni is in preparation (Drewes and Altig); the H. thomensis tadpole has a max- imum tooth row formula of 2(2)/3, but a number of individuals lack the second upper row (LTRF 1/3; nomenclature of Altig 1970). Ecomorphogically the larvae most closely fit the exotrophic, lentic, arboreal type 5 of McDiarmid and Altig (1999). The large number of adults found together along with the various developmental stages of the tadpoles and egg masses suggests that tree-holes are used by multiple individuals at different times; phytotelmata probably represent primary reproductive sites at higher elevations where free water is absent. We assume based on buttress structure and trunk morphologies that there were addition- al holes that were too high to be surveyed in this tree and in other nearby trees of similar size. RES returned to the radio tower site with other CAS team members on successive nights from 5—8 May 2001. On 5 May, beginning around 1930 hr, she heard an estimated nine different males calling from about 4.5 m above ground at locations approximately 30—40 m apart. Three locations within the forest where frogs were heard were marked. On 6 May, she and three others returned to the marked spots at about 2130 hr and noted a distinct reduction in calling from the night before. On this second evening, the moon was full, and there was a lack of mist. At one marked spot she heard a male calling from the same perch as on the night before and noted that a few other males seemed to be calling from sites somewhat lower within the canopy than on the previous occasion. Individuals were extremely wary: if disturbed they ceased calling, and then seemed to resume from another spot. On the night of 7 May, numerous frogs were again heard from “all over the radio tower primary forest area [= Macambrara],” but no calls were heard along the road to the lower plantation areas. Finally, on the night of 8 May at around 1900 hr near the original tree from which the H. thomensis series was initially collected on 9 April, RES heard, recorded and saw a single individual about 7 m above ground at the juncture of a bifurcating tree trunk. The recording is con- sistent with the description and sonogram given in Loumont (1992). Upon descending to Bom Successo she noted that no calls were heard in cultivated areas, but a few were heard in a remnant of primary forest between the plantation and Bom Successo. Our extensive field work on Sao Tomé leads us to the conclusion that H. thomensis is a high- elevation, primary forest, tree-hole breeding species, which is not consistent with collecting local- ities given in Loumont (1992). 576 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 31 II. Localities and Distributions We were puzzled by our inability to confirm some of the H. thomensis localities cited in Loumont’s paper and by the absence of supporting data that one would normally expect to see (e.g., specific collecting dates, voucher numbers associated with sonograms, and behavioral observa- tions). Published sonograms of male advertisement calls by H. thomensis that were supposedly col- lected at Java and Colonia Acoreana indicate that they were recorded from captive males calling in plastic bags (as were the recordings of Phrynobatrachus dispar and H. molleri [Loumont, p. 40]). However, there are no individual voucher numbers provided with these sonograms, and it is not clear from which specimens these calls were recorded, nor where they were made — on-site, in the town of Sdo Tomé or in Geneva. Although the sonogram of H. thomensis 1s accurate based on our recordings and observations noted above, we assume that Loumont failed to note calling sites, times and specific dates because she did not collect nor observe these specimens in the field. Additional inconsistencies lead us to believe that much of her hyperoliid material was brought to her by local collectors. Loumont indicated that specimens of H. thomensis were found in Sao Tomé town (coastal), Colonia Acoreana (coastal), Java (600 m), and Monte Café (680 m) under banana leaves, and in mossy hollows in old walls, close to small reservoirs of water; these represent the entirety of the microhabitat information provided by Loumont (op. cit., p. 51) for this species. We sampled the abandoned plantation at Java on several occasions but found no evidence of H. thomensis, although H. molleri adults, tadpoles and egg masses were readily found in such microhabitats. No suitable sites were found in the town of Sao Tomé in spite of many days of searching there and at many other coastal and lowland localities. We did not survey Monte Café (500+ m below the CAS Macambrara site), but in October-November 2003, a private collector camped for three days at Pousada Boa Vista while actively looking for H. thomensis and other endemics. Pousada Boa Vista is above Monte Café on the same road, ca. 800 m. Five H. thomensis were brought to him by a local boy who indicated they came from a site about an hour away and stated that they were “very hard to find.” (Daniel Hofer, pers. commun.). The CAS expedition did not survey the coastal local- ity of Colonia Agoriana per se but worked many similar sites along the east and southeast coasts (Micondo, vicinity of Rio Angobo, Sao Joao, vicinity of Dona Augusta). The Natural History Museum in Geneva kindly provided by electronic mail all of the field data associated with Loumont’s collections of H. (as Nesionixalus) thomensis and also of Leptopelis palmatus (Principe Id.). Loumont’s collection dates with respect to nearly all of these specimens are recorded inclusively rather than as individual dates. For instance, Loumont’s data for 21 “Nesionixalus” thomensis contain but one specific date: 10.07.1988 for two specimens from Java. Two specimens from Monte Café bear the date 7.1988. Four more from Java are recorded as 1.1990, and the remaining specimens from Sao Tomé town, Gulf of Guinea, Monte Café, Agua Cascada, Colonia Acoreana and Java all bear the collection date 1.12.1989—31.12.1990, a 13- month period. The Agua Cascada specimen (MHNG 2492.084) and the two from “Golfe de Guinée (MHNG 2492.081—082) were not included in her publication, but the MHNG data sheets indicate the same collecting date as the others: 1.12.1989-31.12.1990. Similarly, all Leptopelis palmatus taken from three separate localities on Principe Island are cataloged as having been collected from 1.12.1989—31.1.1990. In her materials and methods section, Loumont (1992) stated that field work took place in early July 1988 and in January 1990. If this is accurate then any and all specimens taken during December of 1989 must have been collected by someone else prior to her arrival. We conclude that Loumont did not publish nor archive individual collection dates or more spe- cific locality site data because she did not know them, and that she relied on local collectors for DREWES AND STOELTING: GULF OF GUINEA AMPHIBIANS 7 collection of the specimens and for their general locality data. Data such as “Golfe du Guinée” is consistent with the broad and technically correct information a researcher might record for a valu- able specimen if more precise locality data was unknown or had been lost. Similarly suspect is the collection locality of Colonia Agoreana. Colonia Agoreana is a lowland site on the southeastern coast at about 200 m and surrounded by old plantations and secondary growth; however, the topog- raphy inland rises to above 600 m within 6 km of the town where primary growth likely still per- sists. We think it is reasonable that a local collector bringing specimens to SAo Tomé city would probably indicate “Colonia Acoreana” as the point of capture for animals that had perhaps been col- lected at an un-named locality high in the mountains above that town. Loumont indicated that most of her localities were within two ecological zones on both islands: a lower altitude zone from 0-500 m, where Ptychadena, Phrynobatrachus and Leptopelis are found, and a middle montane zone (500-1000 m) where “espéces parasylvicoles” (species of disturbed-forest/secondary forest?) such as Nesionixalus (= H. thomenis and H. molleri) occur but they may also descend to the lower plains. She mentions three other zones from which she had no specimens: the northeast savannah on Sao Tomé (grassland with baobab trees [Adansonia digita- ta] in the rain shadow of Pico do Sao Tomé, the primary site of 16'* Century sugar production by the Portuguese), the areas adjacent to the central peak (from 1500 to 2000 m), and the forests of middle elevation on the southern exposures of both islands which are inaccessible by road. During our stay there, the local inhabitants and expatriates on S40 Tomé were consistent in telling us that amphibians were not found above 500 or 600 m, but Schiotz (1999) wrote that Hyperolius (as Nesionixalus) molleri occurs between 500-1000 m, evidently following Loumont (1992). Below, we provide species distributions based on our own data for specimens collected at altitudinal extremes, our interpretation of Loumont’s (1992) data and records by J. Baillie (1999). Hyperolius thomensis Bocage, 1886. This frog is the largest member of the genus Hyperolius and an island giant. H. thomensis is an endemic to Sao Tomé Island and restricted to remnants of primary forest which are usually at higher elevations above 800 m. The five type specimens lost in the Lisbon fire included a male and female from Roca Saudade (800 m), a male and female from “Tle Sao Tomé” (exact provenance unknown), and a single female from the Rio Quiza (= Rio Quya, A. Gascoigne, pers. commun.), a river in the southwestern part of the island with headwaters at 900 m. Specific localities documented by voucher specimens include CAS 219059, above Bom Sucesso on trail to Lagoa Amelia (ca. 1170 m) and CAS 218925—218937, in the tree hole on pri- mary-forested ridge at Macambrara (1280 m.). Probable localities (viz Loumont, 1992) are forest remnants on slopes above Monte Café and Java, and inland from Colonia A¢oreana. We reject the capital city of Sdo Tomé as a locality for this species. It is likely that this species still exists wherever primary forest remains, especially on the steep terrain of relatively inaccessible southern slopes of the island. Hyperolius molleri Bedriaga, 1892. A common Hyperolius on both Sao Tomé and Principe islands, this species appears to be a classic “farmbush” species, sensu Schiotz (1967). It breeds in still or very slow-moving water: egg masses are deposited on overhang- ing vegetation (Fig. 1). On SAo Tomé, H. mol- leri occurs at sites near sea level adjacent to a : j FIGURE 1. Egg mass of Hyperolius molleri. Sao Tomé: Sao Tomé town such as at Praia Melao (CAS — Quisinda. Photograph by D. Lin 578 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 31 219068-219069), is common in both low and mid-elevation abandoned and functioning planta- tions such as those at Caxueira (CAS 218848—218891) and Java (218974—218994), and specimens were found calling from bushes on the banks of Lagoa Amelia in primary forest at 1412 m (CAS 219048-219050). On Principe, H. molleri was found in the same microhabitats as on SAo Tomé. CAS 219203- 219208 were collected while calling from vegetation near pooling areas of a small stream at sea level, just in from our campsite on the beach at Baia das Agulhas on the southwestern coast. Another series was taken at about 150 m near the town of Santo Antonio while males called from vegetation overhanging eddies of the slow-moving Ribeira Doutor. Notably, RES did not encounter H. molleri at higher elevations during her ascent of Pico do Principe. Leptopelis palmatus (Peters, 1868). Endemic to Principe Island, this species has hitherto been considered a lowland treefrog based on three localities reported by Loumont (1992), who suggest- ed it was found up to 300 m. CAS 219370—219383 and 219397—219401 were collected at about 620 m on the central massif of Pico do Principe; BMNH 2000.58 was taken at approximately 700 m during a daytime descent from Pico do Principe by Jonathan Baillie in 1999. (Baillie, personal communication) Because this collection is the basis for the first description of the male of this species, it is treated in further detail in Section III, below. Ptychadena newtoni (Bocage,1886). This Sao Tomé endemic may be the only true lowland amphibian species sensu Loumont (1992); all of her collection localities are in the northern low- lands of Sao Tomé Island (north of a line between SA0 Tomé town and Diogo Vaz on the northwest coast), well below 150 m. We collected two series of adults in town (CAS 219248—219263 in an undeveloped lot, and CAS 219313—219317 in a swampy area of Agua Grande). We also collected a series of P. newtoni tadpoles (CAS-RCD 13682) together with Phrynobatrachus dispar larvae in a road-side puddle at Java (595 m), which would constitute an altitude record for these species. At this site, no adults were seen. A description of the PR newtoni tadpole and others is in progress. P newtoni is the only endemic amphibian species of Sao Tomé and Principe that may be endangered. Since Loumont’s visit, many low-lying areas in the vicinity of Sao Tomé town have been drained for housing. We visited a number of sites at which we were told this species used to be common; in most cases, the areas were dry. Loumont (1999) records a female P. newtoni of 76 mm, snout-vent length. To our knowledge this is the largest known Prvchadena specimen. This indicates that together with Hyperolius thome- nsis, and Leptopelis palmatus, there are three examples of anuran island gigantism inhabiting the Gulf of Guinea Islands. Phrynobatrachus dispar (Peters, 1870). This species was originally described from Principe Island; a second species, Phrynobatrachus feae (Boulenger, 1906) also from Principe, was described later. The differences between descriptions are rather minor, and Loumont (1992) con- cluded that the latter is a synonym of the former. The CAS expedition took tissue samples from a number of populations on both islands but has not yet sequenced them. We treat these taxa as one, Phrynobatrachus dispar. P. dispar is a nearly ubiquitous frog in suitable wet areas in disturbed as well as near-pristine conditions on both islands. CAS 218918—218919 and 219064219077 were collected at 700 m in aqueduct tunnels of the Rio Contador on the west side of Sao Tomé Island; CAS 219047 was taken from the edge of Lagoa Amelia at 1412 m. On Principe Island P. dispar was collected by RES at 620 m (CAS 219385—219393) in a tributary of the Ribeira Banzu, and on the top of Pico do Principe at 948 m (CAS 219393219394). Baillie (1999) records the species as very common on the top of Pico Mesa, ca. 530 m. Schistometopum thomense (Bocage,1873). The endemic Sao Tomé caecilian is perhaps the most unlikely amphibian inhabitant of the island from a dispersal point of view. Wholly fossorial, DREWES AND STOELTING: GULF OF GUINEA AMPHIBIANS Sf) it might be expected to occur principally in the lowlands, but such is not the case. Two specimens were collected on the west side of the island in a vertical rock crevice along the Rio Contador cul- vert at 700 m. (CAS 218914—218915), and a series (CAS 219324—219334) was found in a cultivat- ed field above the arboretum at Bom Sucesso, ca 1180 m. J. Measey (personal communication) has collected specimens of S. thomense within 100 m of the Lagoa Amelia crater rim (which would be at about 1300 m). Ill. The male of Leptopelis palmatus Prior to the CAS Gulf of Guinea expedition, the Principe endemic treefrog Leptopelis palma- tus (Peters. 1868) was represented in collections by the female holotype (ZMB 6067: “ile de Principe”), eight females from three lower-elevation localities reported by Loumont (1992) and deposited in the MHNG, and two males and a female in the collections of BMNH collected by J. Baillie in 1999. Loumont (1992) published a table of measurements of the females including MHNG 2491.82 from Bela Vista with a snout-vent length of 110 mm, indicating that Leptopelis paimatus attains a greater snout-vent length than any other African treefrog species. She included a photograph of one of the females (1992, fig. 7; museum number not given) which illustrated the dorsal pattern described as marbled (“un aspect marbré”’); she described the other specimens as being dark green or black with a number of light patches. A photograph by Loumont of what appears to be the same specimen is presented in Schiotz (1999, fig. 570), which together with the aforementioned comprise the only published images of L. palmatus to date excepting three on-line images by Baillie (1999). Figure 2 illustrates the range of color pattern variation we observed in females. Figure 2A is a 108 mm SVL individual collected in at Tchipique, a locality in the north- western part of Principe Island; Figure 2B is 88 mm SVL from the Rio Papagaio a central island locality. BMNH 2000.58, the smallest mature female known (oviducal eggs present) exhibits a dark brown pattern on a beige background which we interpret as cryptic. Our series of eleven adult male L. palmatus was collected by the second author (RES) over 17-18 May 2002 on the massif of Pico do Principe along a tributary of the Ribeira Banzu at about 620 m elevation. The same locality yielded two females, three juveniles and three post-metamorphs (Appendix I). The locality is a small cascading stream in primary forest dominated by saplings and tree ferns (the endemic Cyathea camerooniana vat. currori). The specimens were collected from 1800—0030 hr on the 17 May and 1800-2000 hr on 18 May. The males were perched on leaves or branches of saplings or leaves of tree ferns from 1.0—1.5 m above ground, although the larger FIGURE 2. (A) CAS 219177, female Leptopelis palmatus. Principe: Tchipique, near Sundi. Photograph by D. Lin; (B) CAS 219351, female L. palmatus. Principe: Rio Papagaio. Photograph by J. Ledford. [ 580 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 31 female (CAS 219401) was found on the ground beneath a rocky overhang near a pool at the base of a small waterfall. A small post-metamorph, CAS 219370, was found perched on the same leaf as an adult male. RES collected an area about 10 m out from both banks of the stream along approximately 30 m of its length and found the tree frogs remarkably numerous: she saw three times as many specimens as she collected (ca. 60 seen). On both nights RES heard calls of Phrynobatrachus dispar which were numerous in the leaf litter near the stream, but did not hear any advertisement call attributable to L. palmatus males. Description of males Adult male Leptopelis palmatus specimens are less than half the snout-vent length of females (male mean = 41.4 mm — Appendix I; female mean = 93.8 mm, 7 = 12, including data from Loumont, 1992). L. palmatus males are remarkably polymorphic for color pattern; male color pat- terns include individuals with uniform bright green dorsum, slightly darker green with random mustard blotches or faint blackish vermiculations, pinkish maroon with indistinct olive blotches and occasional dull-orange spots, mottled light brown and white, and light brown individuals with fairly distinct partial “X”’-shaped patterns on the dorsum and distinct thigh bands. (Fig. 3). The dor- sal and lateral skin of all specimens is heterogeneous and rough, as in females. The male ventrum is not dark as in females, but ranges from off-white to lightly mottled gray. In larger females, there is a conspicuous fold of skin extending across the head between the posterior margins of the tympani. This structure 1s quite obvious in Peters’ drawing of the holotype (1868), in Loumont’s photographs (1992, in Schiotz 1999) and in our own Figure 2A. This feature was not discussed in Perret’s examination of the type (1973). It is absent in all male specimens and also absent in smaller CAS females (219351; 219401) photographed in life. The irises of all males (and females) are a deep ruby-red in life, in contrast to the gold iris of L. rufus (Schiotz, 1999) with which L. pa/matus was compared in an earlier study by Perret (1973) (Figs. 2-3). This ruby iris color is most similar both in hue and extent to that of live specimens of Leptopelis parkeri of East Africa. As suggested by the figures in Schiotz (1999), whereas many species of Lepfopelis have reddish coloration in the dorsal quarter to fifth of the iris, only a few appear to consistently have red irises, and these are of a lighter shade than L. palmatus, e.g. L. kivuensis and Nigerian L. boulengeri. The tympanum in males is conspicuous and measures a little less than one-half the diameter of the eye as in the female (Perret 1973). The eyes are large and protruding, their horizontal diam- eter nearly equal to the interorbital distance. The toes of the males are nearly as fully webbed as in females, but the proximal side of the terminal phalanx of toe IV is free. The fingers of males are less webbed than those of the females; the area between toes IV and III is about one-half webbed, that between HI and I about one-quar- ter webbed (for comparison, see figure of holotype in Boulenger [1882:136]). Males possess poor- ly defined pectoral glands discernible as light patches posterior to the posterior edge of the inser- tion of the forelimbs. Dissection of the gular regions of three males (CAS 219373; 219377 and 219400) revealed that L. palmatus males lack a vocal sac; there is no elaboration of the m. interhyoideus. Vocal sac openings are absent. This is a unique condition within the genus Leptopelis although there have been few studies that include these characters (e.g., Drewes 1984, for 11 species). Male advertisement call The advertisement call of male L. pa/matus has never been recorded nor analyzed; however, } | t } DREWES AND STOELTING: GULF OF GUINEA AMPHIBIANS 581 FIGURE 3. Leptopelis palmatus. A composite of adult males and one juvenile from Principe: Ribeiro Banzu (CAS 219371-219383, 219400). Photographs by J. Ledford. graphs D} we were assured by the local inhabitants of Principe Island that males do call, and the presence of well-developed tympani in both males and females tends to confirm the fact. On the evenings of 5/17 and 5/18, REL heard multiple calls of Phrynobatrachus dispar but no calls referable to Leptopelis. Males did not call in captivity during transfer back to Santo Antonio. In 1999, the male call was described by Jonathan Baillie in his on-line account of his dissertation field research on Principe Island. The following is an excerpt from his account dated | September 1999: 582 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 31 In the dark we took flashlights and cameras to try and find the frogs responsible for the loud night chorus. Phrynobatrachus dispar was also common in the small streams. . . . If you listen closely you can hear the soft repetitive peep-peep-peep of this tiny frog. The more noticeable croaking noise, more like a pop bottle being continuously opened, belongs to the tree frog (Leptopelis palmatus) that is found in high densities (we saw about 50 that night) on the branches and vines in close proximity to the stream. This “popping” sound had been described to us by locals earlier in Principe’s only town, Santo Antonio, and it is consistent with the non-resonant call that might be associated with males lacking a vocal sac. If this call is correctly assigned, it is very different from the call described ono- matopoetically by Schiotz (1999:256) for Leptopelis rufus: “a series — about ten long cries [of] ‘yiin.’” Baillie’s account and personal communications suggest that the campsite near which these observations were made was probably the same site at which RES collected the CAS series of L. palmatus, or at least very close to it (Baillie’s local guide, Monauna, a former parrot hunter, also guided the CAS party nearly two years later). On 5 Sept, Baillie again heard L. palmatus advertise- ment calls at about 700 m in the forest of Morro de Leste, a ridge approximately 3 km ENE of Pico do Principe, but could not locate any individual frogs (Baillie 1999). Island effects Female Leptopelis palmatus are both the largest of that hyperoliid genus and the largest known African treefrog. They are also among the few anuran island giants that have mainland congeners including Hyla vasta and Eleutherodactylus inoptatus, both of Hispaniola (Cochran 1941; Zug et al. 2001). The two islands of Sao Tomé and Principe have a number of endemics that are classic examples of island gigantism (Carlquist 1965). On Sao Tomé, some of these include the largest treefrog of the hyperoliid genus Hyperolius (H. thomensis, Drewes and Wilkinson 2004), the largest member of the widespread African ranid genus Ptychadena, P. newtoni, the largest nec- tariniid sunbird, Dreptes thomenis (Cheke and Mann 2001), and the giant two-meter begonia, B. baccata (Plana et al., 2004). Both islands share the endemic gecko, Hemidactylus greefiii, which 1s the largest African member of that genus (Loveridge 1947). Although it is an obvious and frequent- ly documented phenomenon, island gigantism is not well understood; release from competition for resources and predation have been suggested as causative factors. Our series of male Leptopelis palmatus exhibit what appears to be non-fixed color pattern polymorphism. Some non-cryptic, arboreal African treefrogs, notably populations of the Hyperolius viridiflavus complex, exhibit striking inter- and intrapopulational variation in color pat- tern, which has led to a proliferation of taxonomic names (Schietz 1971). Within such populations, this variation can usually be assigned to one of a number of phases. For instance, Schiotz (1967), described in Hyperolius a PhJ or juvenile phase exhibited by all juveniles and some adult males and a PhF, the female phase, exhibited by all adult females and some males. In our series of male L. palmatus, color and pattern are seemingly random, and there appears to be no selection for cryp- sis (Fig. 3). We know of no other African treefrog with this magnitude of color pattern variation. Females are dull green, brown or black, whether marbled or not (Figs. 2A—B). We considered the possibility of a sexual difference in microhabitat which might account for the less variable, more subdued coloration in females. Although all of the CAS males were collected in arboreal situations, two of the CAS females, 219177, the largest, most heavy-bodied specimen (Fig. 2A), and CAS 219401 were found — at different localities — on the ground in moist areas protected by over- hangs (Appendix, Table 1). BMNH 2000.58 was found on the forest floor. The other two females, CAS 219399 and 219351, were collected while perched. Unfortunately, our sample of females is DREWES AND STOELTING: GULF OF GUINEA AMPHIBIANS 583 limited to five as there are no habitat data associated with the eight females Loumont deposited in Geneva (MHNG). Sexual size dimorphism is the rule among Lepfopelis species, and, based on specimens collect- ed. L. palmatus exhibits the greatest disparity between adult male and female snout-vent lengths of any species of the genus. Schiotz (1999) provides male and female snout-vent maxima for 33 of the 44 species of Leptopelis he recognized and in most species, males range from 8—20 mm small- er in snout-vent length than females. Table 2 (see Appendix) is a compilation of maximum male and female snout-vent lengths recorded from the seven Lepfopelis species in which the females are greater than 20 mm in length than the males. Our data indicate that the largest known L. palmatus male is only 41% of the snout-vent length of the largest female. Mean male size is 45% of mean female SVL. The only other species in which males are less than half as long as females is the widespread West African mainland species, L. macrotis at 45% (based on SVL maxima). In addi- tion, we are unaware of any other anuran species that exhibits such as marked range of SVL among adult females (Appendix I). There is a full order of magnitude difference in L. palmatus between post-metamorphic size (10.5—11.3 mm, Gosner stages 45, 46 — Table 2 in Appendix) and potential adult size (MHNG female 2491.82—110 mm; Loumont 1992). Nothing is known of the breeding biology of L. palma- tus, and the larvae have yet to be discovered. Small size at metamorphosis can be interpreted as an indication that the water in which early development takes place is ephemeral (this would seem unlikely on Principe which has high rainfall especially in the forested higher elevations) or that pre- dation on developing larvae is high. Baillie (1999) noted abundant crabs of about 9 cm in diame- ter feeding in the river at night at the same elevation (and presumably at the same L. palmatus col- lection site; see above), and these may be a significant predator on small frogs. CAS 219370 (11.0 mm) was perched on the same leaf as an adult L. palmatus male (field number not noted), about 1.5 m above ground, suggesting both resource availability for individuals at both sizes and perhaps avoidance of predation by crabs. However, RES observed no crabs during her visit to the site. The CAS Leptopelis palmatus were removed and taken to lower elevation for photography, processing and removal of tissue; as a result an examination of stomach contents was not informative. Reduction of predation pressure may well be a factor in the extreme variation 1n color and pat- tern observed among L. palmatus males (at least for the known sample which was collected at densely-forested higher elevations). Color pattern variation appears much reduced in females sug- gesting that females are under a different selection regime. ACKNOWLEDGMENTS The California Academy of Sciences Gulf of Guinea Expedition would have been impossible without the assistance of Ned Seligman, Director of STePUP, SAo Tomé, Angus Gascoigne of Voice of America, SAo Tomé, 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 Sao Tomé and Principe. Additional thanks go to all of the mem- bers 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. Omar Idris Nur Husein of the University of Texas, Arlington shared measurements of Leptopelis palmatus males on loan to him from CAS. 584 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 31 Andreas Schmitz of MHNG kindly provided archived MHNG data on the Loumont collections; Barry T. Clarke of BMNH kindly loaned specimens of Leptopelis palmatus collected by J. Baillie, and the latter is thanked for providing additional field data. Our thanks to Elisabeth Fourtanier (CAS) for help with translation. Ronn Altig reviewed an early version of the manuscript. Special thanks to Angus Gascoigne, who as a fine field naturalist and resident of Sao Tomé has been a ready and willing source of information for us and many others. This paper represents contribution no. 36 of the Center for Biodiversity Research and Information of the California Academy of Sciences. LITERATURE CITED ALTIG, R. 1970. A key to the tadpoles of the continental United States and Canada. 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Annales del al Faculté des Sciences du Yaoundé 15—16:81—90. PERRET, J.-L. 1976. Revision des amphibians africaines et principalement des types, conserves au Musée Bocage de Lisbonne. Arquivos do Museu Bocage (2 serie) 6:15—34. DREWES AND STOELTING: GULF OF GUINEA AMPHIBIANS 585 Perret, J.-L. 1988. Sur quelques genres d’Hyperolidae (Anura) restés en question. Bulletin de la Société Neuchdateloise des Sciences Naturelles 111:35-48. Perers, W. 1868. Uber eine neue Nagergattung, Chiropodomys penicillatus, so wie tiber einige neue oder weiniger bekannte Amphibien und Fische. Monatsberichte der Akademie der Wissenschaften zu Berlin 1868:445—470. PLANA, V., A. GASCOIGNE, L.L. FoRREsT, D. Haris, AND R.T. HARRINGTON. 2004. Pleistocene and pre- Pleistocene Begonia speciation in Africa. Molecular Phylogenetics and Evolution 31:449-461. RopeEL. M.-O. 2000. Herpetofauna of West Africa. Vol. 1. Amphibians of the West African Savanna. Edition Chimaira, Frankfurt-am-Main, Germany. 283 pp. ROpeEL, M.-O., V.H.W. RUDOLF, S. FROSCHAMMER, AND K.E. LINSENMAIR. (In press.) Life history of a West African tree-hole breeding frog, Phrynobatrachus guineenesis Guibé and Lamotte, 1961 (Amphibia: Anura: Petropedetidae). Special Publications of the University of Michigan Museum of Zoology ScHATTI, B., AND C. LOUMENT. 1992. Ein Beitrag zur Herpetofauna von SaoTomé (Golf von Guinea) (Amphibia et Reptilia). Zooogische Abhandlungen. Staatliches Museum fiir Tierkunde Dresden 47(4):23-36 ScuioTz, A. 1967. The treefrogs (Rhacophoridae) of West Africa. Spolia Zoologica Musei Hauniensis 25:1-346. ScHioTz, A. 1971. The superspecies Hyperolius viridiflavus (Anura). Vedenskabelige Meddelelser fra Dansk Naturhistorisk Forening 134:21—76. ScHioTz, A. 1999. Treefrogs of Africa. Edition Chimaira. Frankfurt-am-Main, Germany. 350 pp. ZuG, GR., L.J. Vitt, AND J.P. CALDWELL. 2001. Herpetology. An Introductory Biology of Amphibians and Reptiles, 2"4 ed. Academic Press, San Diego, San Francisco. xiv + 630 pp. 586 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 31 Appendix TABLE |. Leptopelis palmatus specimens collected on Principe Island by the CAS Gulf of Guinea Expedition including three specimens from the BMNH. Sexual maturity was determined by dissection in male specimens greater than 33 mm SVL. Sex Museum Catalog Locality Snout-vent length (mm) number Males CAS 219371 Ribeira Banzu* 44.2 219372 45.6 D933, 38.4 219374 34.7 219377 41.8 219378 42.8 219379 44.5 219380 43.35 219381 42 219382 41.2 219400 35 BMNH*** 2000.59 “Pico de Principe” 600 m*. 43.8 2000.60 41.4 Juv. CAS 219375 23 219376 I) 219383 22 metamorphs 219370 11.0 — Gosner stage 45 219397 10.5 — Gosner stage 46 219398 11.6 — Gosner stage 46 Females 219399 105 219401 69.2 POI Tchipique 108 219351 Rio Papagaio 88.8 BMNH*** 2000.58 “Pico de Principe” 700 m** 63.99 (oviducal eggs) * We are reasonably certain that these localities are equivalent in spite of differences in our GPS coor- dinates(see text): CAS = 01 35 20.7 N, 007 22 46.3 E; Baillie (1999) = 01 37 11N, 007 22 52 E. ** This specimen was collected on the forest floor by Baillie at about 700 m during his first descent from Pico do Principe. The precise locality is unknown but if the estimate of elevation is correct, this is the highest known specimen (Baillie personal communication). *** Non—CAS Specimens examined: BMNH 2000.58. Female. Pico do Principe, Principe, 700 m. Sept 1, 1999. col. J.E.M. Bailie; BMNH 2000.59-60. Males. Pico do Principe, Principe 600 m. Sept. 1, 1999. col. J.E.M. Baillie. DREWES AND STOELTING: GULF OF GUINEA AMPHIBIANS 587 TABLE 2. Size disparity between male and female Leptopelis in species where females are greater than 20 mm in snout-vent length than males; compar- ative snout-vent maxima taken from Schietz, 1999. Species Maximum male Maximum female Maximum male SVL as SVL (mm) SVL (mm) % of max. female SVL Leptopelis palmatus 45.6 110 4] L. macrotis 46 84 45 L. millsoni 49 87 56 L. vermiculatus 50 85 58 L. boulengeri 48 81 59 L. occidentalis 43 71 60 Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 32, pp. 588-597, 5 figs. December 30, 2004 New Data on Tropical Eastern Pacific Chromodorididae (Nudibranchia: Doridina) with Description of a New Species of Mexichromis Bertsch, 1977 Terrence M. Gosliner!, Jestis Ortea2, and Angel Valdés} 1 Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103; Email: tgosliner@calacademy.org; 2 Departamento de Biologia de Organismos y Sistemas, Universidad de Oviedo, c/ Catedratico Rodrigo Uria, s/n, 33071 Oviedo, Spain; 3 Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007; Email: avaldes@nhm.org Specimens of a small red-spotted species of chromodorid have been collected from southern México, Costa Rica, Panama, Malpelo Island, Colombia and the Galapagos Islands. Comparison with described species reveals that they are juveniles of Chromodoris baumanni Bertsch, 1970. Specimens studied here exhibit considerable variation both ontogenetically and geographically. Mature specimens share charac- teristics with species attributed to the genus Glossodoris, and the systematic status of this species is reviewed and revised. Another species of chromodorid nudibranch is described from the tropical eastern Pacific. Mexichromis tica sp. nov. has been col- lected from Costa Rica and the Galapagos Islands. It differs from other eastern Pacific, Atlantic, and Indo-Pacific species in the genus in several important regards: (1) it has a white body color with a medial opaque white line and successive submar- ginal opaque white and orange marginal bands; (2) it has few large mantle glands distributed along the entire mantle margin; and (3) it has a vestigial rachidian row of teeth and variously denticulate inner, middle and outer lateral radular teeth. RESUMEN Ejemplares de una especie de cromodorido de pequeno tamano con puntos rojos han sido recolectados en el sur de México, Costa Rica, Panama, Isla Malpelo, Colombia e Islas Galapagos. Comparaciones detalladas con otras especies descritas muestran que son especimenes juveniles de Chromodoris baumanni Bertsch, 1970. Los ejem- plares estudiados presentan una considerable variabilidad ontogenética y geografi- ca. Los ejemplares maduros comparten caracteristicas atribuidas al género Glossodoris y el estatus taxonémico de ésta especie es revisado y discutido. Otra especie de nudibranquio cromodorido se describe del Pacifico Este tropical. Mexichromis tica sp. nov. ha sido recolectada en Costa Rica y las Islas Galapagos. Se diferencia de otras especies del género del Pacifico Este, Atlantico e Indopacifico por varios aspectos importantes: (1) Esta especie tiene un cuerpo blanco con una linea blanca opaca media y sucesivas bandas submarginales blanco opaco y naranja; (2) glandulas del manto de gran tamano distribuidas por todo el borde del manto; (3) y una fila de dientes radulares raquideos vestigiales asi como varios dientes medios y marginales denticulados. Nn ioe) CO GOSLINER ET AL.: TROPICAL EASTERN PACIFIC CHROMODORIDIDAE 589 The chromodorid nudibranchs of the eastern Pacific Ocean have been reviewed by Bertsch (1977, 1978a-—c). Since then only four additional species of chromodorid have been described from the eastern Pacific (Gosliner and Bertsch 1988; Ortea, Bacallado and Valdés 1992). Gosliner and Bertsch (1988) described Mexichromis amalguae from the Pacific coast of Baja California. Ortea et al. (1992) described three new species, Berlanguella scopae, Chromodoris ruzafai and Thorunna talaverai from the Galapagos. They also identified specimens of another chromodorid from the Galapagos Archipelago as supposedly Noumea haliclona (Burn, 1957). This species bears a strik- ing resemblance to specimens of Chromodoris baumanni Bertsch, 1970, and its status is discussed here. Our recent field collections from Costa Rica, Baja California, the Galapagos Islands, and Panama have provided additional material of this species. Detailed anatomical examination of vari- ation in this species is warranted in order to evaluate its systematic status. A second species, col- lected from Costa Rica and the Galapagos, appears to be undescribed. Its external morphology dif- fers markedly from other described eastern Pacific chromodorids. This paper describes and reviews the systematic placement of these two species. SPECIES DESCRIPTIONS Mexichromis tica Gosliner, Ortea, and Valdés, sp. nov. (Figs. 1A, 2-3) TYPE MATERIAL.— HoLotyPe: CASIZ 170938, NE side Isla Darwin, Islas Galapagos, Ecuador, 23 m depth, 13 May 1994, leg. T.M. Gosliner. PARATYPES: CASIZ 097511, one specimen, dissected, NE side Isla Darwin, Islas Galapagos, Ecuador, 23 m depth, 13 May 1994, leg. T.M. Gosliner. INBIO CRI 001486639, two specimens, dissected, Bajo del Diablo, Isla del Cafo, Costa Rica, 18 April 1996, leg. E. Mollo. ETyYMOLOGY.— Mexichromis tica is named for the vernacular name “tico,” which signifies a native of Costa Rica. DISTRIBUTION.— This species is known from Costa Rica and the Galapagos Islands (present study). EXTERNAL MORPHOLOGY.— The living animals (Fig. 1A) are translucent white. The pink vis- cera, showing through the translucent white dorsum, give the animal an overall pinkish appearance. A broad, opaque, white longitudinal band extends mid-dorsally from between the rhinophores to the anterior edge of the branchial cavity. The mantle margin is translucent white. A broad, opaque, white band is present submarginally along the mantle edge. Inside the white band, there is a band of orange or yellow that may be either continuous or interrupted. The posterior end of the foot is translucent white with a medial white stripe. An orange spot is present near the middle of the white stripe. The rhinophores have a translucent white base and an orange apex. The gill pinnae are uni- formly translucent white. Living animals are 3-6 mm in length. The dorsal surface of the mantle is covered with minute conical tubercles. The mantle is elongately oval and straight along its entire edge. The margin that overhangs the body is wide, nearly half of the total mantle width. The posterior end of the foot is triangular and extends some distance behind the posterior end of the mantle. Around the mantle margin are several rows of large, irregularly-shaped mantle glands (Fig. 3A). The glands of the largest specimen are more numerous with smaller glands situated between the larger ones. Triangular spicules are visible between the mantle glands around the margins of the mantle. The rhinophore stalk is short and the bulb bears 7-10 lamellae in the specimens examined. There are 4 unipinnate to bipinnate branchial leaves in the material examined. The foot is relatively narrow. The head and mouth are well-developed with a triangular oral tentacle on either side of the mouth. 590 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 32 ee Figure 1. Living animals. A. Mexichromis tica sp. nov, holotype (CASIZ 097511), from Isla Wolf, Galapagos Islands, photo by T.M. Gosliner. B. Glossodoris baumanni (Bertsch, 1970), adult specimen, 33 mm length, from San Pedrillo, Costa Rica, photo by T.M. Gosliner. C. Glossodoris baumanni (Bertsch, 1970), juvenile specimen, 12 mm length, from San Pedrillo, Costa Rica, photo by T.M. Gosliner. BUCCAL ARMATURE.— At the anterior end of the muscular portion of the buccal mass 1s the chitinous labial cuticle, which bears numerous jaw rodlets. The rodlets (Fig. 2D) are short and broad with a broad apex bearing 3-7 irregular shaped denticles. The radular formula is difficult to determine owing to the broadly overlapping teeth. A vestigial rachidian row of teeth is present in one specimen (CASIZ 097511, Fig. 2A), but it appears to be absent in the other two individuals examined (INBIO CRI 001486639). The rachidian row of teeth in the one specimen consists of teeth with only a short narrow cusp. The innermost lateral teeth (Fig. 2A) have a broad base with a series of 4-6 curved, triangular denticles along the inner margin. There is no primary cusp and most of the denticles are equal in size with the exception of the outermost, which is smaller than the others. The lateral teeth from the middle of the radular row are short and curved (Fig. 2C) and bear 4—6 curved, triangular denticles on the outer side of the teeth. The more basal denticles are smaller than the others, but there is no primary cusp on any of these teeth (acuspidate, sensu Bertsch, 1977). The 4-5 outermost lateral teeth (Fig. 2B) are broad and spatulate and are thinly chi- tinized. They bear 16—20 thin, elongate denticles along the margin. REPRODUCTIVE SYSTEM.— (Fig. 3B) The ampulla is short, thick and tubular, narrowing some- what before bifurcating into an oviduct and vas deferens. The short oviduct enters the female gland mass near the albumen gland. The proximal prostatic portion of the vas deferens curves over the bursa copulatrix and narrows into a short ejaculatory segment. This muscular portion narrows and again widens into the short penial bulb. The penial bulb shares a common atrium with the vagina. The distal end of the vas deferens is devoid of penial hooks. The female gland mass consists of the GOSLINER ET AL.: TROPICAL EASTERN PACIFIC CHROMODORIDIDAE 5911 Fy j e 7 FIGURE 2. Mexichromis tica sp. nov., scannning electron micrographs. A. Rachis and inner lateral teeth (paratype, CASIZ 097511), scale bar = 10 um. B. Outer lateral teeth, (paratype, CASIZ 097511), scale bar = 10 um. C. Lateral teeth from the central portion of half-row, (paratype, CASIZ 097511), scale bar = 7.5 um. D. Jaw rodlets, Kparaly pe, INBIO CRI 001486639), scale bar = 5 um. large mucous gland and smaller membrane and albumen glands. Near the exit of the mucous gland is a small, ovoid vestibular gland. The vagina is relatively thin and straight. It is widest nearest its junction with the penis. The elongate, club-shaped receptaculum seminis has a recurved duct that 592 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 32 joins directly to the base of the thin-walled, spherical bursa cop- ulatrix. The vagina emerges near the base of the bursa. The thin uterine duct emerges from the middle of the thin vagina. The uterine duct is short and curved and enters the female gland mass near the albumen gland. DISCUSSION.— Rudman (1984) reviewed the genera of chromodorid nudibranchs and concluded that Mexichromis Bertsch, 1977, represents a valid genus. Mexichromis 1s character- ized by having a few large man- FIGURE Se ears tica Sp. NOV., A. vente of holotype (CASIZ ; 097511) showing distribution of mantle glands, g = mantle gland, scale bar = tle glands, acuspidate (Bertsch 2 mm. B. Reproductive system, am = ampulla; be = bursa copulatrix; ej = ejac- 1977) or multicuspidate (Rud- — ulatory duct; fg = female gland mass; p = penis; pr = prostate; rs = receptacu- man 1984) radular teeth, and a lum seminis; vg = vestibular gland, scale bar = 1 mm. ramified vestibular gland (Rud- man 1984). In the eastern Pacific, Wexichromis is represented by M. antonii (Bertsch, 1976), M. porterae (Cockerell, 1901), VZ tura (Marcus and Marcus, 1967) and M. amalguae Gosliner and Bertsch, 1988. Mexichromis francoisae (Bouchet, in Bouchet and Ortea, 1980) is known from the eastern Atlantic of Senegal (Bouchet and Ortea 1980) and the Cape Verde Islands (Ortea 1988). This species was transferred to the genus Mexichromis by Ortea et al. (1996). Ortea et al. (1996)also described M. molloi Ortea and Valdés, 1996, from Venezuela. These eastern Pacific and Atlantic species all have a blue ground color with yellow or white longitudinal lines or marginal bands. Rudman (1984) suggested that Chromodoris kempfi Marcus, 1970, may be a species of Mexichromis, but little is known of its anatomy. Its greenish body color with a yellow marginal band with black markings differs markedly from the color of M. fica. Several Indo-Pacific species, M. festiva (Angas, 1864), MM. mariei (Crosse, 1872), VM. macro- pus Rudman, 1983, and M. multituberculata (Baba, 1953), have an opaque white body color with large purple tubercles on the notum. Additional purple, yellow or orange pigment may also be pres- ent. The color pattern of MV. fica, with a transluscent white ground color with opaque white and yel- low submarginal bands, is unique for this species. Of the species of Mexichromis that have been studied, only M. amalguae and M. tica are known to possess a vestigial rachidian tooth. However, a rachdian row of teeth may be present or absent in M. fica. Mexichromis tica is unique among members of Mexichromis in having multidenticulate to pectinate outer lateral teeth. In the remain- ing taxa, the outer lateral teeth are similar in shape to the midlateral teeth. The reproductive system has been described in detail for Mexchromis macropus, M. porterae, M. tura, M. francoisae, M. molloi, and M. amalguae (Rudman 1984; Gosliner and Bertsch 1988; Ortea et al. 1996). In these six species, the vestibular gland consists of multiple lobes, whereas in M. tica it is simply ovoid. Inclusion of M. tica in Mexichromis requires modifying the boundaries of the taxon to include species with pectinate outer lateral teeth and a simple vestibular gland. Mexichromis tica is tenta- GOSLINER ET AL.: TROPICAL EASTERN PACIFIC CHROMODORIDIDAE 598 tively placed in Mexichromis until a more comprehensive phylogenetic study of the Chromodoridiae is undertaken. Glossodoris baumanni (Bertsch, 1970) (Figs. 1B—C, 4-5) Chromodoris norrisi Farmer, 1963: Marcus and Marcus, 1967:170—173, fig. 24, misidentification. Chromodoris baumanni Bertsch, 1970:8, figs. 3-13; Sphon and Mulliner, 1972:150; Bertsch, 1978a:307, fig. 3a, 4, 10-12: Bertsch, in Kerstitch, 1991:57; Debelius, 1996:212, lower figure. Noumea haliclona (Burn, 1957): Ortea, Bacallado and Valdés, 1992:62—65, figs. 19—20, pl. 1E, misidentifica- tion. Glossodoris baumanni, (Bertsch, 1970), comb. nov. MATERIAL EXAMINED.— CASIZ 074004, one specimen, dissected, s side of Pulmo Point, Cabo Pulmo, Baja California Sur, México, 13 m. depth, 22 January 1984, leg. H. Bertsch and L. Aguilar. INBIO CRI 001486926, one specimen, Bahia Hachal, Costa Rica, 8 December 1995. INBIO CRI 001486927, two specimens, Punta Uvita, Costa Rica, 14 April 1996. CASIZ 075179, one specimen, Isla del Coco, Costa Rica, 13 m. depth, 24 March 1989, leg. K. Kaiser WAM 102- 95. one specimen, Isla de Cocos, Costa Rica, 28 March 1997, leg. C. Bryce. WAM 106-95, one specimen, Isla de Cocos, Costa Rica, 23 March 1997, leg. K. Kaiser. CASIZ 088147, five speci- mens, one dissected, sea stack, off Isla Ladrones, Gulf of Chriqui, Panama, 14 m. depth, 14 April 1993, leg. T.M. Gosliner. CASIZ 088150, three specimens, Punta David, se side of Isla Jicarén, Gulf of Chiriqui, Panama, 13 m. depth, 17 April 1993, T.M. Gosliner. CASIZ 088203, four speci- mens, point W of anchorage, Islas Secas, Gulf of Chiriqui, Panama, 13 m. depth, 22 April 1993, leg. T.M. Gosliner. CASIZ 088227, one specimen, off Rocas Tiburon, off Isla Brincano, Gulf of Chiriqui, Panama, 23 April 1993, leg. T.M. Gosliner. CASIZ 088161, two specimens, se side of Isla Jicaron, Gulf of Chiriqui, Panama, 16 April 1993, leg. T.M. Gosliner. CASIZ 097512, one speci- men, NE side Isla Darwin, Islas Galapagos, Ecuador, 22 m. depth, 13 May 1994, leg. T.M. Gosliner. CASIZ 071429, one specimen, Isla Santiago, Islas Galapagos, Ecuador, intertidal zone, 19 February 1988, leg. K. Kaiser. WAM S1164, one specimen, Isla Malpelo, Colombia, 13 m depth, 26 March 1998, leg. C. Bryce. One specimen, Puerto Egas, Isla Santiago, Islas Galapagos, 18 March 1991, leg. J. Ortea. One specimen , Isla Genovesa, Islas Galapagos, 8 March 1991, leg. J. Ortea. DISTRIBUTION.— This species is known from the Gulf of California, Baja California Sur (Bertsch, 1970, 1978a; present study) México; Sayulita, Nayarit, México; several localities in Costa Rica, Panama, the Galapagos Islands and from Isla Malpelo, Colombia (present study). NATURAL HISTORY.— Glossodoris baumanni is found in relatively shallow water on rocky reefs. In Costa Rica and Panama, it is one of the most commonly occurring opisthobranchs from the intertidal zone to 20 meters depth. EXTERNAL MORPHOLOGY.— The living animals (Fig. |B—C) are brightly colored with a net- work of purple to red and opaque white pigment. There is a broad yellowish white or light gray band around the mantle, just inside the translucent white margin. The inside edge of the white bor- der is very irregular and slightly convoluted with opaque white, with 1-5 rows of rounded mantle glands extending into the brick red central part of the body (Fig. 1C). There are numerous purplish red spots, each surrounded by a broad purple-gray or yellowish ring all over the central brick red patchwork. Specimens from Baja California generally lack red pigment between the red spots and the notum is opaque white. More mature specimens south of Baja California have dense red pig- ment between the red spots (Fig. 1B). An identical pattern is present on the surface of the foot. The 594 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 32 majority of these spots are joined together forming patches of varying sizes. The rhinophores have an opaque white base and a broad purple band just below the opaque white apex. The gill pinnae are also opaque white with purple apices. Scattered red pigment spots are present on the innner and outer surfaces of the gill rachis. The living animals are 9-65 mm in length. The mantle is elongately ovoid and folds down over the sides of the body forming a series of permanent undulations of the mantle edge. These undula- tions are more pronounced in more mature specimens. The margin that overhangs the body is wide, nearly half of the total mantle width. The posterior end of the foot is triangular and extends some distance behind the posterior end of the mantle. Immediately interior to the mantle margin are 1—5 rows of irregularly distributed, rounded mantle glands (Fig. 5A). The mantle glands are more dense and smaller near the margin. The interior glands are larger and have an irregular lobed shape. The rhinophore stalk has 9-22 lamellae in the specimens examined. There are 6—15 unipinnate branchial leaves in the material examined. In some of the larger speciments a few of the posterior gills are not entirely separate to the base, but have been counted as distinct gill branches. In the larger specimens the branchial leaves form a spiral on either side and the gill has been observed to vibrate slowly. BUCCAL ARMATURE.— The muscular portion of the buccal mass is approximately equal in length to the oral tube. At the anterior end of the muscular portion of the buccal mass is the chiti- nous labial cuticle, which bears numerous jaw rodlets. The rodlets (Fig. 4D) are elongate, curved and have a bifid apex. The radular formula is 61 x 39.1.39. in one 20 mm specimen examined (CASIZ 088203) and a 12 mm specimen 33 x 21.0.20. A rachidian row of teeth is present in one specimen, but absent in the other four individuals examined. Examination of the holotype and paratype radula revealed similar variation in the presence and absence of a rachidian row of teeth, with the holotype lacking a rachdian row, while the paratype has a row of rachidian teeth. The innermost lateral teeth (Fig. 4A) are arched with a short cusp. There are 1—2 triangular denticles on the inner side of the tooth. The outer side has 1—3 denticles. In the radulae with the rachidian tooth present, it is apparent that the rachidian tooth actually represents a portion of the inner lateral tooth where the inner denticle was present. The second through fourth laterals have 3-4 denticles on the outer side of each tooth. The lateral teeth from the middle of the radular row are far more elongate (Fig. 4B) and bear 4—8 small, triangular denticles on the outer side of the teeth. The outermost lat- eral teeth (Fig. 4C) are elongate with 4-5 denticles situated near the top of the tooth. REPRODUCTIVE SYSTEM.— (Fig. 5B) The ampulla is elongate, thick and tubular, narrowing somewhat before bifurcating into an oviduct and vas deferens. The short oviduct enters the female gland mass near the albumen gland. The proximal prostatic portion of the vas deferens is highly convoluted and extensive. It narrows somewhat as it develops into the muscular, ejaculatory por- tion. The ejaculatory portion is initially narrow and gradually widens and curves over the bursa copulatrix. It then contines with another series of loops and widens into the penial bulb. The penial bulb shares a common atrium with the vagina. The distal end of the vas deferens is devoid of any penial hooks. The female gland mass consists of the large mucous gland and smaller membrane and albumen glands. Near the exit of the mucous gland is a small, ovoid vestibular gland. The vagina is relatively thin, elongate and somewhat convoluted. It is muscular in the distal half and becomes glandular in the half closest to the genital aperture. The elongate, club-shaped receptaculum sem- inis joins directly to the base of thin-walled, spherical bursa copulatrix. The uterine duct and vagi- na both emerge near the base of the bursa. The uterine duct is short and curved and enters the female gland mass near the albumen gland. Discusston.— Bertsch (1970) described Chromodoris baumanni from two specimens collect- ed from Isla San Francisco and Isla Cerralvo, Baja California, México. Subsequent authors have D i i GOSLINER ET AL.: TROPICAL EASTERN PACIFIC CHROMODORIDIDAE Sy) FiGure 4. Glossodoris baumanni (Bertsch, 1970), scannning electron micrographs, (CASIZ 088203), Islas Secas, Gulf of Chiriqui, Panama. A. Rachis and inner lateral teeth, scale bar = 10 um. B. Midlateral teeth from the central portion of half-row, scale bar = 10 um. C. Outer lateral teeth, scale bar = 10 um. D. Jaw rodlets, scale bar = 10 tm. Ni ffl identified additional material from other eastern Pacific locali- ties from mainland México and the Galapagos Islands (Sphon and Mulliner 1972; Bertsch 1978a; Debelius 1996). More recently, Ortea et al. (1992) attributed specimens of this species from the Galapagos to Noumea haliclona (Burn, 1957). Noumea haliclona is a variably colored species that may be either pink or yellow in its body coloration and a few darker pink le IGURE 5) Glossodoris baumanni (Bertsch, 1970) A. Dorsal view of juve- nile specimen showing distribution of mantle glands, g = mantle gland, scale euer (Rudman 1983). Its bar = 2 mm.. B. Reproductive system of mature specimen, am = ampulla; be thinophores and gill are the same = bursa copulatrix; ej = ejaculatory duct; fz = female gland mass; p = penis; color as the body. This species iS pr =prostate; rs = receptaculum seminis; vg = vestibular gland, scale bar = 3 endemic to temperate southeast- ™"- erm Australia. In contrast, 596 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 32 Chromodoris baumanni has more numerous, smaller red pigment spots and the rhinophores and gill branches are tipped with purple pigment. Internally, N. haliclona has a much broader innermost lateral teeth than C. baumanni, with much shorter jaw rodlets. Noumea haliclona has a digitate vestibular gland (Rudman 1984), while that of C. baumanni is simple. The anatomy of specimens of C. baumanni examined here is entirely consistent with material described by Ortea et al. (1992) from the Galapagos Islands, with one notable exception. The reproductive system of their speci- men has a small female gland mass, indicating that their specimen was not sexually mature. There is little question that the specimens that Ortea et al. (1992) attributed to N. haliclona are in fact C. baumanni. Rudman (1984) noted that Glossodoris was characterized by having a convoluted mantle mar- gin, spirally arranged gill branches on either side of the gill, a gill that vibrates rhythmically, a radu- lar ribbon with numerous rows that is much longer than wide and an elongate, thin vaginal duct. He also suggested that species of Chromodoris have a straight or temporarily convoluted mantle margin, a gill without spirally arranged branches, a more squarely-shaped radular ribbon and a short muscular vagina. The only species of Chromodoris that is known to vibrate its gill is C. vibra- ta (Pease, 1860) (Bertsch and Johnson 1981). Specimens of Chromodoris baumanni have several characteristics that are similar to species attributed to Glossodoris. For instance, specimens of this species have a permanently, slightly con- voluted mantle margin. Also, juvenile specimens of C. baumanni have fewer gill branches that do not form a spiral (Fig. 1C), whereas mature specimens have well-developed spirals (Fig. 1B). Larger specimens of C. baumanni have been observed to have a gill plume whose branches vibrate slowly (present study), although the gill moves more slowly than in members of Glossodoris. The shape of the radular ribbon of C. bawmanni is not elongate with relatively few (32-84) rows of radular teeth (Bertsch 1978a; present study), as in most species of Chromodoris. Rudman (1986) described several species of Glossodoris that had a shorter radular ribbon with fewer rows of teeth than had previously been observed. Bertsch and Gosliner (1989) also documented the presence of relatively few radular rows in Glossodoris poliahu Bertsch and Gosliner, 1989 and G tomsmithi Bertsch and Gosliner, 1989. The vaginal duct of C. bawmanni 1s thin and elongate as in species of Glossodoris (Rudman, 1984). Based on the fact that most of the anatomical details of C. baumanni are consistent with those described for species of Glossodoris, this species is placed in Glossodoris. Detailed phylogenetic studies are needed to determine monophyletic groups within the Chromodorididae. Hypselodoris is the only chromodorid taxon whose monophyly has been tested with a study of its species-level phylogeny (Gosliner and Johnson 1999). The hypotheses of relationships within the Chromodorididae proposed by Rudman (1984) and Gosliner and Johnson (1999) require addition- al detailed study and phylogenetic analysis to further define monophyletic groups within the high- ly diverse chromodorid nudibranchs. ACKNOWLEDGMENTS The field work in Costa Rica was organized by the Instituto Nacional de Biodiversidad of Costa Rica, generously supported by the program “Development of biodiversity knowledge and sustainable uses in Costa Rica” of the government of The Netherlands and facilitated by the Ministerio de Ambiente y Energia of Costa Rica. We especially wish to thank our colleague Yolanda Camacho Garcia, who coordinated all of the field research in Costa Rica and the dedicat- ed staff of the Department of Malacology at INBio (Socorro Avila and Alcides Berrocal) who assisted in those efforts. The field work in the Galapagos Islands was made possible by the gen- erosity of Paul Humann, John McCosker and the owners and crew of the M/Y Mistral. We thank GOSLINER ET AL.: TROPICAL EASTERN PACIFIC CHROMODORIDIDAE Soy the staff of the Parque Nacional de Galapagos and Charles Darwin Research Station for supporting these research activities. Additional support was provided by National Science Foundation through the PEET grant DEB-9978155, “Phylogenetic systematics of dorid nudibranchs,” to the senior and junior authors. LITERATURE CITED BertscH, H. 1970. Opisthobranchs from Isla San Francisco, Gulf of California, with the description of a new species. Contributions in Science, Santa Barbara Museum of Natural History 2:\—16. BertscH, H. 1977. The Chromodoridinae nudibranchs from the Pacific coast of America— Part I. Investigative methods and supra-specific taxonomy. The Veliger 20(2):107—118. BertscH, H. 1978a. The Chromodoridinae nudibranchs from the Pacific coast of America.— Part II. The genus Chromodoris. The Veliger 20(4):307-327. BertscH, H. 1978b. The Chromodoridinae nudibranchs from the Pacific coast of America.— Part III. The Genera Chromolaichma and Mexichromis. The Veliger 21(1):70-86. BertscH, H. 1978c. The Chromodoridinae nudibranchs from the Pacific coast of America.— Part IV. The genus Hypselodoris. The Veliger 21(2):236—250. BertscH, H., AND T.M. GOSLINER 1989. Chromodorid nudibranchs from the Hawaiian Islands. The Veliger 32(3):247-265. BeERTSCH, H., AND S. JOHNSON 1981. Hawaiian Nudibranchs: A Guide for SCUBA Divers, Snorkelers, Tidepoolers, and Aquarists. Oriental Publishing Co., Hawaii. 112 pp. BOUCHET, P., AND J. ORTEA. (1980) Quelques Chromodorididae bleus (Mollusca, Gastropoda, Nudibranchiata) de |’Atlantique oriental. Annals Institute Oceanographique, Paris 56(2):117-—125. DeBeLtus, H. 1996. Nudibranchs and Sea Snails. Indo-Pacific Field Guide, IKAN — Unterwasserarchiv. 321 pp. GOSLINER, TM., AND H. BeRTSCH. 1988. A new species of Mexichromis (Nudibranchia: Chromodorididae) from the eastern Pacific. The Veliger 31 (1/2):127-134. GOSLINER, T.M., AND R.F. JOHNSON. 1999. Phylogeny of Hypselodoris (Nudibranchia: Chromodorididae) with a review of the monophyletic clade of Indo-Pacific species, including descriptions of twelve new species. Zoological Journal of the Linnean Society 125:1—114. OrTEA, J.A. 1988. Moluscos opistobranquios del Archipielago de Cabo Verde: Chromodorididae. Publicagoes Ocasionais da Sociedade Portugesa de Malacologia 11:1—16. Ortea, J.A., J.J. BACALLADO, AND A. VALDES. 1992. Resultados cientificos del Proyecto Galapagos: patrimo- nio de la humanidad. No. 1 Chromodorididae (Mollusca: Nudibranchia). Resultados Cientificos del Proyecto Galapagos TF MC 0-1:31—70. OrteA, J.A., A. VALDES, AND J.C. GARCiA-GOMEZ. 1996 Revision de las especies atlanticas de la familia Chromodorididae (Mollusca: Nudibranchia) del grupo cromatico azul. [Review of the Atlantic species of the family Chromodorididae (Mollusca: Nudibranchia) of the blue chromatic group.] Avicennia 1996(Suppl. 1):1—165. RUDMAN, W.B. 1983. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris splendida, C. aspersa and Hypselodoris placida colour groups. Zoological Journal of the Linnean Society 78:105—173 RUDMAN, W.B. 1984. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: a review of the genera. Zoological Journal of the Linnean Society 81:115-273. RUDMAN, W.B. 1986. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: The genus Glossodoris Ehrenberg (= Casella, H. & A. Adams). Zoological Journal of the Linnean Society 86(2): 101-184. SPHON, G., AND D. MULLINER. 1972. A preliminary list of known opisthobranchs from the Galapagos Islands collected by the Ameripagos Expedition. The Veliger 15(2):147-152. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 33, pp. 598-612, 4 figs. December 30, 2004 A New Species of Oscarella (Demospongiae: Plakinidae) from California Guilherme Muricy! and John S. Pearse? ! Departamento de Invertebrados — Museu Nacional/UF RJ. Quinta da Boa Vista s/no, Sao Cristovado 20940-040 Rio de Janeiro, Brazil. Email: muricy@aca.uftj.br.; 2 Joseph M. Long Marine Laboratory—Institute of Marine Sciences, University of California, Santa Cruz. 100 Shaffer Road, Santa Cruz, CA 95060, USA. Email: pearse@biology.ucsc.edu. We describe a new species of Oscarella Vosmaer, 1877 (Porifera: Homosclerophor- ida) from a rocky coast and marine aquariums in central California and analyze its spatial distribution and temporal fluctuations to determine if it is a non-indigenous species. We also compare the efficiency of two different fixatives, Bouin’s solution and glutaraldehyde in seawater, and found that glutaraldehyde is better for compar- isons of anatomical and cytological characters that are important for species identi- fication. This is the first record of the genus Oscarella from the Eastern Pacific. The new species is characterized by its light brown to rusty orange color, undulated appear- ance, bumpy, microlobate surface, soft, slimy consistency, two particular kinds of cells with inclusions, and occurrence in the intertidal high zone. It sometimes thrives in laboratory conditions, indicating that predators may limit its vertical distribution on the shore. The new species of Oscarella is clearly different in external morpholo- gy and cell content from all previously described species of Oscarella, thus giving low support to the hypothesis that it is an invasive species. Instead, it appears to have been previously overlooked because it is inconspicuous and relatively uncommon in the field, and because it could not be distinguished from Halisarca sacra de Laubenfels, 1930 or Halisarca sp. in keys to the West Coast sponge fauna. We pres- ent how it can be distinguished from Halisarca, and discuss the present knowledge of that genus in the northeast Pacific in comparison to Oscarella. KEY worbDs: California, Halisarca, Homosclerophorida, intertidal, Oscarella, Porifera, sponge. Marine invasive species often appear first in aquariums, to be later established in the field. A classic marine example is the alga Cau/erpa taxifolia in the Mediterranean, but there are many oth- ers (e.g., Ribera and Boudouresque 1995). In the mid 1980s, an unidentified slimy sponge without skeleton (either spongin or spicules) appeared growing on submerged rocks and the sides of aquar- iums and sea tables at the Joseph M. Long Marine Laboratory of the University of California, Santa Cruz, on the northern edge of Monterey Bay. At first, it was taken to be a species of Halisarca Johnston, 1842, the only similar organism in the main key for intertidal sponges of central California (Hartman 1975). However, in 1990, M. Cristina Diaz examined a piece of the sponge with scanning electron microscopy and identified it as a member of the genus Oscarella Vosmaer, 1887 (Demospongiae: Plakinidae) (Diaz and van Soest 1994), a genus mainly known in the Mediterranean and previously unknown from the eastern Pacific. The sponge was noticed in the 598 ——— MURICY AND PEARSE: NEW SPECIES OF OSCARELLA FROM CALIFORNIA 599 late 1980s in several of the exhibit tanks of the Monterey Bay Aquarium, which is located on the southern side of Monterey Bay, and at Long Marine Laboratory, where it continues to be present (JSP, pers. obs.). It also became established in the exhibit tanks and teaching labs of the Seymour Marine Discovery Center, adjacent to Long Marine Laboratory, shortly after it opened in 2000. After seeing the sponge at these facilitates in early 2004, Scott Nichols (pers. commun.) found the same sponge in an aquarium at the Bodega Marine Laboratory of the University of California, Davis. In addition to the question of its identity, it was puzzling that the sponge had not been seen in the field, especially since field biologists have been working in the rocky intertidal of the Monterey Bay area of central California throughout the 20th century. Moreover, de Laubenfels (1932) had extensively monographed the sponges of California. Finally, in September 2002, one of us (JSP) found the sponge growing on the sides and undersurfaces of intertidal Monterey Bay Pacific Grove Monterey 5 Monterey Peninsula FIGURE |. Map of Carmel Bay and the Monterey Peninsula show- ing the location of 13 intertidal sites examined between September 2002 and May 2004; underlined numbers indicate sites where spec- rocks on the northwest corner of Carmel Point, in Carmel Bay, central California (Figs. 1, 2A). Most species of Oscarella have been described from the Mediter- ranean, but there are also two Indo- Pacific species and a few doubtful Australian and South African records (e.g., Boury-Esnault et al. 1992; Muricy et al. 1996; Muricy and Diaz 2002; Bergquist and Kelly 2004). Moreover, many species of Oscarella occur in Brazil and the Caribbean (GM, pers. obs.), but to a large extent they are uncom- mon and most have yet to be described. The same might be true for California, and this is likely an undescribed species indigenous to this area. However, with no previous records of the genus Oscarella along the Pacific coast of North America, the question arises whether it could have been introduced from elsewhere, possibly via disposal of aquarium water. If it is non-indigenous, it is not a very attractive addition to the Californian fauna: it is a tan, bumpy, slime-like sponge that overgrows other organisms. If it is conspecif- ic with any of the Mediterranean species of Oscarella, that would support the invasion hypothesis, and there is a risk that it could disrupt natural biotas. Our goals were to determine whether this skeleton-less sponge belongs to Oscarella or Halisarca, and whether it is a new species, either indigenous but unnoticed before or introduced but undescribed, or a known species introduced from somewhere else. We also analyzed the spa- tio-temporal distribution of the species in the Monterey Bay region, and compared the efficiency of two different fixatives, Bouin’s solution and glutaraldehyde in seawater, for the histology and taxonomy of these skeleton-less sponges. imens of O. carmela sp. nov. were found. 1. Whaler’s Cove, Point " Lobos State Park; 2. Carmel Point, southwest corner; 3. Carmel Point, northwest corner; 4. Carmel Point, north side, between site 3 and Carmel Beach; 5. Arrowhead Point on east side of Stillwater Cove; 6. Asilomar State Beach State Park, south end next to Spanish Bay Beach; 7. The Great Tide Pool, south of Point Pinos; 8. Point Pinos; 9. Lucas Point, between Coral Street and Esplande Park; 10. Lovers Point, east side, both sides of Lovers Point Beach; 11. Hopkins Marine Station (Cabrillo Point); 12. Monterey Bay Aquarium (Point Alones), underneath building; 13. Monterey Harbor, between the Coast Guard Pier and Fisherman’s Wharf. Square on inserted map of California outlines the Monterey Peninsula and Carmel Bay. Pebble Beach Carmel Carmel Bay Point Lobos MATERIALS AND METHODS Specimens were photographed with an Olympus Camedia 3040 digital camera and collected in November 2002 and May 2003 from rocks in a high intertidal pool at Carmel Point in Carmel 600 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 33 Bay, central California (Figs. 1, 2A). Additional specimens were collected from tanks in Long Marine Laboratory and the Seymour Marine Discovery Center of the University of California, Santa Cruz (UCSC). Small pieces of the sponges were collected with a pocketknife and immedi- ately fixed in Bouin’s fixative (75 ml saturated aqueous solution of picric acid: 25 ml 37-40% for- malin (formaldehyde): 5 ml glacial acetic acid) or freshly prepared 2.5% glutaraldehyde in seawa- ter (25% glutaraldehyde diluted and buffered with seawater). Bouin’s-fixed samples were embed- ded in paraffin for thick sections and epon resin for semithin sections. Glutaraldehyde-fixed sam- ples were post-fixed in 1% osmium tetroxide and embedded in epon resin for semithin and ultra- thin sections. Thick and semithin sections were stained with toluidin blue or acid fucsin, observed in light microscopy (LM), and photographed with a Nikon Coolpix 4500 digital camera. Ultrathin sections were stained with uranyl acetate, contrasted with lead citrate and observed in a Zeiss M900 transmission electron microscope (TEM). SYSTEMATICS Phylum Porifera Grant, 1836 Class Demospongiae Sollas, 1885 Subclass Homoscleromorpha Lévi, 1973 Order Homosclerophorida Dendy, 1905 Family Plakinidae Schulze, 1880 Genus Oscarella Vosmaer, 1887 Type SPECIES: Halisarca lobularis Schmidt, 1862 (by monotypy). [Oscaria] Vosmaer, 1881: 163 (preocc. by Oscaria Gray, 1873 — Reptilia); Oscarella Vosmaer, 1884: pl. 8 (explanation); 1887: 326 (nom. nov. for Oscaria Vosmaer). Octavella Tuzet and Paris, 1964: 88. DIAGNosis (Muricy and Diaz 2002).— Plakinidae without skeleton, with thinly encrusting to lobate shape. Thin ectosome (< 100 um), often limited to pinacoderm; true cortex absent. Mesohyl poorly developed, with a proportion of mesohyl to chambers varying from 0.5:1 to 1.2:1. The aquiferous system has a sylleibid organization, with spherical, eurypylous choanocyte chambers uniformly arranged around large, regular exhalant canals, and a large basal cavity. Oscarella carmela Muricy and Pearse, sp. nov. (Figs. 2-4) DIAGNOosIs.— Intertidal or aquarium-dwelling Oscarella, light brown to tan or rusty orange in color, with bumpy, microlobate surface, wavy appearance, soft, slimy consistency, and two partic- ular kinds of cells with inclusions (types | and 2). MATERIAL EXAMINED.— (12, all from central California, USA) (MNRJ, Museu Nacional, Universidade Federal do Rio de Janeiro; CASIZ, California Academy of Sciences; SBMNH, Santa Barbara Museum of Natural History) HOLotyPE: MNRJ 8033, Carmel Point, Carmel Bay, intertidal, J. Pearse coll., 08 May 2003. PARATYPES: MNRJ 8032, 07 November 2002, CASIZ 168925, 08 May 2003, Carmel Point, Carmel Bay, central California, intertidal, J. Pearse coll.; MNRJ 8034, 8035, CASIZ 168924, 06 November 2002, SBMNH 354351 (three specimens) 29 July 2004, tanks in Long Marine Laboratory, UCSC, Santa Cruz, J. Pearse coll.; MNRJ 8031 (three specimens), 06 November 2002, teaching lab of the Seymour Marine Discovery Center, UCSC, Santa Cruz, J. Pearse coll. DESCRIPTION.— Thinly encrusting, irregular sponges, generally light brown, up to 20-30 cm MURICY AND PEARSE: NEW SPECIES OF OSCARELLA FROM CALIFORNIA 601 in diameter, with extremely soft, slimy consistency. Thickness, color, and surface irregularities variable. Most specimens tan in color, 2—3 mm thick, and lumpy or undulated with conspicuous channels and oscula (Fig. 2B—F). Some specimens gray-tan and smoother, or pale brown to rusty orange, with very bumpy sur- faces and orange blotches. These different morphologies could be ecophenotypes or different stages of growth. In some cases, one morphology graded into another in what appeared to be the same individual. On the glass walls of aquariums, the sponge is nearly transparent and clusters of choanocyte chambers and spheri- cal embryos can be seen inside. The sponge is easy to peel off smooth surfaces. Under low magnification (60x) the colored spherical choanocyte chambers FIGURE 2. Oscarella carmela sp. nov. A. Habitat at the northwest corner of can be seen in living material, Carmel Point, with boulders in the intertidal zone. B-D. Three colonies on the and often, the colorless spherical upturned sides of boulders exposed during low tide; colonies of the bryozoan hollow embrvos. Eurystomela bilabiata partially overgrown in B, polyps of the corallimorphar- ‘ : ian Corynactis californica partially surrounded in C (scale bar = 2 cm; coin in ANATOMY.— Because glu- B isa U.S. quarter). E. Colony growing up to the water line on the side of a 8 taraldehyde provided better fixa- cm high plastic box in an aquarium in Long Marine Lab. F. Colony growing tion for fine structure than onarock in an aquarium in the Seymour Marine Discovery Center; the sponge is covered with seawater (scale bar = 2 cm). Bouin’s solution (see below), the description of anatomy and cytology is based on glutaraldehyde-fixed material. Choanocyte chambers ovoid to spherical eurypilous, 25—65 um in diameter, organized around large exhalant canals (Fig. 3A—G). Mesohy] between the chambers very thin (S—10 um thick). Proportion of mesohyl to chamber volume less than 0.5:1.0. Ectosome thin, 5-10 um thick. Basal cavities large, separated by septa without choanocyte chambers, 10 um thick. Cinctoblastula larvae characteristic of the family Plakinidae, 50-300 um in diameter (Fig. 3H). CyToLoGy.— Choanocytes ovoid to pyramidal, irregular, 2—S um in diameter by 3—5 um high (Fig. 4A—C). Nucleus basal or apical, 1.5—2.2 um in diameter. Cytoplasm with up to eight phago- somes, 0.5—1.5 um wide. Choanocyte collars with 25—33 microvilli. Apopylar cells not observed. Pinacocytes flat or elongated to ovoid, 5-10 um long by 0.5—2 um wide, flagellated, anchored in the mesohyl by short pseudopodia (Fig. 4C—D). Nucleus ovoid, 1—2 um in diameter, nucleolated. Cyto-plasm with numerous small phagosomes, 0.3—0.7 um, and often also one larger vacuole with clear contents. Choanoderm and pinacoderm lined by a basement membrane-like structure, which is a continuous, 5—10 nm thick layer of condensed collagen fibrils in the mesohyl closely adjacent 602 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 33 to the base of the cells. Collagen deposition in the mesohyl het- erogeneous (Fig. 4D). As seen in both LM and TEM, ameboid cells of the mesohyl include archaeocytes and two types of cells with inclusions. Archaeo- cytes roughly ovoid, irregular, 5.0 x 3.5 um, with abundant pseudopodia (Fig. 4E, F). Cyto- plasm filled with phagosomes 0.3—1.5 um wide containing food particles, including bacteria, in various stages of digestion. Nucleus ovoid or irregular, 1.0-2.5 um, nucleolated. Type 1 cells with inclusions ovoid, 5.0 x 3.0 um, irregular, with short pseudopodia (Fig. 4E). Nucleus 2 um in diameter, ovoid or com- pressed by the abundant cyto- plasmic inclusions. Cytoplasm filled with inclusions of two kinds, 0.2-1.0 um wide, one kind with relatively dark and homogeneous contents and the other with clear, heterogeneous, filamentous contents. Type 2 cells with inclusions have irregu- lar shape, 5.0x3.0 wm, rarely with short pseudopodia (Fig. 4F, G). Cytoplasm with 5-15, 0.5—2.0 um wide inclusions with homogeneous contents, and 3—8 larger vacuoles, 1—4 um in diam- eter, with clear and filamentous contents. Nucleus |.2—2.5 um in diameter, ovoid or compressed |. ne oe 2 eee FIGURE 3. Histology of Oscarella carmela sp. nov. (LM). (A=B) transverse section showing the arrangement of choanocyte chambers around exhalant canals and the large basal cavities (scale bar = 100 tum); (C—E) choanosome (scale bars = 50 um); (F—G) choanocyte chambers (scale bars = 25 pm); (H) embryos (scale bar = 100 um). Fixation: A, G and H, Bouin’s fixative; B-F, glutaraldehyde / osmium tetroxide. by abundant cytoplasmic inclusions. This cell type appears to be involved in the secretion of ground substances of the intercellular matrix, because it is often seen liberating the contents of its larger, clear vacuoles in the mesohy! (Fig. 4G). Endobiont bacteria of two kinds (Fig. 4H): type B1 most common, ovoid, 1.0—1.5 by 0.6—1.0 um, with a wrinkled cell wall, a clear and filamentous nuclear region and a darker cytoplasmic region; type B2 rod-like, 1.0 x 0.2 um, with darker and rather homogeneous contents, and without a clear distinction between nuclear and cytoplasmic regions. ECOLoGy.— Specimens of Oscarella carmela occur as thin sheets on the sides of granite out- crops and on the edges and undersurface of boulders in high-zone tide pools of Carmel Bay and the MURICY AND PEARSE: NEW SPECIES OF OSCARELLA FROM CALIFORNIA 603 Monterey Peninsula (Fig. 1, Fig 2A). These sponges apparently need to remain continually immersed, but do well in the variable environment of high- tide pools. The tops of the boul- ders in this habitat supported typ- ical high-zone algae (Si/vetia compressa, Fucus gardneri, Endocladia muricata, Mazzaella affinis, _ Mastocarpus _ spp.) whereas the sides of boulders immersed in the tide pools often were covered with the red alga, Prionitis lanceolata, and en- crusting and upright coralline algae. The undersurfaces of the boulders supported a rich variety of sessile and encrusting ani- mals, e.g., red poecilosclerid sponges, cnidarians (Corynactis californica and Plumularia sp.), bryozoans (Eurystomella bilabi- ata and others), polychaetes (Spirorbis sp.), gastropods (Petaloconchus montereyensis), and ascidians (Aplidium califor- nicum and Distalpia occidental- is). Often individuals of O. carmela were overgrowing other encrusting organisms, especially the bryozoan E. bilabiata (Fig. 2 B). Surprisingly, O. carmela has not been found in mid or low zones, although thorough search- es down to the low zone were made at all the sites where it was found in the high zone. Specimens of O. carmela often grew profusely in aquari- 7) ee FIGURE 4. Cytology of Oscarella carmela sp. nov. (TEM). (A) section through a choanocyte chamber; (B) choanocytes; (C) prosopyle of a choanocyte chamber showing choanocytes (top) and endopinacocytes (bot- tom); (D) pinacocytes; (E) type | cell with inclusions (left), two archaeocytes (middle and right), and the loose junction between two pinacocytes (top right); F, type 2 cell with inclusions (“spherulous-vacuolar cell”, left) and archaeocyte phagocytizing a bacterium (right); (G) type 2 cell with inclusions (“spherulous- vacuolar cell”); (H) endobiont bacteria types 1 (B1) and 2 (B2). Scale bars: A-G =2 um; H=1 um. ums and sea tables of the Long Marine Laboratory—Seymour Marine Discovery Center complex of the University of California, Santa Cruz, covering submerged rocks and plastic trays right up to the edge of the water line (Fig. 2E). In quiet aquarium water, the sponges developed conspicuous folds with bulging excurrent canals converging on large oscula (Fig. 2F). Asteroids (Asterina miniata, Dermasterias imbricata, Pisaster ochraceus) and abalones (Haliotis rufescens) in some of the sea tables and aquariums did not appear to disturb the sponges. Specimens of O. carmela have been noticed in several of the exhibits of the Monterey Bay 604 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 33 Aquarium for many years, especially in the “Crevice Dwellers” tank, where they are particularly abundant during the spring (Veronica Franklin, pers. commun.). They grow on the artificial rock backing of the tank and on the shells of rock scallops (Crassodoma gigantea). The specimens have only been seen in tanks containing local biota and not in tanks with exotic species (e.g., the tank with Mediterranean cuttlefish); all seawater is filtered both before and after it leaves tanks with exotic species. DISTRIBUTION.— Oscarella carmela occurs in aquariums and sea tables of the Long Marine Laboratory-Seymour Marine Discovery Center complex, the Monterey Bay Aquarium, and in an aquarium of the Bodega Marine Laboratory of the University of California, Davis, approximately 150 km north of Monterey Bay (Scott Nichols, pers. commun.). However, specimens of O. carmela are not ubiquitous in marine laboratories and public aquariums along the California coast. For instance, it was not found in any of the aquarium tanks in the Steinhart Aquarium of the California Academy of Sciences in San Francisco nor in aquariums at the Long Beach Aquarium in southern California. Moreover, it has not been found in any of the tanks with running seawater at Stanford University’s Hopkins Marine Station, which is adjacent to and receives all its seawater from the Monterey Bay Aquarium. Specimens of O. carmela were first found in the field by JSP at Carmel Point on September 2002 and sampled on November 2002. The sponges were on boulders in high intertidal pools directly west of the intersection of Scenic Road and Ocean View Avenue, and to the north of the “Sunburst” house on the ocean side of Scenic Road (see: http://limpets.noaa.gov/monitoring/ rockylntertidal/sites/mbnms/carmelpoint.himl). The site at Carmel Point was visited again on January 2003, several weeks after unusually heavy storms. Many of the boulders in the high tide pools had been disturbed and rearranged, but thin colonies of O. carmela were found on the under- surfaces of some of them. On April 2003, a thorough search of the boulders was made throughout the area from the high zone, through the mid zone dominated by the alga Mazzaella flaccida, to well into the low zone dominated by surfgrass (Phyllospadix spp.). Three out of about 20 boulders at the original high-zone site had specimens of O. carmela growing on their undersurfaces. Only one small colony of O. carmela was found elsewhere, on the underside of a boulder in a high-zone shallow tide pool west of the “Sunburst” house, approximately 30 m southwest of the area where the first individuals were seen. The site at the NW corner of Carmel Point was examined six other times: May, August, and December 2003, and January, February, and April 2004. Specimens were found under boulders in the original pools on all these dates, and when looked for again on August 2003 and January 2004, small individuals at the site west of the “Sunburst” house were also found. Between April 2003 and May 2004, thorough searches were made of 12 additional intertidal sites during low tides around Carmel Bay and the Monterey Peninsula (Fig. 1). One or more spec- imens were found at three of these sites: (1) Whaler’s Cove at Point Lobos, rip-rap boulders adja- cent to the boat launch, February and April 2004, three colonies total; (2) Lovers Point Beach, Pacific Grove, east of the beach at base of cliff below the intersection of Grand Avenue and Ocean View Boulevard, 6 February, 15 February, and 7 March 2004 (three colonies total), but not found on 21 February 2004, nor in boulder field just SE of Lovers Point on 7 March 2004; (3) Hopkins Marine Station, directly north of shop building, March, May, and July 2004, four colonies; but none found during thorough searches on November 2002 and May 2003. At all of these sites, specimens of O. carmela were found on the undersurfaces of partially submerged boulders in high-zone pools, similar to where they were found at Carmel Point. Similar habitats occurred at the other eight sites examined, but no individuals of O. carmela were found. Particular attention was given to the area under and around the Monterey Bay Aquarium, including the outflow channels from the facility. —_—_e ~ 7 MURICY AND PEARSE: NEW SPECIES OF OSCARELLA FROM CALIFORNIA 605 which would likely carry propagules from the exhibit tanks. In addition to the Monterey Peninsula area, six sites with boulders in the high zone between Monterey and San Francisco to the north were examined without finding specimens of O. carmela. These sites were: (1) James V. Fitzgerald Marine Reserve, Moss Beach; (2) Pigeon Point, south side; (3) Point Ano Nuevo, south side; (4) Terrace Point adjacent to the outflow from the Long Marine Laboratory—Seymour Marine Discovery Center facilities; (5) Point Santa Cruz (Lighthouse Point), east side; (6) Elkhorn Slough, northwest of Highway | bridge. Although the type locality of another slime sponge, Halisarca sacra de Laubenfels, 1930, was in Elkhorn Slough close to this site, neither it nor O. carmela was found there. On the other hand, after seeing specimens of O. carmela in aquariums at Long Marine Laboratory, J.H.R. Goddard (pers. commun.) told us he had seen it at Cape Arago, Oregon, where it was distinctly different from specimens of Halisarca found there (see below). A photograph he shared with us, taken at North Cove, Cape Arago on 22 May 1985, shows a thin, orange-tan spec- imen, which appears to be identical with specimens of O. carmela around the Monterey Peninsula. In addition, W.C. Austin informed us that he found a slime sponge with spherical choanocyte cham- bers characteristic of Oscarella, and distinct from Halisarca sp., in the Vancouver Aquarium in British Columbia (pers. commun., | June 2004). EtyMo._oGy.— The name carmela derives from the site where this species was first found in the field, Carmel Bay in Central California. DISCUSSION COMPARISON OF FIXATION METHODS.— Proper fixation is important for the identification of Oscarella species. These sponges are very sensitive, and a delay of only a few minutes before plac- ing them in fixative may make them stop filtering, start contracting, and their tissues and cells loose their normal aspect. Bouin’s solution, 4% formalin or 70% ethanol preserve canals and choanocyte chambers well enough to allow their recognition as belonging to Oscarella and not to, for instance, Halisarca (see below). However, these solutions rarely preserve the natural shape of chambers and cells or allow observation of cell contents, which are important taxonomic characters at the species level. We suggest that the best way to fix these sponges for both semithin (LM) and ultrathin sec- tions (TEM) is to put small pieces of the sponge into freshly prepared 2.5% glutaraldehyde in sea- water immediately after collection, preferably while still underwater. In the laboratory, the fixative should be changed to ensure it has the desired concentration. After 24 hrs, the pieces should be post-fixed with osmium tetroxide and then dehydrated until the ethanol concentration is 70%—100%. Following this treatment, they can be stored in ethanol for long periods before mak- ing sections for observation in LM or TEM. Boury-Esnault et al. (1984) compared the results of fixation in four different fixatives for TEM and SEM (scanning electron microscopy) in Oscarella lobularis and Corticium candelabrum, with varying osmolarity (1100 —1440 mosm) and different concentrations of glutaraldehyde (0-3.0 %), seawater, cacodylate buffer (0O—0.4 M), and NaCl (0— 7%). They concluded that their fixative C (2.5% glutaraldehyde in 0.4 M cacodylate buffer, pH 7.4, and seawater 4 vol.: 5 vol., 1120 mosm; rinse in seawater 1100 mosm) was the best fixative for homosclerophorid sponges. TAXONOMY OF OSCARELLA.— The identification of Oscarella at the species level is difficult. The species are homogeneous in most of the few taxonomic characters available: encrusting shape, sylleibid aquiferous system organization, choanocyte chambers spherical-ovoid with 30-60 tm in diameter, cinctoblastula larvae, thin ectosome, etc. (Boury-Esnault et al. 1984, 1992; Muricy et al. 1996: Muricy and Diaz 2002). The differences among species are mostly in external traits: color, 606 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 33 consistency, and aspect of the surface. These characters, however, must be used with care, inas- much as they can vary within a single species. Specimens of Oscarella lobularis (Schmidt, 1862) and O. tuberculata (Schmidt, 1868) from the Mediterranean, for example, may appear in several different colors: white, yellow, green, red, violet, and blue. However, each species is constant and unique in its cell contents (Boury-Esnault et al. 1992). Other species appear in only a single color such as O. viridis Muricy et al., 1996 (green) and O. microlobata Muricy et al., 1996 (brown). The consistency seems to be more constant within species, but it is also more subjective to describe. Finally, the aspect of the surface (smooth, microlobate) is also helpful, but one cannot separate all species on that basis alone. Oscarella carmela varies from light brown to tan or dull orange color, and from a smooth to microlobate surface; when out of the water, especially, the microlobes con- tract and the surface becomes smooth. The new species is similar in color and surface aspect only to Oscarella microlobata. Because species of Oscarella have no skeleton, and histological characters are homogeneous among different species, less usual characters, such as the types of cells with inclusions present, are critical for species identification. Cells with inclusions are special sponge cells with different cytoplasmic inclusions, most of which have unknown functions (e.g., Simpson 1984; Muricy and Diaz 2002). They are abundant and diverse in species of Oscarella, and in the Mediterranean each species has a particular set of cells with inclusions that are useful characters for species identifica- tion (Muricy et al. 1996). It is, therefore, important that the specimens are fixed in glutaraldehyde, so they can be observed under TEM to identify cells with inclusions accurately. Cytological char- acters are also useful for the identification of other sponges without skeleton, such as Halisarca (cf. Vacelet and Donadey 1987). Molecular methods (e.g., allozyme electrophoresis, DNA sequencing) may also help in the definition of species boundaries in Oscarella. The new species differs from Oscarella microlobata and all other Mediterranean Oscarella species in its cell contents, which are simple, with only two kinds of cells with inclusions: spheru- lous cells with two different inclusions, osmiophilic and filamentous, and vacuolar-spherulous cells (Fig. 4E-G). In contrast, O. microlobata has one type of vacuolar cells and three types of cells with inclusions, all different from those of O. carmela: spherulous cells with granular inclusions, spherulous cells with paracrystalline inclusions, and single-inclusion cells (Muricy et al. 1996). Besides being green, O. viridis differs from O. carmela by having two exclusive types of spheru- lous cells, called crescent-shaped cells and microgranular inclusions cells (Muricy et al. 1996). Oscarella imperialis is yellowish-white and has four types of cells with inclusions, two of which with different paracrystalline inclusions and two with large vacuoles with clear or osmiophiolic inclusions (Muricy et al. 1996); none of these cell types occur in O. carmela. Oscarella tubercula- fa is distinguished by the large groups of turgid vacuolar cells that fill most of the choanosome, and O. lobularis has two types of vacuolar cells, one of which is exclusive, and no spherulous cells (Boury-Esnault et al. 1992). The new species, therefore, has a set of cells with inclusions clearly distinct from those of all Mediterranean species of the genus, which are the only other species of Oscarella studied at TEM so far. Oscarella carmela is distinct also in its intertidal habitat: all known Mediterranean species of Oscarel/la occur in caves or in subtidal rocky shores (Boury- Esnault et al. 1992; Muricy et al. 1996). Two other species of Oscarella were recently described from the Indo-Pacific (Bergquist and Kelly 2004). These specimens were fixed in 70% ethanol, and no attempt was made to character- ize their cell composition. However, it is possible to distinguish both from the new species by their external characters: Oscarella stillans Bergquist and Kelly, 2004 forms a series of fused tubes up to 3.5 cm long, some with solid branches, and it is dark honey yellow in color. It also has a char- acteristically high collagen deposition in the mesohyl, giving it a collagenous consistency. MURICY AND PEARSE: NEW SPECIES OF OSCARELLA FROM CALIFORNIA 607 Oscarella nigraviolacea Bergquist and Kelly, 2004 differs from the new species by its dark violet, almost black color, and the oscules situated on top of papillae. According to Bergquist and Kelly (2004), other possibly valid species of Oscarella are O. tenuis Hentschel, 1909, O. membranacea Hentschel, 1909 (both from south Australia), and O. ochreacea Bergquist and Kelly, 2004 (nomen novum for O. lobularis sensu Vacelet et al. 1976) from Madagascar. However, these species must be better characterized before meaningful comparisons can be made. DISTINCTION FROM HALISARCA AND OTHER SPONGES WITHOUT SKELETONS.— With the finding of Oscarella along the California coast, there is now a need to distinguish Halisarca sp. from O. carmela, inasmuch as they belong to two superficially similar but unrelated sponge families. Halisarca sp. is the only tan colored sponge in the current standard keys for sponges in central California (Hartman 1975) and the northeast Pacific (Kozloff 1974) that has neither a collagen nor spicule skeleton. In these keys Oscarel/a would not be distinguished from Halisarca. The complete absence of a skeleton is also shared with Chondrosia Nardo, 1833, another common sponge genus, and three less common genera, Hexadella Topsent, 1896, Bajalus Lendenfeld, 1885, and Pseudo- corticium Boury-Esnault et al., 1995. However, among these genera, only Hexadella is recorded for the West Coast of North America, and it is distinguished from Halisarca sacra and O. carmela by its bright yellow color when live (Kozloff 1996). These genera can also be distinguished through examination of transverse sections to observe the shape and size of the canals, choanocyte chambers, and larvae. In Oscarella, the choanocyte chambers are ovoid to spherical, 30-60 um in diameter, eurypilous, and arranged in a sylleibid organization (the chambers are regularly arranged around the exhalant canals); the larvae of Oscarella are incubated, hollow cinctoblastulae (Fig. 3H). Such larvae are typical of the family Plakinidae (including Oscarella and Pseudocorticium; Diaz and van Soest 1994; Muricy and Diaz 2002). In Halisarca, the choanocyte chambers are elongate, up to 280 um long, tubular, ramified, with a large exhalant opening, and arranged in a leuconoid organization. The larvae are incubated, solid parenchymellae. In Chondrosia, the choanocyte chambers are spherical, 10-30 um in diam- eter, diplodal, and arranged in a leuconoid organization. There is always a thick, easily recogniza- ble collagenous cortex, absent in the other genera. Hexadella has a distinct collagenous ectosome, which provides some mechanical support in the absence of a skeleton. The choanosme is soft, with eurypilous, sac-shaped choanocyte chambers, approximately 60—80 um in diameter. Its spherulous cells are typical of the Verongida (Bergquist and Cook 2002). There are also differences in external morphology among these genera, but such traits are less reliable for identification: Chondrosia has a firm, cartilaginous consistency, and a smooth surface. Both Oscarella and Halisarca have a soft and fragile consistency, but Halisarca has an even, smooth surface in which superficial canals can be seen sometimes. The lobulated surfaces of Oscarella and Hexadella do not have superficial canals, although prominent excurrent canals can often be seen below the surface (Fig. 2F). Species of these genera vary in thickness. Those of Oscarella and Hexadella are usually thicker (5-30 mm thick) than species of Halisarca (usually less than 5 mm). On the other hand, our specimens of O. carmela were usually only 1 mm thick or less, and rarely thicker than 5 mm even when growing in quiet water in the laboratory. In contrast, W. C. Austin (unpublished manuscript, 2004) reports that specimens of Halisarca growing in quiet waters in British Columbia can be up to 1 cm thick (see also Goddard [1984, 2001]) for photo- graphs of thick specimens of Halisarca in Oregon). | STATUS OF HALISARCA ON THE WEST COAST OF THE UNITED STATES AND CANADA.— The only described species of Halisarca in the northeastern Pacific, H. sacra, is based on specimens collect- ed by Ed Ricketts in 1929 from rocks resting on a sheltered mud flat just within the former mouth of Elkhorn Slough, Monterey Bay. However, there has been some question about the validity of the 608 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 33 species (Ristau 1977); Hartman (1975) changed its designation from H. sacra de Laubenfels, 1930 in Hartman and Smith (1954) to Halisarca sp. without comment. Smecher (2004) includes H. sacra in his website but considers it to be synonymous with Halisarca dujardini Johnston, 1842. He also reports that specimens are in the British Columbia Provincial Museum (Smecher 2004), but upon inquiry he replied that the museum does not have the facilities to do proper sections (C. Smecher, pers. commun.). Similarly, Austin (1985) includes Halisarca sacra in his checklist, as does Austin et al. (1999-2001) on their website which includes a photo, but the latter with the caveat that it may be synonymous with H. dujardini Johnson 1842. Austin (unpublished manuscript, 2004) describes specimens of Halisarca collected from southeast Alaska, British Columbia, and northwest Washington and sectioned to reveal the tubular choanocyte chambers, and suggests that they are synonymous with H. sacra de Laubenfels, 1930. Goddard (1981) acknowledged P. Bergquist and W. Hartman for identification from prepared histological sections of Halisarca from Oregon, but they refrained from giving a specific name. There are no confirmed records of Halisarca in California except for the original collections in Elkhorn Slough, the type locality of H. sacra de Laubenfels, 1930 (de Laubenfels, 1932) (types in the National Museum of Natural History, K. Riitzler, pers. commun., 2004). MacGinitie (1935) mentions the species in his classic monograph on Elkhorn Slough, but apparently only on the basis of the original description; it is not mentioned at all in MacGinitie and MacGinitie (1949). In their survey of Elkhorn Slough, Nybakken et al. (1977) list H. sacra only because it was mentioned in MacGinitie (1935), and they did not find it themselves (J. Nybakken, pers. commun., 2004). Similarly, Ricketts and Calvin (1939) reported that H. sacra is known to occur in Elkhorn Slough, apparently on the basis of the original collection and description, and that report is retained unchanged in later editions of their book (Ricketts et al. 1985). The genus is not included in Bakus and Abbott’s (1980) account of Californian intertidal sponges, Thompson et al.’s (1985) list of sponges from San Diego, California, or Bakus and Green’s (1987) account of sponges in the Southern California Bight. On the other hand, the nudibranch Hallaxa chani, which apparently feeds exclusively on Halisarca sp., (Goddard 1981, 1984, 2001), ranges from southeast Alaska to central California (Gosliner and Johnson 1994) and into the Channel Islands (Goddard, pers. commun., 2004). Consequently, Halisarca sp. probably also occurs in California but, if so, it would likely be seen rarely because it is so obscure and perhaps kept at low densities by predation by H. chani and a second nudibranch predator, Cadlina modesta (Goddard 1984, 2001). The Santa Barbara Museum of Natural History has two ethanol-preserved specimens collected from southern California that were identified by R. Given as H. sacra (SBMNH 353403, Moray Reef, Corona del Mar, Orange County, 50 ft, 26 March 1958; SBMNH 353402, Scorpion Anchorage, Santa Cruz Island, Santa Barbara County, 30-40 ft, 24 March 1963). These specimens were examined by one of us (JSP). Although histological sections were not made to reveal the choanocyte chambers, thick sections indicated that the specimens are definitely not O. carmela, based on thickness, surface texture, dis- tinct stiff ectosome, thick collenchymatous choanosome, and large excurrent canals. They almost certainly are specimens of Halisarca, or perhaps of Hexadella. Halisarca sp. 1s included in species lists prepared in the early 1970s during undergraduate stu- dent surveys at Point Ano Nuevo and Pigeon Point, north of Monterey Bay (Pearse, 1980; unpub- lished). However, no vouchers were retained and because the keys used then (a draft of Hartman’s 1975 keys) would not distinguish between Halisarca sp. and O. carmela, the identity of those records is uncertain. Similarly, using available keys, JSP identified skeleton-less sponges as Halisarca that were collected in the early 1970s at the same site at Carmel Point where O. carmela is now found (unpubl. obs.). It is likely that those specimens were O. carmela, which would sug- MURICY AND PEARSE: NEW SPECIES OF OSCARELLA FROM CALIFORNIA 609 gest that the species has been present at the type locality at least since the early 1970s. In support of this continuous presence, M. C. Diaz, who identified O. carmela growing at Long Marine Laboratory, noted it to be present at Carmel Point during a field trip in 1993 (K. Wasson, pers. com- mun., 2004). On the other hand, an undergraduate student survey of the sponges at Carmel Point in 1993 did not list O. carmela (or Halisarca sp.) (JSP, unpublished list), so it is rare and/or easily overlooked even at the type locality. Because of its slimy, non-sponge appearance, which could be confused with encrusting ascidians, and its unusual habitat in the high intertidal, 1t would not be difficult for students to miss. DISTRIBUTION IN THE FIELD AND AQUARIUMS.— The spatial distribution of O. carmela is puz- zling. Despite extensive searching in the vicinity of the Monterey Peninsula of central California, small. scattered individuals were found in only a few localities in the high intertidal zone. It was not abundant anywhere, and there was no evident pattern in the four sites where it was found in this study. On the other hand, it is difficult to find in the field, even when a search image is estab- lished. Thin sheets of encrusting ascidians (e.g., Aplidium spp., Diplosoma listerianum) and sponges (e.g., Zvgherpe hyaloderma, Astylinifer arndti) were misidentified as O. carmela in the field by JSP: only inspection of specimens with a microscope revealed the error. Nonetheless, once noticed, it is quite conspicuous and hard to miss. Outstanding intertidal biologists (e.g., Ed Ricketts, Don Abbott) collected specimens in the intertidal of the Monterey Peninsula for most of the 20" century. Inasmuch as O. carmela is rather unattractive and usually occurs as small, slimy colonies adhering to large boulders, perhaps after de Laubenfels’ 1932 monograph was published, experienced intertidal biologists simply field identified it as Halisarca sacra and then went on to other things. Moreover, they could have been discouraged because of the difficulty in identifying slimy sponges. For example, in reference to H. sacra, Ricketts and Calvin (1939:176) wrote: “Since there are various additional species not mentioned, which, on the basis of field characters alone, could be confused with the above, and with previously treated species, the value of careful laboratory work should be kept in mind.” Another possibility is that Oscarella carmela normally occurs in deeper water, or in caves and hard-to-reach places, as do other species of Oscarella (Muricy and Diaz 2002). After all, it almost always occurs submerged in pools in the intertidal and does well in dimly lit aquariums. However, since being alerted about its appearance, two observant diving biologists (Anthony Draeger and Scott Nichols) have searched for it subtidally on the Monterey Peninsula without success. Although rarely encountered in the field, and then only as small individuals, Oscarella carmela often thrives in aquariums, exhibit tanks, and sea tables in laboratories and public aquar- iums. This difference in abundance between the field and on-shore facilities indicates that there are factors operating to limit the sponge’s establishment and growth in the field. Perhaps it is preyed upon heavily in the field by some unknown predator, which severely limits its growth subtidally and leads to it occurring mainly as small refuge populations under boulders in high intertidal pools. That appears to be the case for Halisarca sp. in the eastern Pacific (Goddard 1981, 1984, 2001), which is also rarely encountered, especially in California. The source of the specimens of Oscarella carmela in aquariums continues to be unknown. It has been found in three on-shore facilities: (1) Long Marine Laboratory-Seymour Marine Discovery Center complex, (2) Monterey Bay Aquarium, and (3) Bodega Marine Laboratory. These are completely separate facilities, and it is unlikely that material was transferred among them. Moreover, Hopkins Marine Station receives all its running seawater from the Monterey Bay Aquarium, and O. carmela has not been found in any of the aquariums there. Furthermore, the sponge has not been found in the intertidal near two of the facilities when those habitats were searched. Consequently, the source is probably not via seawater being brought into the facilities. 610 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 33 Rather, small specimens probably are brought into the facilities on rocks and other material collect- ed from the intertidal. There is also no evidence that the species has escaped from the facilities to colonize nearby intertidal areas. TEMPORAL DYNAMICS.— It is not known whether the colonies of Oscarella carmela are new to Carmel Point and other sites around the Monterey Peninsula, signaling an environmental change, or were simply overlooked in the past. If specimens keyed as Halisarca from the same area at Carmel Point in the early 1970s by JSP were, in fact, O. carmela, then they have probably been there for a long time. They almost certainly were present since 1993, when M.C. Diaz saw them there (K. Wasson, pers. commun.). After specimens of Oscarella carmela were recognized in high zone pools at Carmel Point in September 2002 by JSP, largely because they looked similar to specimens at Long Marine Laboratory, they were found there on each of nine subsequent visits, extending from November 2002 to April 2004. Specific individuals were not followed, mainly because the boulders were sim- ilar in appearance and appeared to be rearranged between visits by storms. In January 2003, after a major winter storm, the specimens were noticeably thin, and in the spring (April and May) they appeared to be particularly robust, as is the case for many intertidal organisms of the northeast Pacific (Foster et al. 1988). The abundance of O. carmela in the Monterey Bay Aquarium was somewhat seasonal too, with an increase in spring months and a die-off during winter (Veronica Franklin, pers. commun.). A similar pattern is found in calcareous sponges in Rio de Janeiro state, Brazil, belonging to the genera Clathrina and Paraleucilla (Michelle Klautau, pers. commun., 2003). INDIGENOUS OR INVASIVE?.— Oscarella carmela has been known for a long time in labs and local public aquariums before being noticed in the field, a pattern suggestive of an invasion. If this sponge were similar to any described species of Oscarella from other region, it could be conclud- ed that it has invaded central California recently. Instead, the species has proved to be new, and it now seems likely that it is reasonably widespread at least between central California and British Columbia, thus suggesting that it is indigenous. This conclusion is consistent with the finding that nearly all introduced marine species in California are found in estuaries and harbors, and only a subset of these occur on the open coast (Wasson et. al., in press). Specimens of O. carmela have not been found in estuaries of central California that are populated by a high number of introduced species (San Francisco Bay, Andy Cohen and Jim Carlton, pers. commun.; Elkhorn Slough, Wasson et al., in press). On the other hand, the taxonomy of the genus Oscarella is so poorly known worldwide that it is nearly impossible to know if it is really indigenous or introduced. Although we believe that there is a high probability that the new species is indigenous of this area, we cannot completely discard the possibility of an invasion from an unknown population of an undescribed species that originat- ed elsewhere. There are no previous records of the genus Oscarella along the Pacific coast of North America, but this may be due to rarity and difficulty in identifying these species. The temporal dynamics of O. carmela shows high seasonal and pluri-annual fluctuations. Therefore, it is likely that, in this case, there has been an increase of abundance at a few localities of a previously rare, resident species, rather than an invasion. ACKNOWLEDGMENTS We acknowledge Maria Cristina Diaz for first recognizing that the sponge was a species of Oscarella, and particularly for bringing us together. We especially thank Bill Austin and Jeff Goddard for generously sharing their knowledge about Halisarca. Jerry Bakus, Jim Carlton, Andy | | MURICY AND PEARSE: NEW SPECIES OF OSCARELLA FROM CALIFORNIA 611 Cohen, Anthony Draeger, Eric Hochberg, Welton Lee, Todd Newberry, Scott Nichols, Jim Nybakken, Kathy Omura, Curt Smecher, Klaus Riitzler, Bob van Syoc, Kerstin Wasson, and Regina Wetzer also all provided useful information. Veronica Franklin and Jonelle Verdugo assist- ed at the Monterey Bay Aquarium, and Patricia Sadeghian sent JSP the specimens from the Santa Barbara Museum of Natural History. Jonathan Krupp (University of California, Santa Cruz) sup- plied glutaraldehyde and osmium tetroxide and helped with the fixation. We are grateful to Dr. Marcia Attias and Noémia Rodrigues (Laboratorio de Ultraestrutura Celular Herta Meyer, Universidade Federal do Rio de Janeiro) for help on histology procedures and TEM. 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Shallow-water Demospongiae of North-central California: Taxonomy and Systematics, Distribution and Fossilization Potential. Ph.D. dissertation, University of California, Davis. vil + 179 pp. SMECHER, C. (undated). The Sponges of BC. http://www. interchg.ubc.ca/csmecher/sponge.htm (accessed 28 May 2004). Simpson, T.L. 1984. The Cell Biology of Sponges. Springer-Verlag, New York. xix + 662 pp. THOMPSON, J.E., R.P. WALKER, AND D.J. FAULKNER. 1985. Screening and bioassays for biologically-active sub- stances from forty marine sponge species from San Diego, California, USA. Marine Biology 88:11—21. VACELET, J., AND C. DONADEY. 1987. A new species of Halisarca (Porifera, Demospongiae) from the Caribbean, with remarks on the cytology and affinities of the genus. Pages 5—12 in W.C. Jones, ed., European Contributions to the Taxonomy of Sponges. Litho Press Co., Midleton, Ireland. VACELET, J., L. VASSEUR, AND C. LEvi. 1976. Spongiaires de la pente externe des récifs coralliens de Tuléar (sud-ouest de Madagascar). Mémoires du Muséum National d'Histoire Naturelle, Paris, ser. A, Zool. 99:1-116. Wasson, K., K. FENN, AND J.S. PEARSE. (in press.) Habitat differences in marine invasions of Central California. Biological Invasions. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34, pp. 613-689, 82 figs., 4 tables (Appendix). December 30, 2004 A Phylogenetic Analysis of the Aegiridae Fischer, 1883 (Mollusca, Nudibranchia, Phanerobranchia) with Descriptions of Eight New Species and a Reassessment of Phanerobranch Relationships Shireen J. Fahey! and Terrence M. Gosliner2 California Academy of Sciences, 875 Howard Street, San Francisco, California 94103; Email: | sfahev@calacademy.org, 2 tgosliner@calacademy.org The phylogenetic relationships among the Aegiridae are examined based upon mor- phological characters of all presently described species and eight new species. The Aegiridae are a monophyletic clade of phanerobranch dorids, a paraphyletic group previously united by the absence of a retractable dorsal gill, which is a pleisomorphic character. There are three traditionally recognized genera within the Aegiridae, Aegires Loven, 1844, Notodoris Bergh, 1875 and Triopella Sars, 1878. Most species of each genus were examined anatomically and some previously undescribed charac- ters were included in the analysis such as details of the central nervous system. Published literature descriptions were utilized in cases where no specimens were available for examination. From the literature review and anatomical examinations, sixty-four characters were considered. These characters were polarized using Bathydoris and the type species of several phanerobranch genera. Additionally, four cryptobranch dorids were included in the analysis for comparative purposes. The phylogeny obtained supports both the monophyly of Aegiridae and that Notodoris is a monophyletic clade nested within Aegires. Triopella incisa, the type species of a monotypic genus, is nested within Aegires and is the sister species of Aegires sublae- vis. This phylogeny necessitates inclusion of Notodoris and Triopella as junior syn- onyms of Aegires to preserve the monophyly of Aegires. This also renders Aegires cit- rinus Pruvot-Fol, 1930 as a junior homonym of Aegires citrinus (Bergh, 1875). The new name Aegires pruvotfolae is given to this former species. The following are new species of Aegires: A. exeches sp. nov., A. flores sp. nov., A. hapsis sp. nov., A. incusus sp. nov., A. lemoncello sp. nov., A. malinus sp. nov., A. ninguis sp. nov. and A. petalis sp. nov. All these new taxa are found in the Indo-Pacific except A. ninguis, which is found in temperate South Africa. Unique combinations of morphological characters distinguish these as new species of Aegires. The present phylogeny supports the notion that the Phanerobranchia represents a paraphyletic group. The Suctoria are the sister group to the Cryptobranchia. Aegiridae is the sister group to Cryptobranchia plus Suctoria. Polyceratidae is the sister group to Cryptobranchia plus Suctoria plus Aegires. Hexabranchus is basally situated to all of these other taxa but is more derived than Bathydoris. Aegiridae Fischer (1883) is one of the families of the traditional group Phanerobranchia (Fischer 1880-1887). The classification of the family Aegiridae (= Notodorididae Eliot, 1910) has an interesting, although convoluted history and is presented in detail below and summarized in 613 614 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 Table 1 (see Appendix). According to the most recently published classification of the Aegiretidae [sic] (Rudman and Willan 1998) there are only two recognized genera within the group: Aegires Lovén, 1844 and Noftodoris Bergh, 1875. However, Triopella Sars, 1878 is also a genus that has been overlooked by most researchers. The primary character that unites all Phanerobranchia, including Aegiridae, is the presence of a non-retractable dorsal gill. As a result of having an unprotected gill, these animals have devel- oped protective appendages for the gill leaves. This adaptation provides some important informa- tion for phylogenetic analyses, which is discussed in detail in a later section. Thompson (1976) described the protective appendage in his characterization of the Superfamily Anadoridoidea (Odhner’s [1939]), Suborder Anadoridacea. Fischer et al. (1968) were the first to equate Anadoridacea with the Phanerobranchia Fischer, 1883. The Phanerobranchia are now thought to be paraphyletic (Valdés 2002). Historically, the Suborder Anadoidacea was divided into two tribes (Bergh 1890), the Suctoria and Non-suctoria. Aegiridae is one of the three currently recognized families within the tribe Non- suctoria along with Triophidae Odhner, 1941 and Polyceratidae Alder and Hancock, 1855 (amend- ed from the original authors’ spelling) (Rudman 1998). While these three families have a widely variable external anatomy there are also some similarities. These include the anus and the gill sit- uated medio-dorsally, a well-developed radula usually lacking a rachidian tooth, the edge of the mantle more or less reduced, and a contractile gill that is not retractile. Some of the internal differ- ences between the families include genital organs that may have small hooks on the vas deferens or be unarmed and either a smooth jaw or a jaw having rods (Fischer et al. 1968). The characters that unite Aegiridae (as noted first by Thiele 1931, although he had called the Family Notodoridinae) include: “‘a hard body with calcareous spicules, gills that have a separate integumental fold, rhinophores that retract and in most cases, are not lamellate, radula without a rachidian tooth and all lateral teeth hook- or arch-shaped.” Several of the species of both Aegires and Notodoris were originally described in very abbre- viated terms, or based on single specimens. Some entire organ systems were never described either initially or by subsequent workers. In the present study, specimens of each species were examined when available and when not available, all published literature was reviewed. Anatomical details are described for all known species of each genus, some of these details for the first time. The present study provides an in-depth look at the Aegiridae and presents a phylogenetic analysis of the placement of this family within the Phanerobranch dorids (the Anadoridoidea). The name Phanerobranchia is the most widely used and recognized name, and it is used here instead of the synonymous Anadoridoidea. Eight new species are described based on examination of material lodged at the California Academy of Sciences and the South African Museum. HISTORY OF AEGIRIDAE CLASSIFICATION Lovén (1844) introduced the new genus Aegires without designating a family affiliation. For the type species of his new genus, he selected Polycera punctilucens (D’Orbigny 1836-1842). Alder and Hancock (1845-1855) labeled the first Suborder in their classification of the Nudibranchia, Anthobranchia. In the early years of the monograph (1845) they placed Aegires within Family #1, Dorididae, (Sub-Family Polycerinae), along with Thecacera, Polycera and Idalia. Later, in 1855, they proposed to use a classification originally suggested by Gray, which now named Family #2 the Dorididae, and which now included Aegires with Doris, Goniodoris and Ceratosoma. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 615 Bergh (1875) introduced the new genus Nofodoris and drew some anatomical features of the type species Notodoris citrina. However, he did not designate a familial association. Sars (1878) described and drew the type species for his new genus Triopella (incisa). He men- tions the radular similarities between Triopella and Aegirus in the discussion of the new species. Fischer (1880-1887) divided the Nudibranchia into 5 suborders: Anthobranchiata, Inferobranchiata, Polybranchiata, Pellibranchiata and Parasita. He further divided the Anthobranchiata into Aglossa and Glossophora. The Aglossa had one family, Doridopsidae, while the Glossophora were subdivided into three groups: the Cryptobranchiata (with one family, Dorididae), the Phanerobranchiata (Family Polyceridae) and the Abranchiata (Family Heterodorididae). Fischer, noting that Triopella and Aegires (which, like Sars, he spelled Aegirus) were closely related, placed both these genera in the Phanerobranchiata (Family Polyceridae) with no mention of Nofodoris. Fischer did acknowledge that his Polyceridae corresponded to Bergh’s “Doris phanerobranchs”, so although Fischer left out Notodoris he was evidently aware of Bergh’s 1875 publication. Fischer further separated the Polyceridae into three sub-families, Acanthodoridinae, Polycerinae and Aegirinae. Bergh (1890) acknowledged Fischer’s group Phanerobranchiata when he placed his two newly described species, Plocamopherus amboinensis Bergh, 1890 and Plocamopherus indicus Bergh 1890 into the Subfamily Phanerobranchiata (Suborder Nudibranchiata Holohepatica, Family Dorididae). Two years later, when Bergh described the systematics of the Nudibranchia (1892:141) he placed Aegires, Notodoris and Triopella within the Polyceridae (Subfamily Dorididae Phanerobranchiatae Non Suctoriae). He used the spelling Aegires not Aegirus as Fischer did in 1883. This may have been in recognition of the indexed generic names published in Nomenclator Zoologicus (Scudder 1882). Eliot (1903) placed his newly described Notodoris gardineri within the Family Dorididae Phanerobranchiatae (his spelling). He described the Phanerobranchiatae as those dorids that have no dorsal cavity into which the branchia can be retracted. Eliot acknowledged Bergh’s (1875) divi- sion of the Phanerobranchiata into the Goniodorididae (which possesses a buccal gizzard) and Polyceradae (no gizzard), but stated that a more “natural” division would be Polyceradae (having limaciform bodies, sometimes bearing appendages) and Pseudodorididae (flat dorid-like forms). Eliot also noted in his study that Notodoris is closely related to Aegires and the little known Triopella because all three had a hard consistency and had branchiae and rhinophores protected by tubercles. Eliot (1910:65) in A Monograph of British Nudibranchiate Mollusca, Part 8, went on to estab- lish the Family Notodorididae for Notodoris, Aegires, and Triopella. He united these genera based upon their simple, hook-shaped teeth, branchial valves and smooth rhinophores. Eliot did not ref- erence Fischer’s 1883 family name Aegiridae at this time. However, thirteen years later, in their classification, Iredale and O’ Donoghue (1923) also used the family name Aegiretidae for Aegires, Notodoris and Triopella but without providing a diagno- sis for this group. Apparently these authors were aware that Aegiretidae (Aegiridae) was an older name than Notodorididae. In 1926, Odhner recognized that the Family Notodorididae = Aegiretidae, Iredale and O’Donoghue, incorrectly attributing the name Aegiretidae to these authors (and maintaining the incorrect spelling of the family name). Odhner provided two distinguishing characters for Notodorididae (Phanerobranchia): “Radular teeth hooked, uniform and tentacles small, inconspic- uous”. Risbec (1928) provided a diagnosis for the Family Aegiridés [sic] using the genus Aegirus 616 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 Lovén 1844 as his model. In this document he also provided a lengthy description of A. leuckartii Verany, 1853 and synonymized A. albopunctatus MacFarland, 1905 with this species. This syn- onymy was later questioned by Schmekel and Portmann as will be discussed later in the present paper. Pruvot-Fol (1930) placed a new species she described as Aegires citrinus into the Family Aegiretidae, with this particular spelling. The following year, Thiele’s classification (1931) placed Aegires, Notodoris and Triopella within the Subfamily Notodoridinae (Stirps Doridacea (= Holohepatica) Family Polyceridae). Thiele’s diagnosis of the Notodoridinae follows: “Body hard, with calcareous spicules; gills with a separate integumental fold; rhinophores retractile, in most cases not lamellate; radula without cen- tral plate, all lateral plates hook- or arch-shaped.” Odhner (1934) established a new genus Anaegires with A. protectus as the type species. However, this new genus was based upon characters that were later found by Wagele (1987) to be within the realm of intraspecific variation. Wagele justifiably synonymized 4. protectus with A. albus Thiele, 1912. Odhner also commented that Iredale and O’ Donoghue (1923) should not have changed the family name to Aegiretidae without justification. Odhner stated that Nofodoris may be kept as the type genus of the Family and thus Eliot’s family name Notodorididae is valid. He com- pletely overlooked the fact that the name Aegiretidae (Aegiridae) was used twenty-seven years prior to Notodorididae. Pruvot-Fol (1954) placed Aegires in Aegiretidae noting that Thiele (1931) erroneously called the subgroup Notodorididae when he grouped Notodoris and Aegires in the family Polyceradae. Pruvot-Fol correctly pointed out that Aegires Loven, 1844 1s an older name than Nofodoris Bergh, 1875. In this paper, Pruvot-Fol also synonymized Aegires hispidus (Polvcera hispida Hesse, 1872) and A. leuckartii Verany, 1853 with A. punctilucens (Polycera d’Orbigny, 1837). However, she did not mention Risbec’s previous (1928, 1953) synonymy of 4. hispidus and A. leuckartii with Aegires albopunctatus. In 1966, MacFarland classified the Notodoridinae (sic) as a subfamily of Polyceridae. Fischer et al. (1968) placed Aegiretidae into the Anadoridacea Odhner (1968) in the tribe non- suctoria (one of two tribes first identified by Bergh [1892]). Aegiretidae were considered Family #2 of the Anadoridacea (Family | = Triophidae Odhner nov and Family 3 = Polyceridae Alder and Hancock 1845). Fischer et al. wrongly attributed the name Notodorididae to Odhner, 1926, when in fact Eliot first established this group in 1910. Nordsieck (1972) established a new genus Serigea naming Aegires (Serigea) sublaevis as the type species. He presented the following brief diagnosis of Serigea: “Features like Aegires, but without tubercles (papillae); rhinophore sheaths low, smooth. Only a few tubercles behind the rhinophores.” Notably, Nordsieck did not mention Nofodoris in his manuscript on European marine molluscs when he grouped Aegires, Triopella and Serigea noy. gen. within the Family Aegiretidae Fischer, 1883. This is most likely because there are no known Nofrodoris species found in European marine habitats. He did pronounce as Pruvot-Fol did in 1954, that Aegiretidae = Notodorididae Odhner, 1926. Nordsieck thus continued the incorrect attribution of the name Notodorididae to Odhner, 1926 and the incorrect spelling of the family name. Thompson (1976) changed the name Anadoridacea to Anadoridoidea, in which he included the Family Notodorididae. Thompson listed only Notodoris and Aegires as the genera in this family. He did not cite the authors of any family names. (see Valdés 2002 for a discussion of the Family name Anadoridacea). Schmekel and Portmann (1982) used Thompson’s name Anadoridoidea but erroneously attrib- FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 617 uted the name to Odhner (1959); a non-existent publication that was also omitted in their literature cited. Gosliner and Behrens (1997) incorrectly attributed the name Notodorididae to Bergh (1897) in their description of the new Nofodoris serenae. Bergh never used this name to categorize the new genus Nofodoris. It wasn’t until Eliot (1910) wrote the text for Part 8 of Alder and Hancock’s monograph of the British nudibranchiate molluscs that this name was first used. In the most recently published classification of Aegiretidae, Rudman (1998) placed Aegires and Votodoris in the Family Aegiretidae, Superfamily Anadoridoidea, suborder Doridina. Triopella was once again left out of the family and the incorrect spelling of the family name was continued. The correct spelling, Aegiridae is discussed in detail by Willan (2000). Table 1 (see Appendix) summarizes the historical classification of Aegiredae. MATERIAL AND METHODS Twenty-two species of Aegiridae were included in the analysis. This includes all known species of the recognized genera, Aegires, Notodoris and Triopella as well as eight additional, pre- viously unidentified species of Aegires. In some cases, the complete published description of cer- tain features of a species allowed the extraction of data from the literature, which was then verified by direct examination of a specimen when available. Type material and additional non-type mate- rial was obtained from the following sources: the California Academy of Sciences (CAS), the Los Angeles County Natural History Museum (LACM), the Natural History Museum of Norway, Oslo (D), the South African Museum (A) and the Museo National de Ciencias Naturales Madrid (MNCN). Specimens were drawn under a dissecting microscope using a camera lucida then dis- sected by dorsal incision. The internal anatomy was drawn as described and then examined either by compound or scanning electron microscope (SEM). External features were examined directly when specimens were available, by photographs, or by literature review. Special attention was given to the reproductive anatomy, the central nervous system and the circulatory system, as some of these features were infrequently (or cursorily) described in the literature. Dorsal tubercles of available specimens were drawn and examined by SEM. Table 2 (see Appendix) shows the list of character states derived from dissections and from the literature reviewed for the present study. Ten additional species were selected for outgroup comparison as discussed below. Phylogenetic analytical methods Phylogenetic analyses were performed using the program Phylogenetic Analysis Using Parsimony (PAUP) version 4.0 (Swofford 2001) using the heuristic algorithm (TBR branch swap- ping option). One hundred random start trees were obtained by stepwise addition. Bremer analyses were performed to estimate branch support (Bremer 1994). Characters were unordered and were polarized using Bathydoris abyssorum Bergh, 1884 as the outgroup. This type species was select- ed based on the analysis of the cryptobranch dorids by Valdés (2002), which showed that Bathydoris is sister to the dorids. Additionally, Wagele (1989a) and Wagele and Willan (2000) demonstrated that Bathydoris is the most basal member of the Anthobranchia and is the sister group to the rest of the dorid nudibranchs. Polycera quadrilineata (Miller 1776), Okenia elegans (Leuckart, 1828), Onchidoris bilamellata (Linnaeus, 1767), Holoplocamus papposus Odhner, 1926, Diaphorodoris luteocinta (Sars, 1870) and Calycidoris guentheri Abraham, 1876 were also tested as outgroup taxa in the analyses. These five additional genera are phanerobranchs and basal- ly situated relative to the rest of the dorids. Four cryptobranch dorids were included in the analy- sis for comparative purposes. The genera were chosen from the most recent analysis of the crypto- 618 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 branch dorids (Valdés, 2002) and are the type species and represent the most basal taxa of their par- ticular clade. Those genera are Actinocyclus Ehrenberg, 1831; Conualevia Collier and Farmer, 1964; Hexabranchus Ehrenberg, 1831 and Mandelia Valdés and Gosliner, 1999, which has been included in the Cryptobranchia in the most recent classification of the Doridina (Rudman 1998). The source of the information on the outgroup species included in the present analysis is shown in Table 3 (see Appendix). Synapomorphies were examined using MacClade and the character-trace option, using the majority rule tree from PAUP analyses. SPECIES DESCRIPTIONS Family Aegiridae Fischer, 1883 Genus Aegires Lovén, 1844 TyPE SPECIES: Polycera punctilucens d’Orbigny, 1837, by monotypy. TyPE SPECIES: Aegires albus Thiele, 1912, by original designation. Synonyms of Aegires: Anaegires Odhner, 1934:241, type species Anaegires albus Odhner, 1934 by monotypy; Notodoris Bergh, 1875:64, type species Notodoris citrina Bergh, 1875 by monotypy, syn. nov.; Serigea Nordsieck, 1972, type species Serigea sublaevis Nordsieck, 1972 by subsequent designation; Triopella Sars, 1878:310, type species Triopella incisa Sars, 1878 by monotypy, syn. nov. DIAGNosis.— Diagnoses of this genus have been provided by multiple authors: Lovén (1844), Schmekel and Portmann (1982), and Thompson and Brown (1984). A summary of these authors’ diagnoses follows: The body 1s firm, with a reduced, indistinct mantle skirt. The dorsum is covered with numerous blunt, pedunculate dorsal tubercles. The rhinophores extend from cylindrical pal- lial sheaths that have tubercles around the rims. The gill is protected by tuberculate lobes. A medi- an dorsal cuticular plate or jaw is present in the buccal mass. The radular teeth are simple, hamate and not bifid. Aegires albopunctatus MacFarland, 1905 (Figs. 1A, 2-6) Aegires albopunctatus MacFarland, 1905:35-54, pl. 18, Figs 5-8. TYPE MATERIAL.— SYNTYPE: Monterey Bay, California, No. 181,281. U.S. National Museum. MATERIAL EXAMINED.— LACM 127330, one specimen, 18 mm, dissected, Rio San Ysidro, Baja California, collected 28 June 1946, C. Johnson. LACM 71-36, one specimen, 18 mm, dissect- ed, Point Dume, Southern California, collected 8 February 1971, G Sphon. LACM 90-2.3, one specimen, 7 mm, dissected, Cabo Colnett, Baja California, collected 10 February 1990, collector not specified. CASIZ 118550, one specimen, 20 mm, dissected, Pacific Coast, Baja California North, collected 3 February 1963, W. Farmer. CASIZ 068347, nineteen specimens, one specimen, 20 mm dissected, Corona del Mar, Southern California, collected 4 May 1946, F. MacFarland. CASIZ 068349, five specimens, one specimen, 20 mm, dissected, Cannery Row, Monterey Bay, California, collected 28 September 1968, A. Smith. FIGURE 1. Living animals. (A) Aegires albopunctatus MacFarland, 1905. CASIZ 118550, photo by A. Ferreira, Monterey Bay, California, 19 mm. (B) Aegires pruvotfolae nom. nov. CASIZ 097449, photo by T. Gosliner, Napili Bay, Maui, Hawaii, 6 mm. (C) Aegires gomezi Ortea, Luque and Templado, 1990. CASIZ 077315, photo by T. Gosliner, Grand Cayman Island, 5 mm. (D) Aegires ninguis.sp. nov. CASIZ 073982, photo by T. Gosliner, Cape Province, South Africa, 8 mm. (E) Aegires lemoncello sp. nov. CASIZ 086465, photo by T. Gosliner, Pig Island, Papua New Guinea, 4 mm. (F) Aegires malinus sp. nov. CASIZ 085889, photo by T. Gosliner, Bebbit, Philippines, 8 mm. (G) Aegires incusus sp. nov. CASIZ 156668, photo by T. Gosliner, Cemetary Beach, Luzon, Philippines, 5 mm. << FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 619 620 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 FIGURE 2. Aegires albopunctatus MacFarland, 1905. CASIZ 118550. Drawing of preserved specimen. (A) Dorsal view. (B) Ventral view of head. Scale = 7 mm. DISTRIBUTION.— West coast of North America from British Columbia to Baja California ({[Behrens 1991] Rudman, SeaSlug Forum, accessed Sept. 2004). EXTERNAL MORPHOLOGY.— MacFarland (1905, 1906, 1966) gave thorough descriptions of the external morphology of this species. The spec- imens examined for the present study matched his descriptions and thus there is no additional infor- mation to present. See Fig. 1A for a photo of a liv- ing animal from the type locality and Fig. 2 for a drawing of a preserved specimen from Baja California. DIGESTIVE SYSTEM.— Aegires albopunctatus shares the same general digestive anatomy as other Aegires species (Fig. 3). The buccal bulb is nearly round, with four large muscles attached; two per side. The buccal bulb is shorter and more round than the oral tube (Fig. 3A). There are two glands at the side of the oral tube, near the mouth (Fig. 3B). The radular sac slightly protrudes from the bulb, under the esophagus. The labial disk frames the triangular opening to the buccal bulb and is FIGURE 3. Aegires albopunctatus MacFarland, 1905. LACM 127330. (A) Buccal bulb: bb = buccal bulb, ens = central nervous system, m = muscle, o = esophagus, rs = radular sac, scale = 0.3 mm. (B) Digestive system: bb = buccal bulb, bga = blood gland artery, ed = collecting ducts, ens = central nervous system, ht = heart, i = intes- tine, o = esophagus, og = oral gland, ot = oral tube, st = stomach, scale = 3 mm. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 621 lined with a thick cuticle. There is a thick plate at the top of the open- ing, with thick rods at the edge (Fig. 4A). The radular formula (CASIZ 118550) i 19 x 21.0.21, with all teeth as described by MacFarland. That is, they are simple, hooked and similar in form (Figs. 4B— D). The esopha- gus is short and connects directly to the stomach eeic: 6). ~ The intestine makes a simple, wide curve along the outside of the -_ oF ft Sas digestive gland FIGURE 4. Aegires albopunctatus MacFarland, 1905. LACM 127330. Buccal morphology. 8 & ; (A) Jaw, scale =100 um. (B) Whole radula, scale = 100 tum. (C) Inner lateral teeth, scale = 10 um. REPRODUC- (D) Outer lateral teeth, scale = 30 um. um TIVE SYSTEM.— The ampulla is large and ovoid (Fig. 5). It branches into the oviduct and the tubular prostate. The her- maphroditic duct enters the ampulla termi- nally. The thin oviduct enters the large female gland mass. The prostate is differen- tiated into two parts, the proximal portion being a flattened mass and the distal portion being a long thickened tube. The distal por- tion folds once after exiting the flattened mass and narrows into the coiled deferent duct. The penis is wider than the deferent duct and terminates into a common genital atrium. The penis contains minute, closely set FigurRE 5. Aegires albopunctatus MacFarland, 1905. penial hooks throughout the length of the LACM 127330. Reproductive system: am = ampulla, be = penis. The vaginal duct is short and wide. It bursa copulatrix, dd = deferent duct, fgm x female gland : : mass, p = penis, pr = prostate, ps = penial spines, rs = recep- was not examined internally and thus the pres- : taculum seminis, v = vagina, vd = vaginal duct, scale = | ence of spines or hooks cannot be confirmed. mm. 622 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 At the distal end, the vagina 1s wide and bulbous. The prox- imal end terminates into the large ovoid bursa copulatrix. From the bursa the oviduct connects to the smaller pyri- form receptaculum seminis. A very short uterine duct leads from the receptaculum seminis into the female gland mass. CENTRAL NERVOUS SYSTEM.— As with other species of Aegires, the cerebral and pleural ganglia are fused together (Fig. 6). The two pedal ganglia are located below the cerebro-pleural complex and are joined by an elongate commissure. The buccal ganglia are placed under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two relatively short nerves. The eyes are sessile at the cerebro-pleural juncture. Farland, 1905. LACM 127330. Central nerv- There are four cerebral nerves leading from each cerebral gus system: bg = buccal ganglia, eple = cere- ganglion including the rhinophoral ganglia, and two large _ bro-pleural ganglia complex, p = pedal gan- pleural nerves leading from the right and left pleural gan- _ glia, r = rhinophoral nerve, vI+pe+ppe = vis- glia. There is a separate abdominal ganglion on the right ceralloop, scale side of the visceral loop. Gastro-esophageal, rhinophoral and optical ganglia are present. ReMARKS.— MacFarland (1905, 1906, 1966) presented thorough external and radular descrip- tions of this northeastern Pacific species. Anatomical information for the present study has been taken from his paper and corroborated by examining specimens from the type locality and the southern California coast. Ernst Marcus (1961) also published additional anatomical details of this species. However, some characters that were necessary for the present study such as the circulato- ry, central nervous or digestive systems were not described. Therefore, additional specimens were examined to complete the data. Risbec (1928) synonymized 4. albopunctatus MacFarland, 1905 with A. leuckartii Verany, 1853 in his lengthy description of A. /euckartii. Marcus (1961) noted some external similarity between 4. albopunctatus and A. punctilucens stating that A. albopunctatus “agrees with the type species A. punctilucens” based upon the papillae on the mantle border of both species. No other subsequent publications referring to A. punctilucens or A. leuckartii recognized the synonymy of either species with 4. albopunctatus (Pruvot-Fol 1954; Fischer et al. 1968; Nordsieck 1972; Schmekel and Portmann 1982; Thompson and Brown 1984). However, Haefelfinger (1968) did synonymize A. /euwckartii with A. punctilucens based upon characters that he thought lie with- in natural variation. But subsequently, Schmekel and Portmann (1982) separated A. punctilucens and A. leuckartii as subspecies, distinguishable by their ecology and reproductive cycles. Further remarks on this synonymy are presented under the sections discussing these two species. The present study confirms that Aegires albopunctatus is a valid species with the following characters distinguishing it from Aegires punctilucens: eee vitpe+ppe tate FIGURE 6. Aegires albopunctatus Mac- 1. EXTERNAL COLOR. Aegires punctilucens has very distinctly colored spots on the dorsum. These spots have been described by various authors (Thompson and Brown 1984) as resembling an ocellus: that is, dark brown oval areas containing an iridescent blue-green spot, with or without additional dark spots on the periph- ery of the ocellated markings. In contrast, MacFarland’s description of A. al/bopunctatus is very clear in that the dorsum only has minute dots of pure white, with or without irregularly scattered small dark brown spots. No ocellus spots are noted on any of his specimens. 2. DORSAL TUBERCLE ARRANGEMENT. Aegires albopunctatus has very densely placed tubercles, closely set in rows and continuing behind the rhinophores as a tuberculate ridge that diminishes in prominence FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 623 (MacFarland 1905). Aegires punctilucens has more widely scattered, larger tubercles that continue as a tuber- culate ridge in a crest along the middle of the back (Schmekel and Portmann 1982). 3. REPRODUCTIVE CHARACTERS. Aegires albopunctatus has two ducts emerging from the base of the bursa copulatrix, while 4. pwnctilucens has only one. The receptaculum seminis duct emerges from the proximal end of the large, bulbous vagina. Additionally, the vagina of 4. albopunctatus is much more elongate than that of A. punctilucens. 4. RADULAR FORMULA. Schmekel and Portmann (1982) reported a radular formula of 16 x 18.0.18 for a 6 mm specimen of 4. punctilucens with the size of the teeth increasing outwards. Thompson’s (1984) 12 mm specimen of 4. punctilucens from the Isle of Man had a radular formula of 23 x 22.0.22 with the teeth from the midline having a hooked tip, but the teeth from the margins were reported as more smoothly hooked. These reports on the radular formulae differ from MacFarland’s reported formula for a 13 mm specimen of A. albop- unctatus of 16-22 x 17.0.17 with the innermost and outermost teeth being similar in size but smaller than the middle lateral teeth. (See also Fig. 4). Aegires albus Thiele, 1912 Anaegires protectus Odhner, 1934:242. Aegires protectus (Odhner) W4gele, 1987:271. TYPE MATERIAL.— HOLOTYPE: No. 63230 (4. albus), McMurdo Sound, Antarctica. Zoologisches Museum zu Berlin; Other material: No. 846 (A. albus) Naturhistoriska Riksmuseet, Stockholm; HoLotyPe: No. 1934.10.5.67 (A. protectus), British Museum of Natural History, London. MATERIAL EXAMINED.— Aegires albus from the Muséum National d’ Histoire Naturelle, Paris: Kerguelen Islands, 1 specimen, collected at 15 m, 16 January 1963, no further collection data pro- vided; MD04 G65 DC 155, | specimen, no further collection data provided; SMK, 10 specimens, 15-144 m, collected 1972 and 1974 by Guille et al. DISTRIBUTION.— This species has only been recorded from the Antarctic Peninsula, the Ross Sea and the Weddell Sea (Wagele 1987). REMARKS.— Aegires protectus is the type species of Anaegires Odhner, 1934. Anaegires is a synonym of Aegires (see Odhner 1934). Wagele (1987) presented a thorough description of Aegires albus, with both external and internal anatomy. She demonstrated that Aegires protectus is a syn- onym of this species. Anatomical information has been taken from this publication and from the specimens noted above for use in the present study. Examination of the reproductive system of specimens listed above confirmed the presence of densely placed penial spines extending the length of the penial bulb as reported by Wagele (1987). Aegires pruvotfolae Fahey and Gosliner, nom. nov. (Figs. 1B, 7-11) Aegires citrinus Pruvot-Fol, 1930:229-232, junior homonym of Aegires citrinus (Bergh, 1875). (See the dis- cussion of Aegires citrinus). TYPE MATERIAL.— Collected in New Caledonia, date and collector not specified. NEOTYPE here designated, CASIZ 157477, one specimen, 6 mm, Layag Layag, Caban Island, Philippines, collected T. Gosliner. MATERIAL EXAMINED.— CASIZ 097449, one specimen, 6 mm, dissected, Napili Point, Maui, Hawaii, no depth available, collected 9 April 1994, T. Gosliner. CASIZ 087056, one specimen, 10 mm, Keahou Beach, Hawaii, no depth available, collected 3 September 1973, T. Gosliner. CASIZ 624 FIGURE 7. Aegires pruvotfolae nom. noy. CASIZ 097449. Drawing of preserved specimen. (A) Dorsal view. (B) Ventral view of head. Scale = 0.4 mm. 087058, three specimens, no depth available, collected 19 February 1986, T. Gosliner. CASIZ 070326, one specimen, 3 mm, dissect- ed, Nosy Tanikely, Madagascar, | m depth, col- lected 14 April 1989, T. Gosliner. CASIZ 088363, one specimen, 8 mm, dissected, Midway Island, Pacific Ocean, 10 m, collected 29 May 1993, T. Gosliner. CASIZ 099286, one specimen, 6 mm, Manahuanja Island, Tanzania, collected 1 Nov 1994, T. Gosliner. DISTRIBUTION.— This species has been recorded from eastern Australia, New Caledonia, the Philippines, Hawaii, Palau, Midway Atoll, Tanzania and Madagascar (Rudman 2004 and present study). EXTERNAL MORPHOLOGY.— The body shape is high and arched (Figs. 1B, 7). The dor- sum has tall, anvil-shaped tubercles with flat tops. Spicules protrude from the tops of all tubercles. The rhinophore pocket is elevated and has one tall tubercle on the outside perime- ter. There are two prominent tubercles on the head. The posterior end of the foot has low PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 FIGURE 8. Aegires pruvotfolae nom. noy. CASIZ 097449. (A) Buccal bulb: bb = buccal bulb, m = muscle, 0 = esopha- gus, ot = oral tube, rs = radular sac, sg = salivary gland, scale = 0.25 mm. (B) Digestive system: bb = buccal bulb, bga = blood gland artery, cg = cerebral ganglia, cns = central nerv- ous system; ht = heart, i = intestine, m = muscle, og = oral gland, ot = oral tube, sg = salivary gland, st = stomach, scale = 0.25 mm. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 625 tubercles that are much smaller than those on the dorsum. The rhinophores are smooth. The gill pocket lies in the posterior third of the dorsum and three large anvil-shaped tubercles protect the anterior side of the gill pocket. The three small gill leaves are tripinnate. The background color is pale to medium yellow. The tubercles are the same color, as are the rhinophores. There are light brown to tan spots that vary in size along the dorsum median, between the tubercles. DIGESTIVE SYSTEM.— Aegires pruvotfolae shares the same general digestive anatomy as other Aegires species. That is, there are two glands at the side of the oral tube, near the mouth (Fig. 8). The buccal bulb is nearly round, with four large muscles attached, two per side. The buccal bulb is shorter and more round than the oral tube. The radular sac slightly protrudes from the bulb, under the esophagus. The labial disk frames the triangular opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate at the top of the opening, with thick rods at the edge (Fig. 9A). The radular formula is 16 x 11.0.11. The teeth are simply hamate and the three innermost lateral teeth are smaller than the remaining teeth (Figs. 9B—D). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the diges- tive gland. REPRODUCTIVE SYSTEM.— The ampulla is relatively small and compact. It branches into a short oviduct and the prostate (Fig. 10). The oviduct enters the female gland mass near its center. = 20 um. (B) Whole radula, scale = 100 um. (C) Inner lateral teeth, scale = 10 um. (D) Outer lateral teeth, scale = 10 pm. 626 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 The prostate is very long, tubular and coiled. It narrows slightly then connects to a wide ejacu- latory duct and terminates at the wide penis. There are densely spaced, small hooks inside the penis at the distal tip only. The vagina is short and wide. It was not examined internally and thus the presence of spines or hooks cannot be confirmed. At its proximal end the vaginal duct joins the bursa copulatrix and the seminal receptacle. The uterine duct also leads from this duct. The bursa is spherical and the recep- taculum seminis is about the same size though elongated: F 10. Aegi ifol CASIZ = : IGURE . Aegires pruvotfolae nom. nov. CIRCULATORY SYSTEM. The heart (Fig. 097449. Reproductive system: am = ampulla, be = bursa 8) is relatively small as compared to most cryp- — copulatrix, dd = deferent duct, fgm = female gland mass, p= tobranch dorids (Valdés 2002). There is one _ penis, pr = prostate, ps = penial spines, rs = receptaculum blood gland situated in front of and to the right seminis, V = vagina, vd = vaginal duct, scale = 0.2 mm. side of the visceral cavity. CENTRAL NERVOUS SYSTEM.— As. with other species of Aegires, the cerebral and pleu- ral ganglia are fused together (Fig. 11). The two pedal ganglia are located below the cere- bro-pleural complex and are joined by pedal commissure, the parapedal commissure and the visceral loop. The buccal ganglia are placed under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two relatively short nerves. There are four cerebral nerves leading from each cerebral ganglion, and three pleural nerves leading from the right and left pleural ganglia. There is a Sepak apo ual pangheonon they 2G side 097449. Central nervous system: bg = buccal ganglia, cple = of the visceral loop. Gastro-esophageal., cerebro-pleural ganglia complex, p = pedal ganglia, r = rhinophoral and optical ganglia are present. rhinophoral nerve, vitpe+ppe = visceral loop, scale = 0.2 REMARKS.— Pruvot-Fol (1930) presented ™m. a very abbreviated description of Aegires citrinus collected from New Caledonia. The only infor- mation she provided on this new species was that the color was lemon yellow and the specimen had pointed tubercles with light brown color at the tops. Pruvot-Fol wrote that 4. citrinus resem- bled Aegires leuckartii Verany, 1853 and may be a variant of the latter. We discuss this species in the comparison section below. Risbec (1953) briefly mentioned Pruvot-Fol’s abbreviated description of A. citrinus. He thought that since no precise detailed description had been published, A. citrinus was probably a variation of A. /euckartii (see discussion below). Rudman (2004) provided additional external morphological information from photographs taken of this species. For the present study, specimens from the Indian Ocean, Hawaii and Midway Island, Pacific Ocean were examined. This study has revealed that 4. pruvotfolae is a valid species having sever- FIGURE 11. Aegires pruvotfolae nom. nov. CASIZ FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 627 al characteristics that differ from the other Aegires species. Those characters are: |. EXTERNAL COLORATION. Aegires pruvotfolae is similar in coloration to A. incusus (present study). That is, both species can have a pale yellow background color and light brown to tan dorsal spots. However, the tubercle color of 4 privotfolae is also yellow with tan apices, whereas A. incusus has brown to tan tubercles with a darker top. The rhinophores of A. priuvotfolae have a tan top whereas those of A. incusus do not. 2. TUBERCLE SPICULES. The spicules that project from the tops of the tubercles of A. pruvotfolae are low and less conspicuous than those of A. incusus. 3. TUBERCLE ARRANGEMENT. There are two prominent flat-topped tubercles on the head of specimens of A pruvotfolae. On A. incusus, there are multiple tubercles on the head. The dorsal tubercles of A pruvotfolae are less numerous than on specimens of A. incusus. 4. RADULAR CHARACTERS. Aegires pruvotfolae has one small, thin inner lateral tooth that has a very small hook; the next 34 inner lateral teeth are smaller than the remainder. All other lateral teeth are the same size. In A. incusus, there is also one small inner lateral tooth but it has a distinct hook. The next 2—3 teeth are small- er than the remaining teeth, which are all the same size. In addition, the radular sac does not protrude from the buccal bulb in 4. citrinus as it does in A. incusus. 5. REPRODUCTIVE CHARACTERS. Aegires pruvotfolae has reproductive characters that distinguish it from the most similar Aegires species. Most noticeably, the vagina is very wide as compared to A. incusus. In addi- tion, the bursa copulatrix of A. pruvotfolae is as large as the ampulla, whereas in A. incusus the bursa is com- paratively much smaller. The prostate of A. pruvotfolae is very long and coiled and does not narrow apprecia- bly before entering the deferent duct. In 4. incusus the prostate 1s not nearly as long and it does narrow notice- ably before entering the deferent duct. Aegires pruvotfolae also bears some external resemblance to early descriptions of A. /euckar- tii. For example, Schmekel and Portmann (1982) reported both white and brown specimens of A. leuckartii. They found that the brown specimens had “somewhat symmetrically arranged, smooth, brown areas, which were not circular”. These spots are similar to those found on A. pruvotfolae. Risbec (1953) also considered Aegires pruvotfolae (formerly Aegires citrinus) a probable variation of A. leuckartii. However, there are external and internal differences that separate these two species: 1. The foot, veil and oral tentacles of A. /euckartii are bluish-white while in A. pruvotfolae the entire body, both dorsal and ventral surfaces are pale to muddy yellow. Aegires pruvotfolae has yellow rhinophores while the rhinophores of A. /euckartii have brown tips and 3 white circles (see Vérany 1853). 2. The dorsal tubercle arrangement differs between these two species. Aegires leuckartii has 4 large tubercles on the rhinophore sheath, whereas on A. pruvotfolae there are only 3. 3. The dorsal tubercle shape also differs between these two. Aegires leuckartii has cylindrical or round- ed tubercles while the tubercles of A. pruvotfolae are anvil or mushroom-shaped. 4. There is also a difference in the radular morphology between these two species. Schmekel and Portmann reported that the teeth of 4. /euckartii are all of similar size, whereas in A. pruvotfolae the inner- most radular tooth is the smallest, then there are 3-4 small teeth and the remainder are all the same size. 5. Vérany did not describe the reproductive morphology of 4. /euckartii but Schmekel and Portmann (1982) considered that the reproductive anatomy of A. leukartii was essentially as found with A. punctilucens. As such, there are differences between the reproductive morphology of A. punctilucens and A. pruvotfolae. We describe the morphology of A. pruvotfolae in an earlier section. But to summarize the differences, the prostate in A. punctilucens is rounded and in two parts. In A. pruvotfolae the prostate is very long, tubular and coiled. The vaginal duct in A. punctilucens is very short whereas in A. pruvotfolae it is very long. The recep- taculum seminis of A. punctilucens connects to the uterine duct on a short duct. But in 4. pruvotfolae this duct is very long and coiled. The differences in external and internal morphology between A. leuckartii, A. punctilucens, A. pruvotfolae and A. incusus distinguish these four as separate Aegires species. 628 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 Aegires gomezi Ortea, Luque, and Templado, 1990 (Figs. 1G, 12C—D, 13-15) Aegires gomezi Ortea, Luque, and Templado, 1990:333, Figs 1-2. | ) \ j j } i i t TYPE MATERIAL.— HOLortyPeE: 15.05/1034, La Habana, Cuba, Museo Nacional de Ciencias — Naturales of Madrid. MATERIAL EXAMINED.— HOLOTYPE: MNCN 15.05/1034, one specimen, 2 mm, La Habana, Cuba, collected July 1988, D. Moreno. CASIZ 077320, one specimen, 5 mm, dissected. Grand Cayman Island, no depth available, collected May 1991, T. Gosliner. CASIZ 077315, one speci- men, 3 mm, South Sound, Cayman Islands, no depth available, collected May 1991, T. Gosliner. LACM 2003-41.1, one specimen, 5 mm, dissected. Key Largo, Florida, collected July 2003, A. Valdés. DISTRIBUTION.— This species was reported from La Habana, Cuba in the original description (Ortea, Luque, and Templado 1990) and from Florida and the Cayman Islands (this study). EXTERNAL MORPHOLOGY.— The external morphology of the specimens collected in the Cayman Islands matches that of the original (1987) description of A. gomezi. The color photo of the Cayman specimens (Fig. 1C) matches the descriptions by Templado et al. (1987) and Ortea et al. (1990) DIGESTIVE SYSTEM.— The digestive system of the 5 mm specimen collected in Cayman Islands in 1991 was examined for the present study. The buccal bulb is nearly round, with the radu- lar sac protruding noticeably from the posterior end. There are two long salivary glands extending from under the esophagus. As found in other species of Aegires, there are four main muscles attached at the midpoint of the buccal bulb (Fig. 13). The radular formula of this specimen is 18 x 12.0.12 (Fig. 14). The teeth are simply hamate as reported for the holotype. The jaw has numerous rodlets (Fig. 14D). REPRODUCTIVE SYSTEM.— One of the specimens from the Cayman Islands was dissected for the present study. The reproductive system is triaulic. The ampulla is elongate. It branches into a short oviduct and the prostate (Fig. 15). The oviduct enters the female gland mass near an edge. FIGURE 13. Aegires gomezi Ortea, Luque, and Templado, 1990. CASIZ 077320. Buccal bulb: bb = buccal bulb, m = muscle, ot = oral tube, rs = radular sac, sg = salivary gland, scale = 0.13 FIGURE 12. Drawings of preserved animals. Aegires ortizi Templado, Luque, and Ortea, 1987. MNCN12-64/1006. (A) Dorsal view. (B) Ventral view of head. 9 ™™ Scale = 0.7 mm. (C) Aegires gomezi Ortea, Luque, and Templado, 1990. CASIZ 077320. Dorsal view. (D) Ventral view of head. Scale = 0.7 mm. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 629 FIGURE 15. Aegires gomezi Ortea, Luque, and Templado, 1990. CASIZ 077320. Reproductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, ps = penial spines, rs = receptaculum seminis, v = vagina, vd = vaginal duct, scale = 0.08 mm. 630 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 The prostate is a thick, coiled tube. It narrows into a small, coiled tube, then connects to a very wide ejaculatory duct and terminates at the glans penis. There are densely packed, small hooks inside the penis extending the entire length of the penial bulb. The vagina is long and narrow. It was not examined internally and thus the presence of spines or hooks cannot be confirmed. At its proximal end it joins the bursa copulatrix. A long separate duct joins the receptaculum seminis to the base of the bursa, and also connects to the female gland mass near the same edge as the oviduct. The bursa is large, spherical and the receptaculum seminis is about one-third the size of the bursa. CENTRAL NERVOUS SYSTEM.— The central nervous system was not available for examination. REMARKS.— Ortea et al. (1990) described this species from a single specimen collected in 1988 from a tidepool in La Habana, Cuba. Their description included the external and radular mor- phology, but the reproductive, central nervous system and digestive systems were not described. Aegires leuckartii Verany, 1853 Aegires leuckartii Verany, 1853:388. Aegires leuckartii (Verany) Bergh, 1883:135. Aegires leuckarti (Verany) Vayssiere 1901:5S5ff. Aegires leuckarti (Verany) Risbec, 1953: 60. Aegires leuckarti (Verany) Pruvot-Fol, 1954:245. Aegires leuckarti (Verany) Haefelfinger, 1960c:355. Aegires leuckarti (Verany) Schmekel, 1968b:116. Aegires punctilucens leuckarti, Schmekel and Portmann, 1982:102. TYPE MATERIAL.— Verany described this species from a single specimen collected from Nice, France. Attempts to locate the type specimen were unsuccessful. EXTERNAL MORPHOLOGY.— In the original description, the external morphology and col- oration of Aegires leuckartii was the only information provided by Verany of the specimen collect- ed. To summarize: The body shape of this species is wedge-shaped, with a straight or flat anterior and terminating in a pointed tail. There is no mantle edge and there is a large frontal veil with rounded lateral lobes. The rhinophore pocket is shaped like a large “horn” that is irregularly tri- lobed. The three gill leaves are very small, bipinnate and at the anterior and lateral edge of the gill pocket are large tubercles. These are conical and concave, larger along the median and smaller along the edge and scattered fairly regularly on the dorsum. The foot, the frontal veil, the oral ten- tacles are all bluish-white and the dorsum is yellowish-brown, light along the edges and darker in the center. There are brown spots on the dorsum. The tips of the rhinophores are brown with three white circles. Verany chose the particular spelling of /ewckartii for this new species. Subsequent authors, beginning with Vayssiere (1901) dropped the last “7”. However, the /nternational Code of Zoolog- ical Nomenclature allows for the Latinate spelling designated by the original author, thus /euckar- fii 1S correct. As mentioned previously, Pruvot-Fol (1930) described a specimen of A. pruvotfolae from New Caledonia. Risbec thought her specimen was A. /euckartii, similar to a specimen he found in New Caledonia. Both these specimens are most likely 4. pruvotfolae. See the comparison of these species under A. pruvotfolae. DISTRIBUTION.— This species is only found in the Mediterranean. REMARKS.— Schmekel and Portmann (1982) described Aegires punctilucens leuckartii from Nice, France. They reported external differences from Risbec’s New Caledonian specimens that he collected and identified as 4. punctilucens. (But, see the discussion under A. pruvotfolae.) The dor- FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 631 sal tubercles on the specimens from France were reported to be cylindrical and/or rounded, as described by MacFarland for A. a/bopunctatus, and not flat tubercles as reported by Risbec. Additionally, Schmekel and Portmann reported three tubercles arranged in a line in the mid-dorsal region, with three large tubercles on the outer edge of the rhinophore sheaths, and several smaller ones on the inner side. Risbec noted four large papillae on the rhinophore sheath. There are some differences in external coloration between Schmekel and Portmann’s speci- mens and Risbec’s. Risbec does not mention the opaque white dots scattered on the dorsum that Schmekel and Portmann observed and also noted by MacFarland for 4. albopunctatus. Both Risbec and Schmekel and Portmann report dark spots on the foot but MacFarland observed only white on the foot. The color of the rhinophores differs in that the specimens from France had a transverse brown ring, and the tips of the gill leaves were opaque white. Risbec reported only grayish rhinophores and translucent gill leaves and MacFarland reported lemon yellow rhinophores and gill leaves with spots. The observations of the radular teeth of specimens from the Mediterranean indicate that all teeth were of similar size. The radular formula reported for a 6 mm specimen was 16 x 15.0.15 (Schmekel and Portmann 1982). Bergh (1883) reported a radular formula of 17 x 15—16.0.15—16 for an 8 mm specimen from Trieste. Schmekel and Portmann’s description of Aegires punctilucens leuckartii most nearly matches that of A. albopunctatus MacFarland, whereas Risbec’s description of specimens collected from New Caledonia most nearly match that of 4. /euckartii Verany. Nordsieck (1972) listed 4. /euckartii as a synonym of A. punctilucens, but without any justifi- cation. The specimens he described externally match 4. punctilucens. Regarding the status of Aegires leuckartii: without having the description of the reproductive morphology from original material, it would be confusing to synonymize this species with 4. punc- tilucens (see Haefelfinger 1968). Descriptions of the background color of A. /euckartii are most similar to A. albopunctatus, not A. punctilucens. The body shape and tubercle arrangement of A. punctilucens as illustrated by d’Orbigny (1837) does not match the original description of A. leuckartii Verany. The radular formula of A. /euckartii noted by Risbec (1928) was 16 x 12.0.12 for a 6mm specimen and Schmekel and Portmann (1982) reported the formula for an 8 mm specimen of A. punctilucens leuckartii as 16 x 15.0.15. Schmekel and Portmann also reported that the teeth increase in size outwards, as they found in 4. punctilucens. Schmekel and Portmann also noted the difference in the gill structure between 4. leuckartii and A. punctilucens. The former has bipinnate gill leaves while the latter has a tripinnate structure. MacFarland described A. albopunctatus as having three small tripinnate gill leaves. Schmekel and Portmann did not agree with Haefelfinger’s synonymy and stated that the mor- phological and ecological differences they observed clearly distinguish 4. punctilucens from A. leuckartii. On the same basis, examination of A. albopunctatus indicates that it too, should be con- sidered distinct from these two as noted in the previous discussion of A. albopunctatus (present study). Photo images of specimens collected from the Mediterranean and identified as 4. leuckartii and A. cf. leuckartii are most likely A. leuckartii (Koehler 2004; Tocino 2004). Aegires palensis Ortea, Luque and Templado, 1990 is a probable synonym of A. /euckartii. Ortea, Luque, and Templado did not compare their specimen to A. /euckartii because they consid- ered A. leuckartii as a synonym of A. punctilucens. Risbec (1928, 1953) synonymized A. leuckartii with A. albopunctatus, in lengthy descriptions of specimens he collected from New Caledonia as discussed above. His specimens are most likely A. pruvotfolae as discussed below and in the Remarks section of A. pruvotfolae. 632 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES © Volume 55, No. 34 Externally, Risbec described his specimens as thin, wedge-shaped and elongate with a point- — ed posterior end of the foot. Aegires albopunctatus in contrast has a high, rounded dorsum with a | rounded tail (MacFarland 1966). The dorsal tubercles described and illustrated by Risbec are all © flattened and less numerous than those illustrated by MacFarland who reported both rounded and ~ flattened tubercles. Risbec illustrates a specimen with three distinct rows of dorsal tubercles, whereas MacFarland describes tubercles that are more numerous and larger toward the end of the dorsum behind the gill. MacFarland describes the tubercles as forming three rows, only behind the gill. Also, the large gill-protective tubercles on Risbec’s specimens are shown as much smaller and more simple than those illustrated by MacFarland. The external coloration also differs between Risbec’s specimens and Aegires albopunctatus. Risbec’s have grayish rhinophores and the rhinophores on A. a/bopunctatus are lemon yellow with tiny dots of white (MacFarland 1966). The gill plumes of Risbec’s specimens are pale yellow with tiny opaque white dots, while those of MacFarland have no dark pigment except for occasional spots (color unspecified by MacFarland). Risbec reports a faint greenish border on the foot, where- as MacFarland mentions white as the only color on the foot of A. albopunctatus. There are radular differences as well. The teeth of Risbec’s specimens (1928) have a straighter hook than does A. albopunctatus. Risbec reported that all radular teeth of his specimens were of similar size. But MacFarland noted that the teeth of A. a/bopunctatus gradually increased in size from the center, with the outermost tooth being smaller than the rest. The radular formula reported for Risbec’s specimen is 16 x 12.0.12 for a 6 mm specimen, whereas MacFarland reports a formu- la of 16-22 x 17.0.17 for A. albopunctatus specimens averaging 13 mm. Although both authors provided a very limited description of the reproductive system, they both noted that the penis is armed with many small hooks. It seems likely that the specimens described by Risbec were not Aegires albopunctatus MacFarland or Aegires punctilucens (d’Orbigny, 1837) but were A. pruvotfolae. The most distin- guishing feature of A. punctilucens is the “ocular” markings on the dorsum. These markings are not present in Risbec’s illustrations or discussion. Aegires ortizi Templado, Luque, and Ortea 1987 (Figs. 12A—B) Aegires ortizi Templado, Luque, and Ortea 1987:306, Figs 1-3. TYPE MATERIAL.— Holotype: 15-05/1006, Cayo Bocas de Alonso, Archipelago de los Conerneos, Cuba, Museo Nacional de Ciencias Naturales of Madrid. MATERIAL EXAMINED.— Holotype MNCN 15-05/1006, one specimen, 5 mm, Cayo Bocas de Alonso, Archipelago de los Canarreos, Cuba, 4 m, collected April 1984, J. Templado. DISTRIBUTION.— Aegires ortizi has only been reported from Cuba by the original authors. REMARKS.— Templado et al. (1987) described this species from four specimens collected in 1984 (holotype) and in 1988 (three additional specimens from Cuba). Their description included the external and radular morphology but the reproductive, central nervous system and digestive systems were not described. We were unable to examine the complete reproductive system or nery- ous system of the holotype, as the specimen appears to have been damaged during the removal of the buccal mass during the original study. The slide prep of the holotype radula was also examined. Additional material is needed to determine the extent of the pointed denticles on the lateral teeth of A. ortizi as described by the orig- inal authors. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 633 Aegires palensis Ortea, Luque, and Templado, 1990 (Figs. 16-17) Aegires palensis Ortea, Luque, and Templado, 1990:336, Figs 1-2. TYPE MATERIAL.— Holotype: 15.05/1035, Cabo de Palos, SE Spain, Museo Nacional de Ciencias Naturales of Madrid. MATERIAL EXAMINED.— Holotype MNCN 15.05/1035, one specimen, 6 mm, Cabo de Palos, Bajo de Dentro, Spain, collected July, 1987, J. Templado. MNCN 15.05/27821, one specimen, 2 mm, Aqua Amarga, Spain, 15 m, June 1995, J. Templado. DISTRIBUTION.— This species has only been reported from Spain (original description). EXTERNAL MORPHOLOGY.— Ortea et al (1990) provided a thorough description and drawings of this species. No additional materi- al was available for examination during the present study. However, drawings were made of the preserved paratype (Fig. 16). REPRODUCTIVE SYSTEM.— The reproduc- tive system examined here is triaulic (Fig. 17). The ampulla is large and bulbous. It branches into the oviduct and the prostate. The prostate is long and tubular and coils once, then narrows only slightly before entering the long, tubular deferent duct. The deferent duct enters the very wide, long penis. The penis of the specimen was broken off and thus the presence of penial hooks cannot be confirmed. The vagina 1s nar- row and was not examined internally. Thus the presence of spines or hooks cannot be con- firmed. The short, narrow vaginal duct enters the bursa copulatrix at the proximal end. The receptaculum seminis connects directly to the vaginal duct via a short duct that bifurcates into FIGURE MI uAcenest palcncin Orca elt cuctmantd the oviduct, which leads into the female gland Templado, 1990. MNCN15.05/1035. Reproductive system: mass. The bursa is nearly round and about two- am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm iaidsithe size of the ampulla. The receptacu- ~ male gland mass, p = penis, pr = prostate, rs = receptac- Mee : ; , ulum seminis, v = vagina, vd = vaginal duct, scale = 0.25 lum seminis is ovoid and is approximately one- half the size of the bursa. CENTRAL NERVOUS SYSTEM.— The central nervous system was not available for examination. REMARKS.— Ortea et al. described this species from a single specimen collected in 1987 at 34 m depth from SE Spain. Their description included the external and radular morphologies, but the reproductive, central nervous and digestive systems were not described. A second specimen was deposited into the Museo Nacional de Ciencias Naturales of Madrid in 1995 and examined for the present study. Photo images of specimens collected from the Mediterranean and identified as A. leuckartii and A. cf. leuckartii are most likely A. leuckartii and A. palensis may be a synonym of A. leuckartii (Koehler 2004; Tocino 2004). Additional material of both A. leuckartii and A. palen- sis is needed to determine whether this is the case. However, there have been no further reports on FIGURE 16. Aegires palensis Ortea, Luque, and Templado, 1990. MNCN15.05/1035. Drawing of preserved specimen. Dorsal view. Scale = 0.06 mm. 634 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 | collections or sightings of A. palensis except for the two specimens noted herein. No specimens of | A. leuckartii were available for examination during this study. Aegires punctilucens (d’Orbigny, 1837) Polycera punctilucens d’Orbigny, 1837:7, pl 106. Aegirus punctilucens (d’Orbigny) Alder and Hancock 1845/55, pl. 21. Aegires hispidus Hesse, 1872:346. Aegires punctilucens (d’Orbigny) Vayssiere 1901:58f. Aegires punctilucens (d’Orbigny) Pruvot-Fol, 1954:243. TYPE MATERIAL.— Collected (1826) at the Port of Brest, France. No further data are available. Attempts to locate the type material for examination were not successful. DISTRIBUTION.— Specimens of Aegires punctilucens have been reported from the Atlantic coast of France (d’Orbigny), Scandinavia and the Mediterranean Ocean (Pruvot-Fol 1954). EXTERNAL MORPHOLOGY.— D’Orbigny provided a thorough accounting of the external mor- phology of Aegires punctilucens. To summarize his description: the body is short but strongly con- vex, bulged in the middle, slightly tough and covered with flattened tubercles. The tubercles are arranged as follows: two between the rhinophores, four lateral and posterior to these, then one large median and many laterally placed tubercles. There is one large tubercle posterior to the gill, along with four round tubercles placed on each side of the tail median. Around the rhinophores are five elevated tubercles, three larger ones and two very small ones. The foot is elongate, ends in a point and is much narrower than the body, though slightly wider at the median. The gill is tri-lobed and multi-pinnate. The body color is a mixture of yellow and violet, except in between the tubercles where there are spots of bright green surrounded by a border of black. In front of the gill and between the rhinophores are definite points of matte white, with four smaller points lateral to these. On the foot there are many spots of color, more intense than the rest of the body color, placed obliquely or ver- tically. The bottom of the foot has a slight border. REMARKS.— Many authors have published descriptions of Aegires punctilucens, the species first described by d’Orbigny (1837) as Polycera punctilucens. The original description was con- fined to the external morphology. Subsequent authors (Schmekel and Portmann 1982; Thompson and Brown 1984; Templado et al. 1987) provided more thorough descriptions, including the radu- lar and reproductive anatomy, however none of the previous authors described the central nervous system or circulatory system. Baba (1974) provided a detailed description of a specimen from Japan, however this specimen is most likely Aegires exeches sp. nov. (See the description further on in the present study). Haefelfinger (1968) synonymized Aegires leuckartii with A. punctilucens. Schmekel and Portmann (1982) also noted the consistency between these two species, and designated them as subspecies. Thompson’s publication (1984) maintained the synonymy of the two (see discussion under 4. albopunctatus). As stated previously, there is no description of the reproductive morphology from the original material. Due to this lack, and the differences in the external morphology between A. punctilucens, A. leuckartii and A. albopunctatus and considering Schmekel and Portmann’s argument for sepa- rate subspecies, we propose to maintain 4. punctilucens from the Mediterranean Sea as a separate species from A. /euckartii and A. albopunctatus. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 635 Aegires sublaevis Odhner, 1932 (Figs. 18-25) 5 Aegires sublaevis Odhner, 1932:39. Serigea sublaevis (Odhner) Nordsieck, 1972:55. TYPE MATERIAL.— Puerto de Orotavo, Tenerife, Canary Islands. MATERIAL EXAMINED.— CASIZ 168921, four specimens, one specimen, 10 mm, dissected, Ponta de Piramede, Azores, no depth available, collected July 1988, T. Gosliner. CASIZ 078393, one specimen, 8 mm, dissected, Punta Cormorant, Isla Floreana, Galapagos Islands, no depth avail- able, collected September 1991, T. Gosliner. CASIZ 072608, three specimens, one specimen, 5 mm, dissected, Ila Sao Miguel, 1 km E of Caloura, Azores, 20 m, collected July 1988, T. Gosliner. CASIZ 072603, one specimen, 9 mm, Ila SAo Miguel, Mosteiros, Azores, 3 m, collected July 1988, T. Gosliner. CASIZ 168923, three specimens; one specimen, 12 mm, dissected. Ponta de Galora, Azores, collected July 1988, T. Gosliner. DISTRIBUTION.— This species has been reported from the Mediterranean Sea (Schmekel and Portmann 1982); the Canary Islands (Odhner 1932; Altimira and Ros 1979; Pérez Sanchez, Bacallado and Ortega 1991; Ortea et al. 1996; Ortea et al. 2000)); Panama (Meyer 1977), Bermuda (Thompson 1981) and the Galapagos Islands, which represents the first record from the Pacific (present study). EXTERNAL MORPHOLOGY.— The external morphology of this species has been described extensively in the literature (Odhner 1932; Meyer 1977; Altimira and Ros 1979; Thompson 1981; Schmekel and Portmann 1982; Templado et al. 1987) and will not be repeated here. However, drawings were made of preserved specimens from the Azores to compare to specimens from the type locality (Fig. 18). These specimens match Odhner’s original description. We noted one differ- 1d0A eon oss a ‘ ae, Ss i + i. + =a PE z ; FIGURE 18. Aegires sublaevis Odhner, 1932. CASIZ FicurE 19. Aegires sublaevis Odhner, 1932. Buccal 168921. Drawing of preserved specimen: (A) Dorsal view. bulbs: (A) CASIZ 072608. (B) CASIZ 078393, bb = buccal (B) Ventral view of head. Scale = 1.43 mm. bulb, cns = central nervous system, m = muscle, oe = esoph- agus, ot = oral tube, rs = radular sac, scale = 0.33 mm. 636 FiGuRE 20. Aegires sublaevis Odhner, 1932. CASIZ 072608. Buccal morphology: (A)Whole radula, scale = 100 um. (B) Inner lateral teeth, scale = 10 um. (C) Outer lateral § teeth, scale = 10 um. (D) Jaw, scale = 20 um. . PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 FIGURE 21. Aegires sublaevis Odhner, 1932. CASIZ 078393. Buccal morphology: (A) Whole radula, scale = 100 um. (B) Inner lateral teeth, scale = 10 um. (C) Outer lateral teeth, scale = 10 um. (D) Jaw, scale = 100 um. ence between specimens examined from the Azores and those from the Galapagos Islands. Specimens from the Azores have one ring of dark pigment on the rhinophores whereas those from the Galapagos have two dark rings. DIGESTIVE SYSTEM.— The arrangement of the digestive sys- tem is as illustrated for other Aegires (see Figs. 3B, 8B). The buc- cal bulb is rounded and the radular sac protrudes noticeably from the posterior side (Fig. 19). There were no oral glands noted. The radular formula is 17 x 17.0.17 for a 12 mm specimen from the Azores (CASIZ 168921)(Fig. 20) and 12 x 13.0.13 for an 8 mm specimen from the Galapagos (CASIZ 078393) (Fig. 21). The jaw is well developed and has a thickened edge. No labial rods were noted (Figs. 20D, 21D). All teeth are simply hamate, with a pointed hook. Rachidian teeth are absent. The first inner lateral tooth is much smaller than the remaining lateral teeth. The next two lateral teeth are slightly larger than the first lateral tooth. The outermost lateral tooth is shorter than the middle teeth. REPRODUCTIVE MORPHOLOGY.— The reproductive mor- phology of the specimens examined for the present study (Figs. 22-24) closely resembles the drawings and descriptions previ- ously reported by Schmekel and Portmann (1982). The reproductive system is triaulic. The ampulla is large and bulbous. It branches into the oviduct and the prostate. The prostate is long and tubular and coils twice, then narrows for a FIGURE 22. Aegires sublaevis Odhner, 1932. CASIZ 168921. Repro- ductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, Vv = vagina, vd = vaginal duct, vg = vestibular gland, scale = 0.33 mm. short distance before entering the wider deferent duct. The deferent duct enters the long penis, which is wider than the deferent duct. The penis has small densely packed penial hooks that extend FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 637 FIGURE 23. Aegires sublaevis Odhner, 1932. CASIZ 072608. Reproductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vd = vaginal duct, scale = 0.5 mm. [NB. Vestibular gland not evident. | throughout the length of the penis. The vagina is narrow and was not examined internally. Thus the presence of spines or hooks cannot be confirmed. The long, narrow vaginal duct enters the bursa copulatrix at the proximal end. The receptaculum seminis connects to the bursa with a longer duct than the oviduct, which leads from the bursa into the female gland mass. The bursa is round and as large as the ampulla. The receptaculum seminis is ovoid and less than one-half the size of the bursa. Specimens examined have a vestibular gland at the genital atrium (Figs. 22 and 24) as described by Schmekel and Portmann. However, in the specimen examined from the Azores, the vestibular gland was not evident. It may have broken off. No other Aegires species has this gland. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural ganglia (Fig. 25). The eyes are small and sessile on the cerebral-pleural complex, but protrude slightly at the sides of the cerebral-pleural com- plex. The pedal ganglia are slightly smaller FIGURE 24. Aegires sublaevis Odhner, 1932. CASIZ 078393. Reproductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagi- na, vd = vaginal duct, vg= vestibular gland, scale = 0.5 mm. Us “¥ ran) mae el cat det ~ eet ce Pinan as FIGURE 25. Aegires sublaevis Odhner, 1932. CASIZ 072608. Central nervous system: bg = buccal ganglia, cple = cerebro-pleural ganglia complex, p = pedal ganglia, r = rhinophoral nerve, vl+pe+ppe = visceral loop, scale = 0.2 mm. than the cerebral-pleural complex and they are joined by the visceral loop-pedal/parapedal com- missures. Three prominent nerves lead from the pedal ganglia and four nerves, including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two buccal ganglia are positioned ventral to the esophagus. 638 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 REMARKS.— The specimen from the Galapagos represents the first record of this species from the Pacific. With the exception of having two pigment rings on each rhinophore, Galapagos spec- imens are identical to Atlantic specimens of A. sublaevis. Odhner’s (1932) description of Aegires sublaevis is quite detailed but did not include a description of the reproductive system. However, Schmekel and Portmann (1982) provided a com- plete description of the reproductive anatomy in their thorough study. Templado et al. (1987) described the external and radular morphology of this species. In both these studies, drawings of the examined features accompany the description. Nordsieck (1972) established a new genus Serigea and designated S. sublaevis (Odhner, 1931) (with an incorrect date cited) as the type species. He placed Serigea in the Family Aegiretidae based on similar characters to Aegires. He described the following Serigea characters: dorsal papil- lae, smooth rhinophores, gill with protective valves, among others. However, Nordsieck did not indicate why a new genus was needed for these characters. Altimira and Ros (1978) published a study on the molluscs of the Canary Islands, declaring that Serigea equals Aegires sublaevis Odhner. Subsequent publications on Aegires sublaevis also recognized this name exclusively (Meyer 1977; Thompson 1981; Schmekel and Portmann 1982; Templado et al. 1987). Aegires villosus Farran, 1905 (Figs. 26-27) Aegires villosus Farran, 1905:329-364, pls 1-6. Aegires spp. Debelius, 1996:192, bottom large photo. Misidentification. TYPE MATERIAL.— Collected from the northwest of Cheval Paar, Ceylon. MATERIAL EXAMINED.— CASIZ 158799, one specimen, 5 mm, dissected. Luzon, Batangas, Philippine Islands, collected 6 May 2001, T. Gosliner. CASIZ 088089, one specimen, 12 mm, dis- sected. Layag Layag, Batangas, Philippine Islands, collected 27 March 1993, T. Gosliner. CASIZ 105658, one specimen, 8 mm, dissected. Kirby’s Rock, Batangas, Philippine Islands, collected 23 February 1995, T. Gosliner. CASIZ 65306, one specimen, 6 mm, Madang, Papua New Guinea, col- lected 22 January 1988, R. Willan. CASIZ 116888, one specimen, 12 mm, dissected. Coral Ledge, Garden Island, Western Australia, collected 6 January 1999, S. Fahey. DISTRIBUTION.— Aegires villosus has been reported from Ceylon (Farran 1905), Japan (Baba 1955), New Caledonia (Risbec 1928), Dar es Salaam, Tanzania (Edmunds 1971), Papua New Guinea (present study), Bali, Malaysia, Samoa and Australia (Rudman 2004). EXTERNAL MORPHOLOGY.— Several authors have described and/or drawn this species (Farran 1905; Risbec 1928; Baba 1955; Edmunds 1971; Rudman 2004). The specimens examined for the present study from various localities match the existing drawings and descriptions of the external morphology of Aegires villosus and no additional detail is necessary. DIGESTIVE SYSTEM.— The arrangement of the digestive system is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is ovoid and the large radular sac does not protrude noticeably from the posterior-ventral side (Fig. 26). There are numerous ovoid oral glands that cover the sides of the oral tube near the mouth and line the posterior edge of the oral tube. Two tubular salivary glands extend from the underside of the esophagus and lie along the top of the buc- cal bulb. The radular morphology was described by previous authors and the present study (Fig. 26) confirms the morphology as described by Farran (1905) and Risbec (1928). Edmunds (1971) described the outer lateral tooth as being elongate, but we did not find this to be the case. The radu- lar formula for the 12 mm specimen dissected is 16 x 17.0.17. All teeth are simply hamate, with the innermost tooth being substantially smaller than the remaining teeth in each row. The second FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 639 FIGURE 26. Aegires villosus Farran, 1905. CASIZ 088089. Buccal morpholo- gy: (A) Jaw, scale = 10 um. (B) Whole radula, scale = 100 um. (C) Inner lateral teeth, scale = 10 um. (D) Outer lateral teeth, scale = 10 um. (E) Buccal bulb, bb = buccal bulb, m = muscle, oe = esopha- gus, og = oral glands, ot = oral tube, rs = radular sac, sg = salivary gland, scale = 0.25 mm. lateral tooth is also reduced in size. REPRODUCTIVE SYSTEM.— The reproductive morphology has not been _ thoroughly described previously. The repro- ductive system is triaulic. The ampulla is small and ovoid. It branches into a short oviduct and fiec-aprostate (Fig. 27). The oviduct enters the female gland mass near an edge. The prostate is a very long, thick, coiled tube. It narrows slightly before widen- ing slightly into the ejaculatory duct and terminates at a slightly bulging penis. There are small hooks inside the penis located at the distal end only. The vagina is long and nar- row. It was not examined internally and thus the presence of spines or hooks cannot be con- firmed. At its proximal end it joins the bursa copulatrix. A long separate duct joins the large ovoid receptaculum seminis to the base of the bursa, and also connects to the female gland mass. The bursa is large, spherical and the receptaculum seminis is over half the size of the bursa. CENTRAL NERVOUS SYSTEM. The central nervous system is as found in other Aegires species. That is, it has fused cerebral and pleu- ral ganglia (not shown). The eyes are large and sessile on the cerebral-pleural complex and do not protrude. The pedal ganglia are slightly FIGURE 27. Aegires villosus Farran, 1905. CASIZ 088089. Reproductive system: am = ampulla, bc = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, ps= penial spines, rs = receptaculum seminis, v = vagina, vd = vaginal duct, scale = 0.3 mm. SAV AN ey 640 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 smaller than the cerebral-pleural complex and they are joined by the visceral loop-pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two buccal ganglia are positioned ventral to the esophagus. REMARKS.— Farran (1905) described Aegires villosus from one specimen collected from Cheval Paar, Ceylon (Sri Lanka). The description included the external morphology and radular morphology, along with adequate drawings of each. Other authors provided further details of the external anatomy, radular morphology, the central nervous system and/or the reproductive system (Edmunds 1971; Risbec 1928). Except for one difference, the specimens examined for the present study matched the previous descriptions in both external and internal morphology. Edmunds illus- trated and described elongate outer radular teeth in the 4 mm specimen he examined. He also stat- ed that the outermost tooth was very small and not illustrated in this immature specimen. The larg- er 8 mm specimen that he collected was not illustrated. The 8 and 12 mm specimens that we exam- ined had simply hamate outer lateral teeth. NEw SPECIES DESCRIPTIONS Family Aegiridae Fischer, 1883 Genus Aegires Lovén, 1844 Aegires ninguis Fahey and Gosliner, sp. nov. (Figs. 1D, 28-32) =Aegires sp. Gosliner, 1987:99, top photograph TYPE MATERIAL.— HOoLotyPe: CASIZ 073982, one specimen, 6 mm, Phillip’s Reef, Algoa Bay, Cape Province, Indian Ocean, South Africa, 10 m, collected January 1991, T. Gosliner. PARATYPES: CASIZ 073230, six specimens, 4-6 mm, Phillip’s Reef, Cape Province, Indian Ocean, South Africa, 10 m, collected May 1984, T. Gosliner. CASIZ 073929, one specimen, 8 mm, dis- sected, Llandudno, Cape Province, Atlantic Ocean, South Africa, 23 m, collected October 1982, T. Gosliner. A35568, one specimen, 6 mm, Miller’s Point, Indian Ocean, South Africa, 8 m, collect- ed June 1980, T. Gosliner. A35569, one specimen, 8 mm, Bakoven, Atlantic Ocean, South Africa, 15 m, collected September 1982, T. Gosliner. DISTRIBUTION.— This species has only been reported from the temperate Atlantic and Indian Oceans of South Africa (present study). ETYMOLOGY.— The specific name ninguis is Latin, meaning snowy, which describes the appearance of the dorsum with the white background and tiny opaque white speckles. EXTERNAL MORPHOLOGY.— The body shape is ovoid, slightly raised and has a rounded pos- terior end of the foot that extends only slightly (Fig. 1D). There is a slightly pronounced oral veil that has a scalloped edge (Fig. 28). The dorsum is covered with short, rounded tubercles, all with rounded tops. There are no tuber- cles on the posterior end of the foot. There are two prominent tubercles on the anterior of the head region, with smaller tuber- cles between them, and two FIGURE 28. Aegires ninguis sp. nov. CASIZ 073982. Drawing of preserved prominent tubercles on the poste- animal. (A) Dorsal view. (B) Ventral view of head. Scale = 1.5 mm. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 641 rior of the dorsum that appear joined together. There are also two prominent tubercles between the rhinophores with smaller tubercles between them. Numerous spicules protrude from the tops of the tubercles giving a fuzzy appearance. The rhinophore sheath is only slightly elevated but has five papillae, 3 large and 2 small, on all sides except the innermost. The rhinophores are smooth. The gill pocket lies at the posterior third of the dorsum and is protected on the anterior side by extra- FIGURE 29. Aegires ninguis sp. nov. CASIZ 073929. branchial papillae that are tri-lobed. The three Buccal bulb: bb = buccal bulb, cns = central nervous system, . ec m= muscle, oe = esophagus, ot = oral tube, rs = radular sac, small gill branches are bipinnate. sg = salivary glands, scale = 0.25 mm. The background color is white to pale yel- low. There are minute white speckles on the dorsum, between the tubercles. There are no addition- al colors found on the specimens examined. The rhinophores are pale yellow on white specimens and deeper yellow on the yellow specimens. The gill matches the background color of the living animal. DIGESTIVE SYSTEM.— The arrangement of the digestive system is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is rounded and the radular sac barely protrudes from the posterior side (Fig. 29). There are two short salivary glands situated beneath the esophagus. There were no oral glands noted. The radular formula is 17 x 16.0.16 for a 8 mm specimen (Fig. 30). The jaw (Fig. 30A) is well developed and has a thickened edge with narrow rod-like elements. All teeth are simply hamate, with pointed hooks. Rachidian teeth are absent. The four inner later- al teeth are smaller than the remaining lateral teeth. The outermost lateral teeth are shorter than the middle teeth. REPRODUCTIVE SYSTEM.— The reproductive system is tri- aulic (Fig. 31). The ampulla is short but very wide and branches into the oviduct and the prostate. The prostate is a long, thick tube and narrows only very slightly before entering the long, wide penis. The penis, which expands at the genital atrium, has small densely packed hooks only at the distal end. The vagina is long and wide. It was not examined inter- nally and thus the presence of spines or hooks cannot be con- firmed. At the proximal end, the short, wide vaginal duct enters the bursa copulatrix. The recep- taculum seminis connects direct- ; : : ee , FIGURE 30. Aegires ninguis sp. nov. CASIZ 073929. Buccal morphology: y to the vagina via a short duct. (A) Jaw, scale = 20 um. (B) Whole radula, scale = 100 um. (C) Inner lateral The oviduct, connected to the teeth, scale = 20 um. (D) Outer lateral teeth, scale = 20 1m. 642 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 FIGURE 31. Aegires ninguis sp. nov. CASIZ 073929. FIGURE 32. Aegires ninguis sp. nov. CASIZ 073929. Reproductive system: am = ampulla, be = bursa copulatrix, Central nervous system: bg = buccal ganglia, cple = cerebro- dd = deferent duct, fgm = female gland mass, p= penis, pr=_ pleural ganglia complex, p = pedal ganglia, r = rhinophoral prostate, ps = penial spines, rs = receptaculum seminis, v = nerve, vl+pce+ppe = visceral loop, scale = 0.14 mm. vagina, vd = vaginal duct, scale = 0.33 mm. receptaculum, enters the female gland mass. The bursa is round and slightly larger than the round receptaculum seminis. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural gan- glia (Fig. 32). The eyes are small and sessile on the cerebral-pleural complex, but protrude slight- ly at the sides of the cerebral-pleural complex. The pedal ganglia are slightly smaller than the cere- bral-pleural complex and they are joined by the visceral loop-pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two buccal ganglia are positioned ventral to the esoph- agus. CIRCULATORY SYSTEM.— (not pictured) The heart is relatively small as compared to most cryptobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Externally, the white form of Aegires ninguis most closely resembles the white form of 4. albopunctatus. Although neither color form (white or pale yellow) of A. ninguis has been found with dark spots on the dorsum, both forms do have minute white dots covering the dor- sum as is found on 4. albopunctatus. Both 4. ninguis and A. albopunctatus have yellow rhinophores. The tubercle arrangement is different between these two species. Aegires ninguis has randomly scattered tubercles that vary in size on the notum. There are two prominent tubercles on both the anterior and posterior ends of the notum. There are no tubercles on the posterior end of the foot. In 4. albopunctatus the tubercles are arranged in two distinct rows like ridges, joined from between the rhinophores and extending to the gill. Behind the gill, the tubercles are arranged in three distinct rows (MacFarland 1966). The tubercles on the rhinophore pocket of each species are different as well. In A. ninguis, there are four tubercles on the outside rhinophore pocket and in A. albopunctatus there are five. The gill morphology differs between the two species. Aegires ninguis has very small gill leaves that are almost completely covered by the large, flat-topped protective tubercles. Aegires FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 643 albopunctatus has larger gill leaves that protrude vertically with a branching lobe protecting each plume (MacFarland 1966: plate 18). Internally, the two species also differ. The most noticeable difference in the reproductive mor- phology is that in 4. ninguis the receptaculum seminis connects directly to the vagina via a short oviduct. In 4. albopunctatus the receptaculum connects to the bifurcating oviduct at the base of the bursa copulatrix and to the female gland mass. Also, 4. a/lbopunctatus has spines throughout the penis while in A. ninguis the spines are just at the apex. The deferent duct of these two species differs markedly. In A. ninguis the deferent duct is short and tubular. In 4. albopunctatus the duct is very long, thin and coiled. The penis of A. ninguis is very tubular and long, while that of A. a/bopunctatus is much smaller relative to the other repro- ductive organs such as the vagina and the bursa. No other species of Aegires has the particular combination of characters displayed by A. ninguis. The most externally similar species, Aegires albopunctatus is found along the Eastern Pacific while A. ninguis has been found only in South Africa. Aegires lemoncello Fahey and Gosliner, sp. nov. (Figs. 1E, 33-37) = Aegires sp. 2, Valda Fraser, 2000 and Rudman in SeaSlug Forum. TYPE MATERIAL.— HoLotyPe: CASIZ 086465, one specimen, 4 mm, dissected, Barracuda Point, Pig Island, Madang, Papua New Guinea, 9 m, collected June 1992, T. Gosliner. DISTRIBUTION.— This species has only been reported from Papua New Guinea (present study), South Africa (photo, V. Fraser) and Eastern Australia (no photo, B. Rudman). EtyMoLoGy.— The specific name /emoncello 1s from the Italian liqueur of the same color as some specimens of this new species. EXTERNAL MORPHOLOGY.— The body shape is elongate, slightly raised and has a very narrow posterior (Fig. IE). The dorsum has elongate papillae with flattened tops. Spicules protrude from the tops of the tubercles. The extended oral veil has 10—11 rounded tubercles on the dorsal surface (Fig. 33). The rhinophore sheath is slightly elevated, smooth and protect- ed by a single elongated papilla on the outer edge. The rhinophores are smooth. The gill pocket lies in the posterior third of the dorsum and is protected on the anterior side by five elongate papillae that project posteriorly. Posterior to the gill are two small papillae near the center-line. The three small gill branches are bipinnate. The background color is pale yellowish white to creamy yellow. The papillae are deep- er yellow, with a single ring of orange approximately half way along the length. The rhinophores are the same deeper yellow as the papillae. The gill branches are pale yellow. DIGESTIVE SYSTEM.— The arrangement of the digestive system is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is round and the radular sac barely protrudes from the FIGURE 33. Aegires lemoncello sp. nov. CASIZ 086465. Drawing of preserved animal. (A) Dorsal view. (B) Ventral view. Scale = 2.25 mm. 644 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES posterior side (Fig. 34). There are two elongate salivary glands situ- ated beneath the esophagus. There were no oral glands noted. The radular formula is 13 x 15.0.15 for a 4 mm specimen. The jaw is well developed and has a thickened edge (Figs. 34B, 35). All teeth are simply hamate, with a short hook. Rachidian teeth are absent. The three inner lateral teeth are much thinner and less hooked than the remaining lateral teeth. The outer lateral teeth have a short hook. REPRODUCTIVE SYSTEM.— The reproductive system is tri- aulic (Fig. 36). The ampulla is elongate and branches into the oviduct and the prostate. The prostate is long and tubular and narrows slightly before entering the elongate deferent duct. The deferent duct widens only slight- ly, then enters the short, wide penis. The penis has small dense- ly packed penial hooks at the dis- tal tip only. The vagina is much narrower than the penis. It was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the proximal end, the long, narrow vaginal duct enters the bursa copulatrix. The receptaculum seminis connects to the bursa with a separate long duct that Volume 55, No. 34 FIGURE 34. Aegires lemoncello sp. nov. CASIZ 086465. (A) Buccal bulb: bb = buccal bulb, m = muscle, oe = esophagus, ot = oral tube, rs = radular sac, sg = salivary gland, scale = 0.5 mm. (B) Labial cuticle, scale = 0.05 mm. a FIGURE 35. Aegires lemoncello sp. nov. CASIZ 086465. Buccal morpholo- gy: (A) Jaw, scale = 10 um. (B) Whole radula, scale = 20 um. (C) Inner later- al teeth, scale = 10 um. (D) Outer lateral teeth, scale = 20 um. bifurcates into the oviduct, which enters the female gland mass. The bursa is ovoid and approx1- mately the same size as the ovoid receptaculum seminis. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural gan- glia (Fig. 37). The eyes are large and sessile on the cerebral-pleural complex, but protrude slightly at the sides of the cerebral-pleural complex. The pedal ganglia are slightly smaller than the cere- bral-pleural complex and they are joined by the visceral loop-pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two buccal ganglia are positioned ventral to the esoph- agus. CIRCULATORY SYSTEM.— (not shown) The heart is relatively small as compared to most cryp- _——————— FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) tobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Externally, Aegires lemon- cello does not resemble any other Aegires species. Although A. /emoncello has elongate tubercles like Aegires villosus, the tubercles in that species are more numerous and more com- plex especially around the rhinophores where there are 4—5 tubercles on a raised pocket. The tubercles of 4. /emoncello look more like soft papillae, and there is only one tubercle that lies at the outer side of the rhinophores. The color of these two species is not similar at all. Aegires lemoncello has a yellow or whitish background color with orange rings around the rhinophores and tubercles. Aegires villosus has a white body color with purple and yellow irregular markings covering the notum. Internally, the morphological characters 645 FIGURE 36. Aegires lemoncello sp. nov. CASIZ 086465. also set this species apart from other Aegires. Aegires lemoncello has three elongate curved inner lateral teeth, a feature found in other Reproductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, ps = penial spines, rs = receptaculum seminis, v = vagina, vd = vaginal duct, scale = 0.3 mm. Aegires (A. ortizi and A. petalis). Aegires pru- votfolae has one elongate inner lateral tooth while both A. ortizi and A. petalis have two. Neither of these species even closely resembles A. /emoncello externally, either in color or tubercle morphology. The reproductive morphology also sets 4. lemoncello apart from other Aegires. The combination of a long, thin deferent duct leading from a narrow prostate and into a slightly wider penis is not found in other Aegires. The species that most closely matches this anatomy, A. hapsis, does not share any other internal or external characters. The large ovoid receptaculum that enters the common oviduct at the base of the bursa copulatrix is different from other Aegires species also. The receptaculum of other Aegires species either enters a common oviduct with the bursa, or enters the vaginal duct or into the vagina directly (A. ninguis ). FIGURE 37. Aegires lemoncello sp. nov. CASIZ 086465. Central nervous system: bg = buccal ganglia, cple = cerebro-pleural ganglia complex, p= pedal ganglia, r= rhinophoral nerve, cl+pc+ppe = visceral loop, scale = 0.25 mm. The combination of characters exhibited by Aegires lemoncello sets it apart as a previously undescribed species of Aegires. Aegires malinus Fahey and Gosliner, sp. nov. (Figs. 1F, 38-42) TYPE MATERIAL.— HoLotyPe: CASIZ 085889, one specimen, 8 mm, Bebbit, Batangas Region, Philippines, collected March 1993, T. Gosliner. PARATYPES: CASIZ 168919, one specimen, 646 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 15 mm, dissected, Bebbit, Philippines, collected March 1993, T. Gosliner. CASIZ 096248, one specimen, 14 mm, Layag-Layag, Batangas Region, Philippines, collected March 1994, M. Miller. DISTRIBUTION.— This species has only been reported from the Philippine Islands (present study). ETyMOLOGY.— The specific name malinus is Latin for apple green, the color of the rhinophore and gill appendages of this species. EXTERNAL MORPHOLOGY.— The body shape is elongate and convex. There are tiny pointed, randomly scattered tubercles on the dorsum (Figs. 1F, 38). The rhinophore pockets are raised and the outer edge is taller with 3 flat-topped tubercles of varying size and a lobed frill that resembles tufts of seaweed. The rhinophores are smooth. The gill pocket is protected by 3 tall, flat tubercles. The gill leaves are feathery, resembling tufts of seaweed and are multi-pinnate. They extend beyond the protective tubercles. The background color of the dorsum is deep reddish-brown. The raised rhinophore pocket is the same as the background color and the lobed frill and tubercles are apple green. The rhinophores are white with concentric bands of brown speckles around the club. The gill leaves are pale yellowish-green and the protective tubercles are bright apple green. DIGESTIVE SYSTEM.— The arrangement of the digestive system is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is elongate and the large radular sac protrudes noticeably from the posterior-ventral side (Fig. 39). There are numerous ovoid oral glands that extend from the ventral side of the oral tube. Two short salivary glands extend from the underside of the esophagus. The jaw is well developed and has a thickened edge with long rods along the edge (Fig. 40A). The radular formula for a 15 mm specimen is 22 x 8.0.8 (Fig. 40). The teeth are simply hamate, and the 3—5 innermost teeth much smaller and thinner than the remaining teeth. Rachidian teeth are absent. The outermost teeth are larger than the middle lateral teeth. REPRODUCTIVE SYSTEM.— The reproduc- tive system is triaulic (Fig. 41). The ampulla is very long and tubular. It branches into the oviduct and the prostate. The tubular prostate is approximately the same length as the ampulla and it coils once before narrowing into the thin, elongate deferent duct. The deferent duct is very long and widens slightly, then enters the long penis. The penis is wider than the deferent duct and it has small densely packed hooks at FIGURE 38. Aegires malinus sp. nov. CASIZ 085889. Drawing of preserved animal. (A) Dorsal view, scale = 1.88 mm. (B) Ventral view of head, scale = 1.88 mm. (C) View of rhinophore pocket from inner edge, scale = 0.5 mm. (D) View of rhinophore pocket from outer edge, scale = 0.5 mm. FIGURE 39. Aegires malinus sp. noy. CASIZ 168919. Buccal bulb: bb = buccal bulb, cns = central nervous system, m = muscle, oe = esophagus, og = oral glands, ot = oral tube, rs = radular sac, scale = 0.4 mm. I FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 647 the distal tip only. The vagina 1s narrower than the penis. It was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the proximal end, the long, narrow vaginal duct enters the bursa cop- ulatrix. The receptaculum semin- is connects to the bursa via a sep- arate longer duct that bifurcates into the oviduct, which enters the female gland mass. The bursa is ovoid and slightly smaller than the ovoid receptaculum seminis. CENTRAL NERVOUS SYS- TEM.— The central nervous sys- tem has fused cerebral and pleu- ral ganglia (Fig. 42). The eyes are large and sessile on the cere- bral-pleural complex and do not protrude. The pedal ganglia are slightly smaller than the cerebral- pleural complex and they are joined by the visceral loop- pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves including the rhinophoral nerve, originate from the cere- (A) Jaw rodlets, scale = 10 um. (B) Whole radula, scale = 30 wm. (C) Inner bro-pleural ganglia. Two buccal _ lateral teeth, scale = 10 pm. (D) Outer lateral teeth, scale = 20 pm. ganglia are positioned ventral to the esophagus. CIRCULATORY SYSTEM.— (not shown) The heart is relatively small as compared to most cryptobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Externally, Aegires malinus does not resemble any other Aegires species in either color or body texture. Along with A. hap- sis, it is one of the two Aegires species that does not have prominent dorsal tubercles. However, like other Aegires, the rhinophore pockets are lobed, particularly on the outer edge. But the lobes are distinctly different from the texture of the notum. The gill is also pro- tected by appendages, as with other Aegires, FiGuRE 41. Aegires malinus sp. nov. CASIZ 168919. Reproductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, ps = penial spines, rs = receptaculum seminis, v = vagina, vd = vaginal duct, scale = 0.86 mm. 648 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 but in A. malinus the appendages are tall, nar- eplc1-4 row, yet shorter than the gill leaves. This is unusual for Aegires, in which the gill is normal- [sf ly small and almost hidden by the appendages. ‘ f Internally, A. malinus shares some repro- ductive characters with other Aegires species, such as a bursa and receptaculum that are near- ly equal in size, a tubular coiled prostate and an elongate penis with spines. The ampulla in A. malinus is extremely elongate, as is found in 4. petalis. But in A. petalis, the penial morpholo- gy is different, in that the penis is very short and wide as compared to A. malinus, in which it is elongate. Externally, these two species share no common features or color. Aegires malinus has numerous oral glands clustered at the base of the oral tube. Aegires FIGURE 42. Aegires malinus sp. nov. CASIZ 168919. villosus also has clustered oral glands, but in ©®™4! nervous system: bg = buccal ganglia, eple — cerebro; ; = pleural ganglia complex, p = pedal ganglia, r = rhinophoral that species the glands are more numerous and jee. vLupe bppe = visceral loop! scale Oe imme there is a second row of glands along the oral tube (see Fig. 26). The other species in which oral glands were observed are A. incusus and A. hap- sis. But the glands in those two species have a different morphology (see Figs. 44 and 63). The particular combination of characters sets Aegires malinus apart as a new, previously unde- scribed species. Aegires incusus Fahey and Gosliner, sp. nov. (Figs. 1G, 43-47) Aegires citrinus Nakano, 2004:117, no. 242, bottom left photograph. Misidentification. = Aegires sp. 1 Rudman, 2004, leading photo for Aegires in SeaSlug Forum. TYPE MATERIAL.— HOovoryPe: CASIZ 070357, one specimen, 4 mm. Patch Reef, Mora Mora, Tulear, Madagascar, collected April 1989, T. Gosliner. PARATYPES: CASIZ 156668, one specimen, 5 mm, dissected, Cemetary Beach, Maricaban Island, Luzon, Philippines, 0.5 m, collected May 2001, Y. Camacho. DISTRIBUTION.— This species has only been reported from Madagascar, Philippine Islands (Rudman 2004 and present study) and from Japan (Nakano 2004). ETYMOLOGyY.— The specific name incusus is taken from the Latin word meaning anvil, the shape of many of the tubercles of this new species. EXTERNAL MORPHOLOGY.— The body shape is elongate and slightly rounded (Fig. 1G). There is no obvious oral veil as is found in other Aegires species. The posterior third of the body is much narrower than the rest of the body. The dorsum has tall, anvil or mushroom-shaped tubercles with flat tops. There are smaller tubercles along the edge of the posterior end of the foot and one promi- nent tubercle on the tip. Spicules protrude from the tops of all tubercles. The rhinophore pocket is slightly elevated and is usually surrounded by three tubercles that vary in size. The rhinophores are smooth. The gill pocket lies in the posterior third of the dorsum and three tubercles protect the ante- rior side of the gill pocket. The small gill leaves are bipinnate. The background color is creamy white and there are a few rounded blotches of tan or light FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) brown spots between some of the tubercles. The tubercle stalks are white and the flattened tops are either all white or all tan. The rhino- phores are creamy white on the specimens with white tubercle tops, or are yellowish on speci- mens with tan tubercle tops. The gill leaves are white. DIGESTIVE SYSTEM.— The arrangement of the digestive system is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is rounded and the large radular sac protrudes noticeably from the posterior-ventral side (Fig. 44). There are two large pear-shaped oral glands that extend from the ventral side of the buccal bulb. The radular formula for a 5 mm specimen is 16 x 13.0.13 (Fig. 45). The jaw (Fig. 45A) is not well developed but has a thickened edge. The teeth are simply hamate, and the innermost tooth is much smaller and thinner than the remaining teeth. Rachidian teeth are absent. The outermost teeth are slight- ly larger than the middle lateral teeth. REPRODUCTIVE SYSTEM.— The reproduc- tive system is triaulic (Fig. 46). The ampulla is large and bulbous. It branches into the oviduct and the prostate. The prostate is long and tubu- lar and coils twice, then narrows for a short dis- tance before entering the wider deferent duct. The deferent duct enters the long penis, which is as wide as the deferent duct. The penis has small densely packed penial hooks at the distal tip only. The vagina is narrow and was not examined internally. Thus, the presence of spines or hooks cannot be confirmed. The short, narrow vaginal duct enters the bursa cop- ulatrix at the proximal end. The receptaculum seminis connects to the bursa with a longer duct than the oviduct, which leads from the bursa into the female gland mass. The bursa is round and half the size of the ampulla. The receptaculum seminis is ovoid and is approxi- mately one-half the size of the bursa. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural ganglia (Fig. 47). The eyes are mid-sized in comparison to other Aegiridae and are sessile on the cerebral-pleural complex. The pedal 649 FIGURE 43. Aegires incusus sp. nov. CASIZ 070357. Drawing of preserved animal. (A) Dorsal view. (B) Ventral view of head. Scale = 0.57 mm. FIGURE 44. Aegires incusus sp. nov. CASIZ 156668. Buccal bulb: bb = buccal bulb, m = muscle, oe = esophagus, og = oral gland, ot = oral tube, rs = radular sac, scale = 0.25 mm. 650 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 ganglia are slightly smaller than the cerebral-pleural complex and they are joined by the visceral loop-pedal/parapedal commis- sures. Three prominent nerves lead from the pedal ganglia and four nerves, including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two buccal ganglia are posi- tioned ventral to the esophagus. CIRCULATORY SYSTEM.— (not pictured) The heart is rela- tively small as compared to most cryptobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Aegires incusus is externally most simi- lar to Aegires pruvotfolae. For a comparison between these two species, see the section under Aegires pruvotfolae in the pres- ent study. The particular combi- FIGURE 45. Aegires incusus sp. nov. CASIZ 156668. Buccal morphology: (A) Jaw, scale = 3 um. (B) Whole radula, scale = 20 um. (C) Inner lateral teeth, scale = 10 um. (D) Outer lateral teeth, scale = 10 um. FIGURE 46. Aegires incusus sp. noy. CASIZ 156668. FIGURE 47. Aegires incusus sp. nov. CASIZ 086465. Reproductive system: am = ampulla, be = bursa copulatrix, Central nervous system: bg = buccal ganglia, cple = cerebro- dd = deferent duct, fgm = female gland mass, p = penis, ps_ pleural ganglia complex, p = pedal ganglia, r = rhinophoral = penial spines, pr = prostate, rs = receptaculum seminis, v_ nerve, vl+pe+ppce = visceral loop, scale = 0.25 mm. = vagina, vd = vaginal duct, scale = 0.25 mm. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 651 FiGURE 48. Photos of living specimens. (A) Aegires flores sp. nov. CASIZ 084277, photo by T. Gosliner, 5 mm. (B) Aegires flores sp. nov. CASIZ 078568, photo by P. Fiene, 9 mm. (C) Aegires flores sp. nov. CASIZ 120931, photo by S. Johnson, 12 mm. (D) Aegires petalis sp. nov. CASIZ 168920, photo by T. Gosliner, 5 mm. (E) Aegires exeches. CASIZ 078629, photo by P. Fiene, 4 mm. (F) Aegires hapsis sp. nov. CASIZ 115721, photo by R. Bolland, 6 mm. (G) Aegires sub- laevis. CAS1Z168923, photo by T. Gosliner, 12 mm. (H) Aegires sublaevis CASIZ, 078393, photo by T. Gosliner, 8 mm. 652 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 nation of external, reproductive and radular characters distinguish A. incusus as a distinct Aegires species. Aegires flores Fahey and Gosliner, sp. nov. (Figs. 48A—C, 49-53) Aegires citrinus Nakano, 2004:117, no. 242, top and right photograph. Misidentification. = Aegires sp. 3, Tanaka, 2001 in SeaSlug Forum. TYPE MATERIAL.— HOLOTYPE: CASIZ 120931, one specimen, 11 mm, Choptop Reef, Enewetak Atoll, Marshall Islands, 8 m, collect- ed August 1988, S. Johnson. PARATYPES: CASIZ 120930, one specimen, 13 mm, dissect- ed, Bubble Butt Pinnacle, Enewetak Atoll, Marshall Islands, collected September 1983, S. Johnson. CASIZ 073059, one specimen, 9 mm, Pinnacle, Madang, Papua New Guinea, 5 m, collected, Oct, 1986, T. Gosliner. CASIZ 168922, Outer Reef, Bagabag Island, Papua New Guinea, collected July 1989, T. Gosliner. CASIZ 075909, one specimen, 4 mm, Barracuda Point, Pig Island, Papua New Guinea, 32 m, collected November 1990, T. Gosliner. CASIZ 075877, two specimens, 4 and 7 mm, dissected, Barracuda Point, Pig Island, Madang, Papua New Guinea, 34 m, collected November 1990, T. Gosliner. CASIZ 078568, one specimen, 9 mm, Bunaken Island, Manado, Sulawesi, 3 m, collected May 1991, P. Fiene. CASIZ 084277, one specimen, 5 mm, Devil’s Point, Maricaban Island, Luzon, Philippines, 10 m, collected February 1992, T. Gosliner. CASIZ SIS 715329 one specimen. 0M mm: Bethlehem, Maricaban Island, Luzon, Philippines, 7 m, collected May 2001, T. Gosliner. DISTRIBUTION.— This species has been reported from the Marshall Islands, Papua New Guinea, Sulawesi, the Philippine Islands (pres- ent study), and from Japan (Nakano and Tanaka 2004). ETYMOLOGy.— The specific name flores is from the Latin word meaning blossom, refer- ring to the shape of the rhinophore and gill pro- tective appendages of this species. FIGURE 49. Aegires flores sp. nov. CASIZ 120930. Drawing of preserved animal. (A) Dorsal view. (B) Ventral view of head. Scale = 1.3 mm. FiGuURE 50. Aegires flores sp. nov. CASIZ 120930. Buccal bulb: bb = buccal bulb, rs = radular sac, oe = esoph- agus, scale = 0.25 mm. EXTERNAL MORPHOLOGY.— The body shape is elongate and slightly rounded (Figs. 48A—C, 49). The dorsum is completely covered by small raised tubercles, some with flat tops and some FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 653 with rounded tops. Spicules pro- trude from the tops of all tuber- cles. The rhinophore pocket is slightly elevated and is surround- ed by approximately five flat- topped tubercles that vary in size. The tubercles on the exterior edge of the rhinophore pocket are large and rounded and look like flower petals. The rhinophores are smooth. The gill pocket lies mid dorsally and is surrounded by approximately twelve flat- tened, paddle-shaped tubercles in varying sizes, arranged like a crown or an opening flower. The small gill leaves are tripinnate. The background color of the living animals is cream. The overlying colors range from ar s oe Oe eae acer eee 2 % IGURE 51. Aegires flores. sp. nov. . Buccal morphology: panei with elon topped (A) Jaw, scale =100 um. (B) Whole radula, scale = 100 wm. (C) Inner lateral tubercles to brown with Orange- teeth, scale = 20 um. (D) Outer lateral teeth, scale = 20 jm. topped tubercles. The paddle- shaped tubercles around the gill can be translucent white with white tips or translucent yellow with darker yellow tips. The rhinophores in all color forms are translucent white as are the gill leaves. DIGESTIVE SYSTEM.— The arrangement of the digestive system is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is more ovoid than round and the radular sac protrudes from the posterior side only very slightly (Fig. 50). The radular formula is 18 x 17.0.17 for a 7 mm FIGURE 52. Aegires flores sp. nov. CASIZ 120930. FiGURE 53. Aegires flores sp. nov. CASIZ 086465. Reproductive system: am = ampulla, be = bursa copulatrix, Central nervous system: bg = buccal ganglia, cplc = cerebro- dd = deferent duct, fgm = female gland mass, p = penis, pap pleural ganglia complex, p = pedal ganglia, r = rhinophoral = papillae, ps = penial spines, pr = prostate, rs = receptacu- nerve, vl+pct+ppc = visceral loop, scale = 0.3 mm. lum seminis, v = vagina, vd = vaginal duct, scale = 0.4 mm. 654 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 specimen (Fig. 51). The jaw (Fig. 51A) is not well developed but has a thickened edge. All teeth are simply hamate, with a short hook. Rachidian teeth are absent. The oldest teeth have a longitu- dinal groove on the outer side, a slight bulge at the posterior midpoint and a longer hook than the remaining teeth. REPRODUCTIVE SYSTEM.— The reproductive system is triaulic (Fig. 52). The ampulla is short and wide. It branches into the oviduct and the prostate. The prostate is relatively short, tubular and curves once before narrowing into the short deferent duct. The deferent duct then widens into the penis, which has small densely packed penial hooks at the distal tip. There are also several rows of papillae inside the penis, at the proximal end. The vagina is narrow and was not examined inter- nally. Thus the presence of spines or hooks cannot be confirmed. The vaginal duct is long and nar- row and at the proximal end, enters a common duct that connects the bursa copulatrix and the receptaculum seminis. The receptaculum is connected to the oviduct, which enters the female gland mass. The bursa is ovoid and much larger than the ampulla. The receptaculum is ovoid and is approximately one-third the size of the bursa. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural gan- glia (Fig. 53). The eyes are small, sessile on the cerebral-pleural complex, but protrude slightly. The pedal ganglia are slightly smaller than the cerebral-pleural complex and they are joined by the vis- ceral loop-pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves, including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two buc- cal ganglia are positioned ventral to the esophagus. CIRCULATORY SYSTEM.— (not shown) The heart is relatively small as compared to most cryp- tobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Aegires flores has a unique external morphology. Although it has an elongate body, a rounded posterior end of the foot, a tuberculate notum, protective tubercles around the rhinophores and gill, and smooth rhinophores, there are several unique characters that set this species apart. Most noticeably, the gill appendages have a unique paddle shape. The tubercles on the outside of the rhinophore pockets also resemble those around the gill, both in shape and in color. The dorsal tubercles are all much lower than the protective tubercles, a feature not shared by other Aegires species. The reproductive morphology is also different from other Aegires species. The short, wide ampulla with a short deferent duct is a combination of characters not seen in other Aegires species. These features combined with the very short duct leading from the receptaculum to the bursa are not found in other Aegires. The unusual combination of external and internal characters distinguishes Aegires flores as a new species. Aegires petalis Fahey and Gosliner, sp. nov. (Figs. 48D, 54-57) TYPE MATERIAL.— HoLotyre: CASIZ 168920, one specimen, 5 mm, dissected, Anemone Reef, Madang, Papua New Guinea, collected February 1988, T. Gosliner. DISTRIBUTION.— This species has only been reported from Papua New Guinea (present study). ETYMOLOGyY.— The specific name pevalis is taken from the Greek word petalon meaning a leaf or petal. This is in reference to the rhinophore sheaths of this new species. EXTERNAL MORPHOLOGY.— The body shape is elongate and the dorsum is high (Fig. 48D). FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 655 Flat-topped raised tubercles are arranged in two lines along the anterior half of the dorsum. On the posterior half of the dorsum, the largest tubercles are arranged in a diamond pattern. Additional large, flat-topped tubercles line the edge of the dorsum. Long spicules protrude from the tops of all tubercles (Fig. 54). The rhinophore pocket is tall and cylindrical and has a petal-like outer edge. The rhinophores are smooth. The gill pocket lies mid dorsally and is protected by three appendages that have flat- tened tops. The small gill leaves are tripinnate. The background color of the living animal is white, as are the tubercles, the rhinophores and the gill leaves. There are no additional col- ors on the specimen examined in this study. . ee DIGESTIVE SYSTEM.— The arrangement of KO Re the digestive system is as illustrated for other ae a Aegires (see Figs. 3B, 8B). The buccal bulb is DY more ovoid than round and the radular sac pro- ST Seay trudes from the posterior side only very slight- FIGURE 54. Aegires petalis sp. nov. CASIZ 168920. Us es =>): The radular formula is 13x Drawing of preserved animal. (A) Dorsal view. (B) Ventral i013) for the 5mm holotype (Figs 56). The view of head. Scale= 0:7 mm. jaw (Fig. 56A) is not well developed but has a thickened edge. All teeth are simply hamate, with a short hook. Rachidian teeth are absent. The three inner lateral teeth are thinner and smaller than the remaining teeth. The oldest teeth have a shallow longitudinal groove on the outer side. REPRODUCTIVE SYSTEM.— The reproduc- tive system is triaulic (Fig. 57). The ampulla is elongate and narrow. It branches into the oviduct and the prostate. The prostate is long, FIGURE 55. Aegires petalis sp. nov. CASIZ 168920. tubular and coils once before narrowing into Buccal bulb: bb = buccal bulb, cns = central nervous system, the long, thin deferent duct. The deferent duct ea esophagus, ot = oral tube, rs = radular sac, scale = 0.3 then widens into the short but wide penis, : which has small, densely packed hooks at the distal tip only. The vagina is narrow and was not examined internally. Thus the presence of spines or hooks cannot be confirmed. The vaginal duct is short and narrow and at the proximal end, enters the bursa copulatrix. A duct from the bursa enters the bifurcating oviduct that originates from the receptaculum. The oviduct enters the female gland mass. The bursa is ovoid and smaller than the round receptaculum. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural gan- glia (not pictured). The eyes are small, sessile on the cerebral-pleural complex, and do not protrude. The pedal ganglia are slightly smaller than the cerebral-pleural complex and they are joined by the visceral loop-pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two 656 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 buccal ganglia are positioned ventral to the esophagus. CIRCULATORY SYSTEM.— (not shown) The heart is relative- ly small as compared to most cryptobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Externally, Aegires petalis most closely resembles A. ortizi. Both species are either white or creamy yellow (A. ortizi) and have dorsal tuber- cles. However, the tubercles of A. ortizi are arranged in four longi- tudinal rows (Templado et al. : : FIGURE 56. Aegires petalis sp. nov. CASIZ 168920. Buccal morphology: 1987) whereas in A. petalis the (A) Jaw, scale =10 wm. (B) Whole radula, scale = 20 um. (C) Inner lateral tubercles are randomly scattered. teeth, scale = 10 xm. (D) Outer lateral teeth, scale = 10 pm. Aegires ortizi also has brown col- oration and minute white spots between the tubercles, which A. petalis does not have. There are three large tubercles on the outside of the rhinophore sheaths of A. ortizi but the rhinophore margins of A. petalis are elevated and multi-lobed. The three gill appendages of A. petalis have flattened tops, but in 4. ortizi the appendages are pointed. The reproductive morphology differs between Aegires petalis and A. ortizi. Although both have an elongate ampulla, the ampulla of A. petalis is relatively longer. The penial mor- phology is noticeably different. The penis of A. petalis is short and thick, and that of 4. ortizi is FIGURE 57. Aegires petalis sp. nov. CASIZ 168920. Reproductive system: am = ampulla, be = bursa copulatrix, very long and wide. In addition, A. petalis only dd = deferent duct, fgm = female gland mass, p = penis; pr = has penial spines at the distal tip while they are __ prostate, ps = penial spines, rs = receptaculum seminis, v = found throughout the penis of A. ortizi. The Y28% v4 = vaginal duct, scale = 0.5 mm. prostate of A. ortizi is also very thick and tubular, whereas in A. pefalis it is much thinner and coiled. The receptaculum of A. peralis is larger than the bursa and connects to the common oviduct. But in A. ortizi the receptaculum is much smaller than the bursa and connects on a very long sep- arate duct to the base of the bursa. This combination of morphological characters sets A. petalis apart from other species of — Aegires. Aegires exeches Fahey and Gosliner, sp. nov. (Figs. 48E, 58-61) Aegires punctilucens Nakano, 2004:117, no. 243. Misidentification. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 657 Aegires punctilucens Imamoto 2002: photos, SeaSlug Forum. Misidentification. Aegires punctilucens Ono, 1999:71, no. 103. Misidentification. Aegires cf. punctilucens Marshall and Willan, 1999:59, pl. 93. Misidentification. Aegires punctilucens Baba, 1974:11-12, Fig. 1. Misidentification. TYPE MATERIAL.— HOLOTYPE: CASIZ 078629, 4 mm, Hekili Point, Maui, Hawaii, 1 m, col- lected May 1991, C. Pittman. PARATYPE: CASIZ 168918, 4 mm, dissected, Hekili Point, Maui, Hawaii, 1 m, collected May 1991, C. Pittman. PARATYPES: CASIZ 109790, 3 mm, Pig Island, Madang, Papua New Guinea, no depth available, collected October 1996, T. Gosliner. CASIZ 068750, 2 mm, Jais Aben, Madang, Papua New Guinea, 15 m, collected July 1989, T. Gosliner. CASIZ 075804, 2 mm, between Wongat and Tabat Islands, Madang, Papua New Guinea, 23 m, col- lected November 1990, T. Gosliner, G. Williams, M. Jebb. CASIZ 079183, 2 mm, Horseshoe Cliffs, Onna Village, Okinawa, 5 m, collected June 1991, T. Gosliner. CASIZ 115394, 4 mm, Maeki-zaki, Seragaki, Okinawa, 46 m, collected, April 1997, R. Bolland. DISTRIBUTION.— This species has been reported from Hawaii, Japan, Papua New Guinea (present study), the Marshall Islands (S. Johnson photo) and the Great Barrier Reef, Australia (Marshall and Willan 1999). EtyMoLoGy.— The specific name exeches is from the Greek exechos meaning jutting out, projecting, prominent, which refers to the shape of the tubercles on this species. EXTERNAL MORPHOLOGY.— The body shape is elongate and the posterior end of the foot ends in a point (Fig. 48E). There are numerous compound tubercles projecting from the dorsum, such that the body appears to be composed completely of tubercles (Fig. 58). The tubercles are elongate and narrow slightly before mushrooming into a flattened plate-like top. From the flattened tops, multiple spicules protrude. The rhinophore pockets are very long and are composed of two main tubercles on the anterior side, two shorter tubercles on the posterior side and much smaller tuber- cles in between. The rhinophores are smooth and have bifid apices. The gill protective appendages are also composed of elaborate tubercles. The gill leaves themselves are small, inconspicuous and tripinnate. ar 4 FIGURE 58. Aegires exeches sp. nov. CASIZ 168918. FIGURE 59. Aegires exeches sp. nov. CASIZ 168918. Drawing of preserved animal. (A) Dorsal view. (B) Ventral Buccal bulb: bb = buccal bulb, oe = esophagus, ot = oral view of head. Scale = 0.7 mm. tube, rs = radular sac, scale = 0.17 mm. 658 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 The background color ranges from white to tannish- white. The tops of the tubercles on the tan specimens have dark spots. The specimen from Enewetak, Marshall Islands is completely white. It has three evenly spaced, pale tan rings around the rhinophores. The tan specimens have three to four blue spots on the dorsum. These spots are arranged symmetrically, with two just posterior to the rhino- phores, one at the centerline in front of the gill and the fourth at centerline just posterior to the gill. DIGESTIVE SYSTEM— The arrangement of the digestive sys- tem is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is more ovoid than round and the radular sac pro- trudes from the posterior side only very slightly (Fig. 59). The radular formula is 13 x 11.0.11 for the 4 mm specimen (Fig. 60). The jaw is not well developed but has a thickened edge (Fig. oy Ww Sa 60A). All teeth are simply FIGURE 60. Aegires exeches sp. noy. CASIZ 168918. Buccal morphology: hamate, with a short hook. (A) Jaw. Scale =100 um. (B) Whole radula. Scale = 100 pm. (C) Inner lateral Rachidian teeth are absent. The _ teeth. Scale = 20 um. (D) Outer lateral teeth. Scale = 20 pm. three inner lateral teeth are thin- ner and smaller than the remaining teeth. The oldest teeth have a shallow longitudinal groove on the outer side. REPRODUCTIVE SYSTEM.— The reproductive system ts triaulic (Fig. 61). The ampulla is a very long, thick tube. It branches into the oviduct and the prostate. The prostate is shorter, but also tubu- lar and does not coil before narrowing into the very long, thin deferent duct. The deferent duct coils once near the entry to the wide penis, which has small irregularly spaced hooks along the entire length of the penis. The vagina is narrow and was not examined internally. Thus the presence of spines or hooks cannot be confirmed. The vaginal duct is long and narrow and at the proximal end, enters the bursa copulatrix. The bifurcating oviduct, which originates from the small ovoid recep- taculum seminis, enters the vaginal duct. The oviduct enters the female gland mass. The bursa is small and nearly round. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural gan- glia (not pictured). The eyes are small, sessile on the cerebral-pleural complex, and do not protrude. The pedal ganglia are slightly smaller than the cerebral-pleural complex and they are joined by the -~ FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) visceral loop-pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves, including the rhinophoral nerve, originate from the cerebro- pleural ganglia. Two buccal ganglia are posi- tioned ventral to the esophagus. CIRCULATORY SYSTEM.— (not shown) The heart is relatively small as compared to most cryptobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Externally, Aegires exeches most closely resembles A. punctilucens from the Mediterranean. However, there are both external and internal differences that separate these two species. The most obvious external differences are the body shape, rhinophoral apices, gill protective structure morphology and tubercle arrangement. Aegires exeches has a very elongate body with extremely elevated, unique tubercles. That is, they are nearly mush- room-shaped, with a flat crown and completely 659 FIGURE 61. Aegires exeches sp. nov. CASIZ 168918. Reproductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, ps = penial spines, rs = receptaculum seminis, v = vagina, vd = vaginal duct, scale = 0.3 mm. cover the dorsum. D’Orbigny also described the body shape of 4. punctilucens as being elongate, with a broadening at the midpoint. He described the tubercles of A. punctilucens as conical, with a flattened top, and a definite symmetry to their arrangement. Schmekel and Portmann (1982) also illustrated and described the tubercles on the specimens from the Mediterranean as being much shorter and wider at the base and sym- metrically arranged, such that there are smooth parts of the dorsum visible between them. In A. exeches, no smooth spaces exist between the tubercles, which are taller, narrower at the base and more densely arranged than on A. punctilucens. The rhinophores of A. exeches have bifid apices whereas the rhinophores of A. punctilucens do not (see Schmekel and Portmann 1982 for an illustration). The gill protective structure of Aegires exeches is also different from that of A. punctilucens. Aegires exeches has a very elaborate, lobed gill structure, while 4. punctilucens has three simple tubercles (see Schmekel and Portmann 1982 for a drawing of the Mediterranean specimens). When comparing the external coloration, both species can be white or light brown. But Schmekel and Portmann describe minute opaque white dots spotted over the brown color, and the iridescent “eye” spots have a brownish-red circular area around them, bordered with black-brown dots. Specimens of Aegires exeches from Okinawa, Hawaii and the Marshall Islands do not have any opaque white dots over the dorsal color, but do have iridescent spots, although in much fewer numbers than is found on A. punctilucens. The “eye” spots on A. exeches can also be surrounded by a border of black-brown dots and a smooth orange circle surrounding the center as seen in pho- tos on the SeaSlug Forum (Rudman 2004). There are significant differences in the reproductive morphology between these two species. Although d’Orbigny didn’t describe or illustrate the reproductive organs of A. punctilucens, Schmekel and Portmann drew the anatomy of specimens from the Mediterranean. Most noticeably, A. exeches has a receptaculum seminis that attaches to the base of the bursa copulatrix. In A. punc- 660 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 | tilucens the receptaculum connects directly to the bulbous vagina. The prostate of A. exeches is — short, tubular and nearly as thick as the ampulla, whereas in A. punctilucens the prostate is very — thick and sausage-shaped and very coiled. Aegires exeches has a very long, tubular ampulla, a nar- row vagina and a long, thin vaginal duct. Aegires punctilucens has a very bulbous ampulla and a very wide vagina, with a thick vaginal duct. There are two differences in the radular teeth between these two species. Schmekel and Portmann illustrate an inner lateral tooth that is substantially smaller than the rest of the teeth in the row. They also state that the size increases outwards. In A. exeches, the first three lateral teeth are smaller than the remaining teeth, which are all the same size. In A. exeches, the radular formula is 14 x 11.0.11 for a 4 mm specimen. Schmekel and Portmann give a radular formula for a 6 mm specimen of 16 x 18.0.18. Baba (1974) described a specimen found in the Sado district of the Japan Sea as Aegires punc- tilucens. Comparison of Baba’s drawings and description of that specimen to our specimens of Aegires exeches leads us to believe that Baba’s specimen is also A. exeches. Whereas A. exeches bears some external similarities to the Mediterranean 4. punctilucens, the Indo-Pacific specimens clearly represent a distinct species. Aegires hapsis Fahey and Gosliner, sp. nov. (Figs. 48F, 62-66) TYPE MATERIAL.— HOLotyPeE: CASIZ 115721, one specimen, 6 mm, dissected, Horseshoe Cliffs, Okinawa, Ryukyu Islands, 3 m, collected March 1998, R. Bolland. DISTRIBUTION.— This species has only been reported from Okinawa (present study). EtTyMOoLoGy.— The specific name /apsis is a Greek word meaning mesh or network, a word that describes the fine white webbing that covers the dorsum of this species. EXTERNAL MORPHOLOGY.— The body shape is elongate (Fig. 48F, 62). The dorsum has small raised tubercles, arranged in a some- what regular fashion on the sides of the dorsum and along the midline behind the gill. Spicules protrude from the tops of all tubercles. The rhinophore pocket is raised and the outer edge is formed by three large tubercles. The rhino- phores are smooth. The gill pocket lies in the posterior third of the dorsum and is protected by a tri-lobed appendage. Each appendage is further divided into 4—5 smaller lobes. The small gill leaves are tri-pinnate. The background color is pale tan and white. There is a fine network of white web- bing covering the dorsum, especially notice- able on the posterior third, behind the gill pock- et. There are small brown dots on either side of the dorsum midline. The tubercles that lie on the edge of the rhinophore pocket have a dark FIGURE 62. Aegires hapsis sp. nov. CASIZ 115721. brown top. The rhinophores are dark brown. Drawing of preserved animal. (A) Dorsal view. (B) Ventral The gill branches are white. ME Ce ae 1 | \ 1 ' FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 661 FicurRE 63 (right). Aegires hapsis sp. nov. CASIZ 115721. Buccal bulb: bb = buccal bulb, cns = central nerv- ous system, m = muscle, oe = esophagus, og = oral glands, ot = oral tube, rs = radular sac, scale = 0.3 mm. FIGURE 64. Aegires hapsis sp. nov. CASIZ 115721. Buccal morphology: (A) Jaw, scale =20 um. (B) Whole radula, scale = 20 um. (C) Inner lateral teeth, scale = 20 um. (D) Outer lateral teeth, scale = 20 um. 662 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 DIGESTIVE SYSTEM.— The arrangement of the digestive system is as illustrated for other Aegires (see Figs. 3B, 8B). The buccal bulb is more ovoid than round and the radular sac protrudes noticeably from the posterior side (Fig. 63). Large oral glands on either side of the oral tube extend markedly from the base of the oral tube. The radular formula is difficult to confirm owing to the very compressed, flattened tooth arrangement, but is approximately 18-20 x 20-21.0.20-21 for the 4 mm specimen (Fig. 64). The jaw (Fig. 64A) has a thickened edge and rounded thickenings at each side. All teeth are hamate, with a long hook on the inner lateral teeth and a shorter hook on the outer teeth. The tips of the teeth are recurved. Rachidian teeth are absent. Some inner lateral teeth have a sharper hook than the remaining teeth. There is no longitudinal groove on the oldest lateral teeth as found in other Aegires species. REPRODUCTIVE SYSTEM.— The reproductive system is triaulic (Fig. 65). The ampulla is very large and bulbous. It branches into the oviduct and the prostate. The prostate is tubular and coils once before narrowing into the very long, thin deferent duct. The deferent duct then widens into the penis. The penis was subsequently lost during preparation, and thus the presence of hooks could not be confirmed. The vagina is narrow and was not examined internally. Thus the presence of spines or hooks cannot be confirmed. The vaginal duct is long and narrow and at the proximal end, enters the bursa copulatrix. The bifurcating oviduct, which originates from the small peanut-shaped receptaculum seminis, enters the vaginal duct. The oviduct enters the female gland mass. The bursa is ovoid and small. CENTRAL NERVOUS SYSTEM.— The central nervous system has fused cerebral and pleural gan- glia (Fig. 66). The eyes are small, sessile on the cerebral-pleural complex, and do not protrude. The pedal ganglia are slightly smaller than the cerebral-pleural complex and they are joined by the vis- ceral loop-pedal/parapedal commissures. Three prominent nerves lead from the pedal ganglia and four nerves, including the rhinophoral nerve, originate from the cerebro-pleural ganglia. Two buc- cal ganglia are positioned ventral to the esophagus. CIRCULATORY SYSTEM.— (not shown) As with other species of Aegires, the heart is relatively small as compared to most cryptobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Externally, Aegires hapsis does not closely resemble any other Aegires species. Only A. malinus has a similarly smooth dorsum with a fine network pattern overall and tiny dorsal pr FIGURE 65. Aegires hapsis sp. nov. CASIZ 115721. FIGURE 66. Aegires hapsis sp. nov. CASIZ 115721. Reproductive system: am = ampulla, be = bursa copulatrix, Central nervous system: bg = buccal ganglia, cple = cerebro- dd = deferent duct, fgm = female gland mass, p = penis, pr= pleural ganglia complex, p = pedal ganglia, r = rhinophoral prostate, r = receptaculum seminis, v = vagina, vd = vaginal nerve, vl+pce+ppc = visceral loop, scale = 0.25 mm. duct, scale = 0.25 mm. ! FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 663 tubercles. But that species has a very different background color (red-brown) and apple green rhinophoral and gill appendages. The radular teeth of A. hapsis are different from other species as well. In A. hapsis the teeth are much more elongate and numerous than found in other species. For example, the inner lateral teeth have a longer hook than the outer lateral teeth and most teeth have a downward bend at the tip. This feature is not found in any other Aegires species. The reproductive morphology has some similarities to other Aegires. The long thin deferent duct and the wider, tubular prostate are features shared with A. exeches, A. incusus and A. malinus. Aegires ninguis and A. incusus also have a large, balloon-shaped ampulla like A. hapsis. However, the combination of a large ampulla, a long thin deferent duct with a wider prostate combined with a receptaculum that leads from a separate duct into the vaginal duct 1s not shared by any of the other Aegires species. The particular combination of morphological characters separates Aegires hapsis as a distinct species. Taxa formerly included in the Genus Notodoris Bergh, 1875 Type spEcIES: Notodoris citrina Bergh, 1875:64, by monotypy. D1aGNosis.— Both Bergh (1875) and Eliot (1906) provided a complete diagnosis for the genus: the body is limaciform, with no distinction between the back and sides. The body is hard, rough and rugose and the surface is filled with spicules. The frontal veil is large. The rhinophores are smooth, retractable and protected by valves. The gill is protected by a large valve, is sometimes guite ramified and not retractable. The radula has no rachidian tooth, but has numerous hamate uni- form lateral teeth, which have a rudimentary secondary denticle below the main hook. REMARKS.— Bergh did not assign the family name Notodorididae to his new genus. The first author to use this name was Eliot (1910). See the earlier discussion on the history of the classifi- cation. Aegires citrinus (Bergh, 1875) (Figs. 67—71) Notodoris citrina Bergh, 1875:53—100 Aegires citrinus (Bergh) comb. nov. TYPE MATERIAL.— Rarotonga, Pacific Ocean. The type material has been lost and therefore, specimens from other Indo-Pacific localities have been examined for this study. MATERIAL EXAMINED.— CASIZ 116864, one specimen, 52 mm, dissected. Rottnest Island, Western Australia, collected December 1998, S. Fahey. CASIZ 113597, one specimen, 23 mm, dis- sected. Laiwan Island, Louisiade Archipelago, Papua New Guinea, collected June 1998, T. Gosliner. DIsTRIBUTION.— Australia, Cook Islands, Fiji, Marshall Islands, Papua New Guinea, Solomon Islands, and New Caledonia (Bergh 1875; SeaSlug Forum, accessed | March 2004 and present study). EXTERNAL MORPHOLOGY.— Bergh (1878) described the external morphology of this species. No differences were found between Bergh’s description and the specimens examined for the pres- ent study. DIGESTIVE SYSTEM.— Aegires citrinus shares the same general digestive anatomy (Fig. 67) as other Aegires (formerly Notodoris) species (Figs. 3B, 8B). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the diges- 664 tive gland. The buccal bulb is nearly round, with four large muscles attached, two per side (Fig. 67). The buccal bulb is shorter and more round than the oral tube. The radular sac slightly protrudes from the bulb, under the esophagus. The labial disk frames the triangular opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate at the top of the opening, with no indication of rods at the edge (Figs. 68A, 69A). The radular teeth of the specimens exam- ined from Western Australia and the Marshall Islands are as described by O’Donoghue (1924). That is, the teeth are so closely packed together that they are difficult to count (Figs. 68-69). The teeth are tall and hamate, with a small denticle under the tip of each tooth. All lateral teeth are similar in size and there is no sign of a rachidian tooth. REPRODUCTIVE SYSTEM.— The reproductive system is triaulic (Fig. 70). The ampulla is very long, thick and tubu- lar. It branches into the oviduct and the prostate. The tubular prostate is thinner, though much longer and coiled than the ampulla and it narrows appreciably into the short deferent duct. The deferent duct enters the very wide penis. The penis has small, sparsely spaced penial hooks near the opening at the genital atrium. The vagina is much narrower than the penis. It was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the proximal end, the long, narrow vaginal duct enters the bursa copula- trix. The receptaculum seminis connects to the bursa via a long separate duct that bifurcates into the oviduct, which enters the female gland mass. The bursa is round and the same size as the large, ovoid receptaculum seminis. CENTRAL NERVOUS SYSTEM.— As with species of both Notodoris and Aegires, the cerebral and pleural ganglia are fused together (Fig. 71). The two pedal ganglia are located below the cerebro-pleural complex and are joined by the pedal commissure, the parapedal commissure and the viscer- al loop. The buccal ganglia are located under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two relatively long nerves as compared to most other Aegires. There are four cerebral nerves leading from each cerebral ganglion, and three large pleural nerves leading from the nght and left pleural ganglia. There is a separate abdominal ganglion on the right side of the viscer- al loop. Gastro-esophageal, rhinophoral and optical ganglia are present. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 FIGURE 67. Aegires citrinus (Bergh, 1875). (A) Buccal bulb: CASIZ 113597. Scale = 0.46 mm. bb = buccal bulb, cns = central nervous system, oe = esophagus, ot = oral tube, rs = radular sac. (B) Digestive sys- tem: CASIZ 116864. Scale = 6.4 mm. bb = buccal bulb, cg = cerebral ganglia, fgm = female gland mass, ht = heart, i = intestine, m = muscles, 0 = esophagus, ot = oral tube, st = stomach, sg = salivary glands. CIRCULATORY SYSTEM.— The heart (Fig. 67B) is relatively small as compared to most cryp- tobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Bergh’s (1875) description of this species was thorough and his drawings FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 665 “¥y Vie mm, FIGURE 68. Aegires citrinus (Bergh, 1875). CASIZ 113597. Buccal morphology: (A) Jaw, scale =10 tm. (B) Whole radula, scale = 30 um. (C) Inner lateral teeth, scale = 10 um. (D) Outer lateral teeth, scale = 10 tm. 20pm ii FH . z Mes FIGURE 69. Aegires citrinus (Bergh, 1875). CASIZ 116864. Buccal morphology: (A) Jaw, scale =100 pm. (B) Whole radula, scale = 100 um. (C) Inner lateral teeth, scale = 20 um. (D) Outer lateral teeth, scale = 20 pm. 666 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 FIGURE 70 (right). Aegires citrinus (Bergh, 1875). CASIZ 116864. Reproduc- tive system: am = ampulla, be = bursa cop- ulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, ps = penial spines, rs = receptaculum seminis, v = vagina, vd = vaginal duct, scale = 1.4 mm. © @f ew an? * ~w E FIGURE 71. Aegires citrinus (Bergh, 1875). CASIZ 116864. Central nervous system: bg = buccal ganglia, cple = cere- bro-pleural ganglia complex, pg = pedal ganglia, r = rhinophoral nerve, vl+pe+ppe = visceral loop, scale = 0.5 mm. vi+pe+ppc pl1-3 FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 667 include the external, radular, reproductive and central nervous system morphology. O’ Donoghue (1924) provided additional details of the anatomy and radular morphology and Gosliner and Behrens (1997) examined all Notodoris species except N. citrinus during their study of the new species, V. serenae. However, none of these recent authors confirmed Bergh’s descriptions of the central nervous system, reproductive or digestive systems of Notodoris citrinus. For the present study, specimens were examined from localities other than the type locality to determine the extent of variation. Aegires gardineri (Eliot, 1906) (Fig. 72) Notodoris gardineri Eliot, 1906: 540-573, pl. 32. Notodoris megastigmata Allan, 1932:103. Aegires gardineri (Eliot) comb. nov. TYPE MATERIAL.— Hulule, Maldive Islands. Holotype: 1919.10.7.47, one specimen, 3.35 cm, collected by Professor J.S. Gardiner, British Museum of Natural History. MATERIAL EXAMINED.— CASIZ 106060, one specimen, 50 mm, dissected. Okinawa, Ryukyu Islands, collected 14 May 1995, R. Bolland. DISTRIBUTION.— Maldive Islands, Indonesia, Papua New Guinea, Okinawa, Western Caroline Islands, Solomon Islands and Australia (Eliot 1906; Marshall and Willan 1999; Coleman 2001; Rudman 2004; present study). EXTERNAL MORPHOLOGY.— The external morphology of Notodoris gardineri was described in detail by several authors (O’ Donoghue 1924; Yonow 1993; Rudman 2004). REPRODUCTIVE SYSTEM.— Gosliner and Behrens (1997) illustrated and described the repro- ductive anatomy of Notodoris gardineri. DIGESTIVE SYSTEM.— Aegires (formerly Nofodoris) gardineri shares the same general diges- tive anatomy as other Nofodoris species (Fig. 67B). The buccal bulb is nearly round, with four large muscles attached, two per side. The buccal bulb is shorter and more round than the oral tube. The radular sac slightly protrudes from the bulb, under the esophagus. The labial disk frames the trian- gular opening to the buccal bulb and is lined with a thin cuticle (see also O’Donoghue 1924 and Yonow 1993). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. CENTRAL NERVOUS SYSTEM.— As with other species of previously included Notodoris, the cerebral and pleural ganglia are fused together (Fig. 72). The two pedal ganglia are located below the cerebro-pleural complex and are joined by the pedal commissure, the para- pedal commissure and the visceral loop. The buccal ganglia are located under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two relatively aN UY é long nerves as compared to most other Aegires. be by There are four cerebral nerves leading from each cerebral ganglion, and three large pleural j : ean’ mus: ee 106060. Central nervous system: bg = buccal ganglia, cple = OM leading from the right and left pleural cerebro-pleural ganglia complex, p = pedal ganglia, r = ganglia. There is a separate abdominal ganglion rhinophoral nerve, vit+pe+ppe = visceral loop, scale = 0.5 on the right side of the visceral loop. Gastro- mm. FIGURE 72. Aegires gardineri (Eliot, 1906). CASIZ 668 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 esophageal, rhinophoral and optical ganglia are present. CIRCULATORY SYSTEM.— (not shown) The heart is relatively small as compared to most cryp- tobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Eliot (1906) described both the external and radular morphology of Notodoris gardineri. However, he did not describe nor illustrate the radular teeth except to note that they fit Bergh’s (1875) plate [X exactly. O’ Donoghue (1924) and Yonow (1993) provided further descrip- tions of the radular morphology. Eliot also did not examine the reproductive anatomy, stating that the specimen he obtained was “too hardened” to examine. Although Gosliner and Behrens (1997) examined all Notodoris species except NV. citrinus, they did not describe the central nervous systems or the digestive systems of the previously included Notodoris species. Therefore, additional specimens were examined to com- plete the data for the present study. Aegires minor (Eliot, 1904) (Figs. 73-74) Notodoris minor Eliot, 1904: 83-105, pls 3-4. Aegires minor (Eliot) comb. nov. TYPE MATERIAL.— Chuaka, East coast of Zanzibar. Type: 1919.9.16.4, one specimen, 13 mm. British Museum of Natural History. MATERIAL EXAMINED.— CASIZ 068668, fourteen specimens; one specimen, 65 mm dissect- ed. Madang, Papua New Guinea, collected August 1989, T. Gosliner. DISTRIBUTION. Zanzibar, Mauritius, Oman, Philippines, Eastern Caroline Islands, Solomon Islands, Papua New Guinea, Manado, Sulawsi, Indonesia, and Australia (Eliot 1904; Gosliner, Behrens, and Williams 1996; Rudman 2004). EXTERNAL MORPHOLOGY.— The external morphology of the specimens examined for this study is as described by Eliot (1904) and corroborated by O’ Donoghue (1924), Yonow (1993) and Gosliner and Behrens (1997). DIGESTIVE SYSTEM.— Aegires minor shares the same general digestive anatomy as other Notodoris species (Fig. 67B). The buccal bulb is nearly round (Fig. 73) with four large muscles attached, two per side. The buccal bulb is shorter and more round than the oral tube. The radular sac slightly protrudes from the bulb, under the esophagus. The labial disk frames the triangular opening to the buccal bulb Sate! us Tinea wane ental inet opnap hss Buccal bulb: bb = buccal bulb, cns = central nervous system, short and connects directly to the stomach. The — jy = muscles, ot = oral tube, rs = radular sac, sg = salivary intestine makes a simple, wide curve along the — glands, scale = 1.75 mm. outside of the digestive gland. CENTRAL NERVOUS SYSTEM.— As with other species of Aegires (formerly Notodoris), the cerebral and pleural ganglia are fused together (Fig. 74). The two pedal ganglia are located below the cerebro-pleural complex and are joined by the pedal commissure, the parapedal commissure and the visceral loop. The buccal ganglia are located under the esophagus, below the central nerv- ous system. They are joined to the cerebral ganglia by two relatively long nerves as compared to FIGURE 73. Aegires minor (Eliot, 1904). CASIZ 068668. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 669 most other Aegires. There are four cerebral nerves leading from each cerebral ganglion, and three large pleural nerves leading from the right and left pleural ganglia. There is a sepa- rate abdominal ganglion on the right side of the visceral loop. Gastro-esophageal, rhinophoral and optical ganglia are present. CIRCULATORY SYSTEM.— (not shown) The heart is relatively small as compared to most eryptobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— Eliot (1904) described the external and the radular morphology of this species, but only described and drew one fea- F 3 FIGURE 74. Aegires minor (Eliot, 1904). CASIZ 068668. ture of the reproductive anatomy: the armed Central nervous system: bg = buccal ganglia, cplc = cerebro- penis. Gosliner and Behrens (1997) examined pleural ganglia complex, p = pedal ganglia, r = rhinophoral all Notodoris species during their study of the _ nerve, vl+pet+ppe = visceral loop, scale = 0.75 mm. new species NV. serenae. But they did not describe the central nervous systems or the digestive systems of Aegires (Notodoris) at that time. Therefore, additional specimens of Aegires minor were examined to complete the data for the pres- ent study. Aegires serenae (Gosliner and Behrens, 1997) (Figs. 75—76) Notodoris serenae Gosliner and Behrens, 1997: 303-307, Figs 7C, 12A—C, 13. Aegires serenae (Gosliner and Behrens) comb. nov. TYPE MATERIAL.— For a complete list of the type material see Gosliner and Behrens (1997). MATERIAL EXAMINED.— Paratype: CASIZ 107229, two specimens; one specimen 57 mm, dis- sected. Madang, Papua New Guinea, collected August 1989, T. Gosliner and D. Behrens. DISTRIBUTION.— Belau, Papua New Guinea, Philippines, Indonesia, Malaysia, Pohnpei, Guam, and Okinawa (Gosliner and Behrens 1997; C. Carlson and P.J. Hoff, pers. commun. ). DIGESTIVE SYSTEM.— Aegires serenae shares the same general digestive anatomy as other Notodoris species (Fig. 67B). The buccal bulb uke is nearly round (Fig. 75) with four large mus- cles attached, two per side. The buccal bulb is Se shorter and more round than the oral tube. The We UNS ree ; radular sac slightly protrudes from the bulb, = aT pee ee LOL under the esophagus. The labial disk frames the a an Ah i : : i : Cg cert a —S ~S C Sea triangular opening to the buccal bulb and is f/f gern" “ts 2 Veg lined with a thin cuticle. The esophagus is short °97" and connects directly to the stomach. The intes- A —_— tine makes a simple, wide curve along the out- Hy side of the digestive gland. FIGURE 75. Aegires serenae (Gosliner and Behrens, CENTRAL NERVOUS SYSTEM.— As with 1997). CASIZ 107229. Buccal bulb: bb = buccal bulb, m = muscles, oe = esophagus, ot = oral tube, rs = radular sac, sg other species of Aegires (formerly Notodoris), _ cent seiay IEG oo Os 08 670 the cerebral and pleural ganglia are fused together (Fig. 76). The two pedal ganglia are located below the cerebro-pleural complex and are joined by the pedal commissure, the para- pedal commissure and the visceral loop. The buccal ganglia are located under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two relatively long nerves as compared to most other Aegires. There are four cerebral nerves leading from each cerebral ganglion, and three large pleural nerves leading from the right and left pleural ganglia. There is a separate abdominal ganglion on the right side of the visceral loop. Gastro- esophageal, rhinophoral and optical ganglia are present. REMARKS.— Gosliner and Behrens (1997) have recently described this Indo-Pacific species. Anatomical information for the present study has been taken from their paper. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 « * oe * . Pa Je | © ‘ 1) ite A A v1+pctppc FIGURE 76. Aegires serenae (Gosliner and Behrens, 1997). CASIZ 107229. Central nervous system: bg = buccal ganglia, cple = cerebro-pleural ganglia complex, p = pedal ganglia, r = rhinophoral nerve, vl+pce+ppe = visceral loop, However, they did not describe some charac- scale = | mm. ters that were necessary for the present study such as the central nervous system and the digestive systems. Therefore, the additional specimens noted above were examined to complete the data. Taxon formerly included in the Genus Triopella Sars, 1878 TYPE SPECIES: Triopella incisa Sars, 1878:310, by monotypy. (Figs. 77-81) Aegires incisus (Sars, 1878) Triopella incisa Sars, 1878:310, pl. 27, Fig. 3. Aegires incisus (Sars, 1878) comb. nov. DIAGNOsIs.— Sars’ (1878) partial description is as follows: “Small body, oblong and angled, elongate spicules throughout the dorsum. At the posterior end of the mantle are two lobes. The dor- sum is highly arched, having superficial dorsal appendages, and longitudinal medial ridges that converge between the tentacles (rhinophores). At the base of the rhinophores there 1s a single, large tubercle, with S—7 smaller tubercles arranged longitudinally along the two medial ridges. There are two small tubercles on the anterior of the dorsum. The oral tentacles are inconspicuous. Dorsal rhinophores are short, perfoliate and retractable. The gill has 3 small and sparse branchae, which are arranged in a semi-circle around the anus. The anus is not completely round. The radula has no rachidian tooth, nor denticulate lateral teeth.” TYPE MATERIAL.— This species was collected from the Norwegian Arctic Sea. No additional collection data are available. Type material not found in the collections at the Natural History Museum, Oslo. MATERIAL EXAMINED.— D 15983, two specimens, 8-9 mm, Bodg, Norway; D 15988, two specimens, 6—8 mm, Sande Fjord, Norway. No additional collection data are available. DIsTRIBUTION.— Norway and Greenland (Sars 1878; Norman 1893; Odhner 1922; Marcus | | FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 671 and Marcus 1969). This species has been men- tioned several times in the literature (Fischer 1880-1887; Bergh 1883, 1892; Norman 1893; Odhner 1907; Eliot 1910a; Thiele 1929-31; Odhner 1939). EXTERNAL MORPHOLOGY.— The speci- mens examined for the present study match the previously published description and drawings mentioned above. A drawing of one of the pre- served specimens from the type locality is pre- sented in Figure 77. DIGESTIVE SYSTEM.— 4egires incisus shares the same general digestive anatomy as other Aegiridae species (Figs. 3B, 8B). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is nearly round, with four large BISU RE), cetes UC Usae asian oue ys NINE f : D15988. Drawing of preserved specimen. (A) Dorsal view. muscles attached, two per side (Fig. 78). The (g) ventral view. Scale = 0.8 mm. buccal bulb is shorter and more round than the oral tube. The radular sac slightly protrudes from the bulb, under the esophagus. The labial disk frames the triangular opening to the buccal bulb and is lined with a thick cuticle. There is a thick jaw plate at the top of the opening, with thick rods at the edge (Fig. 79A). The radular formula is 13 x 16.0.16. The teeth are simply hamate and the two innermost lateral teeth are smaller than the remaining teeth (Figs. 79B-D). REPRODUCTIVE SYSTEM.— The reproduc- tive system is triaulic (Fig. 80). The ampulla is large and bulbous at the distal end and tubular FIGURE 78. Aegires incisus (Sars, 1878). NMNH D15988. at the proximal end. It branches into the Buccal bulb: bb = buccal bulb, m = muscles, oe = esophagus, ; ; : ot = oral tube, rs = radular sac, sg = salivary glands, scale = oviduct and the prostate. The prostate is wide 9 ¢ mm. and narrows before broadening again in the central portion. It narrows before entering into the long sausage-shaped deferent duct. The defer- ent duct leads to the penis, which is only slightly bulbous at the distal end. There are densely packed hooks along the length of the penis. The vagina is very wide and was not examined inter- nally. Thus the presence of spines or hooks cannot be confirmed. The vaginal duct 1s extremely wide and short and at the proximal end, enters the bursa copulatrix. The bifurcating oviduct that originates from the very elongate receptaculum seminis enters the vagina. The oviduct enters the female gland mass. The bursa is round and small. CENTRAL NERVOUS SYSTEM.— The cerebral and pleural ganglia of Aegires incisus are fused together (Fig. 81). The two pedal ganglia are located below the cerebro-pleural complex and are joined by the pedal commissure, the parapedal commissure and the visceral loop. The buccal gan- glia are located under the esophagus, below the central nervous system. They are joined to the cere- 672 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 FIGURE 79. Aegires incisus (Sars, 1878). NMNH DI 5988. aut morphology: (A) Jaw, scale =10p um. (B) Whole radu- la, scale = 100 um. (C) Inner lateral teeth, scale = 20 um. (D) Outer lateral teeth, scale = 10 um. FIGURE 80. Aegires incisus (Sars, 1878). NMNH D15988. Reproductive system: am = ampulla, be = bursa copulatrix, dd = deferent duct, fgm = female gland mass, p = penis, pr = prostate, ps = penial spines, rs = receptaculum seminis, V = vagina, vd = vaginal duct, scale = 0.3 mm. FIGURE 81. Aegires incisus (Sars, 1878). NMNH D15988. Central nervous system: bg = buccal ganglia, eple = cerebro-pleural ganglia complex, p = pedal ganglia, r = rhinophoral nerve, vl+pe+ppe = visceral loop, scale = 0.67 mm. bral ganglia by two relatively short nerves. There are four cerebral nerves leading from each cere- bral ganglion, and three large pleural nerves leading from the right and left pleural ganglia. There is a separate abdominal ganglion on the right side of the visceral loop. Gastro-esophageal, rhinophoral and optical ganglia are present. | FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 673 CIRCULATORY SYSTEM.— (not shown) The heart is relatively small as compared to most cryp- tobranch dorids (Valdés 2002). There is one blood gland situated in front of and to the right side of the visceral cavity. REMARKS.— 4egires incisus (formerly Triopella incisa) has been often overlooked as a species of Aegiridae as discussed in the earlier section on the history of the classification. However, the species examined from the type locality can be confirmed as Triopella incisa Sars, 1878 from the drawings and descriptions provided by Sars and subsequent collectors. Phylogenetic analyses performed for the present study using 58 of the 64 considered morphological characters indicate that 7. incisa is the sister species of Aegires sublaevis. Therefore, Triopella cannot be maintained as a separate taxon without rendering Aegires paraphyletic. We propose to unite Triopella incisa with the Aegires with the name Aegires incisus (Sars, 1878). Further details are provided in the phylogenetic analysis below. PHYLOGENETIC ANALYSIS The outgroup taxa chosen for the present analysis are discussed in a prior section, Material and Methods. Character states for Bathydoris abysscrum were taken from the literature (Bergh 1884; Gosliner and Bertsch 1988; Wagele 1989; Cervera et al. 2000; Valdés 2002). Character states for the additional outgroups were taken both from the literature and from museum material when avail- able. Table 3 (see Appendix) summarizes the information for the outgroup taxa. Morphological data were organized using MacClade, ver 3 (Maddison and Maddison 1998). CHARACTERS.— The following characters were considered for use in the analyses of Aegiridae. The character states are indicated as follows: 0: the presumed plesiomorphic condition; 1.2.3: apomorphic condition. For character states that are not applicable, “-” is used and for miss- ing data, *?” is used. The distribution of plesiomorphic and apomorphic character states is present- ed in Table 2 (see Appendix). Character states for the taxa examined for the present study can be found in Table 4 (see Appendix). “Outgroup taxa” refers to the combination of Bathydoris, the six phanerobranchs, the three cryptobranchs and Hexabranchus. Only the generic name is used for the outgroup taxa. “Ingroup taxa” refers to all Aegires, including those taxa formerly known as Notodoris and Triopella. 1. Body shape: Wide with a distinct mantle (0): all outgroup taxa except for Holoplocamus, Okenia and Polycera. Elongate with a reduced/absent margin (1): all ingroup taxa. . Dorsal ridges: Absent (0): all outgroup taxa and most ingroup taxa. Present (1): Aegires albopunctatus, A. palensis, A. punctilucens, A. sublaevis and Triopella incisa. 3. Dorsal features: Autotomizable (0): Bathydoris. Permanent (1): all other outgroup and ingroup taxa. This character is not applicable to Hexabranchus, which does not have dorsal features. 4. Permanent features: Low (0): all outgroup taxa except Holoplocamus and Okenia and over half the ingroup taxa. Elevated (1). . Dorsal feature shape: Rounded (0): All outgroup taxa and two-thirds of the ingroup species. Flattened (1): one-third of the ingroup taxa. 6. Tubercles: Scattered (0): All outgroup taxa except Okenia and Polycera and over half the ingroup taxa. In rows (1): nearly half the ingroup taxa. 7. Rhinophores: Not retractable (0): Bathydoris and Okenia. Retractable (1): all other taxa. 8. Rhinophore lamellae: Transverse or longitudinal (0): all outgroup taxa except Conualevia. Smooth (1): all ingroup taxa and Conualevia. 9. Rhinophore pocket: Simple hole, slit (0): all outgroup taxa except Hexabranchus. Raised (1): all ingroup taxa. 10. Rhinophore tubercles: None (0): all outgroup taxa. Outer marginal only (1): all ingroup taxa except for A. palensis, A. punctilucens and A. exeches which all have character state 2. All around (2). i) Nn 674 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 11. Rhinophore tubercle number: None (0): all outgroup taxa. Three to five (1): half the ingroup taxa. One tubercle (2): half the ingroup taxa. This character was not included in the final analyses because the char- acter states are included in other characters (#10). 12. Oral tentacles/hood: Tentacles (0): Bathydoris, Calycidoris, Conualevia, Hexabranchus, and Holoplo- camus. Oral hood/veil (1): no ingroup taxa have an oral hood or veil. Neither (2): all remaining outgroup taxa and all ingroup taxa. 13.Oral tentacle size: Large (0): Bathydoris and Hexabranchus. Short/small (1): Conualevia and Holoplocamus. This character is not applicable for the ingroup taxa. 14. Integumentary spicules. Present (0): Actinocyclus, Bathydoris, Hexabranchus, Holoplocamus and Polycera. Absent (1): all other outgroup taxa and all ingroup taxa. 15. Gill retraction: Contractile (0): all ingroup taxa and Bathydoris, Holoplocamus, Okenia, and Polycera. Retractile (1): all other outgroup taxa. 16. Gill branches: Few (0): Most ingroup taxa and Bathydoris, Diaphorodoris, and Holoplocamus. Numerous (1): all remaining outgroup and ingroup taxa. 17. Gill leaf ramification: Multi-pinnate (0): Bathydoris, Hexabranchus, Holoplocamus, and Mandelia. Not multi-pinnate (1): all other outgroup and ingroup taxa. 18. Gill arrangement: Isolated points (0): Bathydoris, Diaphorodoris, Hexabranchus, and Onchidoris. One opening (1): all other outgroup and ingroup taxa. 19. Gill protective structures: Large, firm (0): all Notodoris and A. hapsis. Tubercles (1): Holoplocamus and half the ingroup taxa. Elongate appendages (2): all remaining Aegires species. This character was not included in the final analyses because the character states are included in other characters (#20, #61). 20. Gill protective structures: Simple (0): Holoplocamus and over half the ingroup taxa. Compound, ramified (1): the remaining ingroup taxa. 21.Buccal pump: Absent (0): all ingroup taxa and most outgroup taxa. Present (1): remaining taxa. 22. Oral glands: Absent (0): most taxa included in the study. Present (1): 4. villosus, A. malinus, A. incusus, and A. hapsis. 23. Jaw rodlets: Absent (0): half the taxa included in the study. Present (1): half the taxa included in the study. 24. Jaws: Thick, chitinous (0): Bathydoris, Okenia, Polycera and most ingroup taxa. Thin (1): remaining out- group taxa except Mandelia and all Notodoris. Absent (2): Mandelia. 25. Rachidian: Present (0): Bathydoris and Onchidoris. Absent (1): all remaining taxa included in the study. 26. Lateral teeth size: Outer smallest (0): Bathydoris and Calycidoris. Inner smallest (1): most ingroup taxa and remaining outgroup taxa. Both small (2): Conualevia, A. albus, A. punctilucens, Triopella incisa, and N. citrina. 27.Inner lateral teeth shape: Elongate (0): Actinocyclus and Bathydoris. Hamate (1): Conualevia, Hexabranchus, Holoplocamus, Polycera and most ingroup taxa . Hooked (2): Calycidoris, Diaphorodoris, Okenia, Onchidoris, A. incusus, and A. hapsis. 28.Second lateral tooth: Same size as inner (0): all taxa examined except Actinocyclus, Diaphorodoris, Okenia, and Onchidoris. Markedly reduced (1). 29. Outer lateral tooth: Present (0): all taxa examined except Calycidoris, Diaphorodoris, Okenia, Onchidoris, and Polycera. Absent (1). 30. Outer lateral teeth shape: Hooked (0): all taxa included in the study except for Actinocyclus and Holoplocamus. Not hooked (1). 31.Radular teeth denticulation: No denticles (0): Bathydoris, Conualevia, Holoplocamus, Polycera, and all Aegires species. One spur below main cusp (1): all Notodoris. Denticulate (2):all remaining outgroup taxa. 32. Reproductive system: Diaulic (0) Bathydoris. Triaulic (1): all remaining species in the study. 33. Vagina shape: Elongate/thin (0): all outgroup taxa except Diaphorodoris and Okenia and two-thirds of the ingroup species. Wide, bulbous (1): one-third of the ingroup taxa. This character was not included in the final analyses because the state could not be confirmed in several taxa. 34. Ampulla shape: Elongate, sausage-shaped (0). Rounded, bulbous (1). 35. Penis: Same width as deferent duct (0): all outgroup taxa except Hexabranchus, Mandelia, Okenia, and Onchidoris and over half the ingroup taxa. Wider than deferent duct (1): the remaining ingroup species. 36. Penial hooks: Absent (0): Actinocyclus, Bathydoris, Conualevia, Hexabranchus, and Mandelia. Present FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 675 (1): remaining outgroup taxa and all ingroup taxa. 37. Penial hooks: Few (0): all outgroup taxa. Many (1): all ingroup taxa. This character was not included in the final analyses because it is the same character as #39. 38. Penial spines: Throughout (0): all outgroup taxa and six ingroup taxa. Distal tip only (1): most ingroup taxa. 39. Penial spines: Sparse (0): all outgroup taxa. Dense (1): all ingroup taxa. 40. Vestibular gland: Absent (0): all taxa included in the present study except Aegires sublaevis. Present (1). 41. Bursa copulatrix ducts: One duct (0): Actinocyclus, Bathydoris, Holoplocamus and one-third of the ingroup taxa. Two ducts (1): the remaining outgroup taxa and two-thirds of the ingroup species. 42. Pleural ganglia: Differentiated (0): Bathydoris. Fused with cerebral ganglia (1): all remaining outgroup taxa and all ingroup taxa. 43. Cerebral ganglia: Two ganglia on each side (0): Bathydoris. Fused (1): all remaining outgroup taxa and all ingroup taxa. 44 Cerebral nerve number: Three (0): Bathydoris. Four or more (1): all remaining outgroup taxa and all ingroup taxa.. 45. Eves: Absent (0): Bathydoris. Present (1): all remaining outgroup taxa and all ingroup taxa.. 46. Eye position: Stalks (0): Actinocyclus, Hexabranchus, and Holoplocamus. Prominent (1): all remaining outgroup taxa and all ingroup taxa except Nofodoris. Not prominent (2): Notodoris. 47.Ganglionic tubercles: Absent (0): all taxa included in the study except Hexabranchus. Present (1): Hexabranchus. 48. Inner lateral tooth: Without secondary cusp (0): all taxa included in the study except Holoplocamus and Polycera. With secondary cusp (1): Holoplocamus and Polycera. 49. Gill pocket: Absent (0): all taxa included in the study except Actinocyclus, Conualevia, and Mandelia. Present (1): Actinocyclus, Conualevia, and Mandelia. 50. Inner two lateral teeth: Not elongate (0): all taxa included in the study except Holoplocamus and Polycera. Elongate (1): Holoplocamus and Polycera. 51.Foot corners: Rounded (0): all taxa included in the study except Holoplocamus and Polycera. Prolonged (1): Holoplocamus and Polycera. 52. Foot dimension: Same as mouth (0): all taxa included in the study except Actinocyclus, Conualevia, and Mandelia. Narrower than mouth (1): Actinocyclus, Conualevia, and Mandelia. 53.Rhinophore stalk: Long (0): all taxa included in the study except Actinocyclus, Conualevia, Diaphorodoris, Hexabranchus, and Mandelia. Short (1): Actinocyclus, Conualevia, Diaphorodoris, Hexabranchus, and Mandelia. 54. Dorsal pigment: No dark pigment (0): most outgroup taxa except Actinocyclus, Calycidoris, Mandelia, and Onchidoris. This character is applicable for over half the ingroup taxa. Dark pigment (1): remaining out- group and ingroup taxa. 55. Dorsal pigment: Spots (0): all taxa in the study for which this character is applicable except for Notodoris minor. Lines (1): Notodoris minor. 56. Dorsal tubercle pigment: None (0): all outgroup taxa except Actinocyclus and Polycera. Most ingroup taxa. Dark apex (1): Actinocyclus, Polycera and seven ingroup species. 57.Dorsal ocellae: Absent (0): all outgroup taxa and most ingroup taxa. Present (1): Aegires pruvotfolae, A. punctilucens, A. sublaevis, A. incusus, and A. exeches. 5%. Dorsal rings: Absent (0): all outgroup tax except Mandelia and most ingroup taxa. Present (1): 4. pruvot- folae, A. sublaevis, and A. incusus. This character was not included in the final analyses because the states are included in character #57. 59. Rhinophore color: No dark pigment (0): all ingroup taxa except A. ortizi, A. palensis, A. punctilucens, A. sublaevis, and A. malinus. Dark rings (1): Aegires ortizi, A. punctilucens, and A. sublaevis. Dark specks (2): A. palensis and A. malinus. 60. Rhinophore color: same as body (0): all outgroup taxa except Calycidoris and most ingroup taxa. Different color from body (1): Calycidoris, A. alhopunctatus, Notodoris serenae, A. ninguis, A. malinus, A. incusus, and A. hapsis. 61. Gill protective structures: Simple, digitform (0): one-third of the ingroup taxa. Same as dorsal tubercles 676 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 (1): one-third of the ingroup taxa. Compound structures (2): the remaining third of the ingroup taxa. Fan- shaped (3): only A. flores. 62. Dorsal webbing: Absent (0): all taxa included in the study except A. malinus and A. hapsis. Present (1): Aegires malinus and A. hapsis. 63. Finger-like structures: Simple (0): only four ingroup taxa for which this character is applicable. Lobed (1): only A. albopunctatus and A. lemoncello. 64. Dorsal spicules: Smooth (0): most taxa for which this character is applicable. Furry appearance (1): only A. malinus and A. hapsis. This character was left out of the final analyses because there were not enough data for the taxa included in the study. RESULTS.— Three most parsimonious trees were obtained with 175 steps and had a consisten- cy index (CI) of 0.38, a retention index (RI) of 0.60 and a homoplasy index (HI) of 0.62. All trees were found in one tree island. The strict consensus tree is shown in Fig. 82A. A majority rule tree from the 3 trees was produced and is shown in Fig. 82B with the character numbers and character reversals. The underlined numbers indicate reversals. The trees indicate that the Aegiridae: Aegires, Notodoris and Triopella form a monophyletic clade and that 7riopella is nested in a derived clade of Aegires. Notodoris is nested within Aegires. Bathydoris is the most basal taxon to the study group. The phanerobranchs included in the analysis are basally situated to Aegiridae and the cryp- tobranchs included in the study form a sister clade to some suctorian phanerobranchs. A Bremer support analysis shows that most of the clades are poorly supported (Fig. 82A), with the exception of the clade of species formerly known as Nofodoris and the outgroups. Analyses performed using Bathydoris and the phanerobranchs as outgroups resulted in 6 most parsimonious trees with the same tree scores (trees not shown). The only differences between these trees and those found when using only Bathydoris as the outgroup were at the terminal branches. That is, all the deeper nodes remained the same but two polytomies were unresolved: one for all the phanerobranchs other than Aegires and one polytomy at the node containing A. pruvotfolae and A. incusus. DISCUSSION The results from our phylogenetic analysis of Aegiridae support the monophyly of Aegiridae. Additionally, the phanerobranch and the cryptobranch dorids examined for the present study form monophyletic sister clades (Fig. 82). In our strict consensus tree (Fig. 82A) the cryptobranch clade containing Mandelia, Conualevia and Actinocyclus 1s closely related to the phanerobranchs exam- ined for the present study: Onchidoris, Diaphorodoris and Calycidoris. Valdés (2002) showed the cryptobranchs as more derived than the phanerobranchs, including Aegires. Specifically, Valdés’ phylogeny shows Aegires as the most basally situated taxon in his study group, with the crypto- branchs and Hexabranchus more derived than Aegires. This differs from our results that show Hexabranchus as the most basally situated taxon to the clade containing all other taxa examined and Aegires as the most derived clade. Valdés’ phylogeny also suggests that phanerobranchs are not monophyletic and our results support this view. However, the sequence of branching is distinct. The results of the present study also show that Holoplocamus and Polycera are more closely related to each other and more basally situated to the remaining phanerobranchs and cryptobranchs. Characters that unite this clade are body shape (#1), inner lateral teeth with a secondary cusp (#48), elongate inner two lateral teeth (#50), and prolonged foot corners (#51). Character mapping also revealed interesting relationships between the suctorians and the cryp- tobranchs. For example, retractile gills (415) is the apomorphic state shared by the cryptobranchs and suctorians except for Okenia. This indicates that the suctorians are the sister group to the cryp- | | ' ' FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 677 tobranchs. In the non-suctorians, the gill is truly non-retractile indicating that this character state has been secondarily derived. The synapomorphies that unite all Aegires (including Notodoris and Triopella) are the absence of rhinophore lamellae (#8), the presence of rhinophore tubercles only on the outer margin (#10) and the presence of many penial spines (#39). Our analysis shows that within the study group Aegiridae, there are two major Aegires clades. One contains the taxa not found in tropical Indo-Pacific seas. The species in this clade (Aegires albopunctatus, A. palensis, A. punctilucens, A. sublaevis, A. gomezi, A. ortizi and Triopella incisa) are found only from the Mediterranean, northern Europe, the Caribbean and the west coast of North America. The synapomorphy that supports this clade is dorsal tubercles arranged in rows (#6). The second clade containing Aegires albus, found in the Antarctic, is also basally situated but more closely related to the tropical Indo-Pacific clade that contains all other Aegires species and all for- mer Notodoris species. This Indo-Pacific clade is united by a reproductive synapomorphy, penis wider than the deferent duct (#35). Notodoris clusters together in a monophyletic clade that is basally situated to the Indo-Pacific Aegires. Synapomorphies that unite the Nofodoris clade are the presence of large, firm gill protective structures (#19), radular teeth that have one spur below the main cusp (#31) and reduced eyes (#46). In all analyses performed for the present study, the Notodoris clade is nested within the Aegires sharing multiple synapomorphies with Aegires (see Fig. 82B). The name Aegires Lovén, 1844 is the older name having precedence over Notodoris Bergh, 1875 as discussed in a previous section. Thus, if only considering precedence, the names of the four Notodoris species should be assigned to the genus Aegires. The name Nofodoris has been in gen- eral use by both amateurs and scientific researchers to distinguish these four firm bodied nudi- branchs from other externally similar nudibranchs (Coleman 1989; Wells and Bryce 1993; Gosliner et al. 1996: Marshall and Willan 1999; Coleman 2001; Rudman 2004). In order to maintain the monophyly of Aegires we propose to reassign the four Notodoris species to Aegires. Thus, Notodoris citrina Bergh, 1875 becomes Aegires citrinus (Bergh, 1875). Aegires citrinus Pruvot- Fol, 1930 must be named Aegires pruvotfolae Fahey and Gosliner, 2004, named for Alice Pruvot- Fol who first described Aegires citrinus from New Caledonia (1930). The use of a junior homonym is required since the species name citrinus is already in use within the Aegiridae (Aegires citrinus). Notodoris gardineri becomes Aegires gardineri (Eliot, 1906). Notodoris minor 1s Aegires minor (Eliot, 1904) and N. serenae is Aegires serenae (Gosliner and Behrens, 1997). With regard to Triopella incisa, at this time we propose to name this species Aegires incisus (Sars, 1878). Our proposal takes into consideration the following points: this species is firmly nest- ed within the Aegires clade in all our analyses, the generic name Aegires takes precedence over the name Triopella and the lack of general use and recognition of the name Triopella. In the present analysis, the traditionally recognized clade Phanerobranchia appears to be para- phyletic and no synapomorphies were found to support the group. Wagele and Willan (2000) and Valdés (2002) also found this to be true. In our analysis, the cryptobranch dorids that we included were nested within the phanerobranchs. Analyses that included only the cryptobranchs and phaner- obranchs along with Aegires albus also failed to separate the phanerobranchs as a monophyletic clade (tree not shown). An additional similarity between our analyses and previous studies (Valdés 2002) is the exclu- sion of Hexabranchus from the other cryptobranch or phanerobranch dorids and its placement in a basal position relative to the phanerobranch/cryptobranch clade. Several characters support the position of Hexabranchus in the present analysis (Fig. 82B). In accordance with current perspectives of phylogenetic classification (De Queiroz and 678 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES (1) (1) (1) (1) OT) (1) (1) (1) TT) (1 (1) (1) (1) (1 (1) G (1 (1) (1) (1) A (1) Volume 55, No. 34 Bathydoris Hexabranchus Conualevia Actinocyclus Mandelia Calycidoris Diaphorodoris Okenia Onchidoris A. albus A. flores A. pruvotfolae A. villosus . IncusUus /emoncello petalis exeches . ninguis matinus hapsis serenae . Cltrinus gardineri minor a/bopunctatus pPalensis . punctilucens SESob Toko & ob Boe See SS sublaevis s incisus A. gomezi A. ortizi Holoplocamus Polycera FIGURES 82A-—B. Phylogeny of the Aegiridae. (A) (above). Strict consensus tree, 175 steps. Numbers below the branch- es are Bremer decay values. (B) (right). Majority rule tree of 3 most parsimonious trees. Numbers (100 and 67) above the FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 679 Bathydoris Hexabranchus 9,15,16,23,35,47,53 8.26(2).44 ; 100 = Conualevia 1423,28,30,31(2).41 4656. orerre ka Saree 100 12.27.54 Mandelia 15,16,24 1 oot Calycidoris tas 467 1823 34 21,27,31(2)--> u Okenia 12,28------ > : , 67 lens Diaphorodoris Lend Onchidoris 25.35.54 A. albus 26(2),35 35,41,61(3) aliiores . ree 23,41 725 2627 22.41 42,439.45 100 — Apravevarc & BE : 100 = | 100 (— A. villosus 14117,46 100 4,5 FP all A. incusus 35 27(2),57,60 100 41.5@ * A. lemoncello 34,41 * ; ; 34% 56 petalis 1@(2),20,35,38,54,57 A. exeches 20,35 63 A 5 * my te A. ninguis 100 20.241 59(2 38 23,60 ~|4p— A. malinus 22,61(2),62,64 A. hapsis 27(2),41,544 "PF — A. serenae 100 100 26(2}56,60 *p—2) 4. citrinus 1,8,9,10,12,39 1924.31 ,46(2),61(2) A. 315n83644 16,20] * A. gardineri 41554 minor 23,26(2),60,63 100 A. albopunctatus 59(2) p 3454 |, A. palensis 100 10{2 5 23 26(2) 41.5657 596 A. punctifucens 2,34 40,57,59->) oppo 100 2612) « . SUb/aevis 4,4}- A. incisus 6 100 Fee: gomezi 23,56 A. ortizi 59 100 4,24 ,26,30,41--- Holoplocamus B 1,48,50,51 6.12.16.17.29.56 Polycera branches are the percentage of the 3 consensus trees that agree on this node. The asterisk also denotes 100% agreement on the node. Numbers shown at the branches are character numbers from Table 2. The underlined numbers indicate reversals. 680 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 34 Gauthier 1994) and based on morphological character analysis we propose that the “family | group” Aegiridae contains twenty-three species of Aegires descendent from a common ancestor. BIOGEOGRAPHICAL RELATIONSHIPS The phylogeny of the Aegiridae (Fig. 82) shows a clear-cut case of vicariance and offers some insight into species cladogenesis similar to patterns found in other opisthobranch lineages. There are two conspicuous major clades of Aegires. One clade contains 15 species found in the Southern Hemisphere and in the Indo-Pacific tropics while the second clade contains 7 species found in the temperate and tropical Atlantic, the Mediterranean and the temperate eastern Pacific (a Northern Hemisphere clade). In the clade from the Southern Hemisphere and the Indo-Pacific, the most basally situated species, Aegires albus, is an Antarctic species and sister taxon to the Indo-Pacific clade. The more numerous taxa are found in the tropical Indo-Pacific with the exception of A. ninguis found also from the temperate Indian and Atlantic coasts of South Africa. Given that A. ninguis is a derived member of this clade, it is most likely that these species were derived from Indo-Pacific ancestors rather than from other cold water or temperate taxa. One possible explana- tion for this topology is that there has been a little adaptive radiation of the polar or temperate Aegires species similar to Flabellina and to a lesser extent Hallaxa (Gosliner 1995). The phyloge- ny of Aegires demonstrates a considerable degree of speciation within the Indo-Pacific tropics. Another possible scenario for the distribution pattern seen for this clade of Aegires is that A. albus 1s the last surviving species of a group of cold water Aegires. Other cold water or temperate Aegires that may have previously existed could have radiated from the colder oceans into the warmer waters of the Indo-Pacific. It is notable that within the clade of the Indo-Pacific species is a small clade of all of the Aegires previously known as Notodoris. This clade 1s basally situated to the remaining species and has representative species found along both coasts of Australia, from as far south as the temperate ocean of Perth, Western Australia and extending to latitudes north of the equator. This would provide some evidence for the dispersal of the clade from the Antarctic, north along both coasts of Australia, then into the tropical Indo-Pacific regions. The second major clade is known primarily from the Mediterranean and along the margin of the eastern Atlantic. Only one species of this clade, A. a/bopunctatus, is found along the temperate eastern Pacific, possibly demonstrating vicariance with the closest sister clade of A. palensis and A. punctilucens from the Mediterranean. It is also possible that the ranges of these temperate species are incomplete and additional records may show that there is a higher degree of overlap than is currently recognizable. Finally, species of Aegiridae are extremely small, inconspicuous and infrequently encountered, with the exception of those species previously included in Notodoris. It is likely that not all species have yet been described, and further collections may add to our knowledge of the pattern of species cladogenesis and vicariance. ACKNOWLEDGMENTS The museum material was provided by Kjersti Beate Kvalsvik at the Zoologisk Museum, Oslo (NNM), José Templado at the National Museum of Madrid (MNCN) and Angel Valdés, Natural History Museum of Los Angeles County (LACM; NHM). This study was supported by a National Science Foundation PEET grant DEB 9978155 Phylogenetic systematics of dorid nudibranchs. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 681 LITERATURE CITED ABRAHAM, P.S. 1876. Notes on some genera of nudibranchiate Mollusca, with notices of a new genus and some hitherto undescribed species in the collection of the British Museum. 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Die Gattung Bathydoris Bergh, 1884 (Gnathodoridacea) im phylogetischen System der Nudibranchia (Opisthobranchia, Gastropoda) Zeitschrift fiir Zoologische Systematik und. Evolutionsforschung 27:273-281. WAGELE, H., AND R.C. WILLAN. 2000. Phylogeny of the Nudibranchia. Zoological Journal of the Linnean Society 130:83-181. WELLS, F.E., AND C.W. Bryce. 1993. Sea Slugs of Western Australia. Western Australian Museum, Perth. 184 pp. WILLAN, R. 2000. Family names, particularly Polyceridae and Aegiridae. Nudibranch News 3(3):1-3. Yonow, N. 1993. Opisthobranchs from the Maldive Islands, including descriptions of seven new species (Mollusca: Gastropoda). Revue Francaise d'Aquariologie 20(4):97—130. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 685 Appendix Tables 1—4 TABLE |. Summary of the historical classification of AEGIRIDAE Author and Higher level names used Genus names included Author and Higher level names used Genus names included 1. Alder & Hancock, 1845-1855 SUBORDERS: p. 36-37 ANTHOBRANCHES Dorididae Sub-Family Polycerinae 2. Fischer, 1883 SUBORDERS: p. 519 Anthobranchiata Aglossa: Doridopsidae Glossophora: Cryptobranchiata, Phanerobranchiata POLYCERIDAE: p. 525 Acanthodoridinae Polycerinae Aegirinae 3. Bergh, 1890,1892 Phanerobranchiata Tribe Suctoria Tribe Non Suctoria POLYCERIDAE 4. Eliot, 1910 NOTODORIDIDAE 5. Iredale & O’ Donoghue 1923 AEGIRETIDAE Aegires Lovén, 1844:49 Thecacera Polycera Idalia Aegires Triopella Sars, 1878:310 Aegires Notodoris Bergh, 1875d:64 Triopella Aegires Notodoris Triopella Aegires Notodoris Triopella 6. Thiele, 1929-1935 POLYCERIDAE NOTODORIDINAE p. 696 Aegires Notodoris Triopella 7. Pruvot-Fol, 1954 PHANEROBRANCHIATA Bergh AEGIRETIDAE Aegires (=Notodoridinae Thiele, 1929-35) Notodoris 8. Fischer et al. 1968 PHANEROBRANCHIA Fischer, 1883 Anadoridacea Fischer, 1968 Tribe Non Suctoria Bergh, 1892:141(1133) AEGIRETIDAE Iredale & O’Donoghue 1923 Aegires (= Notodorididae Eliot, 1910) = Anaegires Notodoris Triopella 9. Nordsieck, 1972 AEGIRETIDAE Aegires Anaegires Serigea Nordsieck, 1972:55 Triopella 10. Thompson, 1976 NOTODORIDIDAE Odhner, 1926 Aegires Notodoris 11. Rudman, 1998 AEGIRETIDAE Aegires Notodoris 686 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | | TABLE 2. Characters and states examined for the phylogenetic analysis of Aegiridae. 0 = plesiomorphic; 1,2,3 = apomorphic conditions. Characters Oo WANA MN FW NY n wn ~) oo Lo Ww WwW Ww WwW Wo Wd Ww Lo LW bb N= © Oo — Ann unn Ww No in Amn nn 0. Nn - OR On a (> . Body shape . Dorsal ridges . Dorsal features . Permanent features . Dorsal feature shape . Tubercles . Rhinophores . Rhinophore lamellae . Rhinophore pocket . Rhinophore tubercles . Rhinophore tubercle no. . Oral tentacles/hood . Oral tentacle size . Integumentary spicules . Gill retraction . Gill branches . Gill leaf ramification . Gill arrangement . Gill protective structures . Gill protective structures . Buccal pump . Oral glands . Jaw rodlets . Jaws . Rachidian . Lateral teeth size . Inner lateral teeth shape . Second lateral tooth . Outer lateral tooth Outer lateral teeth shape . Radular teeth denticulation . Reproduction system . Vagina shape . Ampulla shape Penis . Penial hooks . Penial hooks . Penial spines . Penial spines . Vestibular gland . Bursa copulatrix ducts . Pleural ganglia . Cerebral ganglia . Cerebral nerve no. . Eyes . Eye position 7. Ganglionic tubercles . Inner lateral teeth . Gill pocket . Inner two lateral teeth . Foot corners . Foot dimension . Rhinophore stalk Dorsal pigment . Dorsal dark pigment Dorsal tubercle pigment Dorsal ocellae Dorsal rings Rhinophore color 0. Rhinophore color . Gill protective structures . Dorsal webbing . Finger-like structures - Dorsal spicules : wide, distinct mantle; 1: elongate, reduced margin : absent; 1: present : : autotomizable 1: permanent : low; 1: elevated : rounded; 1: flattened; 2: pointed, cones : scattered, 1: in rows : not retractable; 1: retractable : transverse or longitudinal lamellae; 1: smooth : simple hole/slit; 1: raised : none; 1: outer marginal only; 2: all around : many; 1: three to five; 2: one : tentacles: 1: oral hood/veil; 2: neither : large; 1: short/small : present; |:absent : contractile; 1: retractile : few; 1: numerous : multi-pinnate; 1: not multi-pinnate : isolated points; 1: one opening : large, firm; 1: tubercles; 2: elongate appendages : simple; 1: compound/ramified : absent; |: present : absent; 1: present : absent; 1: present : thick, chitinous; 1: thin; 2: absent : present; 1: absent : outer smallest; 1: inner smallest; 2: both small : elongate; 1: hamate; 2: hooked : Same size as inner; 1: markedly reduced : present; 1: absent : hooked; 1: not hooked : no denticles; 1: one spur below main cusp; 2: denticulate : diaulic; 1: triaulic : elongate/thin; 1: wide/bulbous : elongate/sausage-shaped; 1: rounded/bulbous : same width as deferent duct; 1: wider than deferent duct : absent; 1: present : few; 1: many : throughout; 1: tip only : sparse; 1: dense : absent; 1: present : one duct; 1: two ducts : differentiated; 1: fused with cerebral ganglia : two ganglia on each side; 2: fused : three; 1: four or more : absent; 1: present : stalks; 1: prominent; 2: not prominent : absent; 1: present : no secondary cusp; |: with secondary cusp : absent; 1: present : not elongate; 1: elongate : rounded; 1: prolonged : same as mouth; |: narrower than mouth : long; 1: short : no dark pigment; 1: dark pigment : spots; 1: lines none; 1: dark apex : absent; 1: present : absent: 1: present : none; |: dark rings; 2: dark specks : same as body: 1: different from body : absent; 1: present : simple; 1: lobed : smooth; |: furry Volume 55, No. 34 : simple/digitform; 1: same as dorsal tubercles; 2: compound; 3: fan-shaped FAHEY AND GOSLINER: AEGIRIDAE (NUDIBRANCHIA, ANADORIDOIDEA) 687 TABLE 3. Outgroup species and cryptobranch dorids included in the analysis and source of information Literature source Additional material examined Species Actinocyclus verrucosus Ehrenberg, 1831 Bathydoris abyssorum Bergh, 1884 Calycidoris giintheri Abraham, 1876 Conualevia marcusi Collier and Farmer, 1964 Diaphorodoris luteocinta (Sars, 1870) Hexabranchus sanguineus Ehrenberg, 1831; Valdés and Gosliner, 1999; Valdés, 2002 Wagele, 1989; Valdés, 1999, 2002 Abraham, 1876, 1877; Roginskaya, 1972; Valdés, 2002 Valdés, 2002; Collier & Farmer, 1964; Valdés, 2002 CASIZ 018370 (1) Schmekel & Portmann, 1982; Pruvot-Fol, 1954; Valdés, 2002 Cuvier, 1804; Valdés, 2002; Ehrenberg, 1831 CASIZ 072580 (6) CASIZ 071704 (1) (Riippell & Leuckart, 1828) CASIZ 087263 (1) Odhner, 1926; Marcus & Marcus, 1969 Two specimens collected Sept. 1981 by T. Gosliner, Gough Island, Tristan de Cunha Group, So. 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Burgess et al. 6747 (CAS); La Pintada, 49 km S S de Hermosillo, carr. 15, 28°35’N, 110°58'W, A. an Burquez & R. Felger 93-22 (MEXU); Cafion auf Nacapules, ca. 4 km N of Bahia San Carlos, R. : ee ° 3 Felger & B. Straub 85-1316 (CAS, MEXU, TEX); L Ne, ira Mpio. Guaymas, Cafion del Nacapule, 6 km N of are : San Carlos Road, 28°01'N, 111°02'W, R. Felger & F: @ PAphanospeniva'sinalocnsis y *. Reichenbacher 85-1221 (ARIZ, CAS, MEXU, MO, +:27° al eo TEX); Cafion las Barajitas, Sierra el Aguaje, ca. 18 = Yi km NW of San Carlos, ca. 28°03-04'N, 111°11- | aa a8 ee VA 12'W, R. Felger & M. Wilson 95-133 (ARIZ, CAS, =— el MEXU), 95-198 (ARIZ, CAS); Mpio. Guaymas, N FicureE 8. Sonoran distribution of Aphanosperma side of Sierra el Aguaje, Arroyo Las Pirinolas, ca. 5.6 — Sivaloensis. km S from Rancho Las Pirinolas, 28°03’N, 111°03'W, R. Felger et al. 02-246 (CAS); Rio Mayo region, Arroyo Gochico, ca. 4.5 km E of San Bernardo, 27°24'N, 108°47'W, M. Fishbein et al. 2138 (ARIZ, CAS, MEXU); Distr. Alamos, Arroyo Cuchuhacki {Cuchujaqui], H. Gentry 854 (DS, F, MICH); Mpio. Alamos, Sierra de Alamos along Arroyo El Huirotal, ca. 26°S7'N, 108°57'W, V. Steinmann 1290 (ARIZ, CAS, NY); Rio Mayo region, near Tojibampo, 19.5 km NNW of San Bernardo on road to Mesa Colorada, 27°02'N, 108°47'W, T. Van Devender et al. 93-1494 (ARIZ, CAS, MO, TEX). =I Blechum P. Brown Blechum P. Browne, Civ. Nat. Hist. Jamaica 261. 1756. TYPE: Blechum brownei Juss. (= Ruellia blechum L.; = Blechum pyramidatum (Lam.) Utb.). Spreading to decumbent to erect perennial herbs or shrubs with cystoliths. Leaves opposite. Inflorescence of mostly terminal densely bracteate dichasiate spikes; dichasia opposite, 1-3-flow- ered, sessile to subsessile (1.e., peduncles to | mm long), subtended by a bract (or sometimes a dis- tal leaf). Flowers homostylous, sessile, subtended by 2 homomorphic bracteoles. Calyx deeply 5- lobed, lobes homomorphic or subequal in length. Corolla white to blue to purplish, tube expanded distally into a distinct throat, limb subregular and + equally 5-lobed to subbilabiate with upper lip 2-lobed and lower lip 3-lobed, corolla lobes contorted in bud. Stamens 4, didynamous, included in corolla tube or barely exserted from mouth of corolla, anthers 2-thecous, thecae equal in size, par- allel, equally inserted on filament, lacking basal appendages, dehiscing toward lower lip of corol- la (1.e., flower nototribal); pollen (Fig. 3) oblate spheroidal to euprolate, 3-syncolporate, exine fine- ly to coarsely reticulate; staminodes 0. Style included in corolla tube, stigma 2-lobed or with only 1 lobe prominent or evident. Capsule substipitate, ellipsoid, retinacula present, septa with attached retinacula separating from inner wall of mature capsule. Seeds 8-12, homomorphic, lenticular, beset with hygroscopic trichomes on and near margin. (x = 17). This genus of fewer than 10 species is native to tropical America. Blechum pyramidatum, a weedy species, has become naturalized in the Paleotropics. ING PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Blechum pyramidatum (Lam.) Urb., Fedde Repert. Spec. Nov. Regni. Veg. 15:323. 1918. Barleria pyramidata Lam., Encycl. 1:380. 1785. TYPE: SANTO DOMINGO. An illustration of Plumier (Pl. Amer. 2:t. 42, fig. 3. 1756) was cited; specimens, if any exist, not seen. Ruellia blechum L., Syst. Nat., ed. 10, 2:1120. 1759, as “blechnt”. Blechum brownei Juss., Ann. Mus. Natl. Hist. Nat. 9:270. 1807. Blechum blechum (L.) Millsp., Publ. Field Columbian Mus., Bot. Ser. 2:100. 1900. TYPE: illustrations of Sloane (Voy. Jamaica 1:t. 109, fig. 1. 1707) and Plumier (Pl. Amer. 2:t. 42, fig. 3. 1756) were cited (syntypes); see Proc. Calif. Acad. Sci. 48:255. 1995. Phenology. Flowering: March—July, October-November; fruiting: February—July, October— November. Distribution and habitats. Southern United States (Florida), Mexico (Baja California Sur, Campeche, Chiapas, Colima, Guerrero, Hidalgo, Jalisco, México, Michoacan, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosi, Sinaloa, Sonora, Tabasco, Tamaulipas, Veracruz, Yucatan), Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Bolivia, Old World tropics (where presumably introduced); Sonoran plants occur along watercourses and in disturbed areas in regions of tropical deciduous forest at elevations from 150 to 400 m. Illustrations. Figure 9; Fieldiana, Botany 24(10):343. 1974; Flora of the Bahama Archipelago, 1346. 1982; Flora of the Venezuelan Guayana, vol. 2, 344. 1995; Flora of Chiapas 4:17. 1995; Flora del Valle de Tehuacan-Cuicatlan 23:14. 1999; Flora del Bajio 117:13. 2003. Daniel (1995a, 1999a) and Daniel and Acosta (2003) provided descriptions of this widespread and weedy species. Use of the name B. pyramidatum versus B. brownei was discussed by Daniel (1995b). Blechum pyramidatum reaches the northern extent of its distribution in southern Sonora (Fig. 7). Sonoran collections greatly resemble plants described from other regions of Mexico and cleistogamous flowers are sometimes evident among them. SONORAN COLLECTIONS: southeastern border of Alamos (road to mirador), ca. lat. 27°01'N, long. 108°56'W, 7: Daniel 9769 (CAS); Giiirocoba crossing of Rio Cuchujaqui, 12.3 km SSE of Alamos, 26°56'N, 108°53'W, 7: Van Devender 92-514 (ARIZ); Rio Cuchujaqui, Rancho el Conejo, 13.4 km SSE of Alamos, 26°54'N, 108°55'W, T. Van Devender 92-621 (ARIZ); Alamos, 27°01'N, 108°50'W, T: Van Devender 94-204 (ARIZ); El Rancheria crossing of Rio Cuchujaqui, ca. 22.5 km S of Alamos on road to El Chinal, 26°51’N, 108°5S'W, 7: Van Devender & S. Friedman 92-692 (ARIZ, CAS); Rio Cuchujaqui near jct. of Arroyo El Mentidero, 11 km S of Alamos, 26°55’N, 108°55’W, T. Van Devender et al. 92-212 (ARIZ, CAS); Arroyo Alamos between jcts. with Arroyo Potrero and Rio Cuchujaqui, 9 km SE of Alamos, 26°58’N, 108°52'W, T: Van Devender et al. 93-1427 (ARIZ); Rio Cuchujaqui, ca. | km SW of Los Muertos, 26°44'N, 108°55'W, T. Van Devender et al. 94-903 (ARIZ); Rio Cuchujaqui at El Paso, 26°41'N, 108°49'W, 7: Van Devender et al. 95-87 (ARIZ); Alamos, 27°01'N, 108°50'W, 7. Van Devender et al. 2000-13 (MEXU). Carlowrightia A. Gray Carlowrightia A. Gray, Proc. Amer. Acad. Arts 13:364. 1878, nom. cons. LECTOTYPE (Bremekamp, Ind. Nom. Gener. Card 01197. 1956): Carlowrightia linearifolia (Torr.) A. Gray (= Schaueria linearifolia Torr.). Cardiacanthus Nees & S. Schauer in A. de Candolle, Prodr. 11:331. 1847, nom. rej. TYPE: Cardiacanthus neesianus S. Schauer ex Nees. (= Carlowrightia neesiana (S. Schauer ex Nees) T.F. Daniel). Croftia Small, Fl. S.E. U.S. 1088. 1903, non King & Prain (1896). TYPE: Croftia parvifolia (Torr.) Small (= Schaueria parvifolia Torr.). DANIEL: SONORAN ACANTHACEAE 717 718 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Erect to ascending or decumbent perennial herbs or shrubs with cystoliths. Leaves opposite (rarely subopposite). Inflorescence of dichasia in leaf axils, or, more commonly, of dichasiate spikes, racemes, thyrses, or thyrses, the inflorescences sometimes branching and becoming pani- cles; dichasia alternate or opposite, 1-3 (or more)-flowered, sessile or pedunculate, subtended by a leaf or bract. Flowers homostylous, sessile or pedicellate, subtended by 2 homomorphic bracte- oles. Calyx deeply 5-lobed, lobes homomorphic. Corolla white to cream to yellowish to rose-pur- ple to blue, usually with yellow and maroon or purple markings on upper lip, tube cylindric to sub- cylindric, throat indistinct, limb pseudopapilionaceous, 2-labiate, or subactinomorphic, upper lip comprising 2 fused lobes, lower lip consisting of 2 similar lateral lobes and a lower-central lobe that is either similar in form to lateral lobes or + recurved or conduplicate-keeled and enclosing sta- mens and distal portion of style during anthesis, corolla lobes imbricate in bud. Stamens 2, exsert- ed from mouth of corolla, anthers 2-thecous, thecae equal in size, parallel or subsagittate, subequal- ly inserted on filament, lacking basal appendages, dehiscing toward upper lip of corolla (1.e., flow- ers stenotribal); pollen (Fig. 3) prolate spheroidal to perprolate, 3-colporate, 6-pseudocolpate, pseudocolpi 2 per mesocolpium, exine reticulate; staminodes 0. Style exserted from mouth of corolla, stigma 2-lobed. Capsule stipitate, head flattened to nearly spheric, circular to ovate-ellip- tic in outline, retinacula present, septae with attached retinacula remaining attached to inner wall of mature capsule. Seeds 2-4, homomorphic or heteromorphic, flat to concavoconvex, smooth or variously ornamented, lacking trichomes. (x = 18). Twenty-six species are recognized in this New World genus. They occur from the southwest- ern United States throughout Mexico to northwestern Costa Rica and disjunctly to Ecuador and Argentina. With 24 species, Mexico is the center of diversity for Carlowrightia. 1. Corolla 5.5-7 mm long, white with maroon veins on all lobes; dichasia borne in leaf axils throughout plant; bracteoles foliaceous, usually petiolate, narrowly ovate to elliptic-lanceolate, 2—12 mm long, 0.7—5 mm wide; seed margin entire; Chihuahuan desertscrub. ....C. texana 1. Corolla 8-18 mm long, white to yellow with maroon veins (if present) restricted to upper lip; dichasia borne in axils of usually reduced, distal leaves or minute bracts; bracteoles subulate to lanceolate, 1.5—4 (—7) mm long, 0.3—0.8 mm wide; seed margin (entire to) denticulate to dentate to coarsely pectinate; Sonoran desertscrub, thornscrub, and tropical deciduous forest. 2. Leaf blades petiolate, ovate to elliptic, 1.6—2.5 times longer than wide, margin flat; lower- central lobe of corolla conduplicate-keeled, enclosing stamens and style; thecae maroon turning black; capsule glabrous, head 5.5—7.5 mm long; seeds 4, homomorphic 4 ad a cee he aR ee ee ar Tea cite Ci skeathia ale g.clo 6.5.0 6 + C. arizonica 2. Leaf blades sessile to subsessile, linear to narrowly elliptic-lanceolate, 6—29 times longer than wide, margin revolute; lower-central lobe of corolla neither conduplicate nor keel-like, not enclosing stamens and style; thecae golden yellow; capsule pubescent (sometimes only sparsely so at apex), head 3.5—5.5 mm long: seeds 2-4, the third and fourth seeds (when pres- ent) usually markedly different in form from others (i.e., partially conduplicate and with a Conspicuous pectinate marein)s>--.25.22..- 2. «soso o- eee C. pectinata Carlowrightia arizonica A. Gray, Proc. Amer. Acad. Arts 13:364. 1878. TYPE: UNITED STATES. Arizona: Pinal Co., near Camp Grant, 1867, E. Palmer 165 (holotype: GH!). Carlowrightia cordifolia A. Gray, Proc. Amer. Acad. Arts 21:406. 1886. TYPE: MEXICO. Chihuahua: mountains above Batopilas, 1885, E. Palmer 224 (holotype: GH!; isotypes: K!, NY!, US!). Carlowrightia glabrata Fernald, Bot. Gaz. (Crawfordsville) 20:536. 1895. TYPE: MEXICO. DANIEL: SONORAN ACANTHACEAE 719 Sinaloa: Villa Union, Jan 1893, F) Lamb 420 (lectotype, designated by Daniel, 1983a: GH!; isolec- fotypes: DS!, F!, G!, MO!, MSC!, NY!, US!). Carlowrightia californica Brandegee, Zoe 5:172. 1903. TYPE: MEXICO. Baja California Sur: Comondu, 22 Mar 1889, 7. Brandegee s.n. (holotype: UC!). Carlowrightia californica var. pallida 1.M. Johnst., Proc. Calif. Acad. Sci., ser. 4, 12:1169. 1924. TYPE: MEXICO. Sonora: San Esteban Island, 20 Apr 1921, 1. Johnston 3195 (holotype: CAS!: isotypes: GH!, US!). See Daniel (1995a) for a complete listing of synonyms. Phenology. Flowering: February—April, August—October, with a major peak in March-April and a lesser peak in September-October (Fig. 1); fruiting: February—-May, September-November. Distribution and habitats. Southwestern United States (Arizona, California, Texas), Mexico (Baja California, Baja California Sur, Chiapas, Chihuahua, Coahuila, Colima, Guerrero, Jalisco, México, Michoacan, Nayarit, Oaxaca, Puebla, San Luis Potosi, Sinaloa, Sonora), Guatemala, Honduras, Nicaragua, Costa Rica; Sonoran plants occur on slopes, along watercourses, on plains, and in disturbed areas in Sonoran desertscrub (Arizona Upland, Central Gulf Coast, Lower Colorado River Valley, Plains of Sonora), thornscrub, and tropical deciduous forests at elevations from sea level to 800 m. Illustrations. Trees, Shrubs and Woody Vines of the Southwest, 934. 1960; Fieldiana, Botany 24 (10):350. 1974: Desert Plants 5:172. 1984; Trees and Shrubs of Trans-Pecos Texas, 399. 1988; The Jepson Manual, Higher Plants of California, 127. 1993; Flora of Chiapas 4:21. 1995; Flora of the Gran Desierto and Rio Colorado of Northwestern Mexico, 65. 2000. Local names. “Anima aguo” (Mayo, Van Devender et al. 93-1447); “anima ogua” (Mayo, Van Devender et al. 98-2116): “lemilla” (fide Felger 2000); “palo blanco” (Van Devender et al. 93- 1447); “rama toro” (fide Gentry 1942, as C. californica). Use. According to Yetman and Van Devender (2002) the Mayo brew leaves into a tea that is taken for fevers. This tea is considered to be more effective when combined with leaves and inflo- rescences of Elytraria imbricata. Carlowrightia arizonica is widely distrib- >—— = uted in Sonora except for the northeastern por- | i tion of the state (Fig. 10). Although it occurs in. | ~\_ om 100 km all four subdivisions of the Sonoran Desert in | SUN oe the state, it appears to be rare in the Lower ee a Colorado River Valley. It was noted to occur on | ( Isla Tiburon by Moran (1983), but I have no_ | . ry \ record of having seen a collection of it from \ | there. It undoubtedly occurs on that island and | XG je has been collected on the adjacent Isla San AN of” Esteban. Daniel (1983a) provided a description NS HNO ie owe 2 é F \ @ of this widespread and morphologically vari- diy a | ) able species, including all of its known forms. ' Oya gcq Forms occurring in the peninsula of Baja | é : : Fee P J | 1 @ ® { California and Chiapas were subsequently | @ Carlowrightia arizonica Row ey A a described and discussed in more detail by oe \ a Daniel (1995a, 1997). Four more or less dis- | pe tinctive forms have been noted to occur in L_, Bie l aie Ue ak eee Sonora (Daniel 1983a). Many Sonoran plants Figure 10. Sonoran distribution of Carlowrightia ari- do not readily correspond to any of them and = 2enIca. 720 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 others combine characteristics of them in various ways. Perhaps it is more useful to note the range of morphological variation among Sonoran plants of Carlowrightia arizonica than to try to assign specimens to one form or another. Among Sonoran plants, habit varies from spreading perennial herbs to erect shrubs; cauline trichomes vary from evenly to bifariously disposed, retrorsely appressed to retrorse to flexuose to erect, eglandular to glandular (inconspicuous in an understory layer), and 0.05 to 1 mm in length; leaf blades vary from lanceolate to ovate to cordate to elliptic, 3 to 85 mm in length, and | to 65 mm in width; the inflorescence varies from having dichasia in distal leaf axils to having them in bracteate spicate axes, the axes vary from slender (with dichasia 1 per node) to stout (with dichasia opposite at nodes), bracts vary from | to 7 mm in length, and trichomes in the inflorescences vary from eglandular to glandular; calyces vary from 1.5 to 5 mm in length; corollas vary from 9 to 19 mm in length; and capsules vary from 7.5 to 14 mm in length. In any of its guises, this species is readily distinguishable from the other two species of Carlowrightia known from Sonora. SONORAN COLLECTIONS: 9 mi S of Magdalena, L. Abrams 13242 (DS, F); Sierra Lopez Rancho [37 mi NW of Hermosillo], L. Abrams 13321 (DS); 2 mi N of Noria, L. Abrams 13376a (DS, F); ca. 6 mi E of Alamos, F) Almeda 2522 (LA); Cerro El Bachoco, NW of Bahia de Guasimas, 27°54’, 110°40'W, T. Burgess et al. 6941 (ARIZ); 2.5 km S de Hermosillo, 29°01'N, 110°57'W, A. Burquez & A. Quijada 90-211 (MEXU); along road between Arizpe and Ures, 17.1 mi NE of Ures, ca. 29°30’N, 110°08’W, T. Daniel 931 (CAS, ENCB, F, GH, MEXU, MICH, MO, NY, TEX, US); along road between Ures and Hermosillo, 17.1 mi SW of Ures, 7. Daniel 939 (CHAPA, DUKE, MICH, UC); along road between Hermosillo and Kino Bay, 2.5 mi E jet. Son. 16 (Calle 4 Sur) to Guaymas, 7. Daniel 941 (CAS, ENCB, F, GH, MEXU, MICH, US); along road between Hermosillo and Sahuaripa, 24.1 mi W of Mazatan, 7: Daniel 945 (ASU, CAS, DUKE, ENCB, F, GH, K, MEXU, MICH, NY, TEX, UC, US); along road between Hermosillo and Sahuaripa, 3 mi W of Mazatan, T. Daniel 954 (MICH); along road between Mazatan and Sahuaripa, 10.4 mi E of Mazatan, T. Daniel 962 (ENCB, MEXU, MICH, NY, US); along road between Mazatan and Sahuaripa, 5.9 mi E of Rebeiquito and 3.9 mi W of El Novillo, 7) Daniel 965 (CHAPA, MICH); along road between El Novillo and Sahuaripa, 14.4- 17.2 mi E of El Novillo, 77 Daniel 967 (MEXU, MICH), 7: Daniel 971 (ENCB, F, MICH, US); along Son. 16 between Hermosillo and La Colorada, 0.9 mi W of La Colorada, 7; Daniel 974 (MICH); along road between La Colorada and Tecoripa, 12.6 mi E of La Colorada, T Daniel 976-980 (MICH); along road between Tecoripa and Tonichi, 9.3 mi E of Tecoripa, 7; Daniel 983 (ENCB, F, GH, MICH, MO, TEX, UC), 984 (CAS, MEXU, MICH, US), 986 (MICH); along road between Tonichi turnoff and Onavas, 4.4 mi S of Tonichi turnoff, 7) Daniel 987 (MICH), 988 (ENCB, MICH), 989 (ENCB, MICH); along road between Tonichi and Onavas, 7.4 mi N of Onavas, 7. Daniel 990 (MICH); along road between Tonichi and Movas, 2.6 mi S of Rio Chico, 7: Daniel 992 (ENCB, MICH); along hwy. between Esperanza and Rosario de Tezopaco, 3-30 mi E of jet. with road to Presa Obregon, 7. Daniel 996 (CAS, MEXU, MICH, US), 998 (ENCB, MICH, NY); along road between Navojoa and Alamos, 24.6 mi E jet. Hwy. 15 in Navojoa, 7. Daniel 999 (ENCB, F, GH, MICH, MO, NY, UC); along road between Alamos and San Bernardo, 8.1—33.4 mi NE of Alamos, 7: Daniel 1005 (MEXU, MICH, US), 7006 (MICH); along road from San Bernardo to Chihuahua, 1.8 mi from San Bernardo, T. Daniel 1007 (MICH), 1008 (MICH); “Microondas Mountain” overlooking Guaymas, along road which is 1.8 mi from Mex. Hwy. 15 on paved road to airport, N of Guaymas, 7. Daniel 1949 (ASU); Mpio. Soyopa, crossing of Rio Yaqui with Hwy. 16, 8.5 km W of jet. to Onavas, ca. 28°34'N, 109°33’'W, T. Daniel et al. 8509 (CAS, MEXU); SW of Villa de Seris [= Hermosillo on S side of Rio Sonora fide Felger], F’ Drouet et al. 3473 (F); Miramar, NW of Guaymas, R. Felger 5512 (ARIZ); 5.8 mi S of Imuris on Mex. 15, R. Felger 7295 (MEXU); Sierra del Viejo, 50 km W of Los Vidrios on Mex. Hwy. 2, ca. 32°09'N, 113°56’'W, R. Felger & M. Dimmitt 85-723 (ARIZ, MEXU); ca. 5 km SW of Ortiz (SE of Guaymas), R. Felger & N. Gonzales 85-393 (ARIZ, CAS); Pinacate Region, Tinaja de los Papagos, R. Felger & G Joseph 86-485A (ARIZ, MEXU); Canon Nacapules, ca. 4.5 mi N of Bahia San Carlos, R. Felger & D. Valdez Z. 84-614 (ARIZ); 2.7 mi E of Tajitos on Mex. Hwy. 2 (between Caborca and Sonoyta), R. Felger & J. Wright 7420 (ARIZ); El Novillo (E- central Sonora), R. Felger et al. 84-281 (ARIZ, MEXU); Cafion Las Barajitas, Sierra E] Aguaje, ca. 18 km NW of San Carlos, ca. 28°03'N, 111°11'W, R. Felger et al. 95-194 (ARIZ, CAS); Isla San Esteban, vicinity of Limansur, SE side of island, 29°39’N, 112°34'W, R. Felger et al. 17650 (CAS); vic. of Rancho Arivaipa, DANIEL: SONORAN ACANTHACEAE 721 ca. 8 mi ENE of Pozo Coyote, ca. 29°37'N, 112°17'W, R. Felger et al. 17833 (ARIZ); Sierra Seri, 29°17'N, 112°08'W, R. Felger et al. 18131 (ARIZ); 0.2 mi S of San Bernardo, 27°23’N, 108°51'W, M. Fishbein et al. 932 (ARIZ): 5.9 mi SE of San Bernardo toward mine at San Rafael, 27°21'N, 108°49'W, M. Fishbein et al. 938 (ARIZ); Mpio. Hermosillo, La Pintada, 50 km S de Hermosillo, 28°35’N, 110°58’'W, A. Flores M. & J. Sanchez E. 5453 (EB); 15 mi N of Magdalena, R. Fosberg 7901 (LA, PH); 43 km N of Hermosillo toward Nogales, T. Frve & E. Frye 2302 (DS, GH, MO, NY, UC, US); Cd. Obregon, H. Gentry 275 (MICH); Distr. Alamos, Qiiirocoba, H. Gentry 808 (DS, F, MICH); Canyon Sapopa, H. Gentry 1049 (ARIZ, F); KM 180 S of Nogales, H. Gentry 16569 (ARIZ, LL, US); Ures, D. Gold 762 (MEXU); 21 mi E of Tesopaco by old road to Nuri, D. Goldberg s.n. (UC); ca. 30 mi S of Hermosillo, B. Hansen et al. 1379 (LL, MEXU, MICH); Rio El Naranjo, N of Taymuco, 27°15'N, 108°43’W, P. Jenkins & P. Martin 88-253 (ARIZ); Guaymas, I. Johnston 3100 (CAS, US); Onavas road, 28°28-35'N, 109°31-32'W, E. Joyal 1363 (CAS, MEXU); Rancho El Aguilar Noria, N of Ures and Santiago, 29°33’N, 110°25-26'W, E. Joyal 1990 (CAS, MEXU, TEX); 1 mi NW of Navojoa on Hwy. 15, W. Mahler & J. Thieret 6012 (SMU); Mocuzari, La Cruz, 27°13'N, 109°0S'W, P. Martin s.n. (ARIZ, CAS): 5 km N of Rosario de Tesopaco, P. Martin s.n. (ARIZ); 8 km E of Alamos toward Sabinito Sur, P. Martin s.n. (ARIZ); Mazocahui Canyon of the Rio Sonora, 66 mi ENE of Hermosillo near Ures, ca. 110°15'N, 29°28'W, P. Martin & S. Nilsson 38 (S); Estacion Margarita, 10 km NW of Navojoa, 109°29'N, 27°05'W, P. Martin et al. s.n. (ARIZ); 18 mi N of Tesopaco, S. McLaughlin 496 (ARIZ); Mpio. Nacori Chico, El Rio Bonito about La Nopalera, C. Muller 3682 (GH, LL, MICH, UC, US); Agiabampo, FE. Palmer 769 (US): Las Guasimas, B. Perrill 5355 (ARIZ); 21.6 mi S of Santa Ana, B. Perrill 5576 (ARIZ); 8 mi E of Moctezuma, 29°50'N, 109°23'W, F: Reichenbacher 193 (ARIZ); Mpio. Yécora, Arroyo El Encino, ca. 1 km N of Curea, ca. 28°19’N, 109°17'W, A. Reina G. & T. Van Devender 99-450 (CAS); Mpio. Baviacora, ca. 5 km WSW of Mazocahui on Son. 89, 29°32’N, 110°08'W, A. Reina G & T. Van Devender 2000-716 (CAS); Mpio. Arizpe, ca. 19.4 km N of Sinoquipe on Son. 89, 30°19’N, 110°12'W, 4. Reina G. & T. Van Devender 2000-727 (ARIZ, CAS); Mpio. Onavas, Rancho El Palmar, 5 km SE of Rio Yaqui on Mex. 16, 28°30'N, 109°24'W, A. Reina G. et al. 97-93 (ARIZ, CAS, MEXU); Guaymas, H. Ripley 14270 (CAS, NY); vic. of Hermosillo, J. Rose et al. 12352 (NY, US); San Carlos Bay, W of Guaymas, near Cerro Los Algodones, ca. 27°58'N, 111°04'W, A. Sanders et al. 251] (TEX); near Santa Ana, F. Shreve 5957 (F); Rio San Miguel at first crossing S of Horcasitas, F’ Shreve 6695 (ARIZ, MICH, UC); Rio Sonora, 23 mi NE of El Sacaton, F) Shreve 6701 (ARIZ); ca. 2.2 mi NE of Hwy. 15 (toward microwave tower), ca. 6.9 mi SE of Cd. Obregén, W. Stevens & M. Fairhurst 2083 (ENCB); ca. 20 mi SE of Altar on Hwy. 2, R. Straw 2106 (DUKE, SD); Mpio. Yécora, La Concepcion, 29°19'N, 109°02'W, W. Trauba s.n. (CAS); 4 mi S of Mazocahui, R. Turner 59-68 (ARIZ); Isla San Esteban, Arroyo Limantur, 28.7°N, 112.6°W, R. Turner 83-28 (ARIZ); 11 mi E of Hermosillo toward Mazatan, R. Turner & J. Hastings 65-139 (ARIZ, DS, SD); 4 mi S of Torres, R. Turner & J. Hastings 65-179 (ARIZ, DS, SD): 8 mi WSW of Santa Ana, R. Turner & J. Hastings 69-87 (ARIZ); Isla San Esteban, Arroyo Limantur, 7; Van Devender 92-483 (ARIZ, CAS); Mpio. Yécora, El Llano de Curea, 28°18'N, 109°52'W, 7: Van Devender & A. Reina G. 2004-162 (CAS); 21.6 mi S of Santa Ana via Mex. Hwy. 15, 7: Van Devender et al. 84-131 (ARIZ); Rio Cuchujaqui, ca. 8 mi SE of Alamos, T. Van Devender et al. 84-143 (ARIZ); below Guajaray on Arroyo Guajaray, 6.5 km WNW of ject. with Rio Mayo, 27°36'N, 108°56'W, 7. Van Devender et al. 93-455 (ARIZ, CAS); Mpio. Alamos, Cerro El Chorro, 4 km NW of Yocogigua, 26°49'N, 109°03'W, 7: Van Devender et al. 93-1447 (ARIZ); La Gacela, 26°52'N, 108°52'W, 7: Van Devender et al. 95-57 (TEX); Mpio. San Javier, 2.7 mi E of La Barranca on Mex. Hwy. 16, 28°34'N, 109°40'W, T: Van Devender et al. 96- 6 (ARIZ): Mpio. Soyopa, Rio Yaqui bridge on Mex. 16, just S of Tonichi, ca. 28°34'N, 109°33’'W, T. Van Devender et al. 97-1475 (CAS); Ejido Francisco Solis, 7 km (air) ENE of Estacion Luis, ca. 26°35’N, 109°06'W, T. Van Devender et al. 95-2116 (NY); 8 mi S of Nuri, ca. 27°58’N, 109°18'W, G Webster & R. Murphey 24377 (MEXU); S of Magdalena, Whitehead 48 (ARIZ); 17 mi NE of Cajeme on road to Tezopaco, I. Wiggins 6397A (DS, MICH, US); 19 mi NW of Quiriego toward Cajeme, 1. Wiggins 6451 (DS, MICH, US); above second crossing of Rio San Miguel just above Fabrica de Los Angeles, . Wiggins 7294 (ARIZ, DS, MO, TEX, US): 21 mi S of Divisadero, 1. Wiggins 7480 (DS, US); 27 mi W of Hermosillo toward Kino Bay, I. Wiggins & R. Rollins 142 (A, ARIZ, DS, MICH, MO, NY, UC, US). 122 : PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Carlowrightia pectinata Brandegee, Proc. Calif. Acad. Sci., ser. 2, 3:160. 1891. TYPE: MEX- ICO. Baja California Sur: San José del Cabo, 24 Sep 1890, T. Brandegee 452 (holotype: UC!; iso- types: F!, GH!, US!). Carlowrightia fimbriata Brandegee, Proc. Calif. Acad. Sci., ser. 2, 3:161. 1891. TYPE: MEX- ICO. Baja California Sur: San Pedro, 29 Oct 1890, 7’ Brandegee 453 (holotype: UC!; isotypes: CAS!, F!, G!, GH!, MIN!, PH!, US!). Carlowrightia lanceolata Leonard, Kew Bull. 1938:66. 1938. TYPE: MEXICO. México: Distr. Temascaltepec, Tejupilco, 1340 m, 21 Nov 1932, G Hinton 2672 (holotype: K!; isotypes: BNCBIGE! IG!) GE! IMO! INY! PH! IRSA) TEX Us): Phenology. Flowering: September—March; ! fruiting: October—March. Be Distribution and habitats. Western and a southern Mexico (Baja California Sur, Colima, \ México, Morelos, Oaxaca, Sinaloa, Sonora); >3! Sonoran plants occur in canyon bottoms, along \ watercourses, and on slopes in riparian Sonoran | \ desertscrub (Central Gulf Coast), thornscrub, nN and tropical deciduous forests at elevations | 5 Qe from 30 to 850 m. -29° ma e A Illustrations. Figure 9; Proceedings of the | ae Sales o, California Academy of Sciences 49:326. 1997. | ox Local name. “Lemilla” (Mexican, Gentry | “®o tions of Carlowrightia pectinata. The species | attains the northern extent of its distribution in | 114° 12 central Sonora (Fig. 11). Among Sonoran : : plants, color of the corolla varies from white (e.g., Daniel 936) to yellow (e.g., Daniel 3381). SONORAN COLLECTIONS: 6 km E of Navojoa, A. Carter & L. Kellogg 3644 (GH, MEXU, UC); 17.1 mi NE of Ures between Arizpe and Ures, T. Daniel 936 (ASU, CAS, ENCB, GH, MEXU, MICH, NY, US); along Hwy. 16 between Tecoripa and Tonichi, 0.3 mi E of Rio Yaqui, 7) Daniel 3345 (CAS, MEXU, NY, US); Arroyo Cuchujaqui, 7.4 mi SE of Alamos on road to Giiirocoba, 7) Daniel 3381 (ASU, CAS, MEXU); Mpio. Soyopa, crossing of Rio Yaqui with Hwy. 16, 8.5 km W of jet. to Onavas, ca. 28°34'N, 109°33'W, T: Daniel et al. 5500 (BR, CAS, K, MEXU, MO); Canon del Nacapule, ca. 6 km N of Bahia San Carlos, ca. 28°10'N, 111°03'W, R. Felger & A. Burquez 92-1055 (ARIZ, CAS, TEX); Mpio. Guaymas, Cafién del Nacapule, 6 km N of San Carlos Road, 28°01'N, 111°02'W, R. Felger & F: Reichenbacher 85-1212 (ARIZ, CAS, MEXU)); El Novillo, E-central Sonora, R. Felger et al. 84-280 (ARIZ, MEXU, TEX); Cafion Nacapules, ca. 4.5 mi N of Bahia San Carlos, R. Felger et al. 84-603 (ARIZ); Rio Mayo Region, Arroyo Infiernillo, near confluence with Arroyo Tepopa, 27°20'N, 108°44'W, M. Fishbein et al. 993 (ARIZ, CAS); Mpio. Guaymas, Sierra el Aguaje, Aguaje Robinson, ca. 10 km NW de San Carlos, 28°03'N, 111°07'W, A. Flores M. & O. Gutierrez R. 5086 (ARIZ); vicinity of Feo. Sarabia, Cerro Tasirogojo, 5.5 km SE of Melchor Ocampo, 26°31'N, 109°08’W, S. Friedman & E. Espinosa 227-94 (ARIZ); Mesa Masiaca, 6.5 km WNW San José de Masiaca, 26°46'N, 109°18'W, S. Friedman & J. Freeh 376-94 (ARIZ, TEX); vicinity of Sirebampo, 9.5 km S on Mex. Hwy. 15 from Las Bocas road turnoff, 3.5 km W on Sirebampo road, 11.5 km S of San José de Masiaca, 26°39’N, 109°1S'W, S. Friedman & J. Zittere 88-95 (ARIZ); Distr. Alamos, Pichiquate Canyon, H. Gentry 966 (DS), 966M (MICH); San Bernardo, Rio Mayo, H. Gentry 1349 (ARIZ, F, MEXU); Curohui, Rio Mayo, H. Gentry 2144 (ARIZ, F, GH, MEXU, MO, S, UC, US, WIS); Cerro de Bayajori, 12 mi W of Navojoa, H. Gentry 7946 e 1349), @® Carlowrightia pectinata se “¥ Daniel (1983a, 1997) provided descrip- Ware Carlowrightia tevana 110° ! FIGURE 11. Sonoran distributions of Carlowrightia pectinata and C. texana. DANIEL: SONORAN ACANTHACEAE 123, (ARIZ); Sierra Bojihuacame, SE of Cd. Obregon, H. Gentry 14533 (ARIZ, LL); Tepustete microwave tower, W of Alamos, P. Jenkins s.n. (ARIZ, CAS); San Carlos Bay, I. Johnston 4388 (CAS); Onavas road, 28°28- 35'N, 109°31-32'W, E. Joyal 1385 (CAS, MEXU, TEX); Mazocahui Canyon of the Rio Sonora, 66 mi ENE of Hermosillo near Ures, ca. 110°15'N, 29°28'W, P. Martin & S. Nilsson 37 (S); Microondas Masiaca, 7 km W of Masiaca, 26°46°, 109°18'W, P. Martin & M. O'Rourke s.n. (ARIZ); Estacion Margarita, 10 km NW of Navojoa, 27°08'N, 109°29'W, P. Martin et al. s.n. (ARIZ, CAS); turn to La Vifiateria on road between Taymuco and Las Chinacas, P. Martin et al. s.n. (ARIZ); along Hwy. 15 ca. 23 mi N of Navojoa, D. Norris et al. 20083 (CAS, MEXU, MO): W of Guaymas, E. Palmer 259 (ARIZ, C, MICH, S, US); Agiabampo, £E. Palmer 764 (ARIZ, MICH, S, UC, US); Onavas, ca. 28°28’N, 109°32'W, A. Rea 1221 (ARIZ); Mpio. Soyopa, NE side of Rio Yaqui bridge on Mex. 16, just S of Tonichi, ca. 28°34'N, 109°33'W, A. Reina G et al. 97-42 (CAS); Cerro Prieto, 8.6 mi E of Navojoa on road to Alamos, ca. 27°15’N, 109°17'W, A. Sanders et al. 4617 (ARIZ, TEX); Mpio. Alamos, Cerro La Luna, ca. 14 km NW of Alamos, ca. 27°07'N, 109°02'W, A. Sanders et al. 13262 (CAS, TEX); 40 mi NE of Cajeme, F’ Shreve 6156 (ARIZ, F); Cerro Prieto, 1.3 mi up road to microwave tower from road between Navojoa and Alamos, E. Smith 3966 (CAS, MEXU, TEX); Nacopuli Canyon, 28°01'N, 110°03'W, G Starr & C. Starr 216 (ARIZ); Arroyo Gochico, ca. 14 km E of San Bernardo, 27°24'N, 108°41'W, V. Steinmann et al. 601 (ARIZ, CAS, MEXU, MO); Mpio. Navojoa, Cerro Prieto, ca. 14 km E of Navojoa, ca. 27°05'N, 109°17'W, Vo Steinmann 1299 (ARIZ, CAS, IEB, MO, TEX); Cafion La Pintada, ca. 78 km N of Guaymas on Mex. Hwy. 15, Cerro Bola, Sierra Libre, R. Van Devender & T. Van Devender 84-23 (ARIZ); Rio Cuchujaqui, ca. 8 mi SE of Alamos, 7. Van Devender & R. Nishida s.n. (ARIZ); Giiirocoba crossing of Rio Cuchujaqui, 12.3 km SSE of Alamos, 26°56'N, 108°53'W, 7: Van Devender & R. Van Devender 93-156 (ARIZ, CAS): below microwave tower on Cerro Prieto, ca. 9 mi E of Navojoa, 27°05’N, 109°17'W, T. Van Devender et al. 93-243 (ARIZ, CAS); Cerro las Tatemas, below Microondas La Luna, 13.8 km NW of Alamos, 27°07'N, 109°02'W, 7: Van Devender et al. 93-341 (ARIZ, CAS, TEX); Mpio. Alamos, Cerro El Chorro, 4 km NW of Yocogigua, 26°49’N, 109°03'W, T. Van Devender et al. 93-1499 (ARIZ, CAS); La Gacela, 26°52’N, 108°52'W, T. Van Devender et al. 95-57 (ARIZ); Mpio. Soyopa, NE side of Rio Yaqui bridge on Mex. 16 just S of Tonichi, ca. 28°34’N, 109°33'W, T. Van Devender et al. 97-1476 (ARIZ, CAS, MEXU, TEX): Mpio. Ures, SSW of Satebuche, ca. 10 km NW of Nacori Grande, E. slope of Sierra de Mazatan, 29°07'N, 110°08’W, 7: Van Devender et al. 2004-483 (CAS); San Javier, cafion entre los cerros El Potrerito y Los Amoles, 7 km S del poblado, 28°32'N, 109°45'W, L. Varela E. 96-312 (MEXU); 35 mi NE of Cajeme on road to Tesopaco, 1. Wiggins 6415B (DS, MICH, US). Carlowrightia texana Henr. & T.F. Daniel, Madrofio 26:27. 1979. TYPE: U.S.A. Texas: Val Verde Co., along Hwy. 163, 6 mi N of Juno, 19 Jun 1957, D. Correll & M. Johnston 18254 (holo- type: LL!; isotypes: GH!, NY!, SMU!). Phenology. Flowering: September; fruiting: September. Distribution and habitat. Southern and southwestern United States (New Mexico, Texas), northern Mexico (Chihuahua, Coahuila, Nuevo Leon, San Luis Potosi, Sonora); Sonoran plants occur on a roadside in Chihuahuan desertscrub at an elevation of 1215 m. Illustrations. Madrono 26:28. 1979; Trees and Shrubs of Trans-Pecos Texas, 401. 1988. Daniel (1983a) provided a description of Carlowrightia texana. It was only recently (in 2002) collected in Sonora for the first time. The locality in northeastern Sonora (Fig. 11) represents the western extent of the known distribution of the species. It should be sought in similar habitats in nearby Arizona where its occurrence is likely, but has yet to be documented. Sonoran collection: Mpio. Agua Prieta, W edge of Agua Prieta on Mex. 2, ca. 31°19'N, 109°34'W, T. Van Devender et al. 2002-617 (CAS). Dicliptera Juss. Dicliptera Juss., Ann. Mus. Natl. Hist. Nat. 9:267. 1807, nom. cons. TYPE: Dicliptera chinen- sis (L.) Juss. (= Justicia chinensis L.). 724 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Diapedium K.D. Koenig, Ann. Bot. 2:189. 1805 (“1806”), nom. rej. TYPE: Diapedium chi- nense (L.) K.D. Koenig (= Justicia chinensis L.). Solenochasma Fenzl in J. Jacquin, Ecl. Pl. Rar. 2:1. 1844. TYPE: Solenochasma assurgens (L.) Fenzl (= Justicia assurgens L.). Dactylostegium Nees in Martius, Fl. Bras. 9:162. 1847. TYPE: Dactylostegium sparsiflorum Nees. Erect to ascending or decumbent perennial herbs or shrubs with cystoliths, young stems + dis- tinctly 6-angled in cross-section. Leaves opposite. Inflorescence of axillary cymes (= modified dichasia ?) bearing | or more, bracteolate cymules; cymes alternate or opposite, subtended by paired bracts, sessile or pedunculate in leaf axils or in axils of inflorescence bracts forming a ter- minal spikelike thyrse or panicle of thyrses; cymules sessile or pedunculate, comprising an involu- cre of several pairs of bracteoles, outermost pair usually conspicuous and larger than inner (often hyaline) pair(s), cymule bracteoles of a pair equal or unequal in size. Flowers 1-several per cymule, homostylous, sessile. Calyx deeply 5-lobed, usually reduced and hyaline, lobes equal to subequal. Corolla often resupinate (1.e., tube twisted 180°; elsewhere sometimes twisted 360°), pink to pur- ple, red, white, or blue, often with pink to purple markings, tube cylindric to gradually expanded distally but lacking a distinct throat, limb 2-labiate, upper lip comprising 2 fused lobes, entire to emarginate at apex, lower lip entire to shallowly 3-lobed (normal position of lips reversed when corolla resupinate), corolla lobes imbricate in bud. Stamens 2, exserted from mouth of corolla or rarely included in corolla tube, anthers 2-thecous, thecae equal to subequal in size, parallel to per- pendicular, equally to unequally inserted on filament, lacking basal appendages (in Mexican species; elsewhere rarely with lower theca minutely appendaged at base), dehiscing toward lower lip (.e., flower nototribal) in species with corollas either not resupinate or twisted 360°, dehiscing toward upper lip (1.e., flower stenotribal) in species with resupinate corollas; pollen (Fig. 12) euprolate to perprolate, 3-colporate, 6-pseudocolpate, pseudocolpi 2 per mesocolpium, exine retic- ulate; staminodes 0. Style exserted from mouth of corolla or rarely included in corolla tube, stig- ma 2-lobed, lobes equal. Capsule substipitate to stipitate, head ellipsoid to obovoid, retinacula pres- ent, septa with attached retinacula separating elastically and rising from inner wall of mature cap- sule. Seeds 2-4, homomorphic, subreniform to lenticular. (x = 40 in New World taxa; x = 13 or 15? in Old World taxa). Although about 300 species have been described from tropical and temperate regions of the world, the number of species usually given is about one-half that number or fewer. The genus 1s in need of critical taxonomic study. About 20 species occur in Mexico. Dicliptera resupinata (Vahl) Juss., Ann. Mus. Natl. Hist. Nat. 9:268. 1807. Justicia sexangu- laris Cav., Icon. 3:2. 1795, non L. (1753). Justicia resupinata Vahl, Enum. Pl. 1:114. 1804. Diapedium resupinatum (Vahl) Kuntze, Revis. Gen. Pl. 2:485. 1891. TYPE: not designated (see discussion in Daniel, 1997). Dicliptera pseudoverticillaris A. Gray, Proc. Amer. Acad. Arts 20:308. 1885. TYPE: MEXI- CO. Sonora: valley of the Altar, 2 Apr 1884, C. Pringle 27 (lectotype, designated by Daniel, 1997: GH!). Dicliptera torrevi A. Gray, Proc. Amer. Acad. Arts 20:309. 1885. Diapedium torrey (A. Gray) A. Heller, Cat. N. Amer. pl. 7. 1898. TYPE: UNITED STATES. Arizona: unspecified collections of Thurber, Wright, Schott, Rothrock, Lemmon, and Pringle were cited (syntypes, see discussion in Daniel 1997). Dianthera sexangularis Sessé & Moc., Pl. Nov. Hisp. 5. 1887. TYPE: Icones Florae DANIEL: SONORAN ACANTHACEAE : rie ete oS > ata oak be FIGURE 12. Pollen of Sonoran Acanthaceae. decumbens (Jones 24437), apertural view. c. D. - a. Dicliptera resupinata (Daniel 1947), interapertural view. b. Dyschoriste Daniel et al. 8559), interapertural view. e. D. xvlopoda (Wilbur & Wilbur 2137), interapertural view. d. D. hirsutissima hirsutissima (Daniel et al. 8559), apertural view. f. Elytraria imbricata Porter 297), interapertural view. g. Henrya insularis (Daniel 2116), polar view. h. H. insularis (Daniel & Bartholomew 4522), interapertural view. 1. H. insularis (Daniel & Bartholomew 4731), apertural view. Scales 10 pm. 726 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Mexicanae no. 22. Original plate preserved at Hunt Institute for Botanical Documentation, Pittsburgh, Pennsylvania, USA (lectotype, designated by Daniel, 1997). Dicliptera formosa Brandegee, Proc. Calif. Acad. Sci., ser. 2, 3:162. 1891. TYPE: MEXICO. Baja California Sur: summit of Sierra de San Francisquito, 20 Oct 1890, 7: Brandegee 455 (holo- type: UC!; isotypes: GH!, NY). Dicliptera resupinata var. orbicularis B.L. Rob. & Seaton, Proc. Amer. Acad. Arts 28:114. 1893. TYPE: MEXICO. Jalisco: barranca near Guadalajara, Oct 1891, C. Pringle 5169 (holotype: GH; isotype: MEXU!). Phenology. Flowering: throughout the year, with peaks in March-April and September- October (Fig. 1); fruiting: September—June. Distribution and habitats. Southwestern United States (Arizona, New Mexico), western Mexico (Baja California Sur, Chihuahua, Colima, Durango, Guerrero, Jalisco, Michoacan, Nayarit, Sinaloa, Sonora, Zacatecas); Sonoran plants occur on rocky slopes, sand dunes, and along water- courses in Sonoran desertscrub (Arizona Upland, Central Gulf Coast, Lower Colorado River Valley, Plains of Sonora), Chihuahuan desertscrub, palm oases, thornscrub, grasslands, tropical deciduous forests, and oak woodlands at elevations from sea level to 1330 m. Illustrations. Figure 9; Flora of Baja | | California, 190. 1980; Proceedings of the Cali- Se. fornia Academy of Sciences 49:331. 1997. FES ~S Local names. “Alfalfilla” (fide Van SSS Devender et al. 2000; Varela & Cuameca 96- \_3 a 496, 97-56): “himari” (Wahrio, Gentry 1075); | ( O78 “kokpondam” (Pima Bajo, Rea /23/); “puenta | | cita” (Mexican, Rea 1231); “rama del toro” | le (Van Devender et al. 94-226); “yerba de la | Ree mula” (Mexican, Rea 1231). | Daniel (1997) provided a description of this species and discussed some of its morpho- logical variation in northwestern Mexico. In L Sonora, length of the corolla varies from 19-34 | mm, and length of the outer cymule bracteoles | _—_ @ Dicliptera resupinata varies from 6-20 mm. Plants with larger bracteoles and corollas occur in vicinity of Guaymas and San Carlos. Dicliptera resupina- -_1 fe l at fa occurs nearly throughout Sonora (Fig. 13), FIGURE 13. Sonoran distribution of Dicliptera but is not known from the northwestern portion —°“?!74/4- of the state. SONORAN COLLECTIONS: 3 mi S of Magdalena, L. Abrams 13213 (DS, F); Camou, L. Abrams 13273 (DS): 1 mi S of Camou on road to Poza, L. Abrams 13283 (DS); San Miguel River, 15 mi NE of Alamitos near Labor, L. Abrams 13359 (DS); puerto de Hermosillo, G Black 36-6773 (US); Mpio. Guaymas, 12.7 mi S of Puente Douglas across Guaymas Bay, D. Breedlove 1477 (DS); Mpio. Hermosillo, 7 mi W of Hwy. 15 along southern road to Bahia Kino, D. Breedlove 15939 (DS, ENCB):; N end of Sierra Libre, 28°35’N, 110°58'W, 7: Burgess & R. Turner 6722 (ARIZ, MEXU); Microwave Mountain, N of Guaymas, M. Burgess et al. s.n. (ARIZ); San Javier, microondas en el Cerro El Durazno, 28°37’, 109°45'W, A. Burquez 94-009 (MEXU); 2.5 km S de Hermosillo, 29°01'N, 110°57'W, A. Burquez & I. Granillo 93-94 (MEXU), A. Burquez & A. Quijada 91-11 (MEXU), 9/-129 (MEXU); Cafion Tepoca, KM 177 de la Carr. Fed. 16, 28°27'N, 109°16'W, A. Burquez et al. 94-134 (MEXU); Mpio. Benjamin Hill, ca. 22 km N of Benjamin Hill, C. Cowan DANIEL: SONORAN ACANTHACEAE TA et al. 5523 (CAS, MEXU, NY, TEX); 3 mi W of Mazatan, between Hermosillo and Sahuaripa, 7. Daniel 946 (CAS, ENCB); between Mazatan and Sahuaripa, 10.4 mi E of Mazatan, 7. Daniel 963 (ASU, CAS, MEXU); “microondas mountain” overlooking Guaymas, along road to top which is 1.8 mi from Hwy. 15 on road to air- port N of Guaymas, 7. Daniel 1947 (ASU, CAS, NY, US); Rio Cuchujaqui, ca. 8 mi SE of Alamos, 7: Daniel & M. Butterwick 3200 (ASU, CAS); between Esperanza and Rosario de Tezopaco, 3.17 mi SE jct. Hwy. 15, T. Daniel 3360 (CAS); along road to El Tezal, 0.2 mi N hwy. between Navojoa and Alamos, 2.6 mi W of Alamos, T. Daniel 3376 (CAS); near San Carlos (NW of Guaymas), ca. 2 mi E of town, 7. Daniel 3972 (CAS): Mpio. Onavas, Rancho La Mula, near Hwy. 16, 18.9 km SE jet. to Onavas, ca. 28°29’'N, 109° 22’W, T. Daniel et al. S352 (CAS); Mpio. Yécora, along road between San Nicolas and Nuri, just S of Curea, ca. 24 km SW jet. Hwy. 16, ca. 28°19'N, 109°17’'W, T: Daniel et al. 8582 (CAS); Bahia San Carlos, E. Dawson 1071 (F); S of Villa de Seris, F) Drouet & D. Richards 3766 (F); mountain W of Alamos, /) Drowet & D. Richards 4003 (F): S of Villa de Seris, F Drouet et al. 3432 (F); 20 mi NE of Ures, F) Drouet et al. 3591 (DS, F, US); Rancho San Jorge, W of Hermosillo, ca. 29°44'N, 111°51'W, R. Felger 6915 (ARIZ); 0.8 km E of Las Guasimas, R. Felger & R. Devine 85-293 (ARIZ, MEXU); N end of Bahia San Carlos, ca. 0.5 mi inland, R. Fe/ger & C. Lowe 2686 (ARIZ); 9 mi E of Empalme end of Douglas Bridge on Mex. 15, then 2.5 mi S of Hwy. on Playa del Sol road, 27°54'N, 110°41'W, R. Felger & F- Reichenbacher 85-1110 (ARIZ, CAS, MEXU); Cafion las Barajitas, Sierra E] Aguaje, ca. 18 km NW of San Carlos, ca. 28°04'N, 111°12'W, R. Felger & M. Wilson 95- 175 (ARIZ, CAS): El Novillo (E central Sonora), R. Felger et al. 84-265 (ARIZ); Ensenada Grande, ca. 0.5 km inland from S end of bay [NW of Guaymas], R. Felger et al. 11407 (ARIZ); Ensenada Grande (= Bahia San Pedro), R. Felger et al. 11571 (ARIZ, ENCB), //63/7 (ARIZ); Isla Tiburon, ca. 13 mi S of Tecomate, ca. 28°S7'N, 112°27'W, R. Felger et al. 12365 (ARIZ, ENCB), 17309 (ARIZ); Isla Tiburén, ca. 13 mi S of Tecomate, ca. 28°57'N, 112°27'W, R. Felger et al. 17309 (ARIZ); 4 mi W of Alamos toward Navojoa, J. Fish 7 (UC); 3.2 mi SE of San Bernardo to mine at San Rafael, 27°22'N, 108°49'W, M. Fishbein et al. 936 (ARIZ); Mpio. Arivechi, 2 km NW de Arivechi, carr. a Sahuaripa, 28°58’N, 109°10'W, A. Flores M. & M. Arvizo Y. 4756 (ARIZ, MEXU); 75 km N of Hermosillo toward Nogales, 7: Frye & E. Frye 2307 (GH, NY, UC, US); San Lorenzo, H. Gentry 226 (DS, SMU); Distr. Alamos, Canyon Estrella, H. Gentry 379 (ARIZ, DS); Distr. Alamos, Saucito Canyon, H. Gentry 687 (DS, US); Canyon Sapopa, H. Gentry 1075 (ARIZ, US); Canyon Sapopa, Rio Mayo, H. Gentry 1075 (F, GH, MO, WIS); San Bernardo, Rio Mayo, H. Gentry 1160 (ARIZ, F, UC); Alamos, H. Gentry et al. 19347 (ARIZ, US); ca. 0.5 km inland from Algodones Bay, Guaymas area, D. Goldberg s.n. (ARIZ); Granados, C. Hartman 237 (GH, NY, PH, UC, US); 4 mi W of Rancho San Nicolas on road between Nuri and Yécora, P. Hubbell s.n. (ARIZ); N of Cd. Obregon, P. Jenkins s.n. (ARIZ); San Carlos Bay, I. Johnston 4363 (CAS, GH); N of Alamos between Cerros el Cucurucho and El Aguaje, 27°04-05'N, 108°58'W, E. Joyal 1342 (CAS, MEXU); Las Cabras, ESE of Alamos, 27°00'N, 108°55'W, E. Joyal 1522 (CAS); Agua Caliente drainage between Rancho El Alamo and Buena Vista, 29°36'N, 108°56’W, E. Joyal & M. Silva C. 1772 (CAS, MEXU); 4 mi W of Caborca toward Tajitos, D. Keck 4050 (DS, US); 3 mi NW of Tajitos (25 mi NW of Caborca) toward Quitovac, D. Keck 4087 (DS, GH, UC, US); Alamo, Magdalena, P Kennedy 7056 (US); Mpio. Hermosillo, Hwy. 21, 25 km E of Hwy. 15, ca. 3 km E of San Francisco de Batuc, 29°15'N, 110°40’'W, G. Levin 2186 (CAS, MEXU); Badehuachi, C. Lloyd 447 (GH, US); KM 145 on Hwy. 15 between Santa Ana and Hermosillo, M. Luckow 2978 (MEXU, TEX); Cernas, D. MacDougal & F- Shreve 7 (GH, US): 34 mi NE of Cajeme on road to Tesopaco, 7. Mallery s.n. (ARIZ); La Fundicion camp, road to San Pedro, 27°01'N, 108°45'W, P. Martin s.n. (ARIZ); Guaymas, 2 mi below microwave, 27°57'N, 110°54'W, P. Martin s.n. (ARIZ); 1 km NE of Bachoco, 26°44'N, 109°21'W, P. Martin & M. O'Rourke s.n. (ARIZ, MEXU, MO): Micoondas Masiaca, 26°46'N, 109°18'W, P. Martin & M. O’Rourke s.n. (ARIZ); near Mesa La Lagunita on road to Sierra Obscura, 5 mi W of Rancho Agua Blanca, P. Martin & R. Robichaux s.n. (ARIZ); Arroyo San Pablo, ca. 2.5 mi S of Guajaray, 27°35’N, 108°51'W, P. Martin & P. Sundt s.n. (ARIZ); Cerro Verde, along Rio San Javier, 28°34’N, 109°44’W, P. Martin et al. s.n. (CAS); Rancho La Presa (Tetagiosa) and road N 10 km to Alamos-Navojoa road, 27°00’—03'N, 109°04—06'W, P. Martin et al. s.n. (ARIZ); 2 mi SE of Agua Fria on road to La Brisca, NE of Cucurpe, V. McCarten & T. Van Devender 2252 (ARIZ, ENCB); 5 mi N of Tezopaco, S. McLaughlin 488 (ARIZ); Guadalupe Cafion, E. Mearns & E. Merton 2037 (DS, US); San Pedro Bay, 30°03’N, 111°17’W, R. Moran 4039 (DS); Mpio. Nacori Chico, El Rio Bonito about La Nopalera (W slope of Sierra Madre), C. Muller 3632 (F, GH, LL, UC); Magdalena, C. Orcutt 1349 (US); Yaqui River, E. Palmer 5 (GH, NY, PH); Alamos, E. Palmer 632 (GH, NY, US); Yaqui River, E. Palmer s.n. in 1869 (US); 728 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 6 mi N of Obregon, K. Parker 8212 (ARIZ, CAS); 2 mi NE of ariport, NW of Hermosillo, D. Pinkava et al. 942 (ENCB); Torres, C. Purpus 410 (UC); 6 km WNW of San Carlos, 27°57'N, 111°06'W, M. Quinn & P. Sundt 071 (ARIZ); Onvas, along Rio Yaqui, 28°28'N, 109°32'W, A. Rea 1231 (ARIZ); Ensenada Chica, 15.1 mi W of Hermosillo-Guaymas Hwy., 6 mi S of San Augustin Beach turnoff, 28°07'N, 111°17'W, & Reichenbacher 243 (ARIZ); 14 mi by Tecolote road W of Mex. Hwy. 15, 29°48’N, 111°16’'W, & Reichenbacher 1022A (ARIZ); Mpio. Yécora, Restaurant La Palmita, 9.5 km W of Restaurant Puerto de la Cruz on Mex. 16 (KM 258 E of Cd. Obregén), N side of Mesa el Campanero, 28°22'N, 109°04’W, A. Reina G. & T. Van Devender 2000-796 (CAS); Mpio. Opodepe, Arroyo San Cayetano, E side of Querobabi, 30°03’N, 111°O1'W, A. Reina G. & T. Van Devender 2001-595 (CAS); 2.7 km WNW of Tepoca on Mex. 16, ca. 28°28’N, 109°16'W, A. Reina G et al. 98-255 (NY); Playa San Carlos, NW of Guaymas, H. Ripley 14258 (CAS, NY): Arroyo San Xavier between Rancho Lo de Campo and Rancho Panzicola, R. Rondeau s.n. (ARIZ); ca. 5 mi below Minas Nuevas, J. Rose et al. 12674 (NY, US); Sierra de Alamos, vicinity of Alamos, J. Rose et al. 13001 (NY, US); Mpio. Alamos, Alamos Wash on E side of Alamos, ca. 27°02'N, 108°55'W, A. Sanders & G Helmkamp 13306 (TEX); near Mina Sahuarito, S of Cerro San Luis, 20 mi SW of Trincheras, ca. 30°09'N, 111°43'W, A. Sanders et al. 3587 (MEXU); near Sta. Magdalena, Scott s.n. (F); Sta. Magdalena y Imuris, Schott s.n. (F); 14 mi W of Pozo Serna on road to Puerto Libertad, F) Shreve 5452 (ARIZ); near Santa Ana, F. Shreve 5956 (ARIZ, F); Magdalena Canyon, 11 mi NE of Imuris, F) Shreve 6626 (ARIZ, F, GH); 26 mi NW of Caborca, F) Shreve 7556a (ARIZ); ca. 3-4 mi NW of San Carlos, E. Smith 3969 (CAS, MEXU, TEX); 25 mi E of La Motica on road to Tezopaco, W. Spaulding 75-3-16 (ARIZ); Sierra Madre Occidental, ca. 4 mi N of Santa Ana on road to Yepachic, W. Spaulding 75-3-52 (ARIZ); 2.6 mi ESE of Alamos, 27°01'N, 108°54'W, G. Starr & D. Palzkill 322 (ARIZ); 18.5 mi SE of Rio Yaqui crossing on Mex. Hwy. 16, G Starr & C. Starr 189 (ARIZ); Microondas Hill, Guaymas, G Starr & C. Starr 195 (ARIZ); Mpio. Alamos, La Huerta, 1.8 km NNW of Alamos toward San Bernardo, ca. 27°02'N, 108°57'W, V. Steinmann et al. 9324 (ARIZ, MEXU); Rio Yaqui, R. Studhalter 1379 (US); Xorim, R. Studhalter 1447 (US); Agua Prieta, J. Thornber s.n. (ARIZ); Magdalena, G Thurber 1028 (F, GH); San Carlos, L. Toolin & K. Roever 562 (ARIZ); 1.5 mi N of Bavispe, R. Turner & J. Hastings 65-45 (ARIZ, DS); 31 mi E of Hermosillo toward Mazatan, R. Turner & J. Hastings 65-154 (ARIZ, DS); 2.5 mi SW of Atil, 30.8°N, 111.6°W, R. Turner & J. Hastings 72-31 (ARIZ, MEXU); 59 km ESE of Hermosillo on Son. 16, 28°45’N, 110°28'W, T. Van Devender & V. Markgraf s.n. (ARIZ); near Rancho Agua Fria on Rio Saracachi, E of Cucurpe, 7. Van Devender & C. Miksicek s.n. (ARIZ); Playa La Manga, N of San Carlos Bay, 7: Van Devender & F- Nishida s.n. (ARIZ); ca. 4 mi SE of Magdalena, 7. Van Devender & K. Schmidt s.n. (ARIZ); Giiirocoba crossing of Rio Cuchujaqui, 12.3 km SSE of Alamos, 26°56'N, 108°53'W, T. Van Devender et al. 92-940 (ARIZ); El Rancheria crossing of Rio Cuchujaqui, ca. 22.5 km S of Alamos on road to El Chinal, 26°51'N, 108°55’W, 7: Van Devender et al. 93-52 (ARIZ, CAS); Mpio. Navojoa, Arroyo Masiaca, ca. 0.5 km N of Teachive de Masiaca, 26°48’N, 109°14'W, T. Van Devender et al. 93-956 (ARIZ); El Rincon Viejo, ca. 3.4 km N of Alamos, 27°04'N, 108°56'W, T. Van Devender et al. 93- 1063 (ARIZ); Cerro Piedra Boluda, ca. | km NE of El Rincon Viejo, ca. 4.5 km N of Alamos, 27°04'N, 108°56'W, T. Van Devender et al. 94-226 (ARIZ); Mpio. Yécora, El Barranco de la Vinateria Los Sauces, 4 km S of Mex. 16 on road to Santa Ana, 28°23’N, 109°08'W, T. Van Devender et al. 97-228 (MEXU); Mpio. Agua Prieta, Arroyo Gallardo, 8 km E of Agua Prieta on Mex. 2, 31°19'N, 109°23’'W, T. Van Devender et al. 2002-635 (CAS); 3.9 mi SW of Mazocahui, 7. Van Devender et al. s.n. (ARIZ); San Javier, entre los cerros El Potrerito y Los Amoles, 7 km S del poblado, 28°32'N, 109°45'W, L. Varela & E. Cuamea 96-496 (MEXU): San Javier, 28°36’N, 109°45'W, L. Varela & E. Cuamea 97-56 (MEXU); 8 mi S of Nuri, ca. 27°58’N, 109°18'W, G. Webster & R. Murphey 24376 (MEXU); Bavispe, Rio Bavispe, S. White 2886 (ARIZ, GH, US); Hacienda Oquitoa, 6 mi E of Altar, /. Wiggins 5975 (DS, UC, US); Tinaja Picu in Picu Mts., 2 mi N of mon- ument on N side of Libertad Road, /. Wiggins 6052 (DS, US); Rancho San Ignacio, 1. Wiggins 6107 (DS, UC, US); 6 mi S of Carbo, from hwy. to San Miguel, Wiggins 6237 (DS): road to Remedios, 3 mi from jet. Cananea Road, /. Wiggins 7091A (DS); ca. 2 mi E of Rancho Noria de Navaso, /. Wiggins 7254 (DS, US). DANIEL: SONORAN ACANTHACEAE 129 Dyschoriste Nees Dyschoriste Nees in N. Wallich, Pl. Asiat. Rar. 3:75, 81. 1832. LECTOTYPE (Britton & Brown, Ill. Fl. N. U.S., ed. 2, 3:240. 1913): Dyschoriste depressa Nees. Calophanes D. Don in Sweet, Brit. Fl. Gard. ser. 2, 2:t. 181. 1833. TYPE: Calophanes oblongifolia (Michx.) D. Don (= Ruellia oblongifolia Michx.). Linostylis Fenzl ex Sond., Linnaea 23:94. 1850. TYPE: Linostylis ovata Sond. Decumbent to erect perennial herbs with cystoliths. Leaves opposite. Inflorescence of dicha- sia in leaf axils throughout plant or restricted to axils of distal leaves or bracts and forming a spike or thyrse; dichasia alternate or opposite, 1—many-flowered, sessile to pedunculate, subtended by a leaf or bract. Flowers homostylous, sessile to subsessile (in ours), subtended by 2 homomorphic bracteoles. Calyx 5-lobed, tube often as long as or longer than lobes during anthesis, regions between lobes usually subhyaline, often splitting nearly to base in fruit, lobes equal to subequal in length, usually somewhat setaceous. Corolla blue to blue-purple (in ours, elsewhere also pinkish purple, red, white, and yellow), tube gradually or abruptly expanded distally into a + distinct throat, limb subactinomorphic to 2-labiate, upper lip 2-lobed, lower lip 3-lobed, corolla lobes contorted in bud. Stamens 4, didynamous, filaments connate in pairs (1.e., a long and a short stamen connate) proximally, exserted from mouth of corolla (at least longer pair and usually with at least a portion of anthers of shorter pair also), anthers 2-thecous, thecae equal in length, parallel to subsagittate, equally inserted, appendaged at base with awns or stout trichomes (in ours, elsewhere sometimes unappendaged at base), dehiscing toward lower lip (1.e., flower nototribal); pollen (Fig. 12) sub- prolate to perprolate, 3-colporate, irregularly polypseudocolpate, pseudocolpi 4-12 per mesocolpi- um, exine minutely verrucate to microechinate (and often microfoveolate to microfossulate); sta- minodes (0). Style exserted from mouth of corolla, stigma unequally 2-lobed, | lobe greatly reduced, rudimentary, or sometimes not evident. Capsule substipitate, subellipsoid to ellipsoid, retinacula present, septae with attached retinacula remaining attached to inner wall of mature capsule. Seeds 2-4, homomorphic, lenticular, covered with appressed hygroscopic trichomes. (x = 15). Dyschoriste consists of approximately 75 species occurring in tropical and warm-temperate regions of America, Africa, and Asia. The genus is best developed in the New World where it has a discontinuous distribution from the southern United States southward through Mexico and Central America to northern Argentina. Major concentrations of species are found in west-central and southern Mexico and southeastern Brazil. The actual number of Mexican species is likely fewer than the 22 presently recognized. Dyschoriste is a complex genus the described species of which offer few characters for their recognition. Kobuski’s (1928) treatment of the American species is now outdated and inadequate both for delimiting and identifying taxa. 1. Herbage glandular or glands restricted to calyx, external surface of lower lip of corolla, and base GE SIS AG: <6 ce Bene Ce a Le aia ors 1) TCA Gna ee Ae A ORAL Ca ee AS Be D. hirsutissima 1. Plants lacking glandular trichomes. 2. Plants usually decumbent; stems puberulent with trichomes less than 0.05 to 0.2 mm long; calyx puberulent with mostly appressed trichomes 0.5—0.2 mm long; corolla 13—21 (—25) mm long; thecae (including basal appendage) 0.8—-2 mm long. .......... D. decumbens 2. Plants mostly erect to diffuse; stems pubescent with at least some (usually all) trichomes 0.3-1.5 mm long; calyx pubescent with flexuose trichomes 0.5—2 mm long; corolla 20—29 mm long; thecae (including basal appendage) 2—2.6 mm long............. D. xylopoda 730 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Dyschoriste decumbens (A. Gray) Kuntze, Revis. Gen. Pl. 2:486. 1891. Calophanes decum- bens A. Gray, Syn. Fl. N. Amer. 2(1):325. 1878. TYPE: MEXICO. Sonora: “near Santa Cruz, stony soil, in mountain valleys,” 1851, C. Wright 1462 (lectotype, designated by Henrickson, 1999: GH, photo at CAS!; isolectotypes: GH?, NY?). Phenology. Flowering: March—September; fruiting: July-September. Distribution and habitats. Southwestern United States (Arizona, New Mexico, Texas), Mexico (Baja California Sur, Chihuahua, Coahuila, Durango, Hidalgo, Querétaro, San Luis Potosi, Sonora, Zacatecas); Sonoran plants occur on rocky slopes in grasslands, oak woodlands, oak forests, and oak-pine forests at elevations from 1390 to 1800 m. Illustrations. Flora of Baja California, 190. 1980; Desert Plants 5:169. 1984; Proceedings of the California Academy of Sciences 49:337. 1997. In the protologue of Calophanes decumbens, Gray cited unspecified collections of Wright, Thurber, Rothrock, and others from western Texas, southern Arizona, and adjacent Mexico. Henrickson (1999) designated a specimen of Wright 1462 at GH as the lectotype of that name. He further indicated that the lectotype was collected in the United States (Arizona). Whether the lec- totype was collected in Arizona or Sonora (as treated here) depends on the date it was collected. It was undoubtedly collected in September of 1851 when Wright traveled through Santa Cruz Co., Arizona between Patagonia and Canelo Pass to Santa Cruz, Sonora (ca. 16 km S of the internation- al border). Most of collections prior to 23 September were made in Arizona, those of 23 September were probably mostly made in Arizona, and those made from 24 September until 2 October were from Sonora (Shaw 1987). Shaw (1987) provided correlations between Wright’s field numbers and localities, but there is no field number on either the lectotype or the isolectotype at GH. Nor are there any collections listed by Shaw (1987) in the field lists with the exact locality, habitat, and plant habit information as presented on the lectotype (see type citation above and habit described as “stems many, procumbent”). Based on the locality provided on the lectotype and localities in Wright’s field lists (as provided by Shaw 1987), it seems more likely that this plant was collected in the grassland communities near Santa Cruz (in Sonora) rather than in similar habitats in nearby Arizona. Henrickson (1999) also noted that the number “1462” was assigned by Asa Gray to all of Wright’s collections of this taxon, and that Wright 1462 at MO from “valley of the Rio Grande” 1s therefore “not considered type material.”” However, Henrickson (1999) also indicated that Wright 1462 at NY and another collection with this number at GH are isolectotypes. The label on these “isolectotype” specimens bear no locality data and therefore it remains a matter of speculation as to whether they represent type material or not. Daniel (1984b, 1997) provided descriptions of this species. Henrickson (1999) treated D. decumbens as one of several varieties of a polymorphic D. schiedeana (Nees) Kuntze. He studied this difficult genus in considerable detail throughout north-central Mexico and the southwestern United States over many years. As a result, he is far more familiar with Dyschoriste in these regions than | am, and he is probably correct that D. decumbens is not deserving of specific rank. The prob- lems for taxonomic delimitation that he noted (Henrickson 1999) are not restricted to the south- western United States and northern Mexico, however. Study of plants occurring further to the south in Mexico reveals similar complexities involving these and other taxa (e.g., Daniel and Acosta 2003, cf. D. microphylla (Cav.) Kuntze), some of which have older names than D. schiedeana. Until such time as a satisfactory taxonomy has been promulgated for the genus throughout North America, | prefer to use traditionally recognized taxa with familiar names for regional treatments such as this one. The leaves of Van Devender et al. 98-882 vary from linear to narrowly elliptic (vs. elliptic to DANIEL: SONORAN ACANTHACEAE 731 oblanceolate to obovate in most other Sonoran i collections of the species). This species ts like- ly more common in the northern and eastern portions of Sonora than the relatively few known collections from the state (Fig. 14) would suggest. SONORAN COLLECTIONS: Yécora, 0.5 mi E of a * | Arroyo Yécora, 28°23’N, 108°54'W, M. Fishbein et ie ne lew al. 2487 (ARIZ); S of Pena Blanca on Hwy. 16, x | 28°23'N, 109°04'W, P. Martin & G Ferguson s.n. p ZN (ARIZ); Mesa Grande, 28°26'N, 108°59'W, P. aS Martin et al. s.n. (ARIZ); Hwy. 16, 5.6 mi E of Rio | ows Maycoba, 6.5 mi W of Maycoba, 28°23’N, at e\ 108°43'W, S. McMahon et al. 177 (ARIZ); Rancho hike AS La Pinosa, 5.9 mi W of Maycoba on Hwy. 16, | eden Sane a 4 % 28°25'N, 108°43'W, S. McMahon et al. 179 (ARIZ); 27° © Dyschoriste xylopoda —— wey “Sonora,” G Thurber 114 (GH); Mpio. Yécora, 0.2 ensues, km N of Yécora on road to Agua Blanca, 28°23’N, 14° hes ane WY 108°56'W, W. Trauba s.n. (CAS), T; Van Devender et -— l l l | | ___ al. 97-786 (ARIZ, CAS): Mpio. Yécora, Rancho La FIGURE 14. Sonoran distributions of Dyschoriste Pinosa. 9 km E of Rio Mavycoba. 10.3 km W of @ecumbens, D. hirsutissima, and D. xylopoda. Maycoba on Mex. 16, 28°25’N, 108°43’W, T. Van Devender et al. 98-882 (CAS); Mpio. Santa Cruz, 3 km N of Santa Cruz on road to Nogales, 31°16’N, 110°36’W, T. Van Devender et al. 2001-704 (CAS); Mpio. Ures, Canada El Yugo, Sierra de Mazatan, 29°06’N, 110°12’W, T. Van Devender et al. 2004-478 (CAS); KM 223, Rt. 69, 13 km NE of Guasabas, 7. Walker & S. Walker s.n. (ARIZ); region of the Rio de Bavispe, Horconcitos, Arroyo del Salto, S. White 3761 (ARIZ); region of the Rio de Bavispe, El Tajo, Rio Fronteras, S. White 4060 (ARIZ). 100 km Dyschoriste hirsutissima (Nees) Kuntze, Revis. Gen. Pl. 2:486. 1891. Calophanes hirsutis- simus Nees in A. de Candolle, Prodr. 11:109. 1847. TYPE: MEXICO. Hidalgo: “supra Hacienda de Guadalupe” [vicinity of Zacualtipan where Ehrenberg collected in November and December of 1839 fide Urban (1897)], Dec, C. Ehrenberg 1223 (holotype: B, destroyed; fragment and photo: MO). Calophanes bilabiatus Seem., Bot. Voy. Herald. 324. 1856. Dyschoriste bilabiatus (Seem.) Kuntze, Revis. Gen. Pl. 2:486. 1891. TYPE: MEXICO. Sinaloa: “Cerro de Pinal,” Dec 1848, B. Seemann 1513 (holotype: K!). “Calophanes bilobatus Rose” as treated in the synonymy of D. hir- sutissima by Kobuski (1928), is treated as an orthographic error for C. bilabiatus because Rose (Contr. U.S. Natl. Herb: 1:109. 1891) cited the name as “Calophanes bilobatus Seem.” Ruellia viscosa Sessé & Moc., Pl. Nov. Hisp. 100. 1889, non Ruellia viscosa H.B.K. (1817). TYPE: not designated, see below. Phenology. Flowering: March—July, October-December; fruiting: March—April, July, December. Distribution and habitats. Mexico (Chiapas, Chihuahua, Colima, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, México, Michoacan, Morelos, Nayarit, Oaxaca, Puebla, Sinaloa, Sonora, Veracruz), El Salvador; Sonoran plants occur in thornscrub, tropical deciduous forests, oak grasslands, oak forests, and various riparian habitats at elevations from 220 to 1200 m. Illustrations. The Botany of the Voyage of H.M.S. Herald, t. 65. 1856; Flora del Valle de Tehuacan-Cuicatlan 23:29. 1999. W32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Local names. “Ciatica” (fide Van Devender et al. 2000); “valeriana del monte” (fide Van Devender et al. 2000). Ruellia viscosa Sessé & Moc. 1s here treated as a synomym of D. hirsutissima for the first time. In the protologue of the former name, a locality is given as “in aridis Ayacapixtlae agris.” According to McVaugh (2000) this is Yecapixtla, Morelos. In the Sessé and Mocifio herbarium at MA no. 2/56 (CNHM negs. 48997-48999) is labelled with this name and pertains to Dyschoriste hirsutissima (Nees) Kuntze. A painting in the Torner Collection (6331.0634) at the Hunt Institute for Botanical Documentation, Pittsburgh, U.S.A. is also labelled with this name and could repre- sent D. hirsutissima. The description in the protologue, especially the foliar pubescence being “‘vil- loso-viscosa,” would appear to pertain to this species as well. Daniel (1995a, 1999a) and Daniel and Acosta (2003) provided descriptions of Dyschoriste hir- sutissina. The vegetative organs of plants of this species are commonly viscid and have a some- what mephitic odor (similar to that of Ruellia inundata). The species attains the northern and west- ern extents of its distribution in southeastern Sonora (Fig. 14). SONORAN COLLECTIONS: Mpio. San Javier, 2.7 km E of La Barranca on Mex. 16, 28°34’N, 109°40'W, A. Burquez M. 96-135 (CAS, MEXU); along Hwy. 16 between Tecoripa and Tonichi, 7.6 mi W of Rio Yaqui, T. Daniel 3344 (CAS); Mpio. Yécora, along Hwy. 16, 15.5 km NW of Tepoca, ca. 28°28'N, 109°21’W, T Daniel et al. 8559 (CAS): W of Alamos, F? Drowet & D. Richards 3982 (CAS, DS, F, NY, US); 5.9 mi SE of San Bernardo on road up Arroyo Taymuco to mine at San Rafael, 27°21'N, 108°49'W, M. Fishbein et al. 940 (ARIZ, CAS); Sierra de Alamos, Arroyo Uvalama, 27°00'N, 108°59'W, M. Fishbein et al. 1098 (ARIZ); 8.2 mi S of Guajaray on road to San Bernardo, 27°32'N, 108°56'W, M. Fishbein et al. 1592 (ARIZ); KM 196 on Hwy. 16 W of Tepoca, G. Ferguson & C. Moore s.n. (ARIZ); Distr. Alamos, Qiiiricoba, H. Gentry 757 (DS), 757M (F); Arroyo Gochico, Rio Mayo, H. Gentry 3634 (ARIZ, F, MEXU); Mpio. Alamos, Rio Cuchujaqui near San Pedro, 27°02'N, 108°42'W, P. Jenkins 89-427 (ARIZ), 89-428 (ARIZ); Sierra de Alamos, 27°54'N, 108°S7'W, P. Jenkins s.n. (ARIZ); 20 mi E of Rio Yaqui on Mex. Hwy. 16, G Joseph & B. Perrill 83-042 (ARIZ); Puenta La Pila above “El Palmar,” ca. 20 km E of Onavas, 28°29’N, 109°22'W, E. Joyal 1567 (CAS, MEXU); Arroyo Verde, P. Martin s.n. (ARIZ); 12 km E of Alamos, P. Martin s.n. (ARIZ, CAS); Rio Yaqui drainage, Cerro Verde, 28°34'N, 109°44’W, P. Martin & G. Ferguson s.n. (ARIZ, CAS); above Rancho Palo Injerto, 27°03'N, 108°44'W, P. Martin & M. O'Rourke s.n. (ARIZ); Rio Cuchujaqui, between Rancho San Pedro and Cerro El Tarahumara, ca. 27°03'N, 108°42-43'W, P. Martin et al. s.n. (ARIZ); Jorinabo, 27°15'N, 108°46'W, P. Martin et al. s.n. (ARIZ); Alamos, E. Palmer 402 (US); 18 mi SE of Rio Yaqui on Mex. Hwy. 16, R. Perrill 5322 (ARIZ); Mpio. Yécora, 3 km N of Tepoca on Mex. 16, 28°27’N, 109°16'W, A. Reina G & T. Van Devender 97-349 (CAS); Mpio. Alamos, Upper Rio Cuchujaqui, Arroyo Verde, 27°06'N, 108°43'W, R. Rondeau & G. Rodda 90-68 (ARIZ); Sierra de Alamos, vicinity of Alamos, J. Rose et al. 12833 (US); 7 mi N of Giitrocoba, 26°59'N, 108°41'W, J. Salmon s.n. (ARIZ): Sierra de la Cebollita, 6.4 mi S of Nuri along road from Tesopaco, ca. 28°04'N, 109°20'W, A. Sanders 3689 (CAS); Mpio. Alamos, SW edge of Alamos in foothills of Sierra de Alamos, ca. 27°01'N, 108°57'W, A. Sanders et al. 13359 (CAS, TEX); Mpio. Alamos, Canon Los Laureles, E slope Sierra de Alamos, 108°58’W, 26°58'N, 4. Sanders et al. 14343 (CAS, MO); Arroyo Gochico, ca. 10 km E of San Bernardo, ca. 27°24'N, 108°44'W, V. Steinmann et al. 605 (ARIZ, CAS, MEXU, NY); Sierra de Alamos, W of Aduana, ca. 6 km W of Alamos, 27°02'N, 109°O1'W, V. Steinman et al. 93-122 (ARIZ, MEXU); 21 mi E of Tonichi (at Rio Yaqui) on road to Yécora, L. Toolin 296 (ARIZ); Canyon las Piedras, Sierra de Alamos, 3.5 km S of Alamos, 26°59'N, 108°57'W, T. Van Devender et al. 92-743 (ARIZ, CAS); El Guayabo crossing of Rio Cuchujaqui, 14 km ESE of Alamos, 27°00'N, 108°47'W, T: Van Devender et al. 93-170 (ARIZ); 4.5 km ESE of El Caracol on Alamos-Navojoa Road, 27°04'N, 109°02'W, T. Van Devender et al. 93-225 (ARIZ, CAS, TEX): Rancho La Junta, 1 km upstream from Mesa Colorada, 3 km NNE of Burapaco, 27°35'N, 108°52’W, T. Van Devender et al. 93-395 (ARIZ, CAS); near Tojibampo, 19.5 km NNW of San Bernardo on road to Mesa Colorado, 27°02'N, 108°47'W, T. Van Devender et al. 93-1491 (ARIZ): Mpio. San Javier, 2.7 km E of La Barranca on Mex. 16, 28°34’N, 109°40’W, 7. Van Devender et al. 96-3 (ARIZ, CAS, MEXU, MO, NY, TEX); Mpio. Yécora, Agua Amarilla, 15 km WNW of Tepoca, ca. 28°08'N, 109°20'W, T. Van Devender et al. 96-21 (ARIZ); 3 km E turnoff to Santa Ana on Mex. 16, 28°25’N, DANIEL: SONORAN ACANTHACEAE 133 109°07'W, 7. Van Devender et al. 97-248 (MEXU); Mpio. San Javier, Puerto San Juan, Cerro San Juan, near San Javier, 28°35'N, 109°45'W, L. Varela E. 97-55 (CAS): San Javier, Cerro San Juan, 28°36'N, 109°45’W, L. Varela & E. Cuamea 97-80 (MEXU); above Aduana, Sierra de Alamos, 7.5 km ESE of Alamos, 27°02’N, 109°01'W, J. Wiens et al. 93-078 (ARIZ, CAS). Dyschoriste xylopoda Kobuski, Ann. Missouri Bot. Gard. 15:54. 1928. TYPE: MEXICO. Jalisco: near Guadalajara, 19 Jul 1893, C. Pringle 4442 (holotype: MO!; isotypes: F, GH!, MEXU!}, MO!, PR!, PRC!, UC!, US!). Erect to spreading (to decumbent) perennial herbs to 4 dm tall. Young stems subquadrate to guadrate-sulcate, + evenly to + bifariously pubescent with flexuose eglandular trichomes 0.2—2 mm long (sometimes with an understory of shorter, + bifariously disposed, and antrorse trichomes as well). Leaves subsessile to short-petiolate, petioles to 2 mm long, blades linear to narrowly ellip- tic to elliptic to obovate, 10-36 mm long, 2-17 mm wide, 2.3—9 times longer than wide, rounded to acute at apex, acute to subattenuate at base, surfaces sparsely pubescent (sometimes nearly glabrous) with flexuose eglandular trichomes to 1.5 mm long, margin entire, ciliate (at least near base of blade) with similar trichomes. Inflorescence of dichasia borne in axils of leaves along dis- tal portion of stems; dichasia opposite, 1 per axil, 1-many-flowered, pedunculate, peduncles 0.5—1.5 mm long. Bracteoles foliose, linear to narrowly elliptic to obovate, 8-21 mm long, 0.8—4 mm wide, sometimes slightly curved, abaxial surface pubescent like leaves, secondary bracteoles similar to bracteoles except often smaller. Flowers sessile to subsessile (1.e., pedicels to | mm long). Calyx 9.5-17 mm long, tube 3—7 mm long, .27—.44 times as long as calyx, lobes subulate- setaceous, 6.5—10 mm long, |1.3—2.7 times longer than tube, abaxially and marginally (at least near base) pubescent with flexuose eglandular trichomes (sometimes dense) 0.5—2 mm long. Corolla bluish-purple, 20-29 mm long, externally pubescent with flexuose eglandular trichomes 0.1—0.5 mm long, tube expanded just distal to midpoint, 15—19.5 mm long, 2.8—3 mm in diameter near mid- point, limb 15-20 mm in diameter, upper lip 7.5—9.5 mm long, lobes 4.5-6 mm long, 4-5.8 mm wide, lower lip 7.5—10 mm long, lobes 6—8.5 mm long, 4-6 mm wide. Stamens with longer pair 7-13 mm long, shorter pair 6—11.5 mm long, thecae parallel, 2—2.6 mm long (including basal appendage), awned at base, awn 0.2—0.3 mm long. Style 19-22 mm long, pubescent with eglandu- lar trichomes, stigma 1.2—1.6 mm long, only | lobe evident. Capsule 8-10 mm long, glabrous. Phenology. Flowering: July-September; fruiting: August—October. Distribution and habitats. Western Mexico (Aguascalientes, Chihuahua, Durango, Jalisco, Nayarit, Sonora); Sonoran plants occur on slopes in oak woodlands and oak-pine woodlands at ele- vations from 900 to 1600 m. Illustration. None found. Sonoran plants resembling those of Dyschoriste xvlopoda from the Nueva Galicia region to the south are treated as conspecific with them pending a satisfactory taxonomic treatment of the genus in North America. The species attains the northern and western extents of its distribution in the upper drainage of the Rio Mayo in southern Sonora (Fig. 14). Most Sonoran plants are erect to spreading in habit. Pennell 19615 is described as a “depressed herb” and looks to be spreading on the ground with upturned distal shoots. Thus, it has a habit similar to most plants of D. decumbens. The relationship of this species to that traditionally referred to D. ovata (Cav.) Kuntze (see discus- sion of this name in Daniel, 1995a) remains to be resolved. SONORAN COLLECTIONS: ca. 35 km NE of Alamos, between El Chiribo and La Vinateria, 27°18’N, 108°42'W, M. Fishbein et al. 1440 (ARIZ, CAS): 1 km E of Los Llanitos on road to Chiribo from Huicoche, 27°19'N, 108°39'W, P. Martin & D. Yetman s.n. (ARIZ); ridge S of Arroyo Gochico, E of San Bernardo, F: Pennell 19540 (PH, US); Cerro Saguarivo, E of San Bernardo, F! Pennell 19615 (PH, US); Rancho Santa 734 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Barbara, E of Alamos, 27°07'N, 108°43’'W, J. Rascon & C. Smith s.n. (ARIZ); Sierra Sahuaribo, ca. 3 km N of La Lobera toward Huicochic, ca. 27°17'N, 108°37'W, V. Steinmann et al. 93-237 (CAS). Elytraria Michx. Elytraria Michx., Fl. Bor.-Amer. 1:8. 1803, nom. cons. TYPE: Elytraria virgata Michx., nom. illegit. (= Elytraria caroliniensis (J.F. Gmel.) Pers.). Tubiflora J.F. Gmel., Syst. Nat. 2:27. 1791, nom. rej. TYPE: Tubiflora caroliniensis J.F. Gmel. Erect to ascending, acaulescent to caulescent perennial herbs lacking cystoliths. Leaves alter- nate, mostly in basal rosettes or crowded at apices of branches, sometimes + diffuse along stems. Inflorescence of scapose or pedunculate densely bracteate axillary and terminal dichasiate spikes; spikes cylindric, simple or sometimes branched, scapes or peduncles covered with imbricate, cori- aceous, clasping scales; dichasia alternate (spirally arranged), 1-flowered, sessile in axil of a bract. Flowers homostylous, sessile, subtended by 2 homomorphic (often mostly hyaline) bracteoles. Calyx deeply 4-lobed, mostly hyaline, lobes heteromorphic, anterior and posterior lobes external, anterior lobe 2-dentate to + deeply 2-cleft. Corolla relatively small, pinkish or blue or white or yel- low, often with colored markings, tube cylindric or slightly expanded near mouth, throat indistinct or evident only near mouth, limb 2-labiate, upper lip 2-lobed, lower lip 3-lobed, corolla lobes often apically divided or 2-cleft, imbricate in bud. Stamens 2, anthers partially exserted from mouth of corolla, 2-thecous, thecae (covered by stigma during anthesis) equal in size, parallel, equally insert- ed on filament, lacking basal appendages (at least in Mexican species), dehiscing toward lower lip (1.e., flower nototribal); pollen (Fig. 12) euprolate to perprolate, 3-colpate, exine finely reticulate; staminodes 0-2, minute. Style exserted from mouth of corolla, stigma unlobed, expanded, flat, subelliptic to subspatulate, folded over anthers during anthesis and straightening when touched (touch-sensitive) and gradually refolding. Capsule estipitate, subconic to ovoid, sometimes irregu- larly constricted proximally, retinacula absent, placentae minute, papilliform. Seeds numerous (up to 20 per capsule), irregularly shaped (often blocky or cubelike), lacking trichomes. (x = 11 or 12?). Elytraria consists of about 15 species occurring in the tropics and subtropics of both the Old World and the New World. The majority of species are American with four occurring in Mexico. Leonard’s (1934) brief account of the American species is much out of date. Elytraria imbricata (Vahl) Pers., Syn. Pl. 1:23. 1805. Justicia imbricata Vahl, Eclog. Amer. 1:1. 1796. TYPE: not located, see Daniel (1995a). Verbena squamosa Jacq., Pl. Hort. Schoenbr. 1:3. 1797. Tubiflora squamosa (Jacq.) Kuntze, Revis. Gen. Pl. 2:500. 1891. Elytraria squamosa (Jacq.) Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica 8:299. 1895. TYPE: unknown. Elytraria tridentata Vahl, Enum. Pl. 1:107. 1804, nom. illegit. (Justicia imbricata Vahl cited as synonym). See Daniel (1995a) for a complete listing of synonyms for this species. Phenology. Flowering: August—May, but with peak flowering periods in March and September (Fig. 1); fruiting: throughout the year. Distribution and habitats. Southwestern United States (Arizona, New Mexico, Texas), Mexico (Baja California Sur, Campeche, Chiapas, Chihuahua, Coahuila, Colima, Durango, Guanajuato, Guerrero, Jalisco, México, Michoacan, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, Sinaloa, Sonora, Veracruz, Yucatan, Zacatecas), Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Argentina, West Indies, DANIEL: SONORAN ACANTHACEAE 735 Old World (introduced); Sonoran plants occur on slopes, along watercourses, and in disturbed areas in Sonoran desertscrub (Arizona Upland, Central Gulf Coast, Plains of Sonora), thornscrub, grass- lands, tropical deciduous forests, riparian forests, oak forests, oak-pine forests, and pine forests at elevations from sea level to 1680 m. Illustrations. Figure 15; Contributions from the United States National Herbarium 31:91. 1951: Fieldiana, Botany 24(10):365. 1974; Flora of Baja California, 192. 1980; Flora of the Venezuelan Guayana, vol. 2, 347. 1995; Flora of Chiapas 4:38. 1995; Proceedings of the California Academy of Sciences 49:340. 1997; Flora del Valle de Tehuacan-Cuicatlan 23:35. 1999. Local names. “Baiboria” (fide Gentry 1963); “cola alacran” (Felger & Yetman 97-132); “cor- doncillo” (Mexican, e.g., Burquez et al. 94-160, Felger & Yetman 96-193, Palmer 285, Pennington 258, White 2772); “cordonsillo” (Hartman 91); “coronia” (Gentry 1048); “mahili guasira” (Guarijio, Felger & Yetman 96-193, 97-132); “nachachikoh” (Guarijio, Gentry 1133). Uses. Used medicinally to treat scorpion sting (Fe/ger & Yetman 96-193); tea made from root drunk three or more times per day for fevers or influenza (Felger & Yetman 97-132); herbage decocted or infused for fevers (Gentry 1048, 1133, and fide Gentry 1963 in reference to its use by the Guarijio); a medicinal tea (mix with mescal) made from plant taken for colds and fever (Pennington 226): remedy for fevers, venereal diseases, etc. (Palmer 285); a refreshing tea made from leaves (Pennington 258); a tea from the whole plant used to treat diarrhea and the leaves and inflorescence are boiled (often together with leaves of Carlowrightia arizonica) and taken as a tea for fever (including malarial fevers), according to Yetman and Van Devender (2002) in reference to its medicinal uses by the Mayo. Daniel (1995a, 1997, 1999a) and Daniel and Acosta (2003) provided recent descriptions of this species in various regions of Mexico. Elytraria imbricata occurs in diverse communities (noted above) and is widespread in Sonora (Fig. 16). It occurs in three of the four subdivisions of the Sonoran Desert there, but appears to be absent from the Lower Colorado River Valley subdivision in northwestern Sonora. Plants are often leafless or nearly so during the dry season (e.g., January—May) when most flowering occurs; plants collected during the rainy season (e.g., August-September) are very leafy. SONORAN COLLECTIONS: Rio de los Alisos, ca. 6 mi N of Cumeral, L. Abrams 13159 (DS); 1 mi N of Cumeral on Nogales road, L. Abrams 13175 (DS, F); Nacopule Canyon, ca. 4 mi. N of San Carlos Bay, M. Ames 77-67 (ARIZ); Rio Cuchujaqui, near Alamos, M@. Ames 77-85 (ARIZ); Mpio. Opodepe, road to La Colmena Microwave Station at KM 76 N of Hermosillo, B. Bartholomew et al. 2468 (CAS, GH, MEXU, MO, NY); Yaqui River, El Toledo, 28°34’N, 109°33.5'W, G Boyer et al. s.n. (ARIZ); Guaymas, 7: Brandegee s.n. (F); 2.5 km S de Hermosillo, 29°01'N, 110°57'W, A. Burquez & A. Quijada 91-21 (MEXU); El Himalaya (Pena Blanca), entre San Carlos y Tastiota, 28°11’N, 111°20'W, A. Burquez et al. 91-300 (MEXU); KM 295 carr. fed. 16, 2.5 km W de los Vallecitos, 28°23’N, 108°50.5'W, A. Burquez et al. 92-415 (MEXU); Cafion Tepoca, KM 177, Carr. Fed. 16, 28°27'N, 109°15'W, A. Buirquez et al. 94-160 (MEXU); Cerro El Verde, San Javier, 28°33'N, 109°43'W, A. Burquez et al. 96-042 (MEXU); Mpio. Alamos, 53.9 km N of Alamos, 3.5 km NE of San Bernardo toward San Rafael, C. Cowan et al. 5554 (TEX); between Cananea and Arizpe, 27.7 mi N crossing of Rio Bacanuchi, ca. 30°50’/N, 110°20'W, 7. Daniel 922 (CAS); between Tecoripa and Tonichi, 9.3 mi E of Tecoripa, 7; Daniel 982 (CAS); “Microondas Mountain” overlooking Guaymas, along road which is 1.8 mi from Mex. Hwy. 15 on paved road to airport, N of Guaymas, 7. Daniel 1953 (ASU); Nacapuli Canyon, 6.3 mi W of Hwy. 15 on road to San Carlos then 4.4 mi N, 7. Daniel 1986 (ASU); road between Hwy. 16 E of Tonichi and Onavas, 0.5 mi S jct. Hwy. 16, 77 Daniel 3351 (CAS); Rio Cuchujaqui, 7.4 mi SE of Alamos toward Giiirocoba, 7. Daniel 3383 (CAS), 4001 (CAS); southeastern border of Alamos (road to mirador), ca. lat. 27°01'N, long. 108°56'W, T. Daniel 9770 (CAS); Mpio. Soyopa, crossing of Rio Yaqui with Hwy. 16, 8.5 km W of jet. to Onavas, ca. 28°34'N, 109°33’W, T. Daniel et al. 8512 (CAS, MEXU); Mpio. Yécora, between San Nicolas and Nuri, just S of Curea, ca. 24 km SW jet. Hwy. 16, ca. 28°19’N, 109°17’W, T. Daniel et al. 8578 (CAS): W of Rio Magdalena, Magdalena, F) Drouet & D. Richards 3813 (DS, F); moun- 736 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Las FIGURE 15. Sonoran Acanthaceae. a. Holographis virgata. b. Holographis pallida. c. Pseuderanthemum praecox. e. Stenandrium pilosulum. e. Elytraria imbricata. DANIEL: SONORAN ACANTHACEAE W237) ‘| tain E of Guaymas, F: Drouet & D. Richards 3843 : ! (CAS, DS, F, MO, NY, US); 4 km NW of Alamos, F: Lo Drouet & D. Richards 4011 (F); El Cerro de la oe HOD isi | Campana, Hermosillo, - Drouet et al. 3388 (DS, F, GH, US); Rancho La Brisca, 3.5 mi. N of Agua Fria, , 30°24'N, 110°33’'W, R. Duncan s.n. (ARIZ); 6.3 mi. E of Rio Bavispe on Guasabas-El Coyote Road, iN Canon Cruz de Penasco, R. Felger 3645 (ARIZ); Lb e Nacapule Spring, NE of Bahia San Carlos, R. Felger i 4082 (ARIZ): 3 mi W of Nuri, R. Felger 4096 nN (ARIZ): 0.8 mi S of Sabino, ca. 10 mi NE of Alamos, 29° R. Felger 5060 (ARIZ); Agua Caliente, 3.6 mi. S of s A cog La Casita on Mex. 15, R. Felger 5629 (ARIZ, oe e $ ace) ENCB); E end of Bahia San Carlos, R. Felger 6552 + 3 ~ (ARIZ): Rancho San Jorge (W of Hermosillo), ca. . * ee 29°44'N, 111°51'W, R. Felger 6907 (ARIZ); 14.7 Son eiabricnts eo 088 mi. N of Imuris on Mex. 15, R. Felger 7248 (ARIZ); 27 | o° ° Ry Isla Tiburon, SE base of Sierra Kunkaak, R. Fe/ger iy 9315 (ARIZ, ENCB); Agua Caliente, R. Felger 114° ili 110° 13989 (ARIZ, UC); ca. 1 km N of Bahia San Carlos 3 : 3 toward Bahia Algadones, R. Felger & R. Devine 85- 3301 (ARIZ): Canon Nacapuli, ca. 4 km N of Bahia San Carlos, R. Felger & M. Dimmitt 55-862 (ARIZ, CAS); 10.2 mi. S of La Pintada (between Hermosillo and Guaymas) on Mex. 15, R. Felger & C. Lowe 8365 (ARIZ); Cerro de la Vigia, Guaymas, R. Felger & R. Thomas 11832 (ARIZ); Cafion Nacapule, NE of Bahia San Carlos, R. Felger & R. Thomas 11861 (ARIZ, ENCB), 12002 (ARIZ); Mpio. Alamos, Rio Mayo drainage, ca. 16 km N of Mesa Colorado, ca. 27°42'N, 108°50'W, R. Felger & D. Yetman 96-193 (ARIZ, CAS); Mpio. Alamos, ca. 3 km SE of Bavicora, 27°40'N, 108°47'W, R. Felger & D. Yetman 97-132 (CAS); Canon Nacapules, ca. 6 km NE of Bahia San Carlos, R. Felger et al. 84-120 (ARIZ); 0.5 mi SE of Alamos at jct. Giiirocoba and El Fuerte rds., J. Fish 44 (UC); S of Los Molinos, 7.8 mi. N of Los Tanques on road from Alamos to San Bernardo, 27°18.5'N, 108°53'W, M. Fishbein et al. 930 (ARIZ); Mpio. La Colorada, 2.5 km S de El Pilar, 28°39’N, 110°52'W, A. Flores M. & J. Leon G. 5396 (JEB): Distr. Alamos, Qiiirocoba, H. Gentry 760 (DS, US); Canyon Sapopa, Rio Mayo, H. Gentry 1048 (F, GH, MO, S, WIS); Conejos, Rio Mayo, H. Gentry 1133 (F); Guaymas, H. Gentry 4697 (ARIZ); Sierra Bojihuacame SE of Cd. Obregon, H. Gentry 14494 (ARIZ, MEXU, LL, US); 4.3 km SW of Santa Ana de Yécora on road to Nuri, 28°23'N, 109°19'W, D. Goldberg & S. McLaughlin 77-165 (ARIZ); ca. 30 mi S of Hermosillo on Hwy. 15, B. Hansen et al. 1380 (LL, MEXU, US, WIS); Cochuto, C. Hartman 91 (GH, NY, PH, US); Calabasas, C. Hartman 109 (F, GH); Rio El Naranjo crossing, N of Taymuco, 27°15'N, 108°43’'W, P. Jenkins 88-175 (ARIZ); Guaymas, I. Johnston 3092 (CAS); San Carlos Bay, /. Johnston 4389 (CAS); Hermosillo, M. Jones 22576 (MEXU, MO, PH); Las Cabras, ESE of Alamos, 27°00'N, 108°55’W, E. Joyal 1422B (CAS); Rancho El Aguilar Noria, N of Ures and Santiago, 29°33’N, 110°25—26'W, E. Joyal 1819 (MEXU); Tiburon Island, Sierra Kunkook, Knight 1016 (UNM); Tiburon Island, San Miguel Peak, Knight et al. 961 (UNM); ca. 70 mi S of Hermosillo, G Lindsay 1137 (DS); Salsipuede, C. Lloyd 442 (GH); Palm Canyon, 18 mi. SE of Magdalena, P. Martin s.n. (ARIZ); 5 km N of Rosario de Tesopaco, P. Martin s.n. (ARIZ); Mocuzari, La Cruz, 27°13'N, 109°05.5'W, P. Martin s.n. (ARIZ); La Fundicion camp., road to San Pedro, 27°06’N, 108°45.5’'W, P. Martin & G. Ferguson s.n. (ARIZ); Cerro Verde, 28°34'N, 109°44'W, P. Martin & G. Ferguson s.n. (ARIZ); 3 km S of Santini, KM 175 on Obregon-Navojoa road, 27°16’N, 109°39'W, P. Martin & M. O'Rourke s.n. (ARIZ); 1 km NE of Bachoco, P. Martin & M. O'Rourke s.n. (ARIZ); 1 km E of El Chiribo, 27°18’N, 108°42'W, P. Martin & D. Yetman s.n. (ARIZ); Yaqui River, El Toledo, 28°34’N, 109°34'W, P. Martin et al. s.n. (MO, US); Estacién Margarita, 10 km NW of Navojoa, 27°08.5'N, 109°29'W, P. Martin et al. s.n. (ARIZ); Arroyo Santa Rosa, 10 km NE of Tesopaco, 27°56'N, 109°19'W, P. Martin et al. s.n. (ARIZ); Canyon de las Palmas, Sierra Baviso, 16 mi SE of Magdalena, N. McCarten 2264 (ENCB); Cafién Los Anegados, Sierra El Aguaje, Ejido Francisco Villa, 28°02’N, 111°08'W, FIGURE 16. Sonoran distribution of Elytraria imbrica- 1a. 738 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Moreno L. et al. 99-25 (ARIZ); Alamos District, near Minas Nuevas, R. Moran 4030 (DS, UC); Guaymas, E. Palmer 285 (C, GH, NY, US); without locale, E. Palmer 12691 (MO), s.n. [in 1869] (PH, US); Maicoba, C. Pennington 226 (TEX); Onavas, C. Pennington 258 (TEX); 2.5 mi. W and 0.75 mi S of Alamos, R. Perrill & V. Phelps 5167 (ARIZ); Nacopuli Canyon, 3.5 mi. N of San Carlos Bay, A. Phillips et al. 75-158 (ARIZ): 8 mi E of Moctezuma on Huasabas Road, 29°50’N, 109°23'W, F: Reichenbacher 188 (ARIZ); Mpio. Guaymas, old road to Algodones, San Carlos, 27°58’N, 111°04’W, A. Reina G. & T: Van Devender 2000-916 (CAS): Guaymas, J. Rose 1275 (US): vicinity of Guaymas, J. Rose et al. 12588 (NY, US); vicinity of Alamos, J. Rose et al. 12726 (NY, US); vicinity of Puerta de Mano, 26°59’'N, 108°39'W, J. Salmon s.n. (ARIZ); 10 mi N of San Carlos, “Catch-22” site, D. Shaw 29 (MO); Mpio. Villa Pesqueira, Sierra Pinta, ca. 33 mi. S of Moctezuma, ca. 29°35'N, 110°01'W, J. Shortman & M. Wilson 96-58 (ARIZ); 37 mi. NE of Cajeme on road to Tesopaco, F: Shreve 6155 (ARIZ); Sierra Baviso, 19 mi E of Magdalena, F Shreve 6642 (F, GH, MO): 7 mi S of Estaci6n Carbo, F: Shreve 6688 (GH); 25 mi. E of La Motica on road to Tezopaco, W. Spaulding 75- 3-17 (ARIZ); ca. 10 mi W of Tezopaco toward Cd. Obregon, W. Spaulding 75-3-36 (ARIZ); 2.6 mi. ESE of Alamos toward Rio Cuchujaqui crossing at Sabinito Sur, Cerro Las Campas, 27°01'N, 108°54’W, G Starr & D. Palzkill 335 (ARIZ); Mpio. Alamos, Sierra de Alamos, ca. 6 km W of Alamos, ca. 27°02.5'N, 109°O1'W, V. Steinmann & C. Smith 93-123 (ARIZ); along road between Yepachic and Yécora, 19 mi W of Chihuahua border, S. Sundberg & M. Lavin 2837 (MO); Cerro Prieto Microondas, 15 km E de Navojoa, P. Tenorio et al. 10185 (MEXU); Santa Cruz, G Thurber 950 (F); 4 mi. S of Mazocahui, R. Turner 59-69 (ARIZ); 7.4 mi N of Las Chinacas, 27°18'N, 108°37'W, R. Turner & P. Martin 79-396 (ARIZ); Bahia San Pedro, ca. 28.1°N, 111.2°W, R. Turner et al. 79-258 (ARIZ); Giiirocoba crossing of Rio Cuchujaqui, 12.3 km SSE of Alamos, 26°56'N, 108°53'W, T. Van Devender et al. 92-903A (ARIZ); Camahuiroa between Agiabampo and Las Cocas on Gulf of California, 26°31'N, 109°16’W, 7: Van Devender et al. 92-1084 (ARIZ, CAS); Mpio. Alamos, 1 km S of Yocogigua, 26°47'N, 109°02'W, 7. Van Devender et al. 93-1091 (ARIZ); Saucito Canyon, Rancho la Sierrita, E slopes of Sierra de Alamos, ca. 6 km SSW of Alamos, 26°58’'N, 108°58’W, 7. Van Devender et al. 93-1333 (CAS); Mpio. Yécora, 5.3 km W of Yécora on Mex. 16, 28°22'N, 108°59'W, T. Van Devender et al. 96-64 (ARIZ, MEXU); Mpio. Soyopa, near Arroyo Garambullo on NE side of Rio Yaqui bridge on Mex. 16, just S of Tonichi, ca. 28°34'N, 109°33'W, T. Van Devender et al. 98-1119 (NY, TEX); Mpio. Nogales, 1.3 km SE of Rancho Las Borregas headquarters on road to Nogales, 31°12'N, 111°07'W, T. Van Devender et al. 2004-248 (CAS); Palm Canyon, 17.7 mi SE of Magdalena in Cerro Cinta de Plata, 77 Van Devender et al. s.n. (ARIZ); San Javier, cafion lo de Campa entre los cerros El Potrerito y Los Amoles a 7 km S del poblado, 28°32'N, 109°45’'W, L. Varela E. 96-443 (MEXU); San Javier, Cerro San Juan, 28°36'N, 109°45'W, L. Varela & E. Cuamea 97-66 (MEXU); Alamos, F- Vilas 69-VII (WIS); N of Guaymas, U. Waterfall 12848 (F, SMU); 2 mi S of El Coyote, ca. 29°56'N, 109°08’W, G. Webster & R. Murphey 21349 (ARIZ); Mpio. Yécora, ca. 2 km NW of Yécora toward Santa Rosa, 28°23'N, 108°51'W, J. Weins et al. 96-107 (ARIZ); loop of the Rio Bavispe, Cafion de Huépari, N of Aribabi, S. White 2649 (ARIZ, GH); loop of the Rio Bavispe, Puerto de Huépari, NW of Aribabi, S. White 2772 (DS, MEXU); Bavispe, S. White 2863 (GH); Horconcitos, Arroyo del Salto, S. White 3774 (GH); 100 mi. from Magdalena, J. Whitehead M127 (ARIZ); 4 mi N of Nuri, 28°10'N, 109°11'W, A. Whittemore et al. 83-084 (MEXU, TEX); 2 mi E of Rancho San Carlos on road to Noria, /. Wiggins 6146 (DS, US); Rio Sasabe, 10 mi E of Magdalena, /. Wiggins 7106 (DS, US); 10 mi N of Carbo, /. Wiggins 7265 (DS, GH, UC, US); 1 mi E of RR on road from Carbo to Horcasitas, 1. Wiggins 7277 (DS, F, GH, LL, MO, UC, US); Sahuaribo (108°40'W, 27°20'N) to Corogui (108°41.5’W, 27°22'N), D. Yetman et al. s.n. (ARIZ). Henyra Nees ex Benth. Henrya Nees ex Benth., Bot. Voy. Sulphur, t. 49. 1845. TYPE: Henrya insularis Nees ex Benth. Solenoruellia Baill., Hist. Pl. 10:445. 1891. TYPE: Solenoruellia galeottiana Baill. (= Henrya insularis Nees ex Benth.). Baillonacanthus Kuntze in Post and Kuntze, Lex. Gen. Phan. 58. 1903 (“1904”). TYPE: Solenoruellia galeottiana Baill. (= Henrya insularis Nees ex Benth.). Erect or ascending perennial herbs or shrubs with cystoliths. Leaves opposite. Inflorescences i DANIEL: SONORAN ACANTHACEAE 739 of axillary and terminal stout to lax dichasiate spikes (to thyrses) collectively forming leafy term1- nal panicles; dichasia alternate or opposite, 1—3-flowered, sessile or short-pedunculate in axils of bracts. Flowers homostylous, sessile, subtended by 2 homomorphic bracteoles, bracteoles oblance- olate to obovate, usually larger and more conspicuous than bracts, concavoconvex, fused from base to near apex along side adjacent to rachis, rounded to acute at apex, mucronate with straight, api- cal or dorsal, erect or divergent pointed projection, secondary bracteoles, if present, much reduced. Calyx deeply 5-lobed, lobes homomorphic or heteromorphic (i.e., posterior lobe reduced in size). Corolla white, cream, or yellow, with maroon, purple, yellow, and/or white markings on upper lip, tube subcylindric to cylindric, throat indistinct, limb pseudopapilionaceous, upper lip comprising 2 fused lobes, 2-fid at apex, lower lip 3-lobed, lateral lobes similar to one another, lower-central lobe usually larger and keeled, corolla lobes imbricate in bud. Stamens 2, anthers 2-thecous, the- cae subequal, parallel, equally to subequally inserted on filament, lacking basal appendages, dehiscing toward upper lip (1.e., flower stenotribal); pollen (Fig. 12) euprolate to perprolate, 3-col- porate. colpi broad (1.e., far exceeding width of centrally positioned ora), 6-pseudocolpate, pseudo- colpi 2 per mesocolpium, exine reticulate to bireticulate; staminodes 0. Style exserted from mouth of corolla, stigma 2-lobed, lobes equal and often inconspicuous. Capsule stipitate, head subspher- ic to broadly ellipsoidal, retinacula present, septae with attached retinacula separating slightly from inner wall of mature capsule. Seeds 2, homomorphic, planoconvex, subcircular to subelliptic in outline, flat surface smooth to bumpy, convex surface and margin either pubescent with hygroscop- ic trichomes or covered with stout, branched or barbed tubercles. (x = 18). Daniel (1990) treated Henrya as consisting of two species occurring in dry regions from the southwestern United States southward to Costa Rica. Both species occur in Mexico. Henrya insularis Nees ex Benth., Bot. Voy. Sulphur, t. 49. 1845. TYPE: plate 49 of Bentham’s The Botany of the Voyage of H.M.S. Sulphur, see Daniel (Taxon 38:265-270. 1989.) Henrya costata A. Gray, Proc. Amer. Acad. Arts 21:406. 1886. Tetramerium costatum (A. Gray) Millsp., Publ. Field Columbian Mus., Bot. Ser. 1:47. 1895. TYPE: MEXICO. Chihuahua: near Batopilas, Aug-Nov 1885, E. Palmer 211 (holotype: GH!; isotypes: K!, LE!, MEXU!, NY!, PH!, US!). Henrya grandifolia Fernald, Bot. Gaz. (Crawfordsville) 20:537. 1895. TYPE: MEXICO. Sinaloa: Esquinapa, Jan 1895, F. Lamb 505 (holotype: GH!). Henrya costata A. Gray var. glandulosa Brandegee, Zoe 5:171. 1903. TYPE: MEXICO. Baja California Sur: Cape Region, Santa Anita, 1901, C. Purpus 266 (lectotype, designated by Daniel, 1990: UC!; isolectotypes: ARIZ!, MO!, US!). Henrya brevifolia Happ, Ann. Missouri Bot. Gard. 24:547. 1937. TYPE: MEXICO. Sonora: Las Durasnillas, 18 May 1892, 7: Brandegee s.n. (holotype: UC!; isotypes: DS!, GH!, NY!, PH!, US!). Henrya ortegana Happ, Ann. Missouri Bot. Gard. 24:552. 1937. TYPE: MEXICO. Sinaloa: Sind. San Juan, San Ignacio, Mar 1931, J. Ortega 6868 (holotype: MO!; isotypes: CAS!, F!, MIN!). See Daniel (1995a) for a complete listing of synonyms for this species. Phenology. Flowering: January—June; fruiting: March—June. Distribution and habitats. Southwestern United States (Arizona), Mexico (einccaentcs Baja California Sur, Chiapas, Chihuahua, Colima, Durango, Guanajuato, Guerrero, Jalisco, México, Michoacan, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, San Luis Potosi, Sinaloa, Sonora, Tamaulipas, Veracruz, Yucatan, Zacatecas), Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica; Sonoran Plants occur along watercourses and on rocky slopes in thornscrub, tropical 740 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 deciduous forests, oak woodlands, and oak-pine woodlands at elevations from 200 to 1380 m. Illustrations. Figure 9; Botany of the Voyage of H.M.S. Sulphur, t. 49. 1845; Flora of Baja California, 193. 1980; Flora of Chiapas 4:40. 1995; Proceedings of the California Academy of Sciences 49:345. 1997; Flora del Valle de Tehuacan-Cuicatlan 23:41. 1999; Flora del Bajio 117:56. 2003. Local names. “Alfalfillo” (fide Van Devender et al. 2000); “hegregado” (Joyal 1563); “rama del toro” (Burquez & Steinmann 96-110, Joyal et al. 1627, Van Devender et al. 93-1523, 94-220). Brandegee (1893:347) indicated that “Las Durasnillas,” the type locality of Henrya brevifolia, is “a small collection of houses about sixty miles from Hermosillo, near a mountain range known as Sierra Matapan.” I did not encounter these place names on the series of maps available to me. A “Rancho Duraznillas” is located at 28°15'N latitude and 109°46’W longitude (Anonymous 1992), near the railroad between Est. Corral (near Esperanza) and Tonichi. Henrya insularis has been collected in this region, and there is a “La Dura” on most maps near these coordinates. This locality is nearly 100 miles from Hermosillo, however. The American Geographical Society’s (1937) map of this region shows a “Duraznillo” about 60 miles east of Hermosillo along the road to Mazatan (at about 29°02'N, 110°17'W). There is a Rio Matape and a Sierra Mazatan in the gen- eral vicinity. This latter locality would have been more accessible to Brandegee from Hermosillo and it is presumed to be the type locality for both H. brevifolia and Tetramerium abditum. This locale is well within the range of H. insularis and T. abditum has been collected nearby. Daniel (1990, 1995a, 1997, 1999a) and Daniel and Acosta (2003) provided descriptions of Henrya insularis and discussed regional morphological variation throughout its range. Plants from Sonora represent a common and widespread ; 1 T T (Arizona to Central America) form of the liu species with bracts shorter than the bracteoles | He PSSe and the bracteolar mucros apically situated. | Ss —~ Vegetative pubescence of Sonoran plants con- +31 aes \ sists of an understory of eglandular (and usual- \ ee 5 ly glandular as well) trichomes and an oversto- | ) e ry of glandular trichomes. Most Sonoran col- | L 7 lections are noted to have yellowish corollas, | e usually with maroon or reddish markings on | A the upper lip. ah we . | IS A *, The species is widely distributed in Sonora | SS (Fig. 17) and occurs in diverse habitats. It like- > Noy ov ly also enters the Arizona Upland subdivision | ; *. 5 ; @ Henrya insularis 4 of the Sonoran Desert along the Rio Magdalena near Magdalena. Its northernmost known | \@ e 1 occurrence is in the lower reaches of Sycamore | ab see he: V7 Canyon in the Parajito Mountains along the & Arizona-Sonora border, west of Nogales. FIGURE 17. Sonoran distribution of Henrya insularis. SONORAN COLLECTIONS: El Rancheria crossing of Rio Cuchujaqui, ca. 22.5 km S of Alamos on road to El Chinal, 26°51'N, 108°55'W, M. Baker et al. 10373 (TEX); Toro Muerto Canyon, ca. 2.5 mi W of Toro Muerto bridge on Hwy. 118, J. Bowers et al. 2813 (ARIZ, ENCB); San Javier, 1 km de la Carr. Fed. 16, 28°35'N, 109°45'W, A. Buirquez 93-12] (MEXU); San Javier, microondas en el Cerro El Durazno, 28°37'N, 109°45'W, A. Burquez 94-018 (MEXU); 0.5 km NW del Rancho Lo de Campa, San Javier, 28°34’N, 109°44.5'W, A. Burquez & V. Steinmann 96-110 (MEXU); Mpio. Huatabampo, Bocarehui [26°49’N, 109°43'W], M. Cordova & R. Alcaraz s.n. (ENCB); along road between Hwy. 16 E of Tonichi and Onavas, 0.5 mi S jet. Hwy. 16, 7 Daniel 3352 (CAS): along road between Rosario de Tezopaco and Nuri, 7.1 mi S of DANIEL: SONORAN ACANTHACEAE 74] turnoff to Nuri, 7: Daniel 3363 (ASU, CAS); Arroyo Cuchujaqui, 7.4 mi SE of Alamos toward Guiricoba, 7. Daniel 3377 (CAS); Mpio. Yécora, between San Nicolas and Nuri, ca. 6.4 km SW jet. Hwy. 16, ca. 28°24'N, 109°14'W, T. Daniel et al. 8571 (CAS, MEXU); near Rancho La Brisca, ca. 8 mi ENE of Cucurpe, J. Duek s.n. (MEXU): Mpio. Guaymas, Sierra Libre, Cafion Tetabejo, 28°32'N, 110°59'W, R. Felger et al. 02-297 (CAS); 9 mi from Imuris in Magdalena River canyon, R. Ferris 8783 (DS, US); 6.3 mi NW of San Bernardo toward Guajaray, 27°26'N, 108°57'W, M. Fishbein et al. 1546 (ARIZ, CAS); Jopopaco vicinity, 2.2 km NE of Mex. 15 to Jopopaco, 25 km NW of Melchor Ocampo, 26°44’N, 109°16'W, S. Friedman & O. Kiser 242- 95 (ARIZ); Arroyo Camahuiroa, 1.5 km NE of Camahuiroa, 9.5 km WNW of Melchor Ocampo, 26°33’N, 109°16'W, S. Friedman et al. 172-94 (ARIZ); Mpio. Imuris, 1.5 km W and 8 km S of Microondas Las Aguilas, Sierra de La Madera (E of Imuris), 30°40’N, 110°46.5'W, G Ferguson et al. 2338 (CAS); 21.5 mi E of Tecoripa, P. Gallagher et al. 290 (ASU); 16.2 mi E of turnoff to Tonichi and 11.2 mi W of Tepoca along Mex. Hwy. 16, P. Gallagher et al. 294 (ASU, CAS, NY); Alamos, Rio Fuerte, H. Gentry 2200 (ARIZ, F, GH, K, MEXU, MO, S. UC, US): Rancho Agrimincor, Rio Mayo, H. Gentry 3043 (ARIZ, F, GH, K, MEXU, MO, S, UC. US): Curohui, Rio Mayo, H. Gentry 3639 (ARIZ, F); Distr. Alamos, H. Gentry s.n. (MICH); 3.7 mi W of Santa Ana on old road to Yécora, D. Goldberg 76-99 (ARIZ); Rio Cuchujaqui, ca. 5 mi ESE of Alamos, 26°56'N, 108°53’W, D. Goldberg & T. Van Devender 76-45 (ARIZ); Mpio. Alamos, caj6n of the Rio Cuchujaqui, 27°02—03'N, 108°42-42.5'W, P. Jenkins 89-432 (ARIZ), 89-440 (ARIZ); Mpio. Alamos, Rio Cuchujaqui, Guirocoba crossing 5 mi SE of Alamos, P. Jenkins et al. 97-01 (ARIZ, CAS); Puente La Pila, ca. 20 km E of Onavas, 28°29'N, 109°22'W, E. Joyal 1563 (CAS, MEXU); Rio Cuchahaqui, SSE of Alamos, 26°52'N, 108°55’W, E. Joyal et al. 1627 (MEXU); 7.5 mi W of Alamos, R. Krizman 2 (ARIZ); ca. 4 mi NE of Santa Rosa on road to Yécora, E. Lehto & T. Reeves L18746 (ASU); Rancho La Brisca, ca. 4-5 km N of Rancho Agua Fria on Rio Santo Domingo, 30°24'N, 110°33'W, P. Martin s.n. (ARIZ, ENCB); 5 km SW of Santa Barbara, El Palmarito Canyon, 27°05'N, 108°45'W, P. Martin & G Ferguson s.n. (ARIZ, CAS); Ignimbrite Canyon, 3 km N of La Brisca, Rio Santo Domingo, 30°25'N, 110°53’W, P. Martin & M. O'Rourke s.n. (ARIZ): Microondas Masiaca, 7 km W of Masiaca, 26°46.5'N, 109°18'W, P. Martin & M. O’Rourke s.n. (ARIZ); above Las Animas, 27°31.5'N, 108°42'W, P. Martin et al. s.n. (ARIZ); Palm Canyon, 19 mi E of Magdalena, C. Mason et al. 3294 (ARIZ, MEXU); Mpio. Alamos, Arroyo Santa Barbara, 27°07’N, 108°43’'W, S. Mathes & P. Martin s.n. (ARIZ); 1.3 mi S of Cucurpe, L. McGill & D. Pinkava 6491 (ASU, F); 2 mi N of Nuri, S. McLaughlin 512 (ARIZ); 21 mi NE of Rio Yaqui crossing on Mex. 16, R. Perrill 5317 (ARIZ); Onavas, NW of pueblo, ca. 28°28’N, 109°32'W, A. Rea 1052 (ARIZ); Rancho E] Palmar, 22.6 km SE of Rio Yaqui on Mex. 16, 28°30'N, 109°24'W, A. Reina G. et al. 97-104 (MEXU); Mpio. Onavas, 4 km W of Agua Amarilla, 18.9 km W of Tepoca on Mex. 16, 28°29'N, 109°22'W, A. Reina G. et al. 95-603 (CAS); Cerro Redondo, 27°05'N, 108°45'W, R. Rondeau & G. Rodda s.n. (ARIZ); Sierra de Alamos, J. Rose et al. 12829 (NY, US); Rio Cuchujaqui, ca. 7 mi ESE of Alamos, ca. 26°59’'N, 108°50'W, A. Sanders et al. 2573 (ARIZ, ASU, RSA): NW side of Sierra de Alamos along road from Alamos—Navojoa road at Rancho Las Lomas to Promontorios, A. Sanders et al. 2613 (ASU); Sierra Madre Occidental, at three-way jct., 6 mi E of Tepoca, A. Sanders et al. 2668 (ASU); Rio Chico, 2 mi N of Nuri, W. Spaulding 75-3-22 (ARIZ); Las Piedras Canyon, E side of Sierra Alamos, 26°59’N, 108°57.5'W, G Starr & D. Palzkill 363 (ARIZ); 1.8 mi N of Alamos on Cerro Dos Negritos, G Starr & D. Palzkill 368 (ARIZ); 20-25 mi NE of Ures, R. Straw 2118 (RSA, UC); Cucurpe, near Rio Dolores, R. Thompson & O. Davis 82-46 (ARIZ); 21 mi E of Tonichi on Rio Yaqui, L. Toolin 297 (ARIZ); Rio Cuchujaqui at Rancho El Conejo, 12.4 km S of Alamos, 26°55'N, 108°55'W, T: Van Devender 92-561 (ARIZ); Mpio. Santa Ana, Canada El Rincon del Diablo, W slope of Sierra Cucurpe, ca. 13 km W of Cucurpe, 30°20’N, 110°51'W, 7: Van Devender & A. Reina G. 98-446 (CAS); Cafion de la Bota, N end of Sierra el Tigre, ca. 34 km ESE of Esqueda, 7. Van Devender et al. 82-61 (ARIZ); near crossing of Rio Cuchujagui ca. 8 mi SE of Alamos, 7: Van Devender et al. 84-146 (ARIZ); canyon N of Los Aguaros on Arroyo Guajaray, 27°38.5'N, 108°58’W, T. Van Devender et al. 93-485 (ARIZ); El Rincon Viejo, ca. 4 km N of Alamos, Sierra de Alamos, 27°04'N, 108°56’W, TZ. Van Devender et al. 93-1523 (ARIZ); Cerro Piedra Boluda, ca. | km NE of El Rincon Viejo, ca. 4.5 km N of Alamos, 7. Van Devender et al. 94-220 (ARIZ, CAS); Mpio. Yécora, Agua Amarilla (Los Pinitos), 15 km WNW of Tepoca, 24.7 km WNW of San Nicolas on Mex. 16 at KM 200, ca. 28°08’N, 109°20’W, T. Van Devender et al. 96-23 (ARIZ); 1.5 km SW of Santa Ana on road to Guadalupe Tayopa, 28°23’N, 109°09’'W, T. Van Devender et al. 97-201 (MEXU, MO, TEX); 2.7 mi WNW of Tepoca on Mex. 16, ca. 28°28’/N, 109°16'W, T. Van Devender et al. 99-71 (NY); Mpio. Nogales, Sierra Las TAD PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Avispas, 15.9 km W of MEX 15 on SON 43 (to Saric), 31°04'N, 111°05°E, 7: Van Devender et al. 2002-757 (CAS); Mpio. Nogales, 3.8 km NE of El Correo (La Arizona), Rancho Las Borregas, Arroyo Planchas de Plata, 31°11.5'N, 111°10'W, 7: Van Devender et al. 2004-217 (CAS); ca. 17 mi SSE of Magdalena, Cerro Cinta de Plata (=Sierra Babiso), 7: Van Devender et al. s.n. (ARIZ, NY); Rio Saracachi, ca. 5 mi NE of Cucurpe, 7. Van Devender et al. s.n. (ARIZ); San Javier, Cerro San Juan, 28°36'N, 109°45'W, L. Varela & E. Cuamea 97-68 (MEXU), L. Varela & A. Feliz 97-34 (MEXU); Los Banos near Angostura, S. Walker BAN 7 (NY); 13 mi E of Imuris, 7. Wiggins 11665 (DS, MEXU, MICH, TEX, UC, US). Holographis Nees Holographis Nees in A. de Candolle, Prodr. 11:728. 1847. TYPE: Holographis ehrenbergiana Nees. Berginia Harv. ex Benth. & Hook.f., Gen. Pl. 2:1096. 1876. TYPE: Berginia virgata Harv. ex Benth. & Hook.f. (= Holographis virgata (Harv. ex Benth. & Hook.f.) T.F. Daniel). Pringleophytum A. Gray, Proc. Amer. Acad. Arts 20:292. 1885. TYPE: Pringleophytum lance- olatum A. Gray. (= Holographis virgata (Harv. ex Benth. & Hook.f.) T.F. Daniel). Lundellia Leonard, Wrightia 2:1. 1959. TYPE: Lundellia argyrea Leonard (= Holographis argyrea (Leonard) T.F. Daniel). Erect to ascending perennial herbs or shrubs lacking cystoliths. Leaves opposite (rarely sub- opposite) or quaternate. Inflorescence of axillary or terminal dichasiate spikes (reduced to 2 flow- ers in H. ehrenbergiana); dichasia opposite to alternate, 1-flowered, sessile in axil of a bract. Flowers homostylous, sessile, subtended by 2 homomorphic bracteoles. Calyx deeply 5-lobed, lobes homomorphic. Corolla greenish white, white, pinkish to purplish, or yellow, often with col- ored nectar guides, long axis horizontally to vertically oriented, tube subcylindric to + abruptly expanded distally into a throat, limb 2-labiate, upper lip 2-lobed, lower lip 3-lobed, corolla lobes imbricate in bud. Stamens 4, equally inserted to + didynamous, anthers slightly exserted from mouth of corolla, anthers 1-thecous, often connivent, pubescent, lacking basal appendages, dehisc- ing toward lower lip (1.e., flower nototribal); pollen (Fig. 18) euprolate to perprolate, 3-colpate, colpi often bifurcate near poles, exine psilate to verrucate to foveolate to rugulate; staminode 1, borne between posterior pair of stamens, short. Style slightly exserted from mouth of corolla, stig- ma subfunnelform or 2-lobed, lobes equal or unequal. Capsule substipitate, ellipsoid to obovoid, retinacula present, septae with attached retinacula remaining attached to inner wall of mature cap- sule. Seeds 4 (or fewer by abortion), homomorphic, lenticular. (x = 13). This genus of 15 species is restricted to, but occurs nearly throughout, the dry regions of Mexico (Daniel 1983b, 1988b). 1. Leaves opposite (or subopposite); inflorescence of terminal spikes to 15 cm long. . H. virgata 1. Leaves quaternate; inflorescence of axillary spikes to 3.7 cm long............... H. pallida Holographis pallida Leonard & Gentry in H.S. Gentry, Brittonia 6:324. 1948. TYPE: MEXI- CO. Sinaloa: Cerro Llano Redondo, W of Caymanero, 25 Apr 1944, H. Gentry 7022 (holotype: MICH!; isotypes: ARIZ!, F!, GH!, NY!, US!). Phenology. February—April; fruiting: March—April. Distribution and habitats. Northwestern Mexico (Sinaloa, Sonora); Sonoran plants occur on slopes and along watercourses in thornscrub and tropical deciduous forests at elevations from 50 to 600 m. Illustrations. Figure 15; Brittonia 6:325. 1948. 743 DANIEL: SONORAN ACANTHACEAE 18. Pollen of Sonoran Acanthaceae. a. Holographis pallida (Daniel 3382), interapertural view. b. H. pallida FIGURE Daniel 3382), apertural view. c. H. pallida (Daniel 3382), polar view. d. H. virgata (Breedlove 60981), interaperatural view. =. Odontonema cuspidatum (Jenkins & Yetman 91-115), polar view. f. O. cuspidatum (Breedlove & McClintock 23784), Pseuderanthemum praecox (McVaugh 26541), apertural view. h. P. praecox (Daniel 2131), polar view. 1. 10 um. polar view. ¢. P enandrium pilosulum (Moran 21965), interapertural view. Scales 4 744 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Local name. “Ramito chino” (Gentry 1350). Daniel (1983b) provided a description of this species, which is endemic to northwestern Mexico. Its corollas are white to greenish white with light pink stripes on the lobes of the lower lip. Holographis pallida reaches the northern extent of its distribution in east-central Sonora (Fig. 19). SONORAN COLLECTIONS: Arroyo Cuchuja- qui, 7.4 mi SE of Alamos on road to Guiricoba, T. Daniel 3382 (ASU, CAS), 3996 (CAS); Mpio. Soyopa, crossing of Rio Yaqui with Hwy. 16, 8.5 km W of jct. to Onavas, ca. 28°34’N, 109°33'W, T. Daniel et al. 8510 (ASU, BR, CAS, K, MEXU, MICH, MO, US); San Bernardo, H. Gentry 1350(A, [27 © Holographis virgata ARIZ, F, MEXU, MO, UC, WIS), 36/5 (ARIZ, F); Sierra Bojihuacame SE of Cd. Obregon, H. Gentry em 14" | 112" 14492 (ARIZ, US); NE side of Rio Yaqui bridge on FIGURE 19. Sonoran distributions of Holographis pall- Mex. 16, just S of Tonichi, ca. 28°34'N, 109°33"W, jaa and H. virgata. A. Reina G et al. 97-947 (CAS); Rio Cuchujaqui, 8 mi SE of Alamos, ca. 26°56’N, 108°53'W, A. Sanders 3685 (CAS); Mpio. Alamos, Cerro La Luna, ca. 14 km NW of Alamos, ca. 27°07'N, 109°02'W, A. Sanders et al. 13257 (ARIZ, CAS, TEX, US); Rio Cuchujaqui at Rancho E] Conejo, 12.4 km S of Alamos, 26°55’N, 108°55’W, T: Van Devender 92-562 (ARIZ); Cerro Prieto, ca. 9 mi E of Navojoa on Alamos Road, 27°05’N, 109°17'W, 7: Van Devender et al. 92-166 (ARIZ, CAS); below Guajaray on Arroyo Guajaray, 6.5 km WNW of ject. with Rio Mayo, 27°36’N, 108°56'W, 7: Van Devender et al. 93-465 (ARIZ, CAS, TEX): between Rancho La Junta on Rio Mayo and Guajaray on Arroyo Guajaray, 27°35-36'N, 108°52—S6'W, 7: Van Devender et al. 93-504 (ARIZ, CAS); Mpio. Soyopa, Mina Soyopita, | km S of Soyopa, W side of Rio Yaqui, 28°46'N, 109°38’W, T: Van Devender et al. 99-911 (CAS). © Holographis pallida Holographis virgata (Harv. ex Benth. & Hook.f.) T.F. Daniel, J. Arnold Arbor. 64:139. 1983. Berginia virgata Harv. ex Benth. & Hook.f., Gen. Pl. 2:1097. 1876. TYPE: MEXICO. State unknown: “California incola,” 7 Coulter 603 (holotype: K!; isotypes: BM!, GH!, K!). Pringleophytum lanceolatum A. Gray, Proc. Amer. Acad. Arts 20:293. 1885. TYPE: MEXICO. Sonora: 50 mi below Altar and 30 mi from Gulf of California, 13 Mar 1884, C. Pringle s.n. (holo- type: GH!; isotypes: A!, F!, G!, MA!, NY!, P!, US!, WIS!). Phenology. Flowering: December—May; fruiting: March—May. Distribution and habitats. Northwestern Mexico (Baja California, Baja California Sur, Sonora); Sonoran plants occur on rocky slopes and along watercourses in Sonoran desertscrub (Central Gulf Coast, Lower Colorado River Valley, Plains of Sonora) and thornscrub at elevations from 15 to 442 m. Illustrations. Figure 15; Proceedings of the California Academy of Sciences 49:348. 1997. Local name. “?aayaam” (Seri, Felger et al. 17831). Daniel (1983b, 1997) provided descriptions of this species and its infraspecific taxa. Corollas of Holographis virgata have a white tube, light pink to whitish upper lip, and a dark pink lower lip with white veins (and a white area near the mouth) on the lobes of the lower lip. Holographis vir- gata subsp. virgata is the only infraspecific taxon present in Sonora. Both the species and the nom- inate subspecies attain their northern and eastern distributional limits in Sonora (Fig. 19). DANIEL: SONORAN ACANTHACEAE 745 SONORAN COLLECTIONS: 12 mi NW of Hermosillo, L. Abrams 13305 (DS, F); canyon back of Palma, 40 mi S of Hermosillo, L. Abrams 13348 (DS); Isla Tiburon, J. Alcocer & C. Sosa R. 6 (ENCB); ca. 31 mi E of Punta Cirio between Puerto Libertad and Caborca, J. Bowers & S. McLaughlin 1613 (ARIZ); S side of Algodones Bay, 2 km W of Cerro Tetas de Cabra, 27.9°N, 111.0°W, 7. Burgess et al. 6363 (ARIZ, ENCB, MEXU); ca. 3.5 km E of El Caracol, 30°02'N, 112°26'W, T. Burgess et al. 6439 (ARIZ); El Himalaya (Pefia Blanca), entre San Carlos y Tastiota, 28°11’N, 111°20'W, A. Burquez et al. 91-312 (MEXU); 10 mi NW of Hermosillo, C. Carter s.n. (MICH); “Microondas mountain” overlooking Guaymas, along road which is 1.8 mi from Mex. Hwy. 15 on paved road to airport, N of Guaymas, 7? Daniel 1958 (ASU); Nacapuli Canyon, 6.3 mi W of Hwy. 15 on road to San Carlos then 4.4 mi N, T Daniel 1969 (ASU), 1972 (ASU, CAS); near Guaymas, E. Dawson 1075 (F, MICH); San Miguel de Horcasitas, G. Eisen s.n. (PH, US); N end of Bahia San Carlos, R. Felger 267] (ARIZ); 2.7 mi E of Tajitos on Mex. 2 (between Sonoyta and Caborca), R. Felger 7419 (ARIZ); 10.2 mi S of La Pintada on Hwy. 15 (between Hermosillo and Guaymas), R. Felger 8381 (CAS); Isla Turmers (= Datil), island off S shore of Isla Tiburon, 28°43’N, 112°17'W, R. Felger 9082 (ARIZ, CAS); NE end of Bahia San Carlos, R. Felger 9617 (CAS); Isla Turners (=Datil), 28°43'N, 112°17.5'W, R. Felger & J. Cooper 15320 (ARIZ); Sierra del Viejo (SW of Caborca), R. Felger & H. Gentry 7912 (ARIZ); Bahia Colorado, ca. 28°18’N, 111°28’W, R. Felger & E. Hamilton 15683 (CAS); ca. 0.5 mi SE of Cafidn Nacapules, ca. 6 km NE of Bahia San Carlos, ca. 28°10'N, 111°03.5'W, R. Felger & R. Thomas 11957 (ARIZ); ca. 0.8 mi N of Bahia San Carlos toward Bahia Algadones, R. Felger & R. Thomas 12027 (CAS); Cerro San Nicolas, W of S end of Kino estuary, R. Felger et al. 9555 (ARIZ); Ensenada Grande (= Bahia San Pedro), R. Felger et al. 11616 (CAS); Isla Tiburon, ca. 13 mi S of Tecomate, ca. 28°57'N, 112°27'W, R. Felger et al. 12384 (ARIZ), 12397 (ARIZ); Isla Tiburon, vicinity of Palo Fierro, Punta Tortuga, R. Felger et al. 12555 (ARIZ, ENCB); Tiburon Island, Ensenada de la Perra, ca. 28°47’N, 112°16'W, R. Felger et al. 17730 (ARIZ); vicin- ity of Rancho Arivaipa, ca. 8 mi ENE of Pozo Coyote, ca. 29°37'N, 112°17'W, R. Felger et al. 17831 (ARIZ, CAS, MEXU); Canon Nacapules, ca. 6 km N of Bahia San Carlos, ca. 28°10'N, 111°03’W, R. Felger et al. 85-565 (ARIZ, CAS, MEXU); Picu Mts., Altar Distr., H. Gentry 4479 (ARIZ, MICH, MO); Sierra Cajon del Diable, H. Gentry 11622 (ARIZ, LL, MEXU, MICH); Bahia Algadones, San Carlos, Guaymas, D. Goldberg Pitiquito, J. Hastings & R. Turner 64-58 (ARIZ, DS, SD); 8 mi ESE of Vicam along Hwy. 15, 27.6°N, 110.2°W, J. Hastings & R. Turner 69-64 (ARIZ); W of Obregon, P. Jenkins 89-317 (ARIZ, CAS); Guaymas, I. Johnston 3114 (CAS, GH, US); Hermosillo, M. Jones 22531 (MEXU, MO); 8 mi NW of Caborca toward Tajitos, D. Keck 4059 (C, CAS, DS, F, MO, NY); Mpio. Pitiquito, 54 km S of Pitiquito toward Puerto de la Libertad, ca. 11 km SW of Bimuri, 30°17’N, 112°07’'W, G. Levin 2172 (CAS, MEXU); near Pozo Cerna, Long 62 (US); Isla Tiburon, £. Lott & T. Atkinson 2518 (CAS, MEXU, TEX); New Kino, Bahia Kino, W. Mahler & J. Thieret 6070 (SUM); N de Guaymas, F’) Miranda 8943 (MEXU), 8944 (MEXU); Tiburon Island, 28°46'N, 112°18'W, R. Moran 4065 (DS, SD, UC); Guaymas, E. Palmer 307 (GH); Las Guasimas, R. Perrill 5353 (ARIZ); Mpio. Empalme, Rancho las Trincheras, ca. 6 km NW of Empalme, 28°01'N, 110°50'W, A. Reina G. & T. Van Devender 2001-182A (ARIZ, CAS); Guaymas area, near Rancho El Valiente, at jct. Hwy. 15 with bypass to Cd. Obregon, 3.8 mi N of turnoff to San Carlos Bay, ca. 28°02'N, 110°56'W, A. Sanders et al. 8767 (MO); near Rancho Verruga, Distr. Altar, F’ Shreve 5823 (ARIZ, US); 5 mi NW of Caborca, F’ Shreve 7531 (ARIZ); 15 mi NW of Caborca, F. Shreve 7553 (ARIZ, F, MICH, MO, US); ca. 20 mi NE of Obregon, W. Spaulding 75-3-12 (ARIZ); Sierra Libre, 28°34.5’N, 110°58’W, G. Starr 752 (ARIZ); 0.2 mi W of Rancho San Alfroso, 28.2°N, 111.2°W, R. Turner & L. Duek 79-37 (ARIZ); 5.8 mi NW of Kino Bay on road to Punta Chueca, R. Van Devender & T. Van Devender 83-96 (ARIZ, MEXU); Puerto Los Mochos, Rancho Los Mochos, ca. 20 mi NNE of Desemboque de San Ignacio, 7: Van Devender & M. Kearns s.n. (ARIZ); Mpio. Hermosillo, Playa Esthela, just N of Bahia de Kino, 28°52'N, 112°01'W, 7: Van Devender & A. Reina G. 2000- 932 (CAS, MEXU); Nacopuli Canyon, ca. 4 mi N of San Carlos Bay, T. Van Devender et al. 84-239 (ARIZ); 8.6 mi NW of Caborca, |. Wiggins 8268 (ARIZ, DS, TEX); Sierra Seri, ca. 29°17/N, 112°08'W, R. Wilkinson s.n. (ARIZ); Sierra del Viejo, 0.5 mi NE of Mina de la Santa Cruz, 30°23’N, 112°22'W, G Yatskievych & P. Fischer 52-04 (ARIZ). 746 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Justicia L. Justicia L., Sp. Pl. 1:15. 1753, nom. cons. prop. LECTOTYPE (Hitchcock and Green, International Botanical Congress, Cambridge (England), 1930: Nomenclature Proposals by British Botanists. 116. 1929): Justicia hyssopifolia L. Dianthera L., Sp. Pl. 1:27. 1753. TYPE: Dianthera americana L. (= Justicia americana (L.) Vahl). Adhatoda Mill., Gard. Dict. Abr. ed. 4, 1:|Adhatoda in alph. seq.]. 1754. TYPE: Adhatoda “zeylanensium” Mill. fide Graham (1988); Adhatoda vasica Nees = lectotype fide Index Nominum Genericorum (as of information dated 9 Feb 1996 at http://rathbun.si.edu/botany/ing/); both names pertain to Justicia adhatoda L. Beloperone Nees in N. Wallich, Pl. Asiat. Rar. 3:76. 1832. LECTOTYPE (Bremekamp, Index Nom. Gen. Card 00123. 1955): Beloperone amherstiae Nees (= Justicia brasiliana Roth). Sericographis Nees in Martius, Fl. Bras. 9:107. 1847. LECTOTYPE (Bremekamp, Index Nom. Gen. Card 02841. 1957): Sericographis rigida Nees (= Justicia sericographis V.A.W. Graham). Sarotheca Nees in Martius, Fl. Bras. 9:113. 1847. LECTOTYPE (Bremekamp, Index Nom. Gen. Card 01996. 1956): Sarotheca elegans Nees (= Justicia sarotheca V.A.W. Graham). Jacobinia Nees ex Moric., Pl. Nouv. Amér. 156. 1847, nom. cons. TYPE: Jacobinia lepida Nees (= Justicia lepida (Nees) Wassh.). Siphonoglossa Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854:159. 1855. TYPE: Siphonoglossa ramosa Oerst. (= Justicia ramosa (Oerst.) V.A.W. Graham). Decumbent to erect perennial herbs, shrubs, or small trees with cystoliths. Leaves opposite. Inflorescence of dichasia in leaf axils (or in axil of 2 partially fused bracts forming a cupulate involucre) or of axillary or terminal dichasiate spikes, racemes, or thyrses, inflorescences some- times branched and becoming panicles; dichasia alternate or opposite, | (—3)-flowered, sessile or pedunculate, subtended by a leaf or a bract. Flowers homostylous, sessile or pedicellate, subtend- ed by 2 homomorphic bracteoles. Calyx deeply 4-5-lobed, lobes equal or unequal in size, posteri- or lobe sometimes greatly reduced in flowers with 5 calyx lobes. Corolla greenish, white, yellow, orange, pink, red, or purple, usually with white or colored markings (often restricted to lower lip), tube cylindric to expanded distally, usually lacking a distinct throat, limb strongly zygomorphic, 2- labiate, upper lip internally rugulate (i.e., with a stylar furrow), comprising 2 fused lobes, entire to 2-lobed, lower lip 3-lobed, corolla lobes imbricate in bud. Stamens 2, exserted from mouth of corolla, anthers 2-thecous (1 theca rarely sterile), thecae equal or unequal in length, parallel to per- pendicular, equally inserted on filament, unequally inserted (but overlapping for some portion of their length), or superposed (i.e., not overlapping), | or both with a basal appendage or appendages absent, dehiscing toward lower lip (1.e., flower nototribal); pollen (Figs. 20-21) subprolate to per- prolate, 2—4-aperturate (varying from porate to colporate with distinctness of colpi sometimes questionable), apertures flanked on each side by 1-several rows of + circular insulae and/or penin- sulae or by both a solid band of exine and a pseudocolpus, exine variously ornamented but usual- ly reticulate; staminodes 0 (although pubescent thickenings near attachment of filaments rarely FIGURE 20 (right). Pollen of Sonoran Justicia. a. J. californica (Daniel 1542), apertural view. b. J. californica (Daniel 1542), interapertural view. c. J. candicans (Daniel 8355), apertural view. d. J. candicans (Daniel 8355), interapertural view. e. J. hilsenbeckii (Lott & Guadalupe A. 2656), apertural view. f. J. caudata (Breedlove & Daniel 70945), apertural view. g. J. phlebodes (Van Devender et al. 93-483), apertural view. h. J. salviiflora (Raven & Breedlove 20133), apertural view. 1. J. sonorae (Van Devender & Reina G. 98-434), apertural view. j. J. spicigera (Avila B. 28), apertural view. k. J. spicigera (Avila B. 28), interapertural view. Scales = 10 um. 747 SONORAN ACANTHACEAE DANIEL 748 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 FIGURE 21. Pollen of Sonoran Justicia. a. J. longii (Butterwick & Hillyard 7384), apertural view. b. J. longii (Butterwick & Hillyard 7384), polar view. c. J. masiaca (White 3590), apertural view. d. J. masiaca (White 3590), polar view. Scales = 10 um. present). Style exserted from mouth of corolla, stigma lobes indistinct to distinct, when distinct equal to unequal in length (1 lobe sometimes vestigial). Capsule stipitate, retinacula present, septa with attached retinacula remaining attached to inner wall of mature capsule. Seeds 2-4, homomor- phic, lenticular to globose. (x = 11, 14). Justicia is the largest genus of Acanthaceae with estimates of up to 600 species worldwide. It is also the largest genus of the family in Mexico (ca. 100 species) and in Sonora (10 species). Morphological diversity in the genus is extensive and the above generic description 1s derived from North and Central American species only. The generic synonymy given above only includes gen- era in which species from our region have been previously treated. Daniel (1995a) provided a more complete listing of generic synonyms in which American species have been treated. Graham (1988) presented a comprehensive generic account of Justicia that includes generic synonyms on a world- wide basis. 1. Calyx 4-lobed with lobes homomorphic or 5-lobed with lobes heteromorphic (posterior lobe greatly reduced in size with respect to other 4 lobes); capsule pubescent with glandular and eglandular trichomes. 2. Corolla cream or greenish with pink and maroon markings, tube 6—10.5 mm long; both the- cae dorsally pubescent; capsule 14-20 mm long; seeds covered with barbed bristles. Se EL Re RE TINT ERE (ena AOR AREER St eS 6 So 0 0 « J. salviiflora 2. Corolla pink to pale purplish, often with white markings on lower lip, tube 11—23.5 mm long; thecae glabrous (or sometimes with the upper theca pubescent); capsule 7—13.5 mm long; seeds tuberculate (lacking bristles). 3. Corolla tube gradually expanded from near base to apex, 1.8—2.7 mm in diameter near midpoint, lower lip 8-17 mm long; thecae subequally inserted; capsule 10—-13.5 mm HOM ess Soe eho Peas 2 ante «hd Seer mua te MEOR oe eee eeene ee J. sonorae . Corolla tube cylindric (expanded only near mouth), 0.6—1.2 mm in diameter near mid- point, lower lip 4-7 mm long; thecae unequally inserted or superposed; capsule 7-10 mm long. 4. Bracteoles 24.5 mm long, 0.2—0.4 mm wide; calyx 3—5 mm long, lobes subulate to lance-subulate: thecae unequally inserted (overlapping by 0.6—0.8 mm), lacking basal appendages; seeds: 2.2-2351mm long:.. 35555259 eee J. hilsenbeckii 4. Bracteoles 4.8-6.5 mm long, 0.5—0.7 mm wide; calyx (5-) 6.5—9 mm long, lobes lin- ear; thecae superposed (separated by a gap 0.40.7 mm long), lower theca with a basal ws) DANIEL: SONORAN ACANTHACEAE 749 appendages secdseles—le/smimllone yea 2 eaaers aia een genet en ay J. phlebodes 1. Calyx 5-lobed, lobes homomorphic; capsule glabrous, pubescent with eglandular trichomes only, or pubescent with both eglandular and glandular trichomes. 5. Bracts of axillary spikes heteromorphic, sterile bracts greatly reduced in size; calyx white- hyaline along margin; corolla externally pubescent with eglandular and conspicuous glan- dular trichomes, tube 5-8 mm in diameter near midpoint; thecae green; seeds smooth and PLEO RUB Cin beeps ren Sy Ooty oct ey ince AIRE A th Mae aoa het cto enn een J. caudata 5. Bracts (if present) homomorphic, sterile bracts (if present) similar to fertile bracts; calyx not white-hyaline along margin; corolla externally pubescent with eglandular trichomes only (infrequently with a few glandular trichomes in J. californica) or glabrous distally and very inconspicuously glandular proximally (J. spicigera), tube 1-4 mm in diameter near mid- point; thecae not green (often reddish); seeds smooth to tuberculate or covered with sub- conic papillae, lacking trichomes. 6. Perennial herbs to 5 dm tall; corolla tube cylindric (expanded only near mouth), 1—1.5 mm in diameter near midpoint; calyx lobes widest above the base (1.e., narrowed prox1- mally); pollen 3—4-aperturate; seeds bubbly tuberculate. 7. Corolla purple (limb) and white (tube), tube 12.5—33 mm long; style 11-33 mm long; PollemrA-apenturate tee bis as Va gel Sieh tester pat tees iene ets 2 J. masiaca 7. Corolla entirely white, tube 23-43 mm long; style (23-)32—40 mm long; pollen 3-aper- ELIT ACCS aos Nara y's cette ae ht Ag pee ait atoren Meea a atat Re ehh Sets AMO i Nees J. longii 6. Shrubs to 3 (or more) m tall; corolla tube gradually expanded from near base to apex, 1.7-4 mm in diameter near midpoint; calyx lobes widest at base; pollen 2-aperturate; seeds smooth to lumpy or covered with subconic papillae. 8. Inflorescence of axillary panicles of dichasiate spikes; bracteoles abaxially glabrous or inconspicuously glandular-punctate; corolla orange, fusiform in bud, externally glabrous distally and inconspicuously pubescent with a few subsessile glandular tri- chomes to 0.1 mm long proximally, tube 19-32 mm long, lower lip recoiled; seeds covered with subconic papillae; introduced plants (cultivated or escaping). i Ci ghiseisaey Ea aS as POR ee MP RRA Sh Ee RE Ma LE AE Bee SEAN reco J. spicigera 8.Inflorescence of dichasia in leaf axils, condensed dichasiate spikes (often appearing as clusters) in leaf axils, or (axillary or) terminal dichasiate thyrses; bracteoles abaxially pubescent with eglandular and sometimes stipitate glandular trichomes as well; corol- la red (sometimes with white markings), linear-ellipsoid in bud, externally pubescent with eglandular (and sometimes also with stipitate glandular) trichomes to 0.5 mm long, tube 11—20 m long, lower lip not recoiled; seeds smooth to lumpy (i.e., covered with irregular, low, rounded bumps); native plants (rarely cultivated). 9. Young stems + pallid resulting from a dense and even covering of very short tri- chomes, the epidermis not (or but barely) visible; inflorescence of (axillary or) ter- minal dichasiate thyrses; dichasia pedunculate, peduncles 1-14 mm long; corolla entirely red, lobes of lower lip 1—5.5 mm long; thecae dorsally pubescent, lower (and sometimes upper) theca with a prominent basal appendage to 0.6 mm long; capsule pubescent; seeds subglobose to + compressed, not red. ....J. californica . Young stems glabrous or pubescent but not pallid as described above, the epidermis usually clearly visible; inflorescence of solitary dichasia in leaf axils or condensed dichasiate spikes (appearing as clusters) in leaf axils; dichasia sessile; corolla red with white markings on lower lip, lobes of lower lip 4.5—11.5 mm long; thecae dor- sally glabrous, lacking basal appendages (or rarely the lower theca with an incon- \o 750 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 spicuous basal appendage to 0.1 mm long); capsule glabrous; seeds sublenticular, usually;somewhat reddishty ys mm. se inet eer J. candicans Justicia californica (Benth.) D.N. Gibson, Fieldiana, Bot. 34:67. 1972. Beloperone californi- ca Benth., Bot. Voy. Sulphur, 38. 1844. Jacobinia californica Nees in A. de Candolle, Prodr. 11:729. 1847. Sericographis californica (Benth.) A. Gray in Torrey in W.H. Emory, Rep. U.S. Mex. Bound. 2(1):125. 1859. TYPE: MEXICO. Baja California Sur: Cape St. Lucas, 1841, R. Hinds s.n. (holotype: K!). Phenology. Flowering: throughout the year, but with a peak in March(—April) and only spo- radically in other months (Fig. 1); fruiting: throughout the year. Distribution and habitats. Southwestern United States (Arizona, California), northwestern Mexico (Baja California, Baja California Sur, Sinaloa, Sonora); Sonoran plants occur on coastal dunes, along watercourses, and on rocky slopes in Sonoran desertscrub (Arizona Upland, Central Gulf Coast, Lower Colorado River Valley, Plains of Sonora), thornscrub, and grasslands at eleva- tions from sea level to 1046 m. Illustrations. Figure 6; Flora of Baja California, 188. 1980; Trees and Shrubs of the Southwestern Deserts, 3rd. ed., 218. 1981; Desert Plants 5:169. 1984; The Jepson Manual, Higher Plants of California, 127. 1993; Flora of the Gran Desierto and Rio Colorado of Northwestern Mexico, 65. 2000. Local names. “Chuparosa” or “chuparrosa” (fide Felger 2000 and Felger and Moser 1985; Burquez 95-123, Burquez et al. 91-280, Rea 1200, Nabhan et al. 249, etc.); “hummingbird bush” (fide Felger 2000); “koo-me-me” (Guarijio, Gentry 1291); “noj-oopis” (Seri, fide Felger and Moser 1985); ““semaluca” (Mayo, Van Devender et al. 94-386); “sewalulukut” (Yaqui, fide Felger 1999); “vipsumar hiosik” (Pima Bajo, Rea 1200): ““wipisimal” (fide Felger 2000); “wipisomal” (Papago, Bowers & Nabhan 172, Nabhan et al. 249). a eS ae 1 Uses. Source of food (floral nectar) for | Mie ——o pre-Columbian cultures (Hodgson 2001). S se NOD) sie Daniel (1997) provided a description of | SS Ses ~ this species. Its distribution is nearly restricted _3, to the Sonoran Desert of the southwestern United States and northwestern Mexico, where it is usually found in or along watercourses. In Sonora, it is widespread throughout western, central, and southern portions of the state (Fig. | Dp) ar (2 2 at j res cE a e 22). Felger (2000) noted that in northwestern os? 3° most Sonora, the flowers attract Costa’s, Rufous, and Allen’s hummingbirds, as well as + “age honeybees and butterflies. He also noted that : . Justicia californic: im carpenter bees (Xv/ocopa) and sometimes O aap eons aoe ~ 3 ; an 197 2 Justicia hilsenbecki) hummingbirds slit the floral tube and consume nectar. Other collectors have noted humming- ee : wee : = — ay : 114 2° bird visitation to flowers of Justicia californi- = Nl l l l ca from other portions of its range in Sonora FIGURE 22. Sonoran distributions of Justicia californi- ca and J. hilsenbeckii. (e.g., Parfitt & Reeves 2398 notes visitation by Anna’s and either Rufous or Allen’s hummingbirds in west-central Sonora). Turner et al. (1995) noted additional ecological aspects of the species. DANIEL: SONORAN ACANTHACEAE 751 SONORAN COLLECTIONS: Sierra Bacha, ca. 17 km SE of Puerto Libertad, ca. 29°47'N, 112°33.5'W, E. Abbey & R. Felger 20245 (ARIZ); 40 mi S of Hermosillo, L. Abrams 13350 (DS); Guaymas, R. Barr & W. Brewer 63-651 (ARIZ, MEXU); Kino Bay, R. Barr & W. Brewer 63-663 (ARIZ, MEXU); Bahia San Carlos, Perinola Palms, ca. 7 mi N of Bahia San Carlos, F’ Boutin & F) Brandt 2827 (MEXU); Quitovac, between Sonoyta and Caborca, J. Bowers & G. Nabhan 172 (ARIZ); Guaymas, T. Brandegee s.n. (UC); Mpio. San Luis Colorado, 11.4 mi S of Sonoita on Mex. 2, D. Breedlove 1457 (DS, DUKE); Mpio. Hermosillo, 7 mi W of Hwy. 15 toward Bahia Kino, D. Breedlove 15942 (DS, ENCB); Sierra del Viejo, 7 km N and 3 km E of El Plomito, 30°19'N, 112°20'W, T: Burgess 5772 (ARIZ); Cafiada Tetabejo, Sierra Libre, S de Hermosillo, 28°33'N, 110°58'W, A. Burquez 95-123 (MEXU); 2.5 km S de Hermosillo, 29°01'N, 110°57'W, A. Burquez & M. Quijada 90-179 (MEXU), 97-90 (MEXU), A. Burquez & M. Quintana 94-204 (MEXU); Cafion las Cocinas, entre San Carlos y Tastiota, 28°16'N, 111°23'W, A. Buirquez et al. 91-280 (MEXU); 59 km SE de Caborca, 30°37'N, 111°36’W, A. Campos V. et al. 4427 (CAS, MEXU); S Sonoyta, O. Clark 11369 (UNM); N Guaymas, O. Clark 14957 (UNM); Mpio. Hermosillo, S of Moro Colorado, 5 mi SSE of Estero Tastiota, J. Copp 70-10 (CAS); Gran Desierto, SE side of Sierra del Rosario, ca. 32°05'N, 114°10.5'W, H. Coss et al. 20389 (ARIZ, ENCB); Torres, F’ Coville 1651 (NY, US); near KM 74, Calle 4 Sur (diagonal road from Mex. 15 S of Hermosillo to Mex. 16 W of Hermosillo), G Cummins s.n. (ARIZ); between Hermosillo and Kino Bay, 2.5 mi E jet. Son. 16 (Calle 4 Sur) to Guaymas, 7: Daniel 942 (CAS); “Microondas mountain” overlook- ing Guaymas, along road which is 1.8 mi from Mex. Hwy. 15 on paved road to airport, N of Guaymas, T. Daniel 1956 (ASU, CAS); Nacapuli Canyon, 6.3 mi W of Hwy. 15 on road to San Carlos then 4.4 mi N, 7: Daniel 1992 (ASU); 1.6 mi NE of Masiaca toward Alamos, 7.1 mi NE Hwy. 15, 7; Daniel 2547 (CAS); along road between Hwy. 16 E of Tonichi and Movas, 15.8 mi S of Onavas, 7: Daniel 3354 (CAS); W base of Sierra (NW of Guaymas), 7. Daniel 3976 (CAS); Bahia Kino, N of village, F) Drouet & D. Richards 3518 (DS, F, US); NW edge of Guaymas, F’ Drouet & D. Richards 4033 (F); along Hwy. 15 ca. 2 mi N of jet. to Becachibampo Bay, D. Dreyer 215 (CAS); Sierra de Calera E of Villa de Seris, F) Drouet et al. 3398 (F); San Carlos Bay, ca. 7 mi W of Hwy. 15, E. Engle 29 (MEXU); Pinacaté, derrame de lava en Campo Rojo, 31°45’N, 113°27'W, M. Equihua s.n. (ARIZ, MEXU); ca. 2 mi E of Esperanza (several mi N of Cd. Obregon, R. Felger 360 (ARIZ); 1.8 mi NW of town of Bahia Kino, R. Fe/ger 2513 (ARIZ, MEXU); A. Rodriguez Dam, E side of Hermosillo, Rio Sonora, R. Felger 2785 (ARIZ, MEXU, TEX); Sau Sal landing field, S end of Tiburén Island, R. Felger 6436 (ARIZ); NW part of Isla Tiburon, vicinity of Tecomate, Arroyo Agua Dulce, R. Felger 6828 (ARIZ); Isla San Esteban, R. Felger 7057 (CAS), R. Felger et al. 17597 (CAS, MEXU); 2.2 mi W of Tajitos (NW of Caborca), R. Felger 7450 (ARIZ); Isla Cholludo, between Isla Tiburon and I. Turners, R. Felger 9162 (ARIZ); Hourglass Canyon, ca. 2 mi NE of Huarache Tank, W side of Pinacate region, R. Felger 19136 (ARIZ, ENCB, GH); SW part of Pinacate region, ca. | km S of Tule Tank, R. Fe/ger 19228-F (ARIZ); 1.3 mi N of village of Bahia Kino and ca. 1 km inland from shore, R. Felger & J. Cooper 15251 (ARIZ); Isla San Esteban, N side of island, R. Felger & J. Cooper 15408 (MEXU); Bahia Colorado, E base of Morro Colorado, ca. 28°18.5’N, 111°28'W, R. Felger & E. Hamilton 15667 (ARIZ); ca. 5 mi N of El Desemboque San Ignacio, R. Felger & M. Moser 5240 (ARIZ); Rancho San Jorge, ca. 20 mi W of Hermosillo, R. Felger & A. Russell 6892 (ARIZ, MEXU); | mi N of Bahia San Carlos on road to Algadones, R. Felger & A. Russell 9590 (ARIZ, MEXU); Isla Tiburon, ca. 13 mi S of Tecomate, ca. 28°57’N, 112°27'W, R. Felger & A. Russell 12356 (ARIZ, ENCB, MEXU, UC); Isla Cholludo, between Isla Tiburon and I. Turners, R. Felger & O. Soule 13414 (ARIZ); Cafion Nacapules, ca. 4 km N of Bahia San Carlos, R. Felger & R. Thomas 11992 (ARIZ, MEXU); 11.8 mi S of Sonoyta, R. Felger et al. 9824 (ARIZ); Ensenada Grande (= Bahia San Pedro, N of Guaymas), R. Felger et al. 11600 (ARIZ); Isla Tiburon, vicinity of Tecomate, NW part of island, R. Fe/ger et al. 12527 (ARIZ); Isla Alcatraz, Bahia Kino, R. Felger et al. 12723 (ARIZ); Isla San Esteban, E-central side of island, R. Felger et al. 12763 (ARIZ); N side of Cerro Tepopa, SW of El Desemboque San Ignacio, ca. 29°22'N, 112°24'W, R. Felger et al. 14141 (ARIZ, CAS); Pinacate Region, Tinaja Chivos, ca. 31°45'N, 113°36.5'W, R. Felger et al. 15609 (ARIZ); Pinacate Region, E side of Sierra Blanca, ca. 31°34’N, 113°26'W, R. Felger et al. 20210 (ARIZ); Sierra del Rosario, ca. 32°06'N, 114°11.5'W, R. Felger et al. 20738 (ARIZ, ENCB); Sierra del Rosario, ca. 32°06’, 114°11’W, R. Felger et al. 75-15 (ARIZ, ENCB, MEXU); Cafion Nacapules, ca. 6 km N of Bahia San Carlos, ca. 28°10'N, 111°03.5'W, R. Felger et al. 85-580 (ARIZ, CAS, MEXU, TEX); Loma Chomajabires, 1.8 mi N of Huatabampito, ca. 8 mi S of Huatabampo, 26°42'N, TS, PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 109°36'W, M. Fishbein & S. McMahon 2748 (ARIZ, CAS); vicinity of Nescotahueca, 4.5 km E of jet. of south Camahuiroa airstrip and Diez de Abril Road, 4 km W of Tierra y Libertad, 26°34’N, 109°15'W, S. Friedman & G Hall 398-93 (ARIZ); Arroyo Camahuiroa, 2 km NE of Camahuiroa, 9.5 km WNW of Melchor Ocampo, 26°33'N, 109°15.5'W, S. Friedman & J. Zittere 025-95 (ARIZ); 277 km S of Nogales toward Hermosillo, T. Frye & E. Frye 2283 (DS, GH, MO, NY, UC, US); N edge of Hermosillo, near Ganadero Motel, A. Gentry 552 (MO); Cocorit, Yaqui Valley, H. Gentry 885 (DS), 885M (ARIZ); Salitral, Rio Mayo, H. Gentry 129] (ARIZ, F); Cerro de Bavajori, 12 mi W of Navojoa, H. Gentry 7953 (ARIZ, US); Algodones Bay, D. Goldberg s.n. (ARIZ); E end of bay at Port Libertad, E. Graham 3810 (DS); 20 mi N of Port Libertad, H. Green s.n. (PH); ca. 30 mi S of Hermosillo on Hwy. 15, B. Hansen et al. 1375 (MEXU, WIS); 2.1 mi NW of Bahia Kino, J. Hastings & R. Turner 64-32 (ARIZ, DS); 2 mi N of Tastiota, J. Hastings & R. Turner 65-168 (ARIZ); 3 mi SSE of Rancho Las Pefiitas, 29.8°N, 111.8°W, J. Hastings & R. Turner 69-112 (ARIZ); Islas Melisas in N part of Guaymas Bay, J. Hastings et al. 61-39 (ARIZ); Punto Cirio, ca. 7 mi S of Puerto Libertad, J. Hastings et al. 63-7 (ARIZ); “Catch 22” ca. 5 mi N of San Carlos, 27°57'N, 111°05'W, A. Johnson 5013 (DAV); “Playas del Sol” ca. 10 mi S of Guaymas, 27°55'N, 110°45’W, A. Johnson 5052 (DAV); Tiburon Island, Freshwater Bay, I. Johnston 3250 (CAS); Tiburon Island, 3 mi N of Willards Point, 2. Johnston 4245 (CAS); 5 mi NW of Caborca on road to Tajitos, D. Keck 4052 (C); 2 mi N of Horcasitas, M@. Kimnach & Lyons 596 (UC, US); 70 mi S of Hermosillo, G Lindsay 1133 (DS); Isla San Esteban, SE de la isla, E. Lott & T. Atkinson 2458 (MEXU); 3 mi S of Mesquite, F Long 13 (US); Picu, D. MacDougal & F. Shreve 15 (US); Kino Pt., D. MacDougal & F. Shreve 27 (US); Arroyo Desemboque, 20 mi N of Sargento, 77 Mallery & W. Turnage s.n. (DS); mouth of Salitral Creek at Rio Mayo, Rancho Salitral near San Bernardo, P. Martin 12 (ARIZ); 35 km SSW of Caborca on road to Libertad, ca. 30°28'N, 112°20'W, P. Martin & S. Nilsson 9 (S); 7 km S of Libertad at Punta Cirio, ca. 29°50’N, 112°39'W, P. Martin & S. Nilsson 13 (S); Estero Santa Barbara, W of Huatabampito, 26°42’N, 109°38'W, P. Martin & M. O'Rourke s.n. (ARIZ); Sonoyta River, E. Mearns 2790 (DS, NY, US); Sierra Seri, ca. 21°17'N, 112°08’W, E. Moser et al. 18157 (ARIZ); Quitovac between Sonoyta and Caborca, G. Nabhan et al. 249 (ARIZ): without locality, E. Palmer s.n. in 1869 (US); near Guaymas, W. Palmer 1213 (US); 1/4 mi W of Condominios Pilar, 1 mi S of hwy. to San Carlos, ca. 4 mi W of Hwy. 15, B. Parfitt & T. Reeves 2398 (ENCB); along Mex. Hwy. 8, near KM 19, S of US-Mexico border at Lukeville/Sonoyta, L. Parker s.n. (ARIZ); 6 mi N of Obregon, K. Parker 8209 (ARIZ); 6 km WNW of San Carlos, 27°57'N, 111°06'W, M. Quinn & P. Sudt 046 (ARIZ); 6.5 mi W of Rio Yaqui along Hwy. 16 near La Barranca, ca. 28°28’, 109°32'W, A. Rea 1200 (ARIZ); Ensenada Chica, 15.1 mi W of Hermosillo to Guaymas Hwy. by dirt road, 6 mi S of San Augustin Beach turnoff, 28°07'N, 111°17'W, F: Reichenbacher 230 (ARIZ); Mpio. San Pedro de la Cueva, Canada La Pila, | km N of Rio Moctezuma, ca. 9.5 km NE of San Pedro de la Cueva, 29°18'N, 109°43'W, A. Reina G. & T. Van Devender 99-255 (CAS); Mpio. Cd. Obregon, 0.5 km S of Estacion Corral junction on Mex. 15 on road to Cocorit, 27°37'N, 109°58'W, A. Reina G. & T. Van Devender 2001-178 (CAS); Mpio. Huatabampo, 6 km NW of Camahuiroa on road to Las Bocas, near Bachomojaqui, ca. 26°34'N, 109°18'W, A. Reina G et al. 98-2138 (MEXU); Mpio. Caborca, SE of Puerto Lobos, Ejido Manuel Avila Camacho, 30°16'N, 112°49'W, 4. Reina G. et al. 2000-166 (CAS); Mpio. Altar, Rancho El Rincon, 3.5 km S of Sasabe, 31°26’N, 111°34'W, A. Reina G et al. 2003-1291 (CAS); vicinity of Hermosillo, J. Rose et al. 12374 (GH, NY, US); Kino Bay, V. Rudd et al. 3042 (ENCB, MEXU),; ca. 0.5 km S of Carnegie Peak, Sierra Pinacate, NV. Sakaki et al. 1988] (ARIZ, ENCB); Canon del Coyote, ca. 4 mi NE of Desemboque, C. Saravia T. s.n. (ARIZ); near Port Libertad, H. Shantz s.n. (WIS); E trail of Pinacate Peak, 31°45’N, 113°30'W, W. Sherbrooke s.n. (ARIZ); inland from San Carlos Bay, Guaymas, P. Sherwin 215 (DUKE); 5 mi NW of Caborca, F Shreve 7535 (ARIZ, F, MO); Cafién Palma, Sierra Libre, F) Shreve s.n. (MEXU); E of Pinacate Peak, Pinacate Mountains, J. Soule s.n. (ARIZ); Nacopuli Canyon, off road to San Carlos Bay, 28°01'N, 111°03.5'W, G Starr & C. Starr 204 (ARIZ): carretera Bahia Kino—Puerto Libertad, 40 km N de intersecciOn con carretera a Hermosillo, 29°30/N, 112°15'W, H. Suzan 315 (MEXU); Isla Tiburon, NW 20 km de la playa, Sierra Kunkaac, F. Torres 4 (ENCB); Punto Cirio, ca. 6 mi S of Puerto Libertad, R. Turner & C. Lowe 60-31 (ARIZ); ca. 12 mi S of Hermosillo, 7. Van Devender s.n. (ARIZ); 29 mi S of Hermosillo on Hwy. 15, T. Van Devender s.n. (ARIZ); 3.2 mi S of Sasabe, 7. Van Devender s.n. (ARIZ, GH, MEXU); Puerto Los Mochos, Rancho Los Mochos, ca. 20 mi NNE of Desemboque del Rio San Ignacio on road to La Ciénega, T° Van Devender & M. Kearns s.n. (ARIZ, TEX); Sierra Bacha, Punto Cirio, near Libertad, 77 Van Devender & M. Kearns s.n. (ARIZ); Mpio. Hermosillo, Cerrito de la Virgen, 5.4 km S of Hermosillo, 28°59’'N, 110°58’W, DANIEL: SONORAN ACANTHACEAE VSS T. Van Devender & A. Reina G. 2004-2 (CAS); 8 km E of Tecoripa on MEX 16, ca. 28°37’N, 109°50’W, 7: Van Devender & A. Reina G. 2004-27 (CAS); 29.4 mi N of Hermosillo on Hwy. 15, 7. Van Devender et al. 88-838 (ARIZ); mouth of Arroyo Masiaca, Las Bocas on Gulf of California, ca. 52 km S of Navojoa, 26°36/N, 109°20'W, 7: Van Devender et al. 94-386 (ARIZ, CAS); Mpio. Navojoa, Teachive de Masiaca, Arroyo Masiaca, 26°47'N, 109°14'W, T. Van Devender et al. 94-956 (ARIZ); Mpio. Hermosillo, Bahia de Kino, 7: Van Devender et al. 96-238 (ARIZ, MEXU); Sierra Bacha, SE of Punta Cirio, ca. 29°50'N, 112°37-38’W, G. Webster 22457 (DAV, MEXU): near Ures, J. Whitehead M152 (ARIZ); KM 101 on Hermosillo—Moctezuma road, 38.6 mi E of Moctezuma, ca. 29°28’N, 110°15’W, 4. Whittemore et al. 83-100 (CAS, MEXU, TEX); 2 mi W of Libertad, . Wiggins 6085 (DS, US); 10 mi N of Hermosillo, 2. Wiggins 6251 (DS, F, UC, US); 5 mi SE of Torres, /. Wiggins 6286 (DS, UC, US); 17 mi NE of Cajeme toward Tesopaco, /. Wiggins 6390 (DS, UC, US): 5.7 mi NW of Caborca toward Sonoyta, 1. Wiggins 8257 (DS, UC, US); Sierra Bacha, S of Punta Cirio, F: Wiseman & W. Spaulding s.n. (ARIZ). Justicia candicans (Nees) L.D. Benson in L.D. Benson and R.A. Darrow, Trees and Shrubs of the Southwestern Deserts, ed. 3, 218. 1981. Adhatoda candicans Nees in A. de Candolle, Prodr. 11:396. 1847. Dianthera candicans (Nees) Hemsl., Biol. Cent.-Amer., Bot. 2:517. 1882. Jacobinia candicans (Nees) B.D. Jacks., Index Kew. 1:1246. 1893. TYPE: MEXICO. Oaxaca: “Cordillerae Mexicanae,” 5000 ft., November—April 1840, H. Galeotti 9/1 (holotype: K ex hb. Hook.!; isotypes: BR!, G!, P!, see discussion). Jacobinia mexicana Seem., Bot. Voy. Herald, 325, t. 66. 1856. TYPE. MEXICO. State unde- termined (see discussion): “Sierra Madre,” B. Seemann 2115 (lectotype, designated here, see dis- cussion below: K ex herb. Hook.!). Jacobinia ovata A. Gray, Proc. Amer. Acad. Arts 21:405. 1886. TYPE: MEXICO. Chihuahua: near Batopilas, Aug-Nov 1885, E. Palmer 220 (holotype: GH!, photo at DS!; isotypes: G!, NY, US!). Jacobinia ovata var. subglabra S. Wats., Proc. Amer. Acad. Arts 24:67. 1889. Jacobinia can- dicans var. subglabra (S. Watson) L.D. Benson in L.D. Benson and R.A. Darrow, Trees and Shrubs of the Southwestern Deserts, ed. 2, 413. 1954. Justicia candicans var. subglabra (S. Wats.) L.D. Benson in L.D. Benson and R.A. Darrow, Trees and Shrubs of the Southwestern Deserts, ed. 3, 218. 1981. TYPE: MEXICO. Sonora: rocky ravines, Guaymas, Oct 1887, E. Palmer 264 (holotype: GH!; isotypes: C!, DS!, NY!, UC!, US). Justicia mexicana Rose, Contr. U.S. Natl. Herb. 1:348. 1895. TYPE: MEXICO. Sonora: Agiabampo, 3-15 Oct 1890, E. Palmer 7&8 (holotype: US!; isotypes: GH!, NY!, US!). Phenology. Flowering: throughout the year, with peaks in March and November (Fig. 1); fruit- ing: September—April. Distribution and habitats. Southwestern United States (Arizona), Mexico (Baja California Sur, Chihuahua, Colima, Durango, Guanajuato, Guerrero, Jalisco, Michoacan, Nayarit, Oaxaca, Puebla, Querétaro, Sinaloa, Sonora); Sonoran plants occur on rocky slopes and along watercourses in Sonoran desertscrub (Arizona Upland, Central Gulf Coast, Plains of Sonora), palm oases, thorn- scrub, tropical deciduous forests, and grasslands at elevations from sea level to 1330 m. Illlustrations. Figure 6; The Botany of the Voyage of H.M.S. Herald, t. 66. 1856; Trees and Shrubs of the Southwestern Deserts, 3rd. ed., 218. 1981; Flora del Valle de Tehuacan-Cuicatlan 355. 1999, Local names. “Ciatica” (fide Yetman and Van Devender 2002); “flor de chuparosa” (Gentry 861); “maso o’ota” (Mayo, Van Devender et al. 93-1437); “muicle cimarron” (fide Van Devender et al. 2000); “palo de venado” (Van Devender et al. 93-1437); “rama venado” (Van Devender et al. 93-1437): “vipsumar” (Pima Bajo, Rea 1262). 754 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Use. Yetman and Van Devender (2002) noted that the Mayo sometimes brew the leaves into a tea that is drunk as a treatment for malaria. Locality data on the holotype of Justicia candicans at K is as noted above. The first set of Galeotti’s collections is at BR, however, and locality information on the presumed isotype there indicates that the plants were collected in Tehuacan, Puebla at an elevation of 5250 ft. in August 1840. Galeotti 911 at P indicates collection at 5000 ft. in Oaxaca but gives the date as August. Whether these collections represent duplicates of the same gathering in either Oaxaca or Puebla, or whether they represent different gatherings (and thus not duplicates) remains undetermined. Based on information provided by McVaugh (1972), Seemann’s type collection of Jacobinia mexicana was likely made in Sinaloa, Durango, or Nayarit. There are two specimens of Seemann 2115 in Hooker’s herbarium at Kew of about equal quality. The one selected as lectotype has orig- inal drawings on the sheet similar to those in the protologue. Plants of the type specimen have tri- angular bracteoles shorter than (i.e., not surpassing) the calyx thereby resembling the Gentry, Krizman & Soule, and Sanders et al. collections from Sonora noted below. Jacobinia mexicana was treated as a synonym of Justicia candicans by Daniel and Acosta (2003). By citing Galeotti 911 under Jacobinia mexicana Hemsley (1882) also apparently felt that the types of Jacobinia mexi- cana and Justicia candicans represented the same taxon. Bentham and Hooker (1876) noted that “Adhatoda sericea Nees” was also a name that applied to Jacobinia mexicana; however, I can find no evidence that such a name was ever published by Nees. Daniel (1997, 1999a) and Daniel and Acosta (2003) provided descriptions of Justicia candi- cans. The species 1s both widely distributed and morphologically variable. It occurs throughout much of Sonora (Fig. 23), but is especially [ ‘ ' a ! common in regions of thornscrub and tropical Ee eae Pa ata deciduous forest. It occurs in three subdivisions _| ee a me of the Sonoran Desert, but is apparently very | <0 hi rare in the Arizona Upland and entirely absent L31° ye 4 a) from the Lower Colorado River Valley. In | N @ ee i) Sonora, some plants, especially some from near L iN we e °° 3 Guaymas, have stems and leaves either | \ e e. | glabrous or very sparsely pubescent (e.g., , e ef Brandegee s.n., Felger & Thomas 11837, |-29: ry ee H Palmer 264, Phillips et al. 75-152). These have | <¢ a °° cet, been treated taxonomically as J. candicans var. = f subglabra. Other plants, including the type of [ >. ~ Jacobinia ovata from southwestern Chihuahua | 1 re 5 and several collections from Sonora (e.g., | 57 On aes Bee cata Abrams 13218, Daniel 3348, Joyal 1562, Moly ® Landrum et al. 5436, Wiggins 7151) are dense- | 110° li ly and evenly pubescent with flexuose eglandu- * lar trichomes. Most Sonoran collections have a pubescence intermediate between these extremes. Because there appears to be a continuum in veg- etative vesture from glabrous (or with very few trichomes) to densely pubescent, no formal taxo- nomic status is here accorded the various “pubescence forms” of this species. Few Sonoran plants of J. candicans show the well-developed clusters of cobwebby trichomes in axils of major veins on the abaxial surfaces of the leaves that were noted by Daniel and Acosta (2003) for representa- tives of the species in south-central Mexico. These are evident only in Gentry 4879 and Sanders et al. 4619. FIGURE 23. Sonoran distribution of Justicia candicans. DANIEL: SONORAN ACANTHACEAE (DS The types of Jacobinia ovata, J. ovata var. subglabra, and Justicia mexicana resemble the majority of Sonoran plants by having bracteoles lance-subulate to linear to oblanceolate and longer than (or overtopping) the calyx. Several specimens from Sonora (e.g., Gentry 4879, Krizman & Soule s.n., and Sanders et al. 4619) have triangular to subulate to linear bracteoles that are shorter than the calyx. In this respect, they resemble the type of Jacobinia mexicana. Among the presumed type specimens of J. candicans, bracteoles vary in length from shorter than (or at least not surpass- ing) the calyx to equaling the calyx to slightly surpassing it and they vary in shape from lance-lin- ear to linear to linear-elliptic. Landrum et al. 5425 from Sonora also appears intermediate with its somewhat linear bracteoles that are slightly shorter than to nearly equal to the calyx. Because of the overlap in bracteolar form evident among plants occurring outside of Sonora, all of these plants are treated as comprising a single variable species. The inflorescence of Justicia candicans consists of solitary dichasia in the leaf axils or of con- densed dichasiate spikes in the leaf axils. The distal leaves bearing axillary infloresceces some- times become reduced in size and bractlike, thereby resulting in a terminal, compound inflores- cence. This is especially evident in Fe/ger 397. SONORAN COLLECTIONS: 6 mi S of Magdalena, L. Abrams 13218 (DS); 12 mi NW of Hermosillo, L. Abrams 13307 (DS); 3 km SSE of El Chinal Road toward Rio Cuchujaqui, ca. 22.5 km SSE of Alamos, 26°S1'N, 108°54'W, M. Baker et al. 10371 (TEX); Hotel Playa, Guaymas, R. Barr & W. Brewer 63-653 (ARIZ, MEXU): Cerro Bocochibampo, Guaymas, FE. Blakley B-1639 (DS); ca. 16 mi S and 6.5 mi W of Trincheras, 30.2°N, 111.6°W, J. Bowers & R. Turner 2194B (ARIZ); Guaymas, T. Brandegee s.n. (UC); San Javier, microondas on el Cerro El Durazno, 28°37'N, 109°45'W, A. Burquez 94-019 (MEXU); Cafion de Nacapule, Sierra el Aguaje, ca. 20 km N de Guaymas 28°01'N, 111°03’W, A. Burquez 94-251 (MEXU)); 2.5 km S de Hermosillo, 29°O1’N, 110°57'W, A. Burquez & A. Quijada 92-84 (MEXU); W of Magdalena, O. Clark 12642 (GH); village 8 mi W of Torres, between Guaymas and Hermosillo, /) Coville 1643 (US); “Microondas mountain” overlooking Guaymas, along road which is 1.8 mi from Mex. Hwy. 15 on paved road to airport, N of Guaymas, 7. Daniel 1957 (ASU); between Hwy. 16 E of Tonichi and Onavas, 0.5 mi S jet. Hwy. 16, 7; Daniel 3348 (CAS); Rio Cuchujaqui, ca. 8 mi SE of Alamos, 7: Daniel & M. Butterwick 3192 (ASU, CAS); Nacapuli Canyon, 6.3 mi W of Hwy. 15 toward San Carlos, then 4.4 mi N, 7. Daniel et al. 1997 (ASU, CAS), 2336 (ASU, CAS); Mpio. Yécora, Arroyo La Quema, 0.3 km SE of Tepoca on Hwy. 16, ca. 28°26'N, 109°15’'W, T. Daniel et al. 8644 (CAS); Bahia San Carlos, E. Dawson 1065 (DS); Magdalena, F: Drouet & D. Richards 3810 (F, US); mountain on E side of Guaymas, F’ Drouet & D. Richards 3553 (F, US); Guaymas, F: Drouet & D. Richards 3893 (F); mountain W of Alamos, F) Drouet & D. Richards 3939 (F); Rio de Sonora near Union, Hermosillo, F Drouet et al. 3442 (F); 11 mi NE of Baviacora, F.) Drouet et al. 3663 (A, DS, F, MO, US); Palm Canyon, 25 km SSE of Magdalena on road to Cucurpe, 30°28’N, 110°48'W, D. Ducote et al. 76-6 (ARIZ, MEXU-245739); near Navojoa, C. Erlanson & M. Souviron 29 (US); Cafion de los Chapotes, E side of Sierra de Alamos, R. Felger 397 (ARIZ, CAS, MEXU); Tiburon Island, Sierra Kunkaa, ca. 4 mi W of Punta Narragansett, SE part of island, R. Felger 6972 (ARIZ, ENCB); Cerro Tetas de Cabra, Bahia San Carlos, R. Felger 85-365 (ARIZ, MEXU); La Sorpresa, SW of Trincheras, ca. 30°10/N, 111°43'W, R. Felger & A. Russell 67685 (ARIZ); Rancho San Jorge, W of Hermosillo, 29°44'N, 111°51’W, R. Felger & A. Russell 6920 (ARIZ), 6943 (ARIZ); Cerro del Vigia, Guaymas, R. Felger & R. Thomas 11837 (ARIZ, CAS, MEXU); 1.6 km W of Estero Solado, ca. 4.5 mi W of Mex. 15 on road to Bahia San Carlos, R. Felger & D. Valdez Z. 84-541 (ARIZ); Ensenada Grande (= San Pedro Bay, north of San Carlos Bay), R. Felger et al. 11576 (ARIZ, TEX); Cafion de Nacapules, ca. 6 km NE of Bahia San Carlos, R. Felger et al. 84-92 (ARIZ, MEXU, TEX); El Novillo, R. Felger et al. 84-232 (ARIZ); Arroyo Guajaray, between Los Aguaros and Cajon del Ardilla, 27°39'N, 108°58'W, R. Felger et al. 94-58A (ARIZ); Cafion Las Barajitas, Sierra El Aguaje, ca. 18 km NW of San Carlos, ca. 28°03'N, 111°11'W, R. Felger et al. 95-193 (ARIZ); El Reparo—Middle, old road to Bermudez, 28°21'N, 109°0S’W, G Ferguson & C. Moore s.n. (MEXU); ca. 8 mi N of Guaymas on San Carlos Bay, R. Ferris 8724 (DS); 14.5 mi E of Rio Yaqui on Mex. 16, P. Fischer & G Yatskievych 6887 (ARIZ); Rancho La Colorada, ca. 1.5 mi SW of Alamos, J. Fish 189 (UC); 21 mi NE of Los Tanques at jct. with road to La Vinateria, 27°15’N, 108°42’W, M. Fishbein et al. 96 (ARIZ); El Naranjo, Arroyo Taymuco, 756 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 27°15’N, 108°46'W, M. Fishbein et al. 216 (ARIZ); N of Guaymas on Hwy. 15, M. Fishbein et al. 221 (ARIZ); Mpio. Guaymas, Sierra El Aguaje, Aguaje Robinson, ca. 10 km NW de San Carlos, 28°03.5'N, 111°07'W, A. Flores M. & O. Gutierrez R. 5089 (ARIZ, MEXU); Mpio. La Colorada, 4 km antes de San José de Pimas, carr. Hermosillo—Yécora, 28°46'N, 110°37’W, A. Flores M. & J. Sanchez E. 4871 (ARIZ, MEXU); Mpio. Hermosillo, La Pintada, 50 km S de Hermosillo, 28°35'N, 110°58'W, A. Flores M. & J. Sanchez E. 5452 (IEB); 9.5 km S on Mex. 15 from Las Bocas Road turnoff, 3.5 km W on Sirebampo Road, 11.5 km S of San José de Masiaca, 26°39/N, 109°15'W, S. Friedman & J. Zittere 90-95 (ARIZ); 9.8 km W of Mex. 15 on Las Bocas Road, 37 km SE of Huatabampo, 5.6 km E of Las Bocas, 26°39'N, 109°19.5'W, S. Friedman & J. Zittere 149-95 (ARIZ); Distr. Alamos, Quiricoba, H. Gentry 743 (DS); Distr. Alamos, Cuchuhacki Arroyo, H. Gentry 861 (DS); Agua Caliente, Distr. Alamos, H. Gentry 905M (ARIZ); Canyon Sapopa, Rio Mayo, H. Gentry 1029 (ARIZ, MO, US); Caramechi, Rio Mayo, H. Gentry 1206 (A, ARIZ, F, MEXU, MO, WIS, S, UC, US); near Las Guasimas, Navojoa—Alamos road, H. Gentry 4879 (ARIZ, MEXU, MO); Los Cerritos, ca. 40 mi N of Navojoa, H. Gentry 14393 (ARIZ, US); Sierra Bojihuacame SE of Cd. Obregon, H. Gentry 14499 (ARIZ, LL, US); Ures, D. Gold 756 (MEXU, MO); ca. 6 mi W of Santa Ana de Yécora, D. Goldberg 77-78 (ARIZ); 4.3 km SW of Santa Ana de Yécora, 28°23’N, 109°19'W, D. Goldberg & R. Frye 77-271 (ARIZ); Las Cuevas, C. Hartman 159 (GH); Oputo, C. Hartman 211 (A, GH, PH, US); 11 mi S of Nacozari, J. Hastings & R. Turner 65-110 (ARIZ); Lo de Campo Ranch, 6 km S of San Javier on Rio San Javier, S of Hwy. 16, 28°32.1'N, 109°44.6'W, P. Holm s.n. (ARIZ); Mpio. Alamos, Rio Cuchujaqui, 27°02’N, 108°42'W, P. Jenkins 89-430 (ARIZ); Santa Ana de Yécora, P. Jenkins s.n. (ARIZ); Guaymas, I. Johnston 3095 (CAS); San Pedro Bay, [. Johnston 4312 (CAS); Guaymas, M. Jones 23264 (CAS, MO, UC); between cerros El Cucurucho and El Aguaje from Alamos to La Higuera, 27°04—05’'N, 108°58'W, E. Joyal 1344 (CAS, MEXU, TEX); Puenta La Pila, above “El Palmar,” ca. 20 km E of Onavas, 28°29’N, 109°22’'W, E. Joyal 1562 (CAS, MEXU); 1 mi E of Navojoa, R. Kirzman & O. Soule s.n. (ARIZ); road to Sahuaripa, ca. 22 mi E of Hwy. 15 bypass, L. Landrum et al. 5425 (CAS); 2.6 mi E of bridge at Huasabas, L. Landrum et al. 5436 (CAS, GH); Arroyo Gochico, ca. 8 km E of San Bernardo, 27°24'N, 108°47'W, G Levin et al. 2018 (CAS); Tres Marias limestone quarry, 27°07'N, 109°10'W, P. Martin s.n. (ARIZ, C); 8 km E of Alamos on road to Sabinito Sur, P. Martin & G. Ferguson s.n. (ARIZ); Cerro Verde, 28°34’N, 109°44’W, P. Martin & G. Ferguson s.n. (ARIZ); Puerto El Chino, 26°41.3’N, 108°57.8'W, P. Martin & M. O'Rourke s.n. (ARIZ); Guaymas, Microwave Mountain, 27°57'N, 110°54'W, P. Martin & M. O’Rourke s.n. (ARIZ); NE of Alamos toward Cuchujaqui, 27°01—06'N, 108°53—55'W, P. Martin et al. s.n. (ARIZ, CAS); Estacion Margarita, 10 km NW of Navojoa, P. Martin et al. s.n. (ARIZ); Jorinabo, along Rio El Naranjo, 27°15.5'N, 108°46.4’W, P. Martin et al. s.n. (ARIZ); Palm Canyon, 17 mi SE of Magdalena on road to Cucurpe, Sierra Baviso, V. McCarten & R. Bittman 2712 (ARIZ); 30 mi N of Nuri, S. McLaughlin & M. Karpiscak 518 (ARIZ); Rio Yaqui drainage, Cerro Verde to San Javier, 28°35’N, 109°44'W, K. Moore s.n. (ARIZ, CAS); San Pedro Bay, 28°03.5'N, 111°17’W, R. Moran 4038 (DS, UC); Mpio. Guaymas, Canon Los Anegados, ca. 2.8 km del Rancho El Aguaje de Robinson, 28°02’N, 111°08'W, L. Moreno M. et al. 41 (ARIZ); Rio Moctezuma, 10 mi SW Nacozari, 30°15’N, 109°40'W, T: Nash et al. L19447 (CAS, US); ca. 43 mi N of Guaymas along Hwy. 15 near microwave station, D. Norris et al. 20041 (DS, MEXU, MO); 6 mi W of Nuri on road to Movas, M@. O'Rourke & P. Martin s.n. (ARIZ); Guaymas, EF. Palmer 264 (US); Mpio. Hermosillo, Predio Ranchito de la Sierra, A. Peinado et al. A- 05 (MEXU); S of Magdalena, F’ Pennell 20265 (PH); Nacopuli Canyon, 3.5 mi N of San Carlos Bay, A. Phillips et al. 75-152 (ARIZ, UC); region of Rio Bavispe, Valle de Teras, E. Phillips 755 (GH); Onavas, ca. 28°28'N, 109°32'W, A. Rea 1262 (ARIZ); 2.3 mi E of Rio de Bavispe by Huasabas to El Coyote and Huachinera Road, 29°56'N, 109°16'W, F: Reichenbacher 216 (ARIZ); Palm Canyon, 15.5 mi toward Cucurpe from Mex. 15 in Magdalena, F) Reichenbacher 913 (CAS); 2 mi E of Santa Rosa and 17 mi W of Yécora, F Reichenbacher & L. Toolin 1373 (ARIZ); Mpio. Moctezuma, Rio Moctezuma at San Clemente de Terapa, 29°41'N, 109°39'W, A. Reina & T. Van Devender 2003-576 (CAS); Mpio. Soyopa, vicinity of Rio Yaqui bridge on Mex. 16, just S of Tonichi, ca. 28°34'N, 109°33'W, A. Reina et al. 97-09 (ARIZ); Mpio. Nacozari de Garcia, Rancho Agua Caliente (Rio Bavispe drainage), 23.2 km SE of Esqueda, 30°39’N, 109°25’W, A. Reina G. et al. 2004-376 (CAS); Guaymas, H. Ripley 14294 (CAS); vicinity of Hermosillo, J. Rose et al. 12481 (F); Sierra de Alamos, J. Rose et al. 12780 (NY, US); Mpio. Alamos, vic. of Presa Mocuzari, 27 km SW of Alamos, ca. 27°14’N, 109°05'W, A. Sanders & G Helmkamp 13319 (CAS); N end of Sierra El Viejo, 25 mi SW of Caborca, ca. 30°22'N, 112°22’W, A. Sanders et al. 3530 (ARIZ); near Mina Sahuarito, 20 mi DANIEL: SONORAN ACANTHACEAE TT) SW of Trincheras, ca. 30°09'N, 111°43’W, A. Sanders et al. 3582 (ARIZ, CAS, UC); Cerro Prieto (= Cerro Colorado), near summit, 8.6 mi E of Navojoa toward Alamos, ca. 27°15'N, 109°17'W, A. Sanders et al. 4619 (CAS); near Santa Rosa, 25 mi S of Trincheras, F’ Shreve 6049 (ARIZ, F); 37 mi NE of Cajeme on road to Tesopaco, F: Shreve 6153 (ARIZ, MEXU); 7 mi S of Magdalena, F: Shreve 10039 (ARIZ); La Pintada Canyon, Hwy. 15, 32.5 mi S of Hermosillo, E. Smith 3973 (CAS, MEXU, TEX); Palm Canyon, 17.7 mi SE of Magdalena, road to Curcupe, Cerro Cinta de Plata, 30°29’N, 110°48'W, G Starr & K. Birgy 157 (ARIZ); 1.8 mi E of Alamos on road to Guadalupe, 27°02’N, 108°54.5'W, G Starr & D. Palzkill 317 (ARIZ); Cerro Dos Negritos, 1.8 mi N of Alamos, 27°03'N, 108°56'W, G Starr & D. Palzkill 380 (ARIZ); Microondas hill, Guaymas, G: Starr & C. Starr 196 (ARIZ); Nacopuli Canyon, 28°01'N, 111°03.5'W, G Starr & C. Starr 205 (ARIZ): Canyon de la Bota, N end of Sierra El Tigre, ca. 36 mi E of Esqueda, ca. 30°36'N, 109°13'W, G Starr et al. 31 (ARIZ); Mpio. de Alamos, Sierra de Alamos along Arroyo El Huirotal, ca. 26°57'N, 108°57'W, V. Steinmann 1295 (CAS); Cerro La Salada, 29.4°N, 111.0°W, R. Turner & D. Brown 85-131 (ARIZ); Bahia San Pedro, 28.1°N, 111.2°W, R. Turner et al. 79-292 (ARIZ); Sierra Alamos, ca. 4 mi SSW of Alamos, 26°59’N, 108°57'W, R. Van Devender & T. Van Devender 83-126 (ARIZ); 5.2 mi ENE of Cucurpe on road to Rancho Agua Fria, 7; Van Devender s.n. (ARIZ); 17 mi SE of Magdalena in Sierra Babiso (= Cerro Cinta de Plata), T. Van Devender s.n. (TEX); 8.2 mi E of Restaurante La Pintada on Mex. 15, Cafion La Pintada, Sierra Libre, T. Van Devender s.n. (ARIZ); Cafion Babiso, Rancho Babiso, 2.5 mi W of San Carlos Bay, 7: Van Devender & M. Kearns s.n. (ARIZ); Palm Canyon, Cerro Cinta de Plata, 17.7 mi SE of Magdalena, 7. Van Devender & C. Miksicek s.n. (ARIZ); Playa La Manga, N of San Carlos Bay, 7: Van Devender & F: Nishida s.n. (ARIZ); 8.7 km E of Tecoripa on MEX 16, 28°37’'N, 109°48'W, T: Van Devender & A. Reina G. 2004-26 (CAS); ca. 8 mi E of turnoff of Son. Hwy. 12 at Esqueda on road to El Tigre, 30°39'N, 109°25'W, T: Van Devender et al. 82-32a (ARIZ); Cafion de la Bota, N end of Sierra El Tigre, ca. 34 km ESE of Esqueda, ca. 30°36'N, 109°13'W, T. Van Devender et al. 82-52 (ARIZ); Guaymas Microwave Tower road (Microondas El Vigia), 7. Van Devender et al. 84-227 (ARIZ); El Guayabo crossing of Rio Cuchujaqui, 2.6 km NE of Sabinito Sur, 14 km ESE of Alamos, 27°00'N, 108°47'W, T. Van Devender et al. 92-1265 (ARIZ); 8.9 mi ESE of Alamos, Rio Cuchujagui, 26°59’N, 108°49'W, T: Van Devender et al. 92-1333 (ARIZ); El Rancheria crossing of Rio Cuchujaqui, ca. 22.5 km S of Alamos toward El Chinal, 26°51'N, 108°55'W, T. Van Devender et al. 93-29 (ARIZ, CAS); Mocuzari Dam on Rio Mayo, 27°13'N, 109°07'W, 7: Van Devender et al. 93-223 (ARIZ, CAS, MO, TEX); canyon N of Los Aguaros on Arroyo Guajaray, 27°38.5'N, 108°58'W, 7: Van Devender et al. 93- 490 (ARIZ, in part); Laguna Barochipa, ca. 3.3 km SSE of Camahuiroa, 26°31’N, 109°16'W, 7: Van Devender et al. 93-1272 (ARIZ, CAS); Saucito Canyon, Rancho La Sierrita, 6 km SSW of Alamos, 26°58.5'N, 108°58'W, T. Van Devender et al. 93-1366 (ARIZ, CAS); Mpio. Alamos, Cerro El Chorro, 4 km NW of Yocogigua, 26°49’N, 109°03.5'W, T. Van Devender et al. 93-1437 (ARIZ, CAS); El Rincon Viejo, ca. 4 km N of Alamos, Sierra de Alamos, 27°04'N, 108°56'W, 7. Van Devender et al. 93-1509 (ARIZ); 4.3 km SW of Huasaguari on Masiaca—San Antonio de Las Ibarras road, 26°47.5'N, 109°11'W, 7. Van Devender et al. 95- 275A (ARIZ, TEX); 1.5 km SW of Santa Ana on road to Guadalupe Tayopa, 28°23’N, 109°09.5'W, T: Van Devender et al. 97-213 (ARIZ, MEXU); Ejido Francisco Solis, 7 km ENE of Est. Luis (SE of Navojoa on Mex. 15), 26°35’N, 109°06'W, T. Van Devender et al. 98-2113 (MEXU); Mpio. Nogales, 3.8 km NE of El Correo (La Arizona), Rancho Las Boregas, Arroyo Planchas de Plata, 31°11.5’N, 111°10'W, 7: Van Devender et al. 2004-214 (CAS); 20.4 mi SE of Magdalena on Cucurpe road, 7. Van Devender et al. s.n. (ARIZ); E of Restaurante La Pintada (ca. 78 km N of Guaymas on Mex. 15), Cerro Bola in Sierra Libre, 7; Van Devender et al. s.n. (ARIZ); 3.9 mi SW of Mazocahui, 7. Van Devender et al. s.n. (ARIZ); San Javier, puerto San Juan a 1.5 km S del poblado, 28°35'N, 109°45’W, L. Varela & E. Cuamea 97-7 (MEXU); near Bahia San Carlos, W of Guaymas, ca. 27°57'N, 111°04’W, G. Webster 19725 (ARIZ, MEXU); 39 mi E of San José de Pimas, 28°38'N, 110°00'W, G. Webster 23808 (MEXU); Cafiada Motepori, ca. 9 mi NW of Banamichi crossing, ca. 30°04'N, 110°20'W, G. Webster & R. Murphey 21475 (ARIZ, GH, MEXU); above Aduana, Sierra de Alamos, 8 km ESE of Alamos, 27°02’N, 109°02'W, J. Weins et al. 93-092 (ARIZ, CAS); 2 mi E of Rancho San Carlos, on road to Norio, 1. Wiggins 6142 (DS); 23 mi NE of Cajeme on road to Tesopaco, /. Wiggins 6402 (DS, UC, US); E of La Palma, 5 mi N of Guaymas in Sierra Libres, 1. Wiggins 6478 (DS, F, UC, US); SW side of Babiso Mts., 18 mi SE of Magdalena, J. Wiggins 7151 (A, ARIZ, DS, UC, US); N of Los Aguaros on Arroyo Guajaray, 27°38’N, 108°58’W, D. Yetman et al. s.n. (CAS). 758 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Justicia caudata A. Gray, Proc. Amer. Acad. Arts 21:405. 1886. TYPE: MEXICO. Chihuahua: near Batopilas, Aug-Nov 1885, E. Palmer 189 (holotype: GH!; isotypes: K!, US). Phenology. Flowering: March—April, August-November; fruiting: March—April, September— November. Distribution and habitats. Mexico (Chihuahua, Chiapas, Distrito Federal, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, México, Michoacan, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, Sinaloa, Sonora, Zacatecas), Guatemala; Sonoran plants occur on rocky slopes, along watercourses, and in disturbed areas in tropical deciduous forests and oak woodlands at elevations from 240 to 805 m. Illustration. Flora Fanerogamica del Valle de México, 716. 2001. Daniel (1995a, 1999a) and Daniel and Acosta (2003) provided descriptions of Justicia L € caudata and discussed some of its variation. pale The species reaches its northwestern distribu- tional limit in east-central Sonora (Fig. 24). +31 eis r Plants from Sonora are rather homogeneous . with the young stems usually sparsely pubes- h cent with retrorse or downward pointing (rarely L flexuose) eglandular trichomes 0.2—1 mm long, \ the abaxial surfaces of the petioles conspicu- _ 49 ously glandular pubescent, and the corollas pink-purple with white markings on the lower lip and 18-31 mm long. is SONORAN COLLECTIONS: ca. 3 km N de Tepoca sobre Mex. 16, 28°27'N, 109°16'W, A. Burquez M. 94-138 (CAS); southeastern border of Alamos (road to mirador), ca. lat. 27°01'N, long. 108°S6'W, 7. Daniel 9771 (CAS); Rio Cuchujaqui, ; ee | ies | ca. 8 mi SE of Alamos, 7? Daniel & M. Butterwick FIGURE 24. Sonoran distributions of Justicia caudata 3195 (CAS); Alamos area, road to Rio Cuchujaqui, — and J. Jongii. A. Faivre et al. 64 (ARIZ); Arroyo La Barranca de Tonichi, 8.1 mi E of Rio Yaqui along Hwy. 16, M. Fishbein et al. 2448 (ARIZ); 4.3 km SW of Santa Ana de Yécora on road to Nuri, 28°23’N, 109°19'W, D. Goldberg & S. McLaughlin 77-178 (ARIZ, TEX); Rio El Naranjo crossing N of Taymuco, 27°15'N, 108°43'W, P. Jenkins & P. Martin 88-252 (ARIZ); Mpio. Alamos, upper Rio Cuchujaqui, Palmarito Canyon, 27°04'N, 108°45'W, P. Jenkins et al. 92-82 (ARIZ); Rancho las Uvalamas near Alamos, 26°58'N, 108°55—5S6'W, P. Martin & M. McWhorter s.n. (ARIZ, CAS); Arroyo el Mentidero at El Chinal road, 11.3 km S of Alamos, 26°55’N, 108°55'W, S. Meyer s.n. (CAS); Alamos, £. Palmer 402 (US); 2.5 mi W and 3/4 mi S of Alamos, R. Perrill & V. Phelps 5168 (ARIZ); 12 km E of Alamos toward upper crossing of Rio Cuchujaqui, ca. 27°00'N, 108°50'W, A. Sanders et al. 9423 (ARIZ, CAS, TEX); Mpio. Alamos, Arroyo Mentidero, from Alamos—El Chinal road to Rio Cuchujaqui, 12 km S of Alamos, 26°55’N, 108°55’W, A. Sanders et al. 12575 (CAS, MEXU); 1 km W of Son. 12 on road to La Quema, 28°22'N, 109°16'W, T. Van Devender & A. Reina G. 97-1029 (ARIZ, CAS, MEXU, TEX); Mpio. Yécora, Arroyo El Pilladito, near Tepoca, 28°26’N, 109°15'W, 7: Van Devender & A. Reina G. 98-1058 (CAS, MO); Mpio. Yécora, ca. 1.5 km SE of San Nicolas on Mex. 16, 7; Van Devender & A. Reina G. 2000-670 (MO); Arroyo El Cobre, near Choquincahui, 26°59'N, 108°41'W, 7: Van Devender & D. Yetman 94-741 (ARIZ); Arroyo Guirocoba, 2.3 km NE of El Cajon de Sabino, 4.8 km S of Guirocoba, 26°51.5'N, 108°42’W, T Van Devender & D. Yetman 94-763 (ARIZ); Arroyo el Mentidero at El Chinal road, 11.3 km S of Alamos, 26°5S'N, 108°5S5'W, T. Van Devender et al. 92-968 (ARIZ), 93-838 (ARIZ, CAS); El Guayabo crossing of Rio Cuchujaqui, 2.6 km NE of Sabinito Sur, 14 km ESE of Alamos, 27°00'N, 108°47'W, 7. Van Devender et al. © Justicia caudata L979 @ Justicia longii DANIEL: SONORAN ACANTHACEAE (59 92-]224 (ARIZ); El Rincon Viejo, ca. 4 km N of Alamos, Sierra de Alamos, 27°04'N, 108°56’W, 7. Van Devender et al. 94-625 (ARIZ, TEX). Justicia hilsenbeckii V.F. Daniel, nom. nov. Siphonoglossa mexicana Hilsenb., Madrofio 36:198. 1989. TYPE: MEXICO. Sinaloa: Imala, 29 Nov 1939, H. Gentry 5099 (holotype: CAS!; isotypes: GH, MO, NY, US), non Justicia mexi- cana Rose (1895). Phenology. Flowering: December; fruiting: December. Distribution and habitats. Endemic to Mexico (Durango, Guerrero, Jalisco, Michoacan, Morelos, Puebla, Sinaloa, Sonora, and Veracruz); Sonoran plants occur in tropical deciduous forests at an elevation of 500 m. Illustration. Madrono 36:199. 1989. Local name. “Cordoncillo” (fide Yetman and Van Devender 2002). Use. The Mayo make a tea from the branches that is used as a wash for rheumatism (Yetman and Van Devender 2002). A description of this species, as Siphonoglossa mexicana, was provided by Hilsenbeck (1989). Daniel (1995a, 1999b) noted that this species, as well as other species of Siphonoglossa recognized by Hilsenbeck (1989, 1990a), would appear to be best treated in Justicia. Graham (1988) also included the type of Siphonoglossa within Justicia. Anew name is necessary for S. mexicana when it is treated in the latter genus. The distinctions among J. canybi Greenm., J. ramosa (Oerst.) Graham, and J. hilsenbeckii do not appear to be substantial. Indeed, Hilsenbeck (1989) noted over- lap in morphological traits between J. canbyi (as S. canbyi (Greenm.) Hilsenb.) and J. hilsenbeckii (as S. mexicana) and between J. ramosa (as S. ramosa) and J. hilsenbeckii (as S. mexicana). As a result, plants treated by Hilsenbeck as S. mexicana exhibit considerable variation in many quanti- tative characters and in pubescence. The entire complex warrants thorough restudy from through- out its range. Morphological data used above in the key to species of Justicia in Sonora is derived from the only known Sonoran collection and the type, which greatly resemble one another. The species reaches the northern extent of its distribution in southern Sonora (Fig. 22). Among Sonoran Acanthaceae, this species might be confused with the morphologically simi- lar one, J. sonorae. Sonoran representatives of these two species can be distinguished by the fol- lowing couplet: 1. Corolla (15—) 19-24 mm long, tube cylindric (expanded only near mouth), 11-19 mm long, 0.6—0.8 mm in diameter at midpoint, lower lip 4-6.5 mm long; thecae 0.9-1.2 mm long, unequally inserted (overlapping by 0.6—0.8 mm); capsule 7-10 mm long... . . J. hilsenbeckii 1. Corolla 23-35 mm long, tube gradually expanded from near base to apex, 18—23.5 mm long, 1.8-2.7 mm in diameter near midpoint, lower lip 8-17 mm long; thecae 1.2—2.2 mm long, subequally inserted (overlapping by 1—1.5 mm); capsule 10—13.5 mm long. ..... J. sonorae Sonoran collection: Sierra de Alamos, El Rincon Viejo, Arroyo El Aguaje, ca. 4 km N of Alamos, 27°04'N, 108°56’W, T. Van Devender et al. 93-1508 (ARIZ, CAS). Justicia longii Hilsenb., Pl. Syst. Evol. 169:231. 1990. Siphonoglossa longiflora (Torr.) A. Gray, Syn. Fl. N. Amer. 2(1): 328. 1878. Adhatoda longiflora Torr. in W.H. Emory, Rep. U.S. Mex. Bound. 2(1):125. 1859, non Justicia longiflora Vis. (1839, 1840). TYPE: MEXICO. Sonora: road between Zufi and Altar, Sep 1855, A. Schott III no. 3 (Mex. Bound. Surv. 726) (holotype: NY!; iso- types: F!, US). 760 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Perennial herbs to 3.5 (—8) dm tall. Young stems subquadrate to multi-striate, evenly pubes- cent with retrorse to retrorsely appressed eglandular trichomes 0.05-0.8 mm long. Leaves subses- sile to petiolate, petioles to 10 mm long, blades linear-lanceolate to lanceolate to elliptic to ovate, 5-70 mm long, 2—12 (—18) mm wide, 2—10 times longer than wide, (rounded to) acute to acumi- nate to subfalcate at apex, acute to subattenuate at base, surfaces pubescent with mostly antrorse eglandular trichomes 0.05-0.8 mm long, trichomes sometimes becoming restricted to major veins on mature leaves. Inflorescence of axillary sessile to subsessile (i.e., borne on peduncles to 1 mm long) dichasia; dichasia opposite at distal leaf nodes, | per axil, 1-3 (or more)-flowered. Bracteoles often subfoliose, linear to lance-elliptic, 4-24 mm long, 0.5—4.7 mm wide, abaxial surface pubes- cent like leaves; secondary bracteoles, if present, usually similar to bracteoles except smaller (sometimes becoming + subulate). Flowers sessile. Calyx 5-lobed, 4—9 (—12) mm long, lobes lin- ear to lance-elliptic, 3.5—8 (10.5) mm long, 0.7—2.1 mm wide, narrowed proximally (i.e., widest above base), abaxially puberulent with antrorse eglandular trichomes 0.05 mm long. Corolla entire- ly white, 31-55 mm long, externally pubescent with flexuose to retrorse eglandular trichomes 0.1—0.3 mm long, tube cylindric, 2343 mm long, 1—1.5 mm in diameter near midpoint, upper lip 5—12 mm long, 2-lobed at apex, lobes 0.5—1 mm long, 0.8—1.2 mm wide, lower lip 6-13 mm long, lobes 4-12 mm long, 3—5.6 mm wide. Stamens 4-7 mm long, filaments glabrous; thecae 1-2 mm long (including basal appendage), equal to subequal in size, parallel to subparallel, unequally inserted (overlapping by 0.6—1 mm), glabrous, both with a basal appendage or the upper theca lack- ing an appendage, basal appendages 0.2—0.3 mm long; pollen (Fig. 21) 3-aperturate, apertures flanked on each side by 2 (—3) rows of insulae. Style (23—) 32-41 mm long, pubescent with eglan- dular trichomes, stigma 0.2—0.3 mm long, subhemispheric to + equally 2-lobed. Capsule 6.5—10 mm long, glabrous, stipe 2.5—4 mm long, head ovoid to ellipsoid to subglobose. Seeds 4, lenticu- lar, 2—2.7 mm long, 2—2.5 mm wide, surface and margin bubbly-tuberculate. Phenology. Flowering: March, August-December; fruiting: March, September—December. Distribution and habitats. Southwestern United States (Arizona, Texas), northwestern Mexico (Sonora); Sonoran plants occur on rocky slopes in Sonoran desertscrub (Arizona Upland, Central Gulf Coast, Plains of Sonora) and thornscrub at elevations from 380 to 910 m. Illustrations. Figure 25; Plant Systematics and Evolution 169:228. 1990. In the protologue of Adhatoda longiflora Torrey cited: “road between Zuni and Alta Sonora, September; Schort.” Hilsenbeck annotated the sole specimen in Torrey’s herbarium at NY with this information as the lectotype. It is here regarded as the holotype. On a sheet at F, there are four Schott labels (road between Zuni and Altar, 1855, III.3.; near Pozo de Marias, no date, no number; Sta. Magdalena, no date, 1; near Sta. Gertruda del Altar, no date, no number) and seven shoots (all pertaining to J. /ongii), but with no correlation of labels and shoots. Hilsenbeck annotated the “specimen” as an isolectotype of Adhatoda longiflora. Because at least one of these labels agrees with information on what is here considered to represent the holotype at NY, an isotype is present among the various collections on this sheet. Although Hilsenbeck (1990b) provided a recent description of Justicia longii, his circumscrip- tion of the species included /. masiaca as well. Thus, a description of J. Jongii based on plants from throughout its range is provided here. The species is nearly endemic to the Sonoran Desert region and reaches the southern limit of its distribution in central Sonora (Fig. 24). Because the species has been collected in Chihuahuan desertscrub in western Texas, it should be sought in the isolated region of that community in northeastern Sonora. The distinctions between J. /ongii and J. masiaca are apparently all associated with the flow- ers; as a result, non-flowering collections can be difficult to identify. For example, Daniel 981 lacks flowers and is tentatively treated as the former species primarily on the basis of the location where DANIEL: SONORAN ACANTHACEAE 761 FIGURE 25. Justicia longii. a. Habit (Jenkins & McLaughlin 1016), <1.2. b. Node (Daniel 159), 4.5. c. Bracteole (Jenkins & McLaughlin 1016), *2.7. d. Secondary bracteole (Jenkins & McLaughlin 1016), <5.6. e. Calyx (Turner et al. 78-45), “5.6. f. Distal portion of corolla and stamens (Jenkins & McLaughlin 1016), *1.9. g. Anther (Butterwick & Hillyard 7384), x29. h. Stigma (Gould et al. 2799), x42. i. Capsule (Daniel 159), «5.9. j. Seed (Daniel 119), x15. Drawn by Alan ~ Chou. 762 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 it was collected (which is nearer to populations of J. /ongii than to those of J. masiaca). SONORAN COLLECTIONS: between La Colorada and Tecoripa, 12.6 mi E of La Colorada, T. Daniel 98] (CAS); Isla Tibur6n, NE [NW fide Felger, pers. comm.] of Santa Rosa, R. Felger 9349 (ARIZ, MEXU); S end of Sierra Libre, 12.3 mi S of La Palma on Hwy. 15 (at KM 190), then 4.9 mi E of Hwy. to 0.1 mi below sum- mit of Microondas Avispas, 28°29'N, 111°02'W, R. Felger & F. Reichenbacher 85-1099 (ARIZ, MEXU, TEX); La Sorpressa, SW of Trincheras, 3010'N, 11143'W, R. Felger & A. Russell 6756 (ARIZ); 12.3 mi W of Mex. 15 on Tecolote Road (1.2 mi N of El Oasis), 29°47'N, 111°15’W, R. Reichenbacher 1060 (ARIZ); Mpio. Ures, 4 km NNW of Ures on road to Rayon, Rio Sonora drainage, 29°27’N, 110°25'W, A. Reina G & T. Van Devender 2001-07 (ARIZ, MEXU, CAS); Mpio. Nacozari de Garcia, Rancho Agua Caliente (Rio Bavispe drainage), 23.2 km SE of Esqueda, 30°39’N, 109°25'W, A. Reina G. et al. 2004-379 (CAS); near Mina Sahuarito, S of Cerro San Luis on road from Mina San Ignacio, 20 mi SW of Trincheras, ca. 30°09’N, 111°43'W, A. Sanders et al. 3583 (ARIZ, CAS, UC); 7 mi S of Magdalena, F’ Shreve 6657 (ARIZ); 36 mi NW of Caborca and 16.5 mi NW of Tajitos, 31.2°N, 112.5°W, R. Turner & J. Hastings 72-36 (ARIZ, MEXU); between Magdalena and Santa Ana, 7 mi S of Magdalena, 1. Wiggins 7190 (DS, F, GH, MICH, UC, US). Justicia masiaca T.F. Daniel, Brittonia 47:408. 1995. TYPE: MEXICO. Sonora: between Masiaca and Alamos, 1.6 mi NE of Masiaca and 7.1 mi NE of Hwy. 15, ca. 26°47'N, 109°14'W, ca. 60 m, 14 Jan 1983, 7. Daniel et al. 2546 (holotype: CAS!; isotypes: ARIZ!, ASU!, ENCB!, MEXU!, MICH!, NY!, US!). Phenology. January—March; August—October; fruiting: January—March; August—October. Distribution and habitats. Northwestern Mexico (Sonora, Sinaloa); Sonoran plants occur in thornscrub and desert grasslands at elevations from 60 to 1100 m. Illustration. Brittonia 47:409. 1995. r aT Daniel (1995c) provided a description of | ~ eS this species, which reaches its northern distrib- ee So utional extent in the grasslands of northeastern | SSS a Sonora (Fig. 26). L 34° , SONORAN COLLECTIONS: Mpio. Navojoa, ca. | 2 km N of Teachive, SW end of Cerro Terucuchi, 26°48'N, 109°14'W, A. Reina et al. 96-581 (CAS, fF MEXU); ca. | mi SW of Mesa Masiaca, E side of Hwy. 15, 8.8 mi S of jet. with Hwy. 176 near KM 116 (S of Navojoa), 7. Van Devender & A. Sanders 29" Say 92-1058 (ARIZ, CAS, MO, TEX); 1.3 km W of oy tees Hwy. 15 on Huatabampo Rd, ca. 22 km §S of Navojoa, 26°49'N, 109°23'W, T. Van Devender et al. 93-257 (ARIZ, ASU, CAS, DAV, MO, SD; TEX, UCR); 9.3 km E of Masiaca on road to Yocojigua, 26°45'N, 109°08'W, T: Van Devender et al. 93-827 (ARIZ, CAS); region of the Rio de Bavispe, Canon > Justicia masiaca 27 @ Justicia salviiflora de las Bellotas, S. White 3590 (GH); region of the __, ne | Rio de Bavispe, Canon del Agua Amarga, S. White Figure 26. Sonoran distributions of Justicia masiaca 3590 (ARIZ, US). and J. salviiflora. Justicia phlebodes Leonard & Gentry, Brittonia 6:327. 1948. TYPE: MEXICO. Sinaloa: Sierra Tacuichamona, between Culiacan and Mazatlan and S of the Rio San Lorenzo, near Africa, 17 Feb 1940, H. Gentry 5652 (holotype: US: isotypes: DS!, GH!, MICH). Perennial herbs to | m tall. Younger stems subterete to subquadrate, pubescent with flexuose to retrorse to retrorsely appressed eglandular trichomes 0.3—1 (—1.5) mm long, trichomes concen- DANIEL: SONORAN ACANTHACEAE 763 trated in or restricted to 2 lines or + evenly disposed. Leaves petiolate, petioles to 30 mm long, blades ovate-elliptic to elliptic to subcircular, 29-122 mm long, 12-70 mm wide, 1|.1—2.7 times longer than wide, acute to subacuminate at apex, (truncate-subattenuate to) attenuate at base, sur- faces pubescent (trichomes sometimes restricted to major veins) with flexuose to antrorse eglandu- lar trichomes to | mm long, margin entire to subsinuate, usually ciliate. Inflorescence of axillary (in axils of leaves or inflorescence bracts distally) or terminal dichasiate spikes (or panicles of spikes), rachises (of both panicles and spikes) evenly pubescent with erect eglandular (sometimes inconspicuous) and glandular trichomes 0.05—0.3 mm long (glandular pubescent); dichasia oppo- site (to alternate), sessile, | per axil, 1-flowered. Bracts (including those subtending spikes and dichasia) elliptic to obovate, 4-7 mm long, 1.2—3 mm wide, abaxial surface glandular pubescent. Bracteoles lance-linear to linear, 4.8—6.5 mm long, 0.5—0.7 mm wide, abaxial surface glandular pubescent. Flowers sessile. Calyx 4-lobed, (5—) 6.5—9 mm long, lobes linear, (4.5—) 5.2-7.5 mm long, equal to subequal in length, 0.4—0.7 mm wide, abaxially glandular pubescent. Corolla 19-28 mm long, pink-purple, externally glandular pubescent, tube cylindric (expanded slightly only near apex), 13-23 mm long, 0.9—1.2 mm in diameter near midpoint, upper lip 4-6 mm long, entire to 2- fid at apex, lobes to 0.6 mm long, lower lip 4-7 mm long, lobes 34.5 mm long, 2—3.5 mm wide. Stamens inserted near apex of corolla tube, 3.5—5 mm long, filaments glabrous, thecae pink-pur- plish, subparallel to parallel, superposed (gap 0.4—0.7 mm long), unequal in size, upper theca 1.3—-1.5 mm long, glabrous, lacking a basal appendage, lower theca 1—1.2 mm long (including basal appendage), glabrous, with a basal appendage 0.2 mm long; pollen (Fig. 20) 2-aperturate, apertures flanked on each side by 2—2.5 rows of insulae (to peninsulae). Style pink-purplish, 16.5—22 mm long, pubescent (at least proximally) with eglandular (and sometimes a few glandular) trichomes, stigma 0.4—0.5 mm long, only | lobe evident. Capsule 7-8 mm long, externally glandular pubes- cent (or with some of the eglandular trichomes retrorse), stipe 2—2.5 mm long, head subellipsoid with a slight medial constriction. Seeds 4 per capsule, sublenticular, 1.5—1.7 mm long, 1—1.3 mm wide, surface and margin tuberculate. Phenology. Flowering: March, October-November; fruiting: March, October-November. Distribution and habitats. Northwestern Mexico (Chihuahua, Durango, Sinaloa, Sonora); Sonoran plants occur in canyons and along watercourses in tropical deciduous forests and oak-pine forests at elevations from 320 to 1250 m. Illustration. Figure 27. Local name. “Cordoncillo” (Fishbein et al. 178). Justicia phlebodes is endemic to northwestern Mexico and reaches the northwestern extent of its distribution in southern Sonora (Fig. 28). SONORAN COLLECTIONS: trail to Tepopa from El Chiribo, 27°19’N, 108°44'’W, M. Fishbein et al. 178 (ARIZ), 199 (ARIZ); Mpio. Alamos, upper Rio Cuchujaqui, Arroyo Santa Barbara, 27°06.5'N, 108°43'W, P. Jenkins et al. 92-99 (ARIZ, CAS, MO); Rancho San Pedro, El Cajon along Rio Cuchujaqui, P. Martin s.n. (ARIZ); Rancho San Pedro, 23 km E, 4 km N of Alamos, 27°03'N, 108°42’W, P. Martin et al. s.n. (ARIZ); canyon N of Los Aguaros on Arroyo Guajaray, 27°38.5’N, 108°58’W, 7: Van Devender et al. 93-483 (ARIZ, CAS). Justicia salviiflora H.B.K., Nov. Gen. Sp. 2:233. 1818. Sarotheca salviiflora (H.B.K.) Nees in A. de Candolle, Prodr. 11:382. 1847. TYPE: cultivated in a Mexican botanical garden (fide proto- logue) (holotype: P-Bonpl.!; isotype: P!). Justicia paniculata Rose, Contr. U.S. Natl. Herb. 1:348. 1895, non Burm f. (1768) nec Forsk. (1775) nec Sessé & Moc. (1887). TYPE: MEXICO. Colima: vicinity of Colima, 9 Jan—6 Feb 1891, E. Palmer 1143 (holotype: US!; isotypes: K!, US!). 764 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 FIGURE 27. Justicia phlebodes. a. Habit (Gentry 5652), x 0.5. b. Inflorescence node with flower (Van Devender et al. 93-483), x 3.2. c. Calyx (Van Devender et al. 93-483), x 5.3. d. Trichomes on abaxial surface of calyx (Van Devender et al. 93-483), x 18. e. Distal portion of stamen with anther (Van Devender et al. 93-483), x 24. f. Capsule (Jenkins et al. 92-99), x 5.7. g. Trichomes on outer surface of capsule (Jenkins et al. 92-99), x 25. h. Seed (Van Devender et al. 93-483), x 17. Drawn by Nadia Strasser. Phenology. Flowering: March—April; fruiting: March—April. Distribution and habitats. Mexico (Chiapas, Chihuahua, Colima, Durango, Guerrero, Jalisco, Mexico, Michoacan, Morelos, Nayarit, Oaxaca, Sinaloa, Sonora); Sonoran plants occur along watercourses, on rocky slopes, and in disturbed areas in tropical deciduous forests and oak wood- lands at elevations from 310 to 580 m. Illustrations. Figure 9; The Botany of the Voyage of H.M.S. Herald, t. 67. 1856; Flora of Chiapas 4:78. 1995. Daniel (1995a) provided a detailed description of this species, which reaches the northern and western extents of its distribution in southeastern Sonora (Fig. 26). Sonoran plants are perennial herbs or shrubs that sometimes appear somewhat vinelike with secondary rooting at the nodes. Corollas are whitish with a green upper lip and purple striped ridges on the lower lip. DANIEL: SONORAN ACANTHACEAE 765 SONORAN COLLECTIONS: Arroyo Guajaray, Y iw eget a between Los Aguaros & Cajon del Ardilla, 27°39’N, ee 2A 108°58'W, R. Felger et al. 94-58A (ARIZ); Rio Mayo Region, Agua Salado, 15 mi NE of Los Tanques, 27°15’N, 108°46'W, M. Fishbein et al. 85 |_ 3° (ARIZ); Rio Mayo Region, Arroyo Guayjaray, just N f of Los Aguaros, 27°38'N, 108°58'W, M. Fishbein et al. 1565 (ARIZ, CAS); Rio Mayo Region, Tepopa, + H. Gentry 1433 (A, ARIZ, F, MEXU, MO, UC, WIS), 7435 (MEXU); Rio Mayo Region, Arroyo Gochico, H. Gentry 3624 (ARIZ, F, GH, MEXU, }+-29° MO, UC, US); Mpio. Alamos, Upper Rio Cuchujaqui, cajon of the Cuchujaqui, 27°02’N, 108°42'W, P. Jenkins 89-438 (ARIZ, CAS); Mpio. F Yécora, Arroyo del Pilladito, near Tepoca, 28°26'N, 109°15’W, A. Reina G. & T. Van Devender 97-337 (ARIZ, CAS, MEXU, TEX); Mpio. Alamos, 9.5km [27° N of Giiirocoba along road to Choquincahui (El Cobre), ca. 26°58’N, 108°41'W, V. Steinmann et al. ae 114 ao Ce ME xO) WoW or Son. 12 (Tepocas FIGURE 28. Sonoran distributions of Justicia phlebodes Cd. Obregon Hwy.) on road to La Quema, 28°22'N, and J/. sonorae. 109°16'W, W. Trauba 216-98 (CAS). © Justicia phlebodes @ Justicia sonorae Justicia sonorae Wassh., Phytologia 49:65. 1981. TYPE: MEXICO. Sonora: 17.2 mi SSE of Magdalena, palm canyon in Cerro Cinta de Plata (= Sierra Babiso), 13 Feb 1977, 7: Van Devender s.n. (holotype: US!; isotypes: ARIZ!, GH!, UC!). Ascending to erect perennial herbs to 4.5 dm (—1 m) tall. Young stems subterete to quadrate- sulcate to hexagonal, glabrous or evenly to bifariously to unifariously pubescent with flexuose to retrorse (to retrorsely appressed) eglandular trichomes 0.2—1 mm long, sometimes also with scat- tered glandular trichomes (usually restricted to distal few internodes) 0.05—0.3 mm long. Leaves (plants sometimes leafless during anthesis) petiolate, petioles to 5 (-7) mm long, blades lance- ovate to ovate (to cordate), 7-55 mm long, 3—22 mm wide, 1.3—2.8 times longer than wide, acute to acuminate at apex, truncate to rounded to acute at base, surfaces pubescent with mostly erect to flexuose eglandular trichomes to | mm long. Inflorescence of axillary or terminal dichasiate spikes to 15 cm long, these sometimes basally branched (and thus becoming a panicle of spikes), spikes (when several) collectively forming a terminal + leafy panicle, spike rachises evenly pubescent with erect to flexuose to retrorse eglandular and glandular trichomes 0.05—0.3 mm long (glandular pubescent); dichasia (alternate to) opposite (sometimes opposite a fertile branch), sessile, 1 per axil, 1-flowered. Bracts opposite, triangular-subulate to lance-subulate, 1-4.8 mm long, 0.8—1.5 mm wide, abaxial surface glandular pubescent (or the glandular trichomes sometimes absent). Bracteoles lance-subulate, 1.23.2 mm long, 0.3—0.8 mm wide, abaxial surface pubescent like bracts. Flowers sessile. Calyx 4—5-lobed, 3-6 mm long, 4 lobes equal, lanceolate, 2.5—5 mm long, 0.4—0.7 mm wide, abaxially glandular pubescent, posterior 5th lobe (if present) filiform, 0.3—-2 mm long. Corolla pinkish purple with white markings on lower lip, 23-37 mm long, externally glandu- lar pubescent, tube funnelform (gradually expanded distally), 18-25 mm long, 1.8—2.7 mm in diameter near midpoint, upper lip 6.5—12 mm long, entire, lower lip 8-17 mm long, lobes elliptic, 4-9 mm long, 3-6 mm wide. Stamens inserted near apex of corolla tube, 7-11 mm long, filaments glabrous, thecae 1.2-2.2 mm long, subequal, parallel to subparallel, subequally inserted, lacking 766 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 basal appendages, glabrous (or sometimes the upper theca dorsally pubescent with eglandular tri- chomes to 0.1 mm long); pollen (Fig. 20) 2-aperturate, apertures flanked on each side by 2 rows of insulae. Style 19-34 mm long, proximally pubescent with eglandular trichomes, stigma 0.3—0.4 mm long, | lobe 0.3—0.4 mm long, other lobe not evident or up to 0.2 mm long. Capsule 10—13.5 mm long, externally glandular pubescent, stipe 3—5.5 mm long, head ellipsoid with a slight medi- al constriction. Seeds 4, 1.8—2.5 mm long, 1.4—2.3 mm wide, surfaces and margin bubbly tubercu- laten(@2— Til): Phenology. Flowering: September—April; fruiting: November—April. Distribution and habitats. Southwestern United States (Arizona) and northwestern Mexico (Sinaloa, Sonora); Sonoran plants occur along watercourses and on slopes in Sonoran desertscrub (Central Gulf Coast, Plains of Sonora), palm oases, and thornscrub at elevations from near sea level to 1230 m. Illustration. Figure 6; Phytologia 49:67. 1981. Justicia sonorae is known from relatively few collections, but appears to have a broad distri- bution in Sonora (Fig. 28). The species shares several morphological features (e.g., inflorescence form and pubescence, corolla shape and pubescence, pollen, capsule shape and pubescence) with J. hians (Brandegee) Brandegee of Baja California Sur. These species can be distinguished by the following couplet: 1. Single-flowered dichasia alternate along rachis; calyx with 5 conspicuous lobes (posterior lobe only slightly shorter than other four lobes); corolla orange red, upper lip 12.5—16.5 mm long, 2-lobed at apex, lobes of lower lip 1.8—3.5 mm wide; young stems evenly and densely pubes- cent with trichomes 0.05—0.1 (—0.2) mm long; Baja California Sur............... J. hians 1. Single-flowered dichasia mostly opposite along rachis; calyx with 4 conspicuous lobes (posteri- or fifth lobe, if present, inconspicuous and less than 1/2 as long as other 4 lobes); corolla pink- ish purple, upper lip 6.5—11 mm long, entire at apex, lobes of lower lip 4-8 mm wide; young stems glabrous or evenly to bifariously to unifariously pubescent with trichomes 0.2—1 mm longAnizona;, Sonora, and Sinaloa. . jeer > sos). coos ce ee eee J. sonorae Justicia sonorae is also similar to the Oaxacan species J. santelisiana Acosta & T.F. Daniel. The latter species differs by the eglandular (only) trichomes on the external surface of the corollas and capsules, smaller (2—3.7 mm long and 1.8—3.3 mm wide) lobes of the lower lip of the corolla, longer (12—16 mm) stamens, longer (13.5—15 mm) capsules, and longer (2.7—3 mm) seeds. Other plants that resemble J. sonorae are represented by two collections from the vicinity of Bahia Topolobampo in northwestern Sinaloa (e.g., Reina G. et al. 98-2070 at ARIZ, ASU, CAS, MEXU, NY: Gentry 14312 at US). These plants have red corollas and few glands on the inflores- cence rachis, but appear otherwise indistinguishable from pink-purple-flowered individuals of J. sonorae, in which species they are tentatively included as a red-flowered form. Rose et al. 13291 (US) was also collected from the vicinity of Topolobampo. Color of the corolla is neither noted nor preserved on this specimen, but it more closely resembles the majority of collections of J. sonorae by having a conspicuously glandular rachis. SONORAN COLLECTIONS: 0.8 mi W of abandoned mine in pass between Cerro San Luis and Sierra Santa Rosa, 30.1°N, 111.6°W, J. Bowers & R. Turner 2196 (ARIZ); Palm Canyon, 17 mi SE of Magdalena in Cerro Cinta de Plata (Sierra Babiso), MZ. Fay 7/4 (ARIZ); Cerro Tetas de Cabra, Bahia San Carlos, R. Felger 85-366 (ARIZ, MEXU, TEX): Rancho San Jorge, W of Hermosillo, ca. 29°44'N, 111°51'W, R. Felger & A. Russell 6909 (ARIZ); Canon Nacapules, ca. 4 km N of Bahia San Carlos, R. Felger & B. Straub 85-1301 (ARIZ); Bahia San Carlos, H. Gentry 1134] (ARIZ, LL); 2.6 mi E of bridge at Huasabas, L. Landrum et al. 5435 (CAS, GH, MO, NY); Algadones Bay area, 4 mi W of San Carlos Bay, P. Martin s.n. (ARIZ); Palm DANIEL: SONORAN ACANTHACEAE 767 Canyon, 17 mi SE of Magdalena on road to Cucurpe, Sierra Baviso, N. McCarten & R. Bittman 2693 (ARIZ); 7.7 mi S of Cucurpe, L. McGill et al. 6534 (ENCB); Palm Canyon, 15.5 mi SE of Mex. 15 at Magdalena toward Cucurpe, F: Reichenbacher 916 (ARIZ, CAS); N end of Sierra Santa Rosa, 22 mi SSW of Trincheras, ca. 30°06'N, 111°39'W, A. Sanders et al. 3600 (CAS, UC); 10 mi N of San Carlos, D. Shaw 30 (MO); Palm Canyon, 17 mi SE of Magdalena, Cerro Cinta de Plata (Sierra Babiso), D. Steadman & K. Schmidt s.n. (ARIZ); near Palm Canyon, 17 mi E of Magdalena, R. Thompson & O. Davis 82-44 (ARIZ); Mpio. Santa Ana, Rancho E] Otate, Arroyo Los Otates, 30°18'N, 110°53’W, 7: Van Devender & A. Reina G. 98-434 (ARIZ, CAS, MEXU, NY, TEX); 20.4 mi SE of Magdalena on Cucurpe road, 7. Van Devender et al. s.n. (ARIZ); KM 101 on Hermosillo-Moctezuma road, 38.6 mi E of Moctezuma, 29°28'N, 110°15'W, A. Whittemore et al. 83- 099 (TEX). Justicia spicigera Schitdl., Linnaea 7:395. 1832. Jacobinia spicigera (Schltdt.) L.H. Bailey, Standard Cyclopedia of Horticulture, 1715. 1915. TYPE: MEXICO. Veracruz: Jalapa, May 1829, A. Schiede s.n. (syntype: B, destroyed; isosyntypes: BR!, GH; probable isosyntypes: E!, P!); Veracruz: cultivated at Hacienda de la Laguna, Jul 1829, 4. Schiede s.n. (syntypes: B, destroyed, P!; isosyntype: BM!). See Daniel (1995a) for a complete list of synonyms. Phenology. Flowering: March, July; fruiting: unknown. Distribution and habitats. Mexico (Baja California Sur, Chiapas, Guerrero, Hidalgo, Jalisco, Mexico, Michoacan, Nayarit, Oaxaca, Puebla, Querétaro, Veracruz, Yucatan), Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica; Sonoran plants are cultivated (and possibly persist- ing after or escaping from cultivation) at elevations from 330 to 700 m. Illustrations. Fieldiana, Botany 24(10):401. 1974; Flora of Baja California, 193. 1980; Flora of the Bahama Archipelago, 1352. 1982; Mexican Roadside Flora, 25. 1987. Local name. “Niple” (Joyal 2059). Use. Leaves are boiled with cinnamon to yield a clear, red tea that is taken for stomach pain (Joyal 2059). Daniel (1995a) and Daniel and Acosta (2003) provided descriptions of this species. Rose et al. 13020 was noted to have been “apparently” cultivated and Joyal 2059 was noted to be planted in a “house garden.” Justicia spicigera is widely cultivated throughout Mexico such that the actual native distributional range of the species is not known with certainty. It is undoubtedly consider- ably smaller than suggested by the states listed above. SONORAN COLLECTIONS: Mpio. Onavas, Rancho El] Palmar, 20 km E of Onavas, 28°29-30'N, 109°23'W, E. Joyal 2059 (CAS); vicinity of Alamos, J. Rose et al. 13020 (US). Odontonema Nees Odontonema Nees, Linnaea 16:300. 1842, nom. cons. TYPE: garden specimen without date or collector (GZU), type cons. Thyrsacanthus Nees in Martius, Fl. Bras. 9:97. 1847, nomen superfl. (= Odontonema). Diateinacanthus Lindau, Bull. Herb. Boissier, ser. 2, 5:369. 1905. TYPE: Diateinacanthus hondurensis Lindau (= Odontonema hondurense (Lindau) D.N. Gibson). Erect perennial herbs or shrubs (sometimes clambering) with cystoliths. Leaves opposite. Inflorescence of mostly terminal and sometimes basally branched dichasiate spikes, racemes, or thyrses; dichasia alternate to opposite to whorled, 1-many (i.e., 18 or more)-flowered, variously contracted or expanded, sessile or pedunculate, subtended by a bract. Flowers homostylous or het- erostylous, usually pedicellate, subtended by 2 homomorphic bracteoles. Calyx deeply 5-lobed, lobes equal in length. Corolla infundibular, white, yellow, purple, pink, or red, tube usually expand- 768 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 ed distally into a prominent throat, limb subactinomorphic to 2-labiate, upper lip 2-lobed, lower lip 3-lobed, corolla lobes imbricate in bud. Stamens 2, thrum stamens exserted from mouth of corol- la, pin stamens usually included in corolla tube, anthers 2-thecous, thecae equal in size, parallel, equally inserted on filament, lacking basal appendages, dehiscing toward lower lip (i.e., flower nototribal); pollen oblate spheroidal to spherical to euprolate, 3(—4)-colporate, 6(—8)-pseudocol- pate, pseudocolpi 2 per mesocolpium, exine foveolate to rugulate to reticulate to bireticulate; sta- minodes 2. Thrum style included in corolla tube, pin style exserted from mouth of corolla, stigma + funnelform or 2-lobed, lobes equal or unequal in length. Capsule stipitate, head obovoid to subel- lipsoid (often with a medial constriction), retinacula present, septa with attached retinacula remain- ing attached to inner wall of mature capsule. Seeds 4, homomorphic, lenticular, lacking trichomes. (x = 21). This genus of 29 species is restricted to the New World tropics and subtropics with eight species in Mexico (Daniel 1995d). Species with tubular, red corollas (like ours) are visited, and appear to be well adapted for pollination, by hummingbirds (see Daniel 1995d). Odontonema cuspidatum (Nees) Kuntze, Revis. Gen. Pl. 2:494. 1891. Thyrsacanthus cuspi- datus Nees in A. de Candolle, Prodr. 11:323. 1847. TYPE: MEXICO. Oaxaca: Sierra S. Pedro Nolasco, Talea, etc., 1843-1844, C. Jiirgensen 985 (lectotype designated by Daniel, 1995a: K!; isolectotype: BM!). Phenology. Flowering: November; fruiting: unknown. Distribution and habitats. Mexico (Chiapas, Guanajuato, Hidalgo, Morelos, Oaxaca, Querétaro, San Luis Potosi, Tabasco, Veracruz), Honduras, West Indies; Sonoran plants are culti- vated and likely escaped (see below). Illustration. Contributions from the University of Michigan Herbarium 20:157. 1995. Daniel (1995a, 1995d) provided descriptions of Odontonema cuspidatum. This species of shrubs with red flowers is often cultivated for ornament. The sole Sonoran specimen from a natu- ral habitat is rather fragmentary, but it does permit a tentative species identification. On the basis of the color of the corolla noted on the label (red) and the pubescence of the inflorescence rachis (mostly erect and evenly disposed eglandular trichomes 0.05—0.2 mm long) this collection resem- bles O. cuspidatum, a species of eastern and southern Mexico (Daniel 1995d). It differs from that species, and resembles O. glaberrimum (M.E. Jones) V.M. Baum, by its mostly sessile to subses- sile dichasia (although some dichasia are borne on peduncles 0.5-1 mm long). Odontonema glaberrimum is known from northern Sinaloa, but differs from Jenkins & Yetman 91-115 by its rachises that vary from glabrous to pubescent with flexuose to antrorse eglandular trichomes 0.05- 0.5 mm long. The number of apertures on pollen grains can be used to distinguish O. cuspidatum (pollen 4-aperturate; Fig. 18) from other Mexican species of the genus (all of which have 3-aper- turate pollen; see palynological information in Daniel 1995d). Examination of pollen from Jenkins & Yetman 9]-115 (Fig. 18) reveals somewhat aberrant grains with four or more apertures. I suspect that Jenkins & Yetman 91-115 represents an escaped occurrence of O. cuspidatum (with somewhat unusual sessile to subsessile dichasia and slightly deformed pollen), which is known to be cultivat- ed for ornament in Sonora. Although Jenkins & Yetman 91-115 was not collected near human dwellings, Jenkins (pers. comm.) indicates that there is a small settlement upstream from which propagules of this species could have washed downstream. SONORAN COLLECTIONS: cultivated grounds of La Ciudadela just off Plaza de Armas in Alamos, ca. lat. 27°01'N, 108°56'W, T. Daniel 9774 (CAS); near jct. of Arroyo Santa Barbara with Arroyo Verde, 27°05’N, 108°43'W, P. Jenkins & D. Yetman 91-115 (ARIZ, CAS). DANIEL: SONORAN ACANTHACEAE 769 Pseuderanthemum Radlk. Pseuderanthemum Radlk., Sitzungsber. Math.-Phys. KI. Akad. Wiss. Miinchen 13:282. 1883. LECTOTYPE (Leonard, Contr. U.S. Natl. Herb. 31(2):292. 1953; see J. Adelaide Bot. Gard. 9:141. 1986): Pseuderanthemum alatum (Nees) Radlk. (= Eranthemum alatum Nees). Siphoneranthemum (Oerst.) Kuntze, Revis. Gen. Pl. 2:494. 1891. Eranthemum L. subg. Siphoneranthemum Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854:166. 1855. TYPE: Siphoneranthemum fasciculatum (Oerst.) Kuntze (= Eranthemum fasciculatum Oerst.). Buceragenia Greenm., Proc. Amer. Acad. Arts 32:303. 1897. TYPE: Buceragenia minutiflora Greenm. Decumbent to erect herbs or shrubs with cystoliths. Leaves opposite. Inflorescence of dicha- sia in leaf axils or of axillary or terminal dichasiate spikes, racemes, thyrses, or panicles; dichasia opposite or alternate, 1—-many-flowered, sessile or pedunculate, subtended by a leaf or bract. Flowers chasmogamous and/or cleistogamous, homostylous or heterostylous, sessile or pedicellate, subtended by 2 homomorphic bracteoles. Calyx deeply 5-lobed, lobes homomorphic. Corolla white, pink, blue, purplish, or red, the lower lip sometimes with colored markings, + salverform (i.e., with a prominent slender tube and a rotate limb), tube cylindric, sometimes + expanded dis- tally into a short throat, limb subactinomorphic to bilabiate, upper lip 2-lobed, lower lip 3-lobed, corolla lobes imbricate in bud, corollas of cleistogamous flowers remaining budlike. Stamens 2, included in corolla tube or exserted from mouth of corolla, anthers 2-thecous, thecae equal to sube- qual in size (or in cleistogamous flowers becoming unequal or with | theca suppressed), parallel, subequally inserted, lacking basal appendages, dehiscing introrsely toward lateral lobes of corolla (1.e., flower pleurotribal); pollen (Fig. 18) oblate spheroidal to euprolate, 3-colporate (sometimes syncolpate), 6-pseudocolpate, pseudocolpi 2 per mesocolpium, the 2 sometimes fused near poles into pseudocolpal ellipses, exine (rugulate to) reticulate; staminodes 0 or 2. Style included 1n corol- la throat or exserted from mouth of corolla, stigma 2-lobed, lobes equal. Capsule stipitate, head ellipsoid with a medial constriction, retinacula present, septa with attached retinacula remaining attached to inner wall of mature capsule. Seeds 4, homomorphic, plano-convex to lenticular, lack- ing trichomes. (x = 21) This genus of 40-60 species occurring in the tropics worldwide is in need of considerable study. Its relationships to Odontonema, Chileranthemum, Pulchranthus and Oplonia are especial- ly problematic in the New World. Cleistogamy and heterostyly are both present in Pseuderanthemum and have added to taxonomic confusion in the genus. Species previously treat- ed in Buceragenia represent plants of Pseuderanthemum with cleistogamous flowers (Daniel 1995b). There are 10-15 species in Mexico. Sonoran plants of Pseuderanthemum are indicative of the problems involved in delimiting and identifying Mexican species of the genus. Pseuderanthemum praecox (Benth.) Leonard, J. Wash. Acad. Sci. 31:99. 1941. Eranthemum praecox Benth., Pl. Hartweg. 291. 1848-1849. Siphoneranthemum praecox (Benth.) Kuntze, Revis. Gen. Pl. 2:497. 1891. TYPE: MEXICO. Guanajuato: Santa Rosa near Guanajuato, 1837, 7: Hartweg 1614 (holotype: K!). Perennial herbs to 5.5 dm tall. Young stems subquadrate to + compressed, sometimes + mul- tistriate, glabrous or + evenly (to + bifariously) pubescent with retrorse to flexuose eglandular tri- chomes 0.05—0.2 (0.3) mm long. Leaves generally absent when chasmogamous flowers present in spring, present along with cleistogamous flowers in late summer, membranaceous, sessile to 770 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 petiolate, petioles to 3 (-12) mm long, blades either linear to lanceolate, 66-200 mm long, 4-15 mm wide, 7.7—23 times longer than wide, acute to attenuate at base, attenuate at apex, or blades narrowly elliptic to elliptic to lance-ovate to ovate to obovate, 37-100 mm long, (7—) 16-37 mm wide, mostly 1.7—-4.7 (—5.7) times longer than wide, acute to attenuate at base, attenuate to acute to rounded at apex, surfaces of all blades glabrous or pubescent (especially along veins) with flexu- ose to retrorse to retrorsely appressed eglandular trichomes 0.05—0.2 mm long, margin flat, entire to subsinuate. Inflorescence of + congested dichasia in axils of leaves and sometimes also in axils of bracts distally, the latter forming a terminal spikelike raceme or thyrse, rachis (if present) glabrous or pubescent like young stems; dichasia opposite or alternate, sessile to pedunculate, 1-4- flowered, peduncles to 3 (-50) mm long. Bracts (if present) subulate to linear, 2-14 (43) mm long, 1—1.2 (-2) mm wide, abaxial surface glabrous or pubescent with flexuose to antrorse to retrorse eglandular trichomes 0.05—0.1 mm long. Bracteoles and secondary bracteoles usually triangular to subulate, 1.2-S mm long, 0.7—1.2 mm wide (sometimes foliose, lanceolate to linear to elliptic to ovate, to 36 mm long and 17 mm wide), abaxial surface glabrous or pubescent with flexuose to antrorse to retrorse eglandular trichomes 0.05—0.1 mm long. Flowers subsessile to pedicellate, pedicels 0.05—3.5 (—5) mm long. Calyx 3.5—7 (—9.3 in fruit) mm long, lobes subulate, 2.5—5.5 (—8.5) mm long, 2.5—4.5 (—5.5) times longer than tube, abaxially nearly glabrous or pubescent like bracteoles (occasionally with a few sessile or stipitate inconspicuous glands to 0.1 mm long as well), or pubescent with erect to flexuose eglandular trichomes and conspicuous glandular tri- chomes 0.05—0.2 mm long. Corolla of chasmogamous flowers salverform, pink to light purplish with white area on lower lip near mouth, 20-35 mm long, externally sparsely pubescent near apex of lobes with flexuose eglandular trichomes, otherwise glabrous, tube subcylindric, 19-21 mm long, 0.9—1.1 mm in diameter near midpoint, expanded slightly above middle into a + inconspicu- ous throat, limb subactinomorphic (lobes similar, elliptic to ovate-elliptic), 17-24 mm in diameter, upper lip 7-13 mm long, lobes 7-13 mm long, 3.5—5.7 mm wide, lower lip 10—14 mm long, lobes 8.7—13 mm long, (4—) 4.7—7 mm wide; corolla of cleistogamous flowers budlike, white with pur- ple at apex, 2—2.5 mm long, apically pubescent with flexuose eglandular trichomes 0.05—0.1 mm long. Stamens of chasmogamous flowers included, 3.5—4 mm long, thecae 2—2.5 mm long; stamin- odes up to 0.6 mm long. Style of chasmogamous flowers included (or with stigma barely exserted from mouth), 17-18 mm long, glabrous, stigma lobes 0.2—0.3 mm long, equal; style of cleistoga- mous flowers 1—1.2 mm long, glabrous, curved to recurved apically, stigma subequally 2-lobed, 0.2 mm long. Capsule 13—18 mm long, glabrous, stipe 5—7 mm long, head 8—11 mm long. Seeds plano- convex to concavo-convex, 2.9-4.5 mm long, 2.5—3.6 mm wide, surfaces irregularly reticulate- ridged, ridges sometimes reduced to knoblike projections on mature seeds. Phenology. Flowering March—May (chasmogamous flowers), August-September (cleistoga- mous flowers); fruiting: March—May, August—October. Distribution and habitats. Mexico (Chiapas, Distrito Federal, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, México, Michoacan, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, Sinaloa, Sonora, Tamaulipas, Zacatecas), Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica; in Sonora plants occur in along watercourses and on rocky slopes in oak woodlands, oak forests, oak- pine woodlands, and oak-pine forests at elevations from 1100-1580 m. Illustrations. Figures 15, 29; Flora de Valle de México, fig. 297B. 1979; Flora del Valle de Tehuacan-Cuicatlan 23:66. 1999; Flora del Bajio 117:118. 2003. Descriptions of this species from other regions of Mexico have been provided by Daniel (1995a, 1999a) and Daniel and Acosta (2003). A description based exclusively on Sonoran plants is presented here because they exhibit morphological variation not observed in these other regions. All Sonoran plants of Psewderanthemum are interpreted as pertaining to a single species, P. prae- | DANIEL: SONORAN ACANTHACEAE gel FIGURE 29. Pseuderanthemum praecox. a. Habit (Reina G. & Van Devender 99-529), x 0.58. b. Leaves, x 0.58: nar- row leaf on right (Reina G & Van Devender 98-1700), wider leaf on left (Reina G& Van Devender 99-527). c. Bracteole from lower node (Reina G. & Van Devender 99-666), x 7.5. d. Bracteole from upper node (Reina G. & Van Devender 99- 666), x 10.4. e. Capsule (Reina G & Van Devender 99-666), x 4.1. f. Seed (Reina G. & Van Devender 99-666), x 8.1. Drawn by Nadia Strasser. cox. It is evident from the above description that P. praecox in Sonora exhibits considerable mor- phological diversity. This variation appears to be attributable to seasonality, cleistogamy, and mor- phological plasticity. Generally leafless plants with chasmogamous flowers have been collected in the spring (March—May) whereas leafy plants with only cleistogamous flowers have been collected only dur- ing the late summer (August-September). Based on the many collections of plants from riparian communities in a region of oak-pine woodland and forest (between Yécora and the Chihuahua bor- der) and the summary of climatic data for the region by Van Devender et al. (2003), the life cycle VI. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 of P. praecox there can by hypothesized as follows: new, leafy shoots appear from the underground caudex in the late summer in response to the summer (June—October) rains; these shoots bear cleis- togamous flowers that self-pollinate and yield fruits that mature prior to winter; the leaves are deciduous and fall sometime during the winter months when extended periods of freezing temper- atures are common; following the winter rains (November—February) and the end of freezing tem- peratures in the spring, the old (and now leafless) shoots bear new growth from the axillary buds that bears chasmogamous flowers; these flowers are potentially available for cross-pollination, but likely self-pollinate as the corolla falls from the flower 1f pollination has not been otherwise effect- ed; fruits mature prior to the death (and collapse) of these shoots during the dry period of late spring and summer. The two seasonal phases of Sonoran plants of PR. praecox can be distinguished by the following couplet: 1. Chasmogamous flowers present, corollas salverform 20-35 mm long; leaves mostly or entirely absent; calyx pubescent with eglandular and conspicuous glandular trichomes. . spring phase 1. Chasmogamous flowers absent, only cleistogamous flowers present, corollas budlike, 2—2.5 mm long; leaves present; calyx nearly glabrous or pubescent with eglandular (and often incon- Spicuoussslandular) trichomes ae tsa aiks aes oie eee eee late summer phase I have not observed the late summer phase in other portions of the range of P. praecox in Mexico (for example, 41 of the 42 Mexican (non-Sonoran) specimens of P. praecox at MEXU are entirely or nearly leafless, have chasmogamous flowers, and were collected between December and June-but mostly in March and April; the other collection was leafless, but lacked corollas), but it may be represented by one or more species previously treated under Buceragenia (see Daniel 1995b). Sonoran plants of Pseuderanthemum praecox are unusual in having two foliar forms (although with some intermediacy between them) and two pubescence forms, all growing together. Most col- lections have the leaf blades either very narrow, linear, and 7.7—23 times longer than wide (e.g., Gentry et al. 19402; Reina G. & Van Devender 98-1700, 99-666, 2000-629; Reina G. et al. 2000- 379) or wider, elliptic to obovate, and 1.3—5.7 times longer than wide (e.g., Reina G & Van Devender 98-1863, 99-529, 2000-623; Reina G. et al. 2000-380; Van Devender & Reina G 98- 1832). Plants with both types of leaf blades or with the blades somewhat intermediate between the two forms occur in a population west of Maycoba. For example, Reina G. & Van Devender 99-525 has both linear (22.5 times longer than wide) and lance-ovate (4.9 times longer than wide) leaf blades. Reina G. & Van Devender 99-527 has leaves varying from 3.3—6.1 times longer than wide. It is not known whether plants with linear leaves represent a riverine morphological form, with the broader leaved plants occurring on somewhat drier sites. Plants with narrow, linear leaves appear to have been first collected by Howard Gentry in October 1961 from the Sierra Tecurahui in south- eastern Sonora (east of Alamos, near the Chihuahua border) at an elevation between 1200 and 1360 meters. In their explorations in the Municipio de Yécora, Ana Lilia Reina G. and Tom Van Devender located similar plants in a tributary of Arroyo Pilares (west of Maycoba) at 1300 meters elevation, and recollected it there many times in efforts to document the foliar variation and flower type. Some plants from both of these populations have stems, leaves, and bracteoles glabrous (e.g., Gentry et al. 19402 at ARIZ, Reina G et al. 99-525 at CAS), whereas other plants at these sites have these organs conspicuously pubescent (e.g., Gentry et al. 19420 at US, Reina G et al. 99-525 at CAS). The structure of the inflorescence is somewhat variable in P. praecox. In most individuals, the reduced dichasia form a dense cluster of flowers in the axils of the leaves. Sometimes the leaves are replaced by bracts distally, resulting in a terminal spikelike raceme to thyrse. The transition DANIEL: SONORAN ACANTHACEAE 73 from leaves to bracts is either abrupt or gradual. On some individuals, especially at the proximal {| nodes of plants with the broader leaf blades, dichasia are borne on elongate peduncles (to 50 mm long) and bear foliose bracteoles. It is [3! sometimes difficult to interpret whether such lateral growth consists of long-pedunculate dichasia (that sometimes continue to elongate sympodially) or lateral branches bearing lateral dichasia. Both appear to be present. 29° Pseuderanthemum praecox reaches the northwestern extent of its range in southeastern Sonora (Fig. 30), from the vicinity of Yécora southward to the vicinity of Alamos. It © Pestidereuterinmn praccon undoubtedly occurs in nearby regions of [7° © Ruellia leucantha Chihuahua, but it has not yet been collected in that state. The entirety of variation described 14° 112° above for this species in Sonora has been doc- SESE : 3 : FIGURE 30. Sonoran distributions of Pseuderanthemum umented at only a single site (1.e., 24.7 km W of praecovandl Rueliaieucante Maycoba). The species is also somewhat diffi- cult to circumscribe in other portions of its range. I have taken various approaches to treating Pseuderanthemum praecox and its relatives (e.g., P. fasciculatum (Oerst.) Leonard and P. hispidu- lum (Nees) Radlk.) in Chiapas (Daniel 1995a), south-central Mexico (Daniel and Acosta 2003), El Salvador (Daniel 2001), and Honduras (Daniel 2004a); none of them seems very satisfactory. SONORAN COLLECTIONS: Mpio. Yécora, Ciénega de Camilo, ca. 14 km E of Maycoba near Hwy. 16, ca. 28°26'N, 108°34'W, 7: Daniel et al. 5617 (CAS, K, MEXU); Rcho. Agrimincor, Rio Mayo, H. Gentry 304] (ARIZ); Sierra Tecurahui, H. Gentry et al. 19420 (ARIZ, US); Mpio. Alamos, Santa Barbara, 27°07'N, 108°43'W, P. Martin s.n. (ARIZ); Mpio. Yécora, tributary of Arroyo Los Pilares near bridge, 24.7 km W of Maycoba on Mex. 16, 28°24'N, 108°48'W, A. Reina G & T. Van Devender 98-1700 (ARIZ, CAS), 98-1863 (CAS), 99-525 (ARIZ, CAS), 99-527 (CAS), 99-529 (CAS), 99-666 (CAS), 2000-623 (CAS), 2000-629 (CAS), A. Reina G. et al. 2000-379 (CAS), 2000-380 (CAS); Mpio. Yécora, Arroyo Los Pilares, 21 km E of Yécora, 28°24’N, 108°47’'W, A. Reina G. et al. 96-59 (ARIZ, MEXU); Mpio. Alamos, Sierra de Alamos, ca. 4 km SW of Alamos at La Huerta, ca. 26°59'N, 108°58’W, V. Steinmann 94-35 (ARIZ, CAS); Mpio. Yécora, Rio Maycoba at Mex. 16 (20.5 km W of Maycoba, 28.6 km E of Yécora), 28°22.5'N, 108°45'W, 7. Van Devender & A. Reina G. 95-458 (ARIZ); Mpio. Yécora, Ciénega de Camilo, 6.3 km E of El Kipor, 11.2 km W of Chihuahua border, 28°25.5'N, 108°34’W, T. Van Devender & A. Reina G. 97-402 (ARIZ, CAS), 98-1832 (CAS). Ruellia L. Ruellia L., Sp. Pl. 634. 1753. LECTOTYPE (Britton and Brown, Ill. Fl. N. U.S., ed. 2, 3:241. 1913): Ruellia tuberosa L. Dipteracanthus Nees in Wallich, Pl. Asiat. Rar. 3:75, 81. 1832. LECTOTYPE (Bremekamp and Nannenga Bremekamp, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 45(1): 15. 1948): Dipteracanthus prostratus (Poiret) Nees (= Ruellia prostrata Poiret). Aphragmia Nees in Lindley, Intr. Nat. Syst. Bot., ed. 2, 444. 1836. TYPE: Aphragmia haenkei Nees (= Ruellia inundata H.B.K.). Decumbent to erect subcaulescent to caulescent perennial herbs, shrubs, or (rarely) trees with 774 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 cystoliths, vegetative and floral surfaces sometimes beset with sessile lenticular glands mostly 0.05—0.2 mm in diameter (glandular punctate). Leaves opposite. Inflorescence of reduced or expanded dichasia in axils of leaves or bracts, sometimes forming dichasiate spikes, thyrses, or panicles; dichasia alternate or opposite, |-many-flowered, 1 or more per axil, sessile or peduncu- late, subtended by a leaf or a bract. Flowers chasmogamous and/or cleistogamous, homostylous, usually subtended by 2 homomorphic bracteoles (sometimes absent or vestigial). Calyx deeply 5- lobed, lobes homomorphic or heteromorphic. Corolla of chasmogamous flowers blue to blue-pur- ple (in ours, elsewhere also pink, red, white, yellow, or blackish), generally concolorous, tube usu- ally funnelform, often curved, gradually or abruptly expanded distally into a + distinct throat (rarely saccate), limb 2-labiate (often appearing subactinomorphic), upper lip 2-lobed, lower lip 3- lobed, corolla lobes contorted in bud, corolla of cleistogamous flowers (if present) small and bud- like or tubular, often whitish. Stamens 4, (homodynamous to) didynamous, included in corolla tube or exserted from mouth of corolla, anthers 2-thecous, thecae equal in length, parallel to subsagit- tate, equally inserted, lacking basal appendages, dehiscing toward lower lip (.e., flower nototrib- al); pollen (Fig. 31) spherical to subspheroidal, 3-porate, exine coarsely reticulate; staminodes 0 or 1. Style included in corolla tube or exserted from mouth of corolla, stigma 2-lobed, lobes equal or unequal, often with | lobe greatly reduced, rudimentary, or not evident. Capsule substipitate or stip- itate, retinacula present, septa with attached retinacula usually remaining attached to inner wall of mature capsule (separating from inner wall in R. ervthropus). Seeds 4-28 per capsule, lenticular, pubescent with hygroscopic trichomes (sometimes restricted to margin). (x = 17). Ruellia comprises approximately 250 species worldwide. Species occur primarily in the trop- ics and subtropics. It is the second largest genus of Acanthaceae in Mexico with about 65 species in the country. The generic synonymy given above only includes genera in which Sonoran species have been treated previously. Ezcurra (1993) provided a more complete list of generic synonyms in which American species have been treated. 1. Corolla white; young stems, leaves, bracteoles, calyx, corolla, and capsule with conspicuous sessile patellitorm slands (glandular punctate); <2. .\-. =. co ee R. leucantha 1. Corolla blue; young stems, leaves, bracteoles, calyx, corolla, and capsule lacking sessile patel- liform glands (stipitate glands sometimes present, however). 2. Leaves with at least some trichomes branched or stellate; plants of northeastern Sonora in Chihuahuan) desertscrub:,mi2rsncrolts hs os Ae Ws te a oe Ae eee R. parryi 2. Leaves lacking any branched or stellate trichomes; plants not restricted to Chihuahuan desertscrub in northeastern Sonora. 3. Limb of corolla 10-15 mm in diameter; calyx lobes linear to oblanceolate, unequal with one conspicuously longer and wider than others; thecae 1.7—2.4 mm long; capsule often spotted withired,,7-S—10 mm long; elabrous: 0-344. @ aa eee R. inundata 3. Limb of corolla 17-46 mm in diameter; calyx lobes subulate to lance-subulate to linear, equal to subequal; thecae 2.8—-5 mm long; capsule never spotted with red, 11-29 mm long, pubescent (sometimes only at apex and inconspicuously so). 4. Plants woody throughout (shrubs); stipe of capsule 4.5—7.5 mm long; seeds 4 per cap- Sule sors cic dale: cuentas ot cick e = ecb? weds 2, ee a ae R. californica 4. Plants woody only at or near base (perennial herbs); stipe of capsule 2—4 mm long; seeds 8—20 per capsule. 5. Those dichasia in axils of distal leaves or leaflike bracts collectively forming a ter- minal glandular-pubescent paniculiform thyrse; peduncles of distal dichasia, abaxi- al surface of bracteoles, pedicels, and capsules pubescent with conspicuous glandu- DANIEL: SONORAN ACANTHACEAE VIS FIGURE 31. Pollen of Sonoran Ruwellia. a. R. californica (Wiggins 11411). b. R. intermedia (Jenkins et al. s.n.). c. R. inundata (Daniel & Butterwick 6952). d. R. leucantha (Brandegee s.n.). e. R. malacosperma (Daniel 9773). £. R. nudiflo- ra (Wiggins & Rollins 136). g. R. parryi (Reina G. et al. 2003-957). Scales = 10 um. Nn lar trichomes, at least some of which are longer than 0.2 mm; capsule pubescent proximal yic wipes oe Sah oe in) SE ee ee R. nudiflora . Dichasia pedunculate in axils of leaves, never forming a terminal glandular paniculi- form thyrse; peduncles of dichasia, abaxial surface of bracteoles, pedicels, and cap- sules glabrous or inconspicuously glandular with glandular trichomes up to 0.2 mm long; capsule glabrous proximally. 6. Leaves 1.1—1.9 times longer than wide, corolla externally pubescent with eglan- dular trichomes only; style essentially glabrous (sometimes with 1—3 scattered tri- chomes present); capsule 12-23 mm long, stipe 2—2.5 mm long... R. intermedia 6. Leaves 2.3—5.8 times longer than wide; corolla externally pubescent with glandu- lar and eglandular trichomes; style proximally pubescent with eglandular tri- chomes; capsule 24-29 mm long; stipe 4 mm long.......... R. malacosperma 776 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Ruellia californica (Rose) 1.M. Johnst., Proc. Calif. Acad. Sci., ser. 4, 12:1171. 1924. Calophanes californica Rose, Contr. U.S. Natl. Herb. 1:85. 1890. TYPE: MEXICO. Baja California Sur: Santa Rosalia, 24 Feb-3 Mar 1889, E. Palmer 190 (holotype: US; isotypes: F!, GH!, NY, UC). Phenology. Flowering: throughout the year, with a peak in March (Fig. 1); fruiting: through- out the year. Distribution and habitats. Northwestern Mexico (Baja California, Baja California Sur, Sonora); Sonoran plants occur on slopes and flats in Sonoran desertscrub (Central Gulf Coast, Plains of Sonora), palm oases, and thornscrub at elevations from sea level to 300 m. Illustrations. Figure 9; Flora of Baja California, 194. 1980; People of the Desert and Sea, 219. 1985; Proceedings of the California Academy of Sciences 49:382. 1997. Local names. “Hupa chumi” (Yaqui, fide Felger 1999); “rama parda” (fide Felger 1999; Felger and Moser 1985); “‘satooml” (Seri, fide Felger and Moser 1985); “stoms” (Seri, Whiting 9067). Uses. A poultice made from the plant is used as a remedy for headache (Whiting 9067). Felger and Moser (1985) noted a diversity of uses by the Seri, including: food (nectar sucked from the base of the corolla), medicine (tea made from the leaves ingested for dizziness or headache, put on the face of one who is “tired out” or for dizziness, applied to the eyes as eye drops, and used as a shampoo to relieve headache; tea made from boiling the root (after removal of the epidermis) ingested as a remedy for colds or stuffy nose), and smoking (leaves or corollas dried and smoked in clay pipe to induce hallucinations). Daniel (1997) provided a description of | T es. | Ruellia californica, which is endemic to north- | / SS | western Mexico and attains the northern and SS —— eastern extents of its distribution in central and es southern Sonora, respectively (Fig. 32). Within -31° the Sonoran Desert of Sonora, the species is | especially common in the Central Gulf Coast subdivision near Guaymas. It has been less fre- | L 7 quently collected in the Plains of Sonora subdi- | ; vision. A collection from near Magdalena, Loe r} about 150 kilometers north of its occurrences in | that subdivision, appears to be from the | Sexe Arizona Upland subdivision. The species is > * sometimes cultivated, and until R. californica © (Ricliiacalifornica , && at is recollected in the Arizona Upland subdivi- » © Ruellia inundata Ree als * Ruellia parryi \ O Zz @ O64 sion, its native occurrence there is treated as SUSDECE- ; : 114 112° 110° Plants from Sonora are all referable to sub- = aa aaa : ay é ate . SURE 32. S ¢ istributi : j- species californica as delimited by Daniel Fic URE 32. Sonoran distributions of Ruellia californi ae ae ; ca, R. inundata, and R. parryi. (1997). Coville 1672 from near Guaymas great- ly resembles subsp. peninsularis (central and southern Baja California Sur) by the length of its corolla (32 mm), length of its calyx (7 mm), and its glutinous herbage. In this specimen (and similar specimens from this region) there are sparse stipitate glands on the stems and leaves and conspicuous glands on the calyx. These latter charac- teristics confirm the placement of these plants into subspecies californica. An apparently rare occurrence of white corollas among Sonoran plants of this species has been noted (e.g., Phillips et DANIEL: SONORAN ACANTHACEAE 777 al. 76-183). The herbage is often somewhat viscid and typically mephitic (somewhat skunklike) in odor. SONORAN COLLECTIONS: Isla Tiburon, J. Alcocer F: & C. Sosa R. 10 (ENCB); San Carlos Bay, R. Barr 61-107 (ARIZ); San Francisco Bay near Guaymas, R. Barr 66-3 (ARIZ); Cerro Bocochibampo, Guaymas, E. Blakley B-1640 (DS); Guaymas, T. Brandegee s.n. (UC); Mpio. Guaymas, 6.5 mi N of road to Bahia San Carlos, ca. 10 mi N of Guaymas on Mex. 15, D. Breedlove 1464 (DS); Mpio. Hermosillo, 7 mi W of Hwy. 15 along southern paved road to Bahia Kino, D. Breedlove 15950 (DS, ENCB); Isla Melliza Oeste, near SW end of Guaymas Bay, 27°54'N, 110°54'W, 7; Burgess et al. 6903 (ARIZ); N of Estero El Rancho, 3.9 km N and 1.4 km W of E end of Empalme causeway, 27°59'N, 110°50’W, 7: Burgess et al. 9885 (ARIZ); Sierra Libre, Cafiada Tetabejo, S de Hermosillo por la Carr. Fed. 15, 28°33'N, 110°58’W, A. Burquez 95-128 (MEXU); La Pintada, 49 km S de Hermosillo, por Carr. Fed. 15, 28°35’N, 110°58'W, A. Burquez & D. Yetman 97-431 (MEXU); near San Juan de Guaymas, 4. Carter & D. Noack 4362 (UC); 5 mi S of Guaymas, G Chan & J. Folkner s.n. (ARIZ, MEXU); S Merimar, O. Clark 14978 (UNM); Guaymas, F Coville 1672 (US); ca. 4 mi N of San Carlos at movie site “Catch-22,” 7. Daniel 120 (CAS); “Microondas mountain” overlooking Guaymas, 7: Daniel 1963 (ASU, CAS); Nacapuli Canyon, 6.3 mi W of Hwy. 15 toward San Carlos then 4.4 mi N, 7. Daniel 2548 (CAS); W base of Sierra Bojihuacame E of Cd. Obregon, 1.2—2.2 mi SE of Bachoco, 7: Daniel 3366 (CAS, MEXU, NY); ca. 2 mi E of San Carlos, NW of Guaymas, 7. Daniel 3978 (CAS); between San Carlos Bay and Catch-22 airstrip NW of Guaymas, 7: Daniel 3987 (CAS); Guaymas, E. Dawson 1002 (DS); Isla Tiburon, R. Delgado B. s.n. (ENCB); Isla Tiburon C. Diaz L. 2247 (ENCB); Guaymas, F: Drouet & D. Richards 3816 (DS, F, US); Isla Tibur6n, Ensenada de la Cruz, S end of island, R. Felger 2589 (ARIZ); Isla Tiburon, SE base of Sierra Kunkaak, R. Fe/ger 9328 (ARIZ, CAS); Las Guasimas, R. Felger & R. Devine 85-277 (ARIZ, MEXU); E end, Bahia San Carlos, R. Felger & C. Lowe 2471 (ARIZ); Isla Tiburon, vic. of Palo Fierro landing field, Punta Tortuga, R. Felger & A. Russell 8933 (ARIZ, CAS), Felger et al. 11075 (CAS); road to Bahia San Carlos, 4 mi W of Hwy. 15, R. Felger & R. Thomas 11917 (ARIZ); ca. 0.5 mi SE of mouth of Nacapule Canyon, ca. 4 mi NE of Bahia San Carlos, R. Felger & R. Thomas 11954 (ARIZ); Sierra El] Aguaje, ca. 18 km NW of San Carlos, ca. 28°03’N, 111°13'W, R. Felger & M. Wilson 95-238 (ARIZ, CAS); E end of Bahia San Carlos, R. Felger et al. 10308 (ARIZ); Isla Tibur6n, Tecomate, NW part of island, R. Felger et al. 11126 (ARIZ); Ensenada Grande (= Bahia San Pedro), R. Felger et al. 12099 (ARIZ); San Carlos Bay, ca. 8 mi N of Guaymas, R. Ferris 8721 (DS, NY, US); Bahia de Guasimas, just W of Cerro El Bachoco, 15 km E of Empalme, 4.6 mi S of Hwy. 15, 27°54'N, 110°40'W, M. Fishbein et al. 921 (ARIZ, CAS); Mpio. Guaymas, Sierra E] Aguaje, Aguaje Robinson, ca. 10 km NW de San Carlos, 28°03.5'N, 111°07'W, A. Flores M. & O. Gutierrez R. 5090 (ARIZ, MEXU); N of Magdalena toward Cucurpe, ca. 0.6 mi W of fork, J. Fryxell 78024A (TEX); Ciudad Obregon, H. Gentry 276 (DS, US); Mutica, Rio Yaqui, H. Gentry 2193 (F, MO); Bachoco, 12 mi E of Cajeme, H. Gentry 2973 (ARIZ, MEXU, S); Guaymas, H. Gentry 4710 (ARIZ); Isla Tiburon, D. Gold 386 (MEXU); 1 mi E of Bahia San Carlos, ca. 12 mi N of Guaymas, B. Hansen et al. 1389 (LL, MEXU, US, WIS); 10.2 mi N of turnoff to Bahia San Carlos, J. Hastings & R. Turner 64-18 (ARIZ, DS); ca. 35 mi N of Guaymas and 7 mi W of Hwy. 15 toward Kino Bay, J. Hastings & R. Turner 65-160 (ARIZ, DS); 2 mi N of San Carlos Bay, P. Hutchison 2447 (MEXU, US); ca. 18 mi N of Hermosillo, E. Jaeger 5 (DS); island in bay near Guaymas, /. Johnston 3088 (CAS); Tiburon Island, /. Johnston 4268 (CAS, NY, UC, US); Ensenada Grande (San Pedro Bay), O. Kleine et al. 11588 (ARIZ); Tiburon Island, Knight et al. 1101 (UNM); 70 mi S of Hermosillo, G Lindsay 1134 (DS); Isla Tiburon, EF. Lott & T. Atkinson 2513 (CAS), 2514 (CAS); road to Bahia San Carlos, C. Lowe & R. Felger 1671 (ARIZ); Algodones Bay area, 4 mi WNW of San Carlos Bay, P. Martin s.n. (ARIZ); Guaymas, Microwave Mountain, 27°57’, 110°54’W, P. Martin & M. O'Rourke s.n. (ARIZ); N de Guaymas, F: Miranda 8945 (MEXU); Ardilla Island, Guaymas Harbor, 27°55’N, 110°54’W, R. Moran 4020 (DS); Tiburon Island, 28°46’N, 112°18’W, R. Moran 4059 (DS, UC); Tiburon Island, Arroyo de la Cruz, 28°46’N, 112°22’W, R. Moran 13005 (ARIZ, CAS); Isla Tiburon, Arroyo Sauzal, S end of island, E. Moser & M. Moser s.n. (ARIZ); ca. 18 mi S of Guaymas, D. Norris et al. 20057 (CAS, MEXU); Guaymas, E. Palmer 196 (C, NY, UC, US); near Guaymas, W. Palmer 1222 (US); 6-8 mi NE of Cajeme (Cd. Obregon), F. Pennell 19470 (MEXU, PH, US); entrance to Nacopuli Canyon, 3 mi N of San Carlos Bay, A. Phillips et al. 75-175 (UC); Nacopuli Canyon road, 4.5 mi N of San Carlos Bay, A. Phillips et al. 76-183 (ARIZ, CAS); Rte. 15, ca. 5 mi N of Navojoa, D. Pinkava et al. P12928 (ARIZ); Ensenada Chica, 15.1 mi W of Hermosillo to Guaymas Hwy., 28°07’N, 111°17'W, F. Reichenbacher 224 (ARIZ); Mpio. Empalme, Rancho las 778 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Trincheras, ca. 6 km NW of Empalme, 28°01’N, 110°50’W, A. Reina G. & T. Van Devender 2001-182B (CAS); camino Bahia de Kino—Punta Chueca, KM 16.3, 28°55’N, 112°02'W, A. Reina et al. 95-62 (ARIZ, CAS); Mpio. Guaymas, 27.4 km S of Restaurant Los Arrieros, KM 143 on Mex. 15, 28°05’N, 110°57'W, A. Reina G. etal. 98-01 (CAS, MEXU, TEX); Guaymas, Colonia Peninsular, H. Ripley 14266 (NY); Tiburon Island, J, Rose 16790 (US); vicinity of Guaymas, J. Rose et al. 12585 (US); vicinity of Empalme, J. Rose et al. 12626 (NY, US); vicinity of Bahia San Carlos, J. Russell s.n. (ARIZ); near Rancho El Valiente, Hwy. 15 at jct. with bypass to Cd. Obregon, 3.8 mi N of turnoff to San Carlos Bay, ca. 28°02’N, 110°56'W, A. Sanders et al. 8766 (CAS, MO); Isla Tiburon, 3 km W Punta Tormenta, NV. Scott P5 (UNM); near Ortiz, F’ Shreve 6104 (ARIZ, F); San Carlos Bay, W of Guaymas, F: Shreve 6405 (ARIZ, F); 17 mi S of Guaymas on Mex. 15, W. Spaulding 75-3-32 (ARIZ); 3 mi E of Mex. 15 on road to La Pintada Canyon, 28°35.5’N, 110°58.5'W, G Starr 754 (ARIZ); 5.8 mi N of turnoff to Bahia San Carlos on Mex. 15, G Starr & D. Palzkill 290 (ARIZ); Nacopuli Canyon near Bahia San Carlos, G Starr & C. Starr 713 (ARIZ); entrance to Nacopuli Canyon, G Starr et al. 27 (ARIZ); near Hwy. 15 ca. 12.2 mi E of Vicum Station, W. Stevens 1506 (ENCB, MO); Isla Tiburon, Ensenada de Perros, 28°46'N, 112°16’W, P. Tenorio L. et al. 9523 (F, MEXU); Tiburon Island, Sau Sal drainage, S side of island, Tewksbery et al. s.n. (ARIZ); Isla Tiburon, F) Torres s.n. (ENCB); 2.3 mi NW of Guaymas, R. Turner & C. Lowe 148 (ARIZ); 7.5 mi W of jct. with Mex. 15, along Kino Bay Road cutoff, 28°17'N, 110°08'W, R. Van Devender & T. Van Devender 83-116 (ARIZ); Cation Babiso, Rancho Babiso, 2.5 mi W of San Carlos Bay, 7; Van Devender & M. Kearns s.n. (ARIZ); Nacopuli Canyon, ca. 4 mi N of San Carlos Bay, 7: Van Devender et al. 84-238 (ARIZ); 10.2 mi W of Mex. 15 toward El Sahuaral (southern road to Bahia Kino), 28°19'N, 110°12'W, 7: Van Devender et al. 92-11 (ARIZ, CAS); Canada El Tetabejo, Sierra Libre, 28°32.5’N, 110°59'W, T. Van Devender et al. 95-1042 (ARIZ); 75 mi S of Hermosillo on Hwy. 15, J. Walker 9S (TEX); 1 mi N of Las Guasimas, 19 mi S of Guaymas along Mex. 15, 27°55'N, 110°34’'W, R. Warren & D. Goldberg s.n. (ARIZ); ca. 15 mi SE of Guaymas, G. Webster & S. Lynch 17003 (MEXU, MO); Tiburon Island, vicinity of Tecomate on N shore of island, 4. Whiting 9067 (ARIZ); 3 mi S of Ortiz toward Guaymas, /. Wiggins 6324 (DS, UC, US); Isla Tiburon, La Sauzal, 28°49’N, 112°35'W, I. Wiggins 17174 (DS, MEXU); Coralitos, shore at S side of Isla Tiburon, R. Wilkinson 193 (MEXU); 10 mi NW of Bahia San Carlos, N. Yensen 740319-1 (ARIZ). Ruellia intermedia Leonard, J. Wash. Acad. Sci. 17:512. 1927. TYPE: MEXICO. Jalisco: Bolanos, 10-19 Sep 1897, J. Rose 2915 (holotype: US!; isotype: NY!). Phenology. Flowering: July-September; fruiting: August—October, February—March. Distribution and habitats. Mexico (Baja California Sur, Chiapas, Chihuahua, Colima, Guerrero, Jalisco, México, Michoacan, Morelos, Nayarit, Sinaloa, Sonora); Sonoran plants occur in Sonoran desertscrub (Plains of Sonora), thornscrub, tropical deciduous forests, and in mesquite bosques along streams at elevations from 120 to 1000 m. Illustration. Journal of the Washington Academy of Sciences 17:513. 1927. Local names. “Conivari” (Pima, Pennington 249); “papachili” (Mayo, fide Van Devender et al. 2000); “rama del toro” (Sa/mon s.n.); “suspinda” (Pima, Pennington 249); “trikis sipunim” (Pima Bajo, Rea 1154); “tronando” (Rea 1/54): “yerba del toro” (fide Gentry 1942, as R. tuberosa). Use. The seeds are used in preparing a “refreshing drink-soak” (Pennington 249). Daniel (1995a, 1997) provided descriptions of this species, which reaches the northern and western extents of its distribution in north-central Sonora (Fig. 33). Although most collections were made in thornscrub and tropical deciduous forest, a few collections have been made in regions of Sonoran desertscrub. Among Sonoran plants, the calyces are pubescent with mostly glandular tri- chomes (e.g., Van Devender et al. 98-1276), pubescent with exclusively eglandular trichomes (e.g., Reina G. et al. 98-862 and most collections), or nearly glabrous (e.g., Van Devender et al. 94- 654A). Trauba s.n. contains a shoot with glandular pubescent calyces and other shoots with calyces lacking glandular trichomes; Daniel 993 shows both glandular- and eglandular-pubescent calyces on the same plant. DANIEL: SONORAN ACANTHACEAE TS SONORAN COLLECTIONS: Mpio. San Javier, ica a ay 2.7 km E of La Barranca on Mex. 16, 28°34'N, L~ ell | 109°40'W, 4. Buirguez M. 96-261 (CAS); between a Mose Onavas and hwy. between Esperanza and Rosario de } Tezopaco, 5 mi S of Movas, 7. Daniel 993 (CAS); El | 3y° Coyote, E-central Sonora, R. Felger 3571 (ARIZ); Mpio. Guaymas, Sierra Libre, Cation Tetabejo, 28°32'N, 110°59'W, R. Felger et al. 02-292 (CAS); Lb ca. | km §S of El Tabelo on Alamos—San Bernardo ee road, 27°09'N, 108°58'W, M. Fishbein et al. 1872 i (ARIZ, CAS); Arroyo Seco, SW of Los Camotes | 29° e | along Los Tanques—Las Chinacas road, 27°16'N, 108°51'W, M. Fishbein et al. 1880 (ARIZ); Dist. “tog Alamos, Canyon Estrella, H. Gentry 353 (DS); high- — F a 4 way between Alamos and Navojoa, 4. Gibson & L. Gibson 2081 (ENCB, TEX): 4.3 km SW of Santa @ Ruellia intermedia ¢° Ana de Yécora on road to Nuri, 28°23'N, 109°19’W, [27 ee D. Goldberg & S. McLaughlin 77-173 (ARIZ); Lo de Campo, near San Xavier mine turnoff on Hwy. 114° 112° | ~#p AUD MBO Bepuoal Ue dilenttio ae Clb it (CNA FIGURE 33. Sonoran distribution of Ruellia intermedia Arroyo Gochico, E of San Bernardo, F) Pennell ; 19516 (GH); Onavas, C. Pennington 249 (TEX); NE of Onavas at mine, 28°28’N, 109°32’W, 4. Rea 1154 (ARIZ); Mpio. Onavas, Rancho la Mula, 5 km W of Agua Amarilla, 28.2 km SE of Rio Yaqui on Mex. 16, 28°29'N, 109°22'W, A. Reina G. 97-576 (CAS); Mpio. Yécora, Arroyo Santa Ana at SON. 12, 8.5 km W of Guadalupe Tayopa, 28°21'N, 109°16'W, A. Reina G. et al. 98-862 (CAS); Mpio. Yécora, Arroyo La Quema, near Tepoca, 28°26’N, 109°15’W, 4. Reina G. et al. 98- 871 (CAS, MEXU): Mpio. Soyopa, near Arroyo Garambullo on NE side of Rio Yaqui bridge on Mex. 16, just S of Tonichi, ca. 28°34'N, 109°33'W, A. Reina G et al. 99-33 (CAS); Mpio. Yécora, base of Cerro El Pilar, above Arroyo Santa Ana, 5.5 km N of Curea on Son. 12, 28°20'N, 109°15'W, A. Reina G, et al. 99-797 (CAS); Mpio. Sahuaripa, 8 km N of Rio Yaqui, 25.5 km N of Sahuaripa on road to Tepache, 29°15’N, 109°19’W, A. Reina G. et al. 2003-933 (CAS); 7 mi N of Guirocoba, J. Salmon s.n. (ARIZ, CAS); Mpio. Alamos, Rancho Esmeralda, NW outskirts of Alamos toward San Bernardo, 27°02'N, 108°56'W, A. Sanders et al. 12501 (CAS); 7 mi S of Baviacora, F. Shreve 6734 (ARIZ, UC); 190 km E of Hermosillo on Yécora, P. Sundt s.n. (ARIZ); Rancho El Carrizo, 100 mi S of Nogales, R. Tomelson s.n. (ARIZ); Mpio. Yécora, 1.6 km E of Rancho Viejo toward Santa Ana, ca. 28°20'N, 109°15'W, W. Trauba s.n. (CAS); Alamos, 27°01'N, 108°50'W, IT. Van Devender 94-619 (ARIZ); Mpio. Onavas, Rancho La Mula, 28.2 km SE of Rio Yaqui on Mex. 16 (KM 195 E of Hermosillo), 28°28.5'N, 109°22'W, T. Van Devender & A. Reina G. 2000-509 (ARIZ); Mpio. Soyopa, Loma Maderista, 3.5 km S of Tonichi, ca. 28°34’N, 109°33’W, 7: Van Devender & A. Reina G. 2001-200 (CAS); Mocuzari (Adolfo Ruiz Cortinez) Dam on Rio Mayo, WNW of Alamos, 27°13'N, 109°06.5'W, 7: Van Devender & R. Van Devender 92-1385 (ARIZ, CAS); Las Lajitas crossing of the Rio Cuchujaqui, 8.9 mi ESE of Alamos, 26°59’N, 108°49'W, T. Van Devender et al. 92-1377 (CAS); El Rincon Viejo, ca. 4 km N of Alamos, Sierra de Alamos, 27°04'N, 108°56'W, 7: Van Devender et al. 94-654 (ARIZ), 94-654A (ARIZ, TEX); Mpio. Yécora, Curea, 28°19’N, 109°17'W, T. Van Devender et al. 98-1276 (CAS); Mpio. Arivechi, Arroyo Agua Caliente, 10.2 km (air) WNW of Arivechi, 28°57'N, 109°17'W, 7: Van Devender et al. 99-880 (CAS); Mpio. Yécora, Rancho La Palma Agujerada, 7.1 km N of Mex. 16 on road to Sahuaripa, 28°28.5’N, 109°11'W, T. Van Devender et al. 2000-359 (ARIZ); Mpio. Moctezuma, 20.5 km SSE jet. with Moctezuma—Huasabas hwy. on road to Tepache (Son. 117), 29°39'N, 109°35'W, T. Van Devender et al. 2004- 413 (CAS); Mpio. San Javier, Cerro el Halcon, near San Javier, 28°37'N, 109°43'W, L. Varela E. 96-209B (CAS); ca. 1 mi E of RR on road to Horcasitas, 1. Wiggins 7275 (ARIZ, DS, GH, MICH, US); E slope of Sierra Batuc, several mi NW of Batuc, /. Wiggins & R. Rollins 292 (ARIZ, DS, GH, MO, UC, US), 292a (DS). 110° | 780 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Ruellia inundata H.B.K., Nov. Gen. Sp. 2:239. 1817. Aphragmia inundata (H.B.K.) Bremek., Verh. Kon. Ned. Akad. Wetensch. Afd. Natuurk., Tweede Sect. 45:10. 1948. TYPE: COLOMBIA. “In ripa fluminis Magdalenae prope Mompox et Badillas,’ May, (fide protologue), 4A. von Humboldt & A. Bonpland mss. n. 3711 (holotype: P-Bonpl.!). Ruellia albicaulis Bertero ex Spreng., Syst. Veg. 2:822. 1825. TYPE: COLOMBIA. “Ad fl. Magdalen.” (fide protologue), no specimens have been seen. Aphragmia haenkei Nees in Lindley, Intr. Nat. Syst. Bot., ed. 2, 444. 1836. Dipteracanthus haenkei (Nees) Nees in A. de Candolle, Prodr. 11:141. 1847. TYPE: MEXICO. Guerrero: Aca- pulco, 1791, 7: Haenke s.n. (no type designated nor place of deposition noted; probable type mate- rial: K, PR!, PRC!). Ruellia galeottii Leonard, Kew Bull. 1938:59. 1938. TYPE: MEXICO. Oaxaca: Sola, Oct 1844, H. Galeotti 510J (holotype: US!; isotypes: NY!, W!). Phenology. Flowering: October—April; fruiting: October—April. Distribution and habitats. Mexico (Baja California Sur, Campeche, Chiapas, Colima, Guerrero, Jalisco, México, Michoacan, Morelos, Oaxaca, Puebla, Quintana Roo, Sinaloa, Sonora, Veracruz, Yucatan), Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, and Brazil; Sonoran plants occur in canyons and disturbed sites in tropical deciduous forests at elevations from 260 to 740 m. Illustrations. Carnegie Institution of Washington Publication 461:210. 1936; Fieldiana, Botany 18:17. 1986. Descriptions of Ruellia inundata were provided by Daniel (1995a, 1997). Sonoran plants are noted as having bluish corollas. Vegetative organs of plants of this species are usually viscid and have a mephitic odor (similar to that of Dyschoriste hirsutissima). The species reaches the north- ern extent of its distribution in southern Sonora (Fig. 32). SONORAN COLLECTIONS: San Bernardo, Rio Mayo, H. Gentry 1068 (ARIZ, F, MO, S, US); near Tojibampo, 19.5 km NNW of San Bernardo toward Mesa Colorada, 27°02'N, 108°47'W, T. Van Devender et al. 93-369 (ARIZ, CAS, TEX), 93-1495 (ARIZ, CAS, MO, NY, TEX), 93-/495A (ARIZ); ca. 4 km NNW of Nahuibampo, W slopes of Cordon Topiyecas, 27°24’N, 108°55’W, 7: Van Devender et al. 94-353 (ARIZ); western Sierra de Alamos, 7.7 km S of Tetajiosa toward Las Rastras, 26°56'N, 109°04'W, 7: Van Devender et al. 95-158 (ARIZ, CAS); Cerro las Tatemas, below Microondas La Luna, 13.8 km NW of Alamos, 27°07’N, 109°02'W, T: Van Devender et al. 95-1158 (ARIZ, CAS); 4.7 km SSE of Piedras Verdes toward Tepustete, ca. 12 km NNW of Alamos, 27°08'N, 108°56'W, 7. Van Devender et al. 95-1180 (CAS). Ruellia leucantha Brandegee subsp. postinsularis (Gentry) T.F. Daniel, Polibotanica 2:7. 1996. Ruellia leucantha var. postinsularis Gentry, Brittonia 6:323. 1948. TYPE: MEXICO. Sinaloa: Cerro Tecomate, W of Pericos, 28 Feb 1940, H. Gentry 5739 (holotype MICH!; isotypes ARIZ, DS!, GH, MEXU!, MO, NA, NY, RSA!, UC!, US). Dyschoriste candida Brandegee, Zoe 5:242. 1906. Ruellia candida (Brandegee) Kobuski, Ann. Missouri Bot. Gard. 15:60. 1928. TYPE: MEXICO. Sinaloa: vicinity of Culiacan, Yerba Buena near Altata, 10 Oct 1904, 7: Brandegee s.n. (holotype UC!; isotypes GH!, US!). Shrubs to 8 dm tall. Young stems quadrate to quadrate-sulcate, evenly puberulent with erect to flexuose to retrorse eglandular trichomes 0.05—0.2 mm long and sessile patelliform glands (glan- dular punctate), the trichomes sometimes very dense and + obscuring epidermis. Leaves petiolate, petioles to 7 mm long, blades ovate to elliptic, 22-57 mm long, 11-33 mm wide, 1.7—2.2 times longer than wide, rounded to acute at apex, rounded to subattenuate at base, surfaces glandular punctate and pubescent (sometimes very densely so) with erect to flexuose to antrorse eglandular DANIEL: SONORAN ACANTHACEAE 781 trichomes 0.1—0.5 mm long, margin ciliate with similar trichomes. Inflorescence of axillary dicha- sia: dichasia sessile, | (—2) per axil, 1-flowered, alternate or opposite at vegetative nodes; flowers sessile to subsessile (1.e., borne on pedicels to | mm long). Bracteoles prominent to inconspicuous (to obsolete), sometimes caducous, petiolate and linear-elliptic to linear or sessile and triangular to triangular-subulate, 0.2—-23 mm long, 0.2—-5 mm wide, pubescent like young stems. Calyx 5-lobed, (5—) 7-14 mm long, tube 1—2 mm long, lobes subulate to linear-subulate, subequal to unequal in length (1.e., 1 lobe up to 1 mm longer than others), (4—) 6.5—13 mm long, 4—8.3 times longer than tube, abaxially pubescent like young stems. Corolla white, 38-68 mm long, externally glandular punctate and pubescent with erect to flexuose to retrorse eglandular trichomes 0.1—0.3 mm long, tube funnelform, 40-52 mm long, 2.2—3 mm in diameter near midpoint, narrow proximal portion of tube 17-30 mm long, distally abruptly expanded into a throat 13—23 mm long, limb (22—) 37-40 mm in diameter, lobes 7-16 mm long, 8-16 mm wide. Stamens included to + exserted, didyna- mous, longer pair 12-18 mm long, shorter pair 10-16 mm long, filaments proximally pubescent with eglandular trichomes and glandular punctate, distally glabrous, thecae (3—) 4.5—5S.5 mm long. Style (25—) 33-47 mm long, pubescent with erect to antrorse eglandular trichomes and glandular punctate; stigma unequally 2-lobed, | lobe 2—2.8 mm long, other lobe 0.2—0.4 mm long. Capsule 8—12 mm long, externally glandular punctate and pubescent with erect to flexuose to retrorse eglan- dular trichomes 0.05—0.2 mm long, stipe 2 mm long, head ovoid ellipsoid to ellipsoid-subcircular, 8—10 mm long. Seeds 4 (—6) per capsule, 3.5-4.5 mm long, 3.24.5 mm wide, surfaces and mar- gin covered with appressed hygroscopic trichomes to 0.3 mm long, margin + swollen. Phenology. Flowering: August; fruiting: April, October. Distribution and habitats. Northwestern Mexico (Sinaloa, Sonora); Sonoran plants occur in desert oases in riparian Sonoran desertscrub (Plains of Sonora) at elevations from 250 to 300 m. Illustration. None found. The collections cited below represent the first records of this species in Sonora. Daniel (1996, 1997) recognized two subspecies of Ruellia leucantha. Plants from Sonora pertain to R. leucantha subsp. postinsularis and greatly resemble the type from Sinaloa. The description of that subspecies provided by Daniel (1996) has been augmented with data from Sonoran plants and 1s provided above. The nominate subspecies is restricted in distribution to the Cape Region of Baja California Sur (Daniel 1997). The subspecies of R. /eucantha can be distinguished by the following couplet: 1. Cauline trichomes erect to flexuose, 0.3-1.5 mm long; capsules 12-15 mm long; Baja SAMOA SURE Fe ee ees ee ble ee re ers oe Pua lo See etnies subsp. leucantha 1. Cauline trichomes erect to retrorse, 0.05—0.2 mm long; capsules 9-12 mm long; Sonora and SITE Cis Sh See cack eet ahs BRIN aA a che ae eae gerne LE ae aS ae subsp. postinsularis The northern distributional extents of both the species and subspecies postinsularis are in west-central Sonora (Fig. 30). The collector of all known Sonoran plants of R. /eucantha noted that the desert oasis in which the species occurs in that state is located in a region of deep canyons (with permanent water) in which numerous tropical species occur. Its occurrence within the Sonoran Desert, like that of Aphanosperma sinaloensis, is restricted to riparian corridors in which more tropical elements extend their ranges. According to T. Van Devender (pers. commun.), the canyon slopes are dominated by thornscrub. Some confusion with respect to corolla color is present among the label data on Sonoran col- lections of this species. The sole flowering specimen (Burquez & Yetman 97-431), identified as R. californica, notes that corollas were purple (1.e., as in R. californica). A fruiting collection (Burquez et al. 97-497), identified as Justicia candicans, noted that corollas were red (i.e., as in J. candi- cans). | suspect that corolla colors as provided on the labels of these collections pertain to the 782 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 species as identified, not to the plants collected. If | am incorrect, and Sonoran representatives of R. leucantha do not have white corollas like their counterparts in Sinaloa and Baja California Sur, then the glandular punctate pubescence still distinguishes this species from other Sonoran Ruellia. SONORAN COLLECTIONS: La Pintada, 49 km S de Hermosillo por la Carr. Fed. 15, 28°35'N, 110°58’W, A. Burquez & R. Felger 93-32 (MEXU), A. Burquez & D. Yetman 97-431 (MEXU); Sierra Libre, Cafiada Prieta, S de Hermosillo por la Carr. Fed. 15, 28°34'N, 110°58’W, A. Burquez et al. 97-497 (MEXU). Ruellia malacosperma Greenm., Proc. Amer. Acad. Arts 34:572. 1899. TYPE: MEXICO. Tamaulipas: Tampico, 30 Apr 1898, C. Pringle 6806 (lectotype, designated here, see discussion below: GH, photo at CAS!; isolectotypes: BM!, F!, L!, LL, MEXU, MIN!, NSW!, NY, PR!, PRC!, Si, WEL WIS) Perennial herbs to | m tall. Young stems quadrate to quadrate-sulcate to + flattened, internodes mostly glabrous or sometimes + evenly pubescent with flexuose eglandular trichomes 0.5—2 mm long, nodes pubescent with flexuose eglandular trichomes 0.5—2 mm long. Leaves petiolate, peti- oles to 20 mm long, blades narrowly elliptic to elliptic, 45-163 mm long, 8-38 mm wide, 2.3—5.8 times longer than wide, (rounded to) acute to attenuate at apex, acute to attenuate at base, surfaces sparsely pubescent with a few scattered flexuose eglandular trichomes along midvein, midvein conspicuously caniliculate on adaxial surface (at least near proximal portion of blade), margin sub- sinuate to crenate-undulate. Inflorescence of pedunculate ascending expanded dichasia to 120 mm long from leaf axils; dichasia (opposite to) alternate, 1 per axil, (1—) 3—many-flowered, peduncles 26-70 mm long, subquadrate to quadrate, glabrous or sparsely pubescent with inconspicuous glan- dular trichomes 0.05—0.1 mm long (glandular-puberulent). Bracteoles sometimes caducous, lance- olate to lance-linear, 5-10 mm long, I1—1.5 mm wide, abaxial surface glabrous, secondary bracte- oles similar to bracteoles except smaller. Flowers pedicellate, pedicels 4-12 mm long, + densely glandular-puberulent. Calyx 13-21 mm long, tube 2—3 mm long, lobes lance-subulate, subequal (1.e., longest lobe 1.1—1.3 times longer than shortest lobe), 10.5—18 mm long, 3.8—7.5 times longer than tube, 1.2—1.5 mm wide, abaxially glandular-puberulent (glands up to 0.2 mm long), margin ciliate with antrorse eglandular trichomes 0.05—0.6 mm long and/or glandular-puberulent. Corolla blue-purple, 45-60 mm long, externally pubescent with flexuose eglandular and glandular (some- times sparse) trichomes 0.1—0.2 mm long, tube 33-38 mm long, narrow proximal portion 9-15 mm long, + abruptly expanded into throat, throat 18-25 mm long, longer than narrow proximal portion, 6.5—14 mm in diameter near midpoint, limb 32-46 mm in diameter, lobes 12—17 mm long, 14-18 mm wide. Stamens included, longer pair 12—15 mm long, shorter pair 8-10 mm long, thecae 3.7-4 mm long. Style 20-21 mm long, pubescent (at least proximally) with upward-pointing eglandular trichomes, stigma unequally 2-lobed, | lobe 0.2 mm long, other lobe 1.5—2.2 mm long. Capsule 24-29 mm long, proximally glabrous, inconspicuously pubescent near apex with erect to antrorse glandular and eglandular (sometimes absent) trichomes 0.05—0.2 mm long, stipe 4 mm long, head linear-ellipsoid. Seeds 16—20 per capsule, 3.1—3.2 mm long, 2.4-2.5 mm wide, surfaces and mar- gin covered with appressed hygroscopic trichomes. Phenology. Flowering: June, October; fruiting: March, June, October. Distribution and habitats. Presumably native to Mexico where it has been recorded from Guerrero, Nuevo Leon, San Luis Potosi, Sinaloa, Tamaulipas, and Veracruz (reports from some of these states undoubtedly are based on cultivated plants); Sonoran plants are cultivated or occur in margins of dried pools in coastal thornscrub with Prosopis, Acacia, Cyperus, and Lycium at eleva- tions near sea level (naturalized or possibly native plants) or up to 415 m (cultivated plants). Illustration. Carnegie Institution of Washington Publication 461:208. 1936. Three collections, from three Mexican states, were cited by Greenman in the protologue: DANIEL: SONORAN ACANTHACEAE 783 Pringle 6806 (Tamaulipas), Palmer 570 (Guerrero), and Botteri s.n (Veracruz). The latter collec- tion was cited somewhat incidentally, and it was noted to be at GH. Although their herbaria of deposit were not noted in the protologue, specimens of both Palmer 570 and Botteri s.n. have been located to date only at GH. Pringle’s collection was widely distributed with a printed label bearing Greenman’s name for this taxon. It remains unknown, however, whether Greenman actually stud- ied all of the duplicates. Therefore, only the specimens at GH of these three collections are here considered to be syntypes. Tharp and Barkley (1949) indicated that the “type” of R. malacosper- ma was Pringle 6806. This would appear to be sufficient under the current International Code of Botanical Nomenclature (Greuter 2000) to effectively lectotypify the species (1.e., with the only syntype of Pringle 6806 being at GH). If it is argued, or can be demonstrated, that Greenman used duplicates of this collection in composing his protologue, then this would not appear to have con- stituted a valid lectotypification under Article 7.11 of the ICBN because the “type element” is not clearly indicated (1.e., there are numerous syntypes of Pringle 6806). To avoid further confusion, Pringle 6806 at GH, a complete specimen with flowers and fruits, is here designated as the lecto- type for this species. It is likely that Tharp and Barkely’s “lectotypification” was unintentional and that mine is superfluous, but neither rationale would prevent the lectotypification of this name under the ICBN. This species is cultivated in Sonora (e.g., Daniel 9773) and either escapes/naturalizes or is also native to the state (e.g., Sanders et al. 13579). There is no indication on Rose 1288 whether the plant was cultivated or not in Guaymas. The above description is based on the type and other Mexican collections resembling it. Sanders et al. 13579, consisting of fruiting rhizomatous plants collected in March in thorn scrub, resembles R. malacosperma in most respects and is tentatively treated in that taxon. Corollas are not present on this collection, however, and it differs from oth- ers of R. malacosperma by its lower stature (2 vs. up to 10 dm tall), shorter leaves (20-50 vs. 45—135 mm long), shorter inflorescences (e.g., peduncles 2—7 vs. 30-65 mm long), and shorter capsules (15-18 mm long). Although the distinctions between R. malacosperma and R. coerulea Morong are not great (Daniel 1995a; Turner 1991-with R. coerulea treated as R. brittoniana Leonard), specimens from Sonora more closely resemble the former rather than the latter taxon by their relatively wider (2.9—6.2 times longer than wide) leaf blades (vs. 5.6—30 times longer than wide in R. coerulea). Both species are widely cultivated, and their origins and/or native distribu- tional ranges are not known with certainty. SONORAN COLLECTIONS: garden of La Ciudadela just off Plaza de Armas in Alamos, ca. 27°01'N, 108°56'W, cultivated, 77 Daniel 9773 (CAS); Mpio. Huatabampo, roadside ca. | km N of Maroncarit, 10 km NW of Yavaros, ca. 26°44’N, 109°37'W, A. Sanders et al. 13579 (ARIZ, CAS, MO); Guaymas, J. Rose 1288 (US). Ruellia nudiflora (Engelm. & A. Gray) Urb., Symb. Antill. 7:382. 1912. Dipteracanthus nud- iflorus Engelm. & A. Gray, Boston J. Nat. Hist. 5:229. 1845. TYPE: UNITED STATES. Texas: Sim’s Bayou near Houston, May—Jul (fide protologue) 1843, F) Lindheimer 157 (lectotype, desig- nated here, see discussion below: GH!; isolectotypes: GH!, UC!). Ruellia tuberosa L. var. occidentalis A. Gray, Syn. Fl. N. Amer. 2(1):325. 1878. Ruellia nudi- flora var. occidentalis (A. Gray) Leonard, J. Wash. Acad. Sci. 17:516. 1927. Ruellia occidentalis (A. Gray) Tharp & F.A. Barkley, Amer. Midl. Naturalist 42:25. 1949. TYPE: collections of Berlandier, Wright, Rothrock, and Coulter from the southwestern United States were cited by Gray; Leonard (J. Wash. Acad. Sci. 17:516. 1927) indicated that the type was collected in Texas by Berlandier; it will require study of these collections to determine whether Leonard effectively lec- totypified the species at that time. 784 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 Ruellia nudiflora var. glabrata Leonard, J. Wash. Acad. Sci. 17:518. 1927. Ruellia glabrata (Leonard) Tharp & F.A. Barkley, Amer. Midl. Naturalist 42:13. 1949. TYPE: MEXICO. Sonora: Ciénega, Aug 1874, J. Rothrock 560 (holotype: US; isotypes: GH!, NY). Ruellia gooddingiana A. Nelson, Amer. J. Bot. 18:437. 1931. TYPE: MEXICO. Sonora: La Ciénega, 18 Jul 1911, L. Goodding 959 (holotype: RM!). See Daniel (1995a) for a complete listing of synonyms for this species. Phenology. Flowering: March, July—October; fruiting: July—October. Distribution and habitats. United States (Arizona, Louisiana, Texas), Mexico (Baja California Sur, Campeche, Chiapas, Chihuahua, Coahuila, Colima, Durango, Morelos, Nuevo Leon, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosi, Sinaloa, Sonora, Tabasco, Tamaulipas, Veracruz, Yucatan), Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, and West Indies; Sonoran plants occur in swampy meadows, flats, swales, along watercourses, and in disturbed areas in Sonoran desertscrub (Arizona Upland, Central Gulf Coast, Lower Colorado River Valley, Plains of Sonora), Chihuahuan desertscrub, palm oases, thornscrub, mesquite grass- lands, mesquite bosques, and tropical deciduous forests at elevations from near sea level to 1180m. Illustrations. Figure 34; Carnegie Institution of Washington Publication 461:211. 1936; Flora of Texas 1(3):240. 1966; Desert Plants 5:172. 1984; Fieldiana, Botany 18:18. 1986. Local names. “Cuetito” (fide White 1949): “papachili” (fide Yetman and Van Devender 2002); “rama del toro” (Van Devender et al. 95-1074); “tronador” (fide Yetman and Van Devender 2002). Use. Yetman and Van Devender (2002) noted that the Mayo use the leaves to brew a tea that is taken to alleviate fever. Dipteracanthus nudiflorus was based primarily on Lindheimer 157 from Sims Bayou (the locality does not appear on syntypes at GH or UC, but is noted by Engelmann and Gray in the pro- tologue). In the protologue Engelmann and Gray also cited Drummond coll. 2, no. 221 (seen at GH and K) and Drummond coll. 3, no. 257 (seen at GH and K), both from unknown localities in Texas. Leonard (1927) indicated that Lindheimer had collected the type at Sim’s Bayou. Because he did not cite a specific specimen, and as there appear to be multiple syntypes, his choice would not appear to constitute a lectotypification. Potential syntypes of Lindheimer 157 include a specimen at GH with the plant name in Gray’s handwriting, a specimen now at GH that came from the Boston Society of Natural History (in whose journal the species was published), and a specimen at UC with the name written apparently by Gray. All of these specimens bear the number “157” which was likely assigned to Lindheimer’s plants as they were studied. The three potential syntypes noted above were all collected in 1843 as part of Lindheimer’s “fasc. I.’ Other specimens of this species bearing the number “157” were noted to have been collected by Lindheimer as “fasc. III’ in 1846 (e.g., at UC). It is unknown to me whether the specimens of Lindheimer’s no. 157 from 1846 were also collected at Sim’s Bayou, but because they were collected subsequent to the publication of this name, they would not figure in its typification. To firmly establish a type for this species, a lecto- type is here designated for Dipteracanthus nudiflorus: Lindheimer (Fasc. 1) 157 (GH-Harvard University Herbaria no. 00093956). The lectotype was collected in 1843 and is mounted on a sheet with a Drummond collection. Daniel (1995a, 1997, 1999a) and Daniel and Acosta (2003) provided descriptions of this species. Some collections note the presence of white corollas. While it is possible that some plants FiGuRE 34 (right). Ruellia nudiflora. a. Habit of base of plant (Wiggins & Rollins 136), x 0.49. b. Habit of apex of plant (Mearns 737), x 0.43. c. Leaf (Wiggins & Rollins 136), x 0.51. d. Corolla (Daniel & Butterwick 2980), x 1.3. e. Distal portion of stamen with anther (Fe/ger et al. 0/-459), x 11.3. f. Distal portion of style with stigma (Felger et al. 01-459), x 17.8. g. Capsule with calyx (lobe in front partially removed) (Mearns 1999), x 3.1. h. Dry seed (Daniel & Marrs 1808), x 6.9. 1. Moistened seed (Daniel & Marrs 1808), x 6.9. Drawn by Nadia Strasser. 785 DANIEL: SONORAN ACANTHACEAE 786 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 may be albinic in corolla color, most collec- tions purporting to have white corollas also L indicate that mature flowers were not present or lack them on the specimens. Corolla buds of R. nudiflora appear whitish but are invariably blue ;3! internally when opened. This species occurs nearly throughout Sonora (Fig. 35), including occurrences in each subdivision of the Sonoran Desert in the state. Flowering of Ruellia nudiflora mostly coin- -29° cides with the summer rains, but occasional plants have been collected in flower during the spring. Plants are usually found associated with watercourses (i.e., on stream banks, in arroyo bottoms, or in riparian flood zones) or flats or | 37° swales where water accumulates following @ Ruellia nudiflora rains. 114° 112° SONORAN COLLECTION: 2.5 km S de Hermo- 7 sillo, 29°O1'N, 110°57'W, A. Burquez & A. Quijada 91-14 (MEXU); from Hwy. 15 to Bahia Agiabampo, ca. 5 km NE of Bahia Agiabampo, ca. lat. 26.41°N, long. 109.20° W, 7: Daniel 9768 (CAS); 4.6 mi N of Fronteras, R. Felger 4039 (ARIZ, ENCB); 10.2 mi S of La Pintada (between Hermosillo and Guaymas) on Mex. 15, R. Felger 8364 (ARIZ), R. Felger & C. Lowe 8364 (ARIZ, MEXU); ca. 7 mi S of Pitiquito, R. Felger & E. Abbey 20271 (ARIZ); Bahia Algadones, ca. 2 km N of Club Med, R. Felger & R. Devine 85-333 (ARIZ); 0.8 km E of Las Guasimas, R. Felger & N. Gonzales 85-382 (ARIZ, MEXU); 9 mi E of Empalme end of Douglas Bridge on Hwy. 15, then 2.5 mi S of Hwy. on Playa del Sol Road, 27°54'N, 110°41'W, R. Felger & F. Reichenbacher 85-1106 (ARIZ, CAS, MEXU, TEX); road to Bahia San Carlos, 4 mi W of Hwy. 15, R. Felger & R. Thomas 11941 (ARIZ); SW of Los Camotes on Arroyo Seco, along Los Tanques-Las Chinacas road, 27°16'N, 108°51'W, M. Fishbein et al. 1880 (ARIZ); Mpio. Magdalena de Kino, Rancho La Tinaja Colorada, ca. 15 km NE de Magdalena de Kino, 30°42'N, 110°48'W, A. Flores M. et al. 4584 (ARIZ, IEB, MEXU); Mpio. San Pedro de la Cueva, 3 km SW de San José de Batuc, carr. Mazatan—San Pedro de la Cueva, 29°18'N, 109°54’W, A. Flores M. & J. Leon 5276 (MEXU); Mpio. La Colorada, 2.5 km S de El Pilar, 28°39'N, 110°52'W, A. Flores M. & J. Leon G. 5356 (IEB, MEXU); 15 mi S of Hermosillo, H. Gentry 4562 (ARIZ); Ciénega on the Rio Saracahi, 1.7 mi NW of Agua Fria, 30°22'N, 110°35’W, D. Goldberg 77-126 (ARIZ, MEXU); La Ciénega, L. Goodding 959 (ARIZ); Logulea [?] to Altar, D. Griffiths 6895 (MO); vic. of San Pedro, C. Hartman 955 (MO, UC, US); Nacori Chico drainage, E of Buena, 29°37'N, 108°59’W, E. Joyal 1722 (MEXU); 5.1 mi NW of San Carlos Yacht Club on road to Algadones, R. Krizman & O. Soule s.n. (ARIZ); without locality, E. Palmer 98 (C, MEXU, NY, US); Bacum Station, near Rio Yaqui, F’ Pennell 20202 (MEXU, PH, US); 14 mi by Tecolote road W of Mex. 15 (1.4 mi N of El Oasis), 29°48’N, 111°16’W, F Reichenbacher 1017 (ARIZ); Mpio. Imuris, Agua Caliente, 19 km N of Imuris, 30°57’N, 110°51'W, A. Reina G. et al. 2001-657 (CAS); Mpio. Agua Prieta, Rancho San Bernardino, 29 km E of Agua Prieta on Mex. 2, 31°20'N, 109°17'W, A. Reina G et al. 2003-1005 (CAS); Mpio. Huatabampo, Camahuiroa, 27 km SE of Yavaros, 26°31'N, 109°16'W, A. Sanders et al. 12713 (CAS); Distr. Altar, Bahio de Aquituni, between Pitiquito and Rancho Verruga, F) Shreve 6023 (ARIZ); Ejido Manual Caudillo, 7.4 mi S of Luis (ca. 10 mi N of Sinaloa border), O. Soule & R. Krizman s.n. (ARIZ); Palm Canyon, SE of Magdalena in Cerro Cinta de Plata, L. Toolin & T. Van Devender 435 (ARIZ); across Rio Bacanuchi from Tauichopa (2.7 mi N of Arispe), R. Turner & C. Lowe 185 (ARIZ); Alamos, 27°01'N, 108°50'W, 7: Van Devender 94-619 (ARIZ); Ciénega near Rancho Agua Fria on Rio Saracachi, E of Cucurpe, 7. Van Devender & C. Miksicek s.n. (ARIZ, ENCB); Mpio. Saric, Rancho Los Nogales, Arroyo El Silencio, 19.5 km N of Saric on old road to Sasabe, 31°15’N, 111°21'W, 7: Van Devender & A. Reina G. 2002-915 (CAS); Mpio. Navojoa, Teachive de Masiaca, Arroyo (eaaee FiGuRE 35. Sonoran distribution of Ruellia nudiflora. DANIEL: SONORAN ACANTHACEAE 787 Masiaca, 26°47'N, 109°14'W, 7: Van Devender & D. Yetman 94-665 (ARIZ); Mouth of Arroyo Masiaca, Las Bocas on Gulf of California, ca. 52 km S of Navojoa, 26°36'N, 109°20'W, T. Van Devender & D. Yetman 94- 703 (ARIZ); Palm Canyon, 17.7 mi SE of Magdalena in Cerro Cinta de Plata, 7) Van Devender et al. s.n. (ARIZ); Mpio. Navojoa, Arroyo Masiaca, ca. 0.5 km N of Teachive de Masiaca, 26°48’N, 109°14'W, 7. Van Devender et al. 93-968 (ARIZ, CAS); El Rincon Viejo, ca. 4 km N of Alamos in Sierra de Alamos, 27°04'N, 108°56'W, T. Van Devender et al. 94-654 (ARIZ); 1 km SSW of Sirebampo, 26°37’N, 109°15'W, 7 Van Devender et al. 95-1074 (CAS); 28 mi SE of Guaymas, U. Waterfall 12830 (TEX, UC, US); 65 mi SE of Sonoita, ca. 31°10'N, 112°20'W, G. Webster & G Breckon 15572 (GH, MEXU); near Colonia Oaxaca, Arroyo del Pulpito, S. White 759 (GH, US); region of the Rio de Bavispe, Cafion de las Bellotas, 8S. White 3565 (ARIZ, GH, MEXU): 9 mi W of La Angostura, S. White 4023 (ARIZ, NY, US); Colonia Morelos, S. White 4111 (ARIZ, NY, US); 7 mi W of Pozo Cerna, /. Wiggins 6040 (DS); 20 mi W of Norio, /. Wiggins 6173 (DS, US); along Rio de los Alisos, 31 mi S of Nogales, /. Wiggins 7016 (DS, US); 27 mi W of Hermosillo, 1. Wiggins & R. Rollins 136 (ARIZ, DS, MO, NY, UC, US). Ruellia parryi A. Gray, Syn. Fl. N. Amer. 2(1):326. 1878. Dipteracanthus suffruticosus Torr. in W.H. Emory, Rep. U.S. Mex. Bound. 2(1):122. 1859, non Ruellia suffruticosus Roxb. (1814). TYPE: MEXICO. Chihuahua: “Presidio del Norte,” 7 Jul 1852, C. Parry “722” (lectotype, desig- nated here, see discussion below: NY [plant at lower right on sheet], photo at http://sidsite.nybg.org/herbim/2970/v-297-00311935big.jpg!). Erect subshrubs to 6 dm tall. Young stems quadrate-sulcate, internodes glabrous or evenly pubescent with retrorse eglandular trichomes 0.05—0.2 mm long, nodes often with a cluster of flex- uose eglandular trichomes to 1.3 mm long. Leaves petiolate, petioles to 8 mm long, blades ovate to lance-elliptic to elliptic to oblanceolate to obovate, 9-25 (—35) mm long, 3—15 mm wide, |.7—2.8 (—7) times longer than wide, acute to rounded at apex, acute to attenuate at base, surfaces pubes- cent (sometimes sparsely so) with erect to flexuose (and with at least some branched or stellate) tri- chomes to 0.8 mm long, margin entire to undulate, usually conspicuously ciliate with trichomes like those of leaf surfaces (or these usually somewhat longer, 1.e., to 1.2 mm long). Inflorescence of pedunculate dichasia in leaf axils; dichasia alternate or opposite, 1 per axil, | (—3)-flowered, peduncles 0.5—6 mm long, glabrous or pubescent like young stems. Bracteoles subfoliose, petio- late, ovate to lanceolate to elliptic to obovate, 6.5—18 mm long, 2—6.5 mm wide, pubescent like leaves. Flowers sessile. Calyx 5—11 mm long, abaxially pubescent with erect to flexuose (some- times branched or stellate) eglandular trichomes 0.05—0.2 mm long, tube 1-2 mm long, lobes unequal in size, anterior segment lanceolate to elliptic, 4-9 mm long, 1—1.8 mm wide, apically 2- lobed, lobes ovate to subulate (sometimes asymmetric), 2-7 mm long (sometimes divided nearly to or to the tube), lateral lobes lance-subulate to lanceolate, 2.5—7 mm long, 0.5—0.7 mm wide, pos- terior lobe lanceolate, 3.5—9 mm long, 0.7—1.2 mm wide, entire. Corolla blue, 20-41 (—50) mm long, externally pubescent with erect to flexuose to retrorse eglandular trichomes 0.05—0.2 mm long, tube 16-38 mm long, narrow proximal portion 8-23 mm long, abruptly expanded distally into throat, throat obconic, 7-13 mm long, shorter than tube or + equal in length to tube, 2.3—7 mm in diameter near midpoint, limb 12—32 mm in diameter, lobes 5-13 mm long, 5-14 mm wide. Longer stamens 7-9 mm long, shorter stamens 5—6 mm long, thecae 1.8—2 mm long. Style 19-26 mm long, proximally pubescent with eglandular trichomes, distally glabrous; stigma unequally 2-lobed, 1 lobe 0.5 mm long, other lobe 1.2 mm long. Capsule ellipsoid, 9-11 (—13) mm long, (glabrous or) entirely or distally pubescent with retrorse eglandular trichomes 0.1—0.2 mm long. Seeds 6 (or more) per capsule, lenticular, 2.5—-3 mm long, 1.9—2.2 mm wide, surfaces and margin pubescent with appressed trichomes. Phenology. Flowering: August; fruiting: August. Distribution and habitat. Southwestern United States (Arizona, New Mexico, Texas) and 788 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 northern Mexico (Chihuahua, Coahuila, Durango, Nuevo Leon, Sonora, Zacatecas); Sonoran plants occur on limestone in Chihuahuan desertscrub at an elevation of 1287 m. Illustrations. Trees, Shrubs and Woody Vines of the Southwest, 934. 1960; Flora of Texas 1(3):247. 1966; Trees and Shrubs of Trans-Pecos Texas, 396. 1988. In the protologue of Dipteracanthus suffruticosus, Torrey cited the following collection data: “Presidio del Norte; July-August; Parry. Sides of rocky hills, valley of the Pecos. No. 1461, Wright.” A lectotype is designated above from among the syntypes in Torrey’s protologue. In Torrey’s herbarium at NY, there are three pertinent collections: Wright 1461 from the valley of the Pecos, a collection made by Parry at Presidio del Norte on 7 July 1852 (with “722” also written on the label), and another collection (without the name of the collector) made at Presidio del Norte in August of 1852. The two collections from Presidio del Norte are mounted on the same sheet. Because Torrey referred to unspecified collection(s) by Parry from July and August, he undoubt- edly attributed both of these collections to Parry and was including both in his citation of syntypes. Thus there are three syntypes and all of them pertain to what is now treated as R. parryi. Tharp and Barkley’s (1949) indication of the type as “at Presidio del Norte, Parry s.n.” would not appear to constitute lectotypification (either intentional or unintentional) because it does not distinguish between the two syntypes presumably collected by Parry at that locality. The lectotype designated above possesses an open and intact flower (unlike the other syntypes). The type locality, Presidio del Norte, was noted in the protologue to be at the junction of the Rio Conchos and Rio Grande and is today called Ojinaga. Ruellia parryi 1s here reported for the first time from Sonora where it reaches the western extent of its distribution (Fig. 31). It is one of at least three Mexican species of Ruellia known to have stellate or dendritic trichomes on mature organs (e.g., leaves, bracteoles, calyx). The other two, R. hirsutoglandulosa (Oerst.) Hemsl. and R. lanatoglandulosa (Nees) Lindau, occur to the south of the Chihuahuan Desert and differ from R. parryi by their longer calyces ((8—) 11-34 mm), usually longer corollas ((32—) 42-85 mm) with the external pubescence including glandular tri- chomes, and capsular pubescence that includes glandular trichomes. Branched trichomes are some- times present on the youngest vegetative growth in other Mexican Ruellia (e.g., R. leucantha Brandegee). Most plants of Ruellia parryi, including those of the sole Sonoran collection, are pubescent more or less throughout. In some (e.g., Breedlove & Mahoney 71916 from Coahuila) the internodes are glabrous and trichomes on the leaves and bracteoles are largely restricted to the midvein and margin. SONORAN COLLECTION: Mpio. Agua Prieta, NE of Sierra Anibacachi, Rancho La Calera, ca. 10 km SW of Agua Prieta, 31°14'N, 109°38’'W, A. Reina G. et al. 2003-957 (CAS). Stenandrium Nees Stenandrium Nees in Lindley, Intr. Nat. Syst. Bot., ed. 2, 444. 1836, nomen cons. TYPE: Stenandrium mandioccanum Nees. Gerardia L., Sp. Pl. 610. 1753, pro parte, nomen rej. vs. Stenandrium Nees (1836). LECTO- TYPE (Pennell, Bull. Torrey Bot. Club 40:120. 1913): Gerardia tuberosa L. Acaulescent (arising from a woody rhizome or caudex) or caulescent erect to spreading peren- nial herbs lacking cystoliths. Leaves opposite or quaternate. Inflorescence of axillary or terminal elongate or headlike usually pedunculate dichasiate spikes; dichasia opposite or alternate, 1-flow- ered, sessile, subtended by a bract. Flowers homostylous, sessile, subtended by 2 homomorphic bracteoles. Calyx deeply 5-lobed, lobes equal or subequal in size. Corolla pink, purple, or white, DANIEL: SONORAN ACANTHACEAE 789 tube expanded distally into a short throat, limb 2-labiate or appearing subactinomorphic, upper lip 2-lobed, lower lip 3-lobed, corolla lobes imbricate in bud. Stamens 4, subdidynamous, included in corolla tube, anthers 1-thecous, pubescent, lacking basal appendages, dehiscing toward lower lip (i.e., flower nototribal); pollen (Fig. 18) euprolate to perprolate (to spherical), 3-colpate (rarely inaperturate), exine psilate to foveolate to fossulate to verrucate to gemmate; staminode 0—1, borne between posterior pair of stamens, usually shorter than filaments. Style included in corolla tube, stigma asymmetrically funnelform. Capsule estipitate to substipitate, ellipsoid to obovoid, retinac- ula present, septa with attached retinacula remaining attached to inner wall of mature capsule. Seeds 4, homomorphic, lenticular, often with barbed or branched trichomes. (x = 13). Stenandrium has traditionally been treated as a New World genus with some 40 to 50 species. Vollesen (1992) included Stenandriopsis S. Moore from Africa and Madagascar within Stenandrium. Although Stenandriopsis in the Old World appears morphologically indistinguishable from Stenandrium in the New World, recent molecular studies (McDade et al. 2003) show the for- mer genus to be the basal member of an otherwise New World clade that includes Stenandrium. These studies do not support the inclusion of Stenandriopsis within Stenandrium, but additional research will be necessary to resolve generic boundaries between these and among other genera of tribe Acantheae. Daniel (1985) recognized nine species of Stenandrium in Mexico. Concentrations of species are also found in the West Indies (ca. 15 species) and South America (15—25 species). Stenandrium pilosulum (Blake) T.F. Daniel, Ann. Missouri Bot. Gard. 71:1039. 1984. Gerardia pilosula Blake, Contr. Gray Herb. 52:101. 1917. TYPE: MEXICO. Chihuahua: vicinity of Madera, 27 May-3 Jun 1908, E. Palmer 317 (holotype: GH!; isotypes: MO!, US!). Acaulescent perennial herbs to 7.5 cm tall from a + woody caudex or rhizome, the caudex or rhizome bearing numerous fleshy roots along its length. Leaves (plants often leafless or nearly so during anthesis) ascendant, petiolate, petioles to 18 mm long, glabrous or pubescent with a few eglandular trichomes to 0.3 mm long, blades oblanceolate to narrowly elliptic to lanceolate (to ovate), 11-55 mm long, 3—7 (—11) mm wide, 2.8—6.9 (—11) times longer than wide, attenuate to decurrent at base, acute to rounded at apex, surfaces glabrous (although the lower surface incon- spicuously glandular along the basal portion) and punctate-pitted, margin red, entire, eciliate. Inflorescence of pedunculate spikes to 30 mm long, peduncles 5-45 mm long, nearly glabrous or pubescent with retrorse to erect trichomes 0.05—0.2 mm long; dichasia opposite to subopposite along spike axis, sessile. Bracts ovate to narrowly elliptic to obovate, 6-12 mm long, 1.5—4.5 mm wide, abaxial surface pubescent like peduncles although with trichomes more numerous, margin inconspicuously ciliate with trichomes 0.05—0.3 mm long. Bracteoles linear to linear-lanceolate, 6—11.5 mm long, 0.5—1.1 mm wide, abaxial surface pubescent like bracts. Calyx 7-14 mm long, lobes linear-subulate to linear-lanceolate, subequal, 6.5-13.5 mm long, pubescent like bracts. Corolla pink with tube and proximal portions of lobes of lower lip white, 10-23 mm long, tube 6-15 mm long, upper lip 3-10 mm long, lower lip 4-10 mm long, lobes subequal, 3—10 mm long, lobes and tube glabrous or sparsely pubescent on external surface, lower-central lobe often dense- ly pubescent on external surface. Stamens 2.5—3 mm long, anthers 1.5 mm long. Capsule 9-12 mm long, pubescent over the entire surface. Seeds 3-4 mm long, 3—3.5 mm wide, densely pubescent with long bristlelike trichomes bearing lateral barbs or branches. Phenology. Flowering: March—July; fruiting: March. Distribution and habitats. Sierra Madre Occidental of northwestern Mexico (Chihuahua, Sonora); Sonoran plants occur on gravely slopes, in pastures, and along watercourses in oak-pine communities at elevations from 1540 to 2250 m. Illustration. Figures 15, 36. 790 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 FIGURE 36. Stenandrium pilosulum. a. Habit (Daniel et al. 8614), x 1. b. Leaf (Laferriere 439), x 2. c. Bract (Daniel et al. 8607), x 4.3. d. Bracteole (Daniel et al. 8607), x 4.3. e. Calyx (Daniel et al. 8607), x 3.4. f. Distal portion of stamen with anther (Daniel et al. 8607), x 15.8. g. Distal portion of style with stigma (Daniel et al. 8614), x 22.7. h. Capsule (Correll & Johnston 21612), x 4.4. i. Capsule valve (Correll & Johnston 21612), x 4.4. j. Seed (Correll & Johnston 21612), x 8.4. k. Trichome from seed (Correll & Johnston 21612), x 24. Drawn by Jenny Speckels. DANIEL: SONORAN ACANTHACEAE 791 Although Daniel (1985) provided a ! ae ae description of this species, sufficient additional collections from Sonora have become available in the intervening years to warrant an augment- ed, and more complete one. The species is endemic to the highlands of eastern Sonora (Fig. 37) and western Chihuahua. Corollas are pink with the internal surfaces of the tube and proximal portions of the lobes of the lower lip white. The ventral portion of the tube near its mouth is densely pubescent with yellowish gold trichomes. Sonoran collections: Mpio. Yécora, ca. 0.5 km NW of cemetery in Yécora, ca. 28°23’N, e iageraee Peas 108°50'W, T: Daniel et al. 8607 (CAS, MEXU); 57° @ Tetramerium glandulosum Mpio. Yécora, El Kipor, 7.4 km E of Maycoba along Hwy. 16, ca. 28°24’N, 108°35’W, ZT: Daniel et al. 8614 (CAS, MICH); Rancho La Pinosa,5.9miW of LL , | aie | Me Maycoba on Hwy. 16, 28°25’N, 108°43.5’W, M. FIGURE 37. Sonoran distributions of Stenandrium pilo- Fishbein et al. 1839 (ARIZ); Yécora, just NW of — suum, Tetramerium fruticosum, and T. glandulosum. cemetery, M. Fishbein et al. 1859 (ARIZ); 28°22.5'N, 108°55.5'W, ca. 5 mi W of Yécora, P. Hubbell s.n. (ARIZ); 7 mi NW of Yécora, ca. 28°26'N, 109°00'W, R. Moran 21965 (CAS, ENCB); Mpio. Yécora, El Kipor (Quipur), Arroyo El Kipor, 28°24'N, 108°36'W, T: Van Devender & A. Reina G. 95-364 (ARIZ, CAS, MEXU), 97-434 (MO, TEX); Mpio. Yécora, 34 km NNW of El Kipor (Quipur), 28°25.5’N, 108°36'W, 7: Van Devender & A. Reina G. 95-405 (ARIZ); ca. 5 mi W of Maicova, 28°22'N, 108°44’W, G. Webster 23849 (CAS). Tetramerium Nees Tetramerium Nees in G. Bentham, Bot. Voy. Su/phur, 147. 1846, nom. cons., non C.F. Gaertn. (1806). TYPE: Tetramerium polystachyum Nees (= Tetramerium nervosum Nees). Averia Leonard, J. Wash. Acad. Sci. 30:501. 1940. TYPE: Averia longipes (Standl.) Leonard (= Drejerella longipes Standl.; = Tetramerium nemorum Brandegee). Decumbent to erect perennial herbs or shrubs with cystoliths. Leaves opposite (rarely ternate). Inflorescence of terminal conspicuously and usually densely bracteate 4-sided unbranched dichasi- ate spikes; dichasia opposite, 1—3-flowered, sessile, subtended by a bract. Flowers homostylous, sessile, subtended by 2 homomorphic bracteoles. Calyx deeply 4- or 5-lobed, lobes homomorphic or, if 5, | usually + reduced in size. Corolla white, cream, yellow, blue, or red, often with maroon and purplish markings on upper lip, tube subcylindric or gradually expanded distally, throat indis- tinct or distinct only near mouth, limb pseudopapilionaceous, upper lip comprising 2 fused lobes, entire to emarginate at apex, lower lip 3-lobed, lower-central lobe (in ours) conduplicate and enclosing stamens and often style during anthesis, corolla lobes imbricate in bud. Stamens 2, exserted from mouth of corolla, anthers 2-thecous, thecae equal to subequal in size, parallel to sub- sagittate, equally inserted, lacking basal appendages, dehiscing toward upper lip (1.e., flower stenotribal); pollen (Fig. 38) subprolate to perprolate, 3-colporate, 6-pseudocolpate, pseudocolpi 2 per mesocolpium, exine (rugulate to) reticulate to bireticulate; staminodes 0. Style exserted from mouth of corolla, stigma 2-lobed, lobes equal. Capsule stipitate, head ellipsoid to obovoid, retinac- ula present, septae with attached retinacula slightly separating from inner wall of mature capsule. 792 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 OER GO eae 4s 925. St ratere)! T. abditum (Daniel 3364), view. b. itum (Daniel 3364), apertura FIGURE 38. Pollen of Sonoran Jetram a interapertural view. c. T. fruticosum (Daniel & Butterv 6869), apertural view. d. TZ. nervosum (Carter & Moran 5324), apertural view. e. 7. nervosum (Carter & Moran 5324), polar view. f. T. glandulosum (Van Devender et al. 94-946), inter- apertural view. g. 7. tenuissimum (Jenkins 88-320), apertural view. h. T. tenuissimum (Jenkins et al. 88-320), interap- l uianum (Van Devender et al. 92-1109), interapertural view. Scales = 10 um. ertural view. 1. 7) yaguianu DANIEL: SONORAN ACANTHACEAE 793 Seeds 4, homomorphic, plano-convex to concavo-convex), lacking trichomes. (x = 18). This genus of 29 species occurs from the southwestern United States southward throughout most of Mexico, Central America, and northwestern South America to southeastern Bolivia (Daniel 1986). With 22 species, Mexico is the center of diversity of Zetramerium. 1. Corollas red to reddish orange, 29-42 mm long, upper lip lacking maroon markings, tube fun- nelform, 18-25 mm long, 2.84.8 mm in diameter at apex; thecae 2.9-3.6 mm long; style PAG PARTUM IN AMR crap ee ala RPE TN PA ENTER cod Che GHRNG Sha ete meaeas eta oes oa aL aR T. abditum 1. Corollas white, cream, or yellow, 7-26 mm long, upper lip with maroon markings, tube sub- cylindric, 2.5—9 mm long, 1—2.5 mm in diameter at apex; thecae 0.6—2.6 mm long; style 4-26 mm long. 2. Viscid shrubs to 3 m tall; herbage conspicuously glandular with glands up to | mm long (at least some glands longer than 0.3 mm); corollas bright yellow with 2 maroon stripes on upper lip, 17-26 mm long; thecae |1.7—2.6 mm long; capsules 6—9 mm long. I edly cpr yee, Leng Wea me Lok aN ay T. glandulosum 2 Perennial herbs to 5 dm tall; herbage eglandular or + inconspicuously glandular with glands to 0.3 mm long; corollas white, cream, or creamy yellow and with a maroon (and usually purple) chevron on upper lip, 7—-17.5 mm long; thecae 0.6—1.5 mm long; capsules 4.5—6.5 mm long. 3. Calyces 4-lobed; apical portion of bract usually conspicuously twisted... . 7) nervosum 3. Calyces 5-lobed; apical portion of bract straight (sometimes slightly twisted in T. yaquianum). 4. Capsules glabrous; bracteoles 5—10 mm long..................... T. fruticosum 4. Capsules pubescent; bracteoles 3—-6.5 mm long. 5. Bracts usually widely spreading (to horizontal or nearly so) from rachis, pubes cence of abaxial surface includes glandular trichomes 0.2—0.3 mm long, margin ciliate with at least some trichomes longer than 0.5 mm; calyx 3.54.5 mm long; corolla 13-6 mmilong,tubei6-8:mm' longs)... sc) .2c)s. - ee: T. yaquianum 5. Bracts erect, closely appressed to rachis, pubescence of abaxial surface includes glandular trichomes 0.05—0.2 mm long, margin ciliate with trichomes to 0.5 mm long; calyx 2-3 mm long; corolla 7-13.5 mm long, tube 2.5—5 mm long. 5 2 aida bys/aId dra ena NBL EM Sc eee asic ae ail eae i ae asim nil ie T. tenuissimum Tetramerium abditum (Brandegee) T.F. Daniel, Syst. Bot. Monogr. 12:113. 1986. Anisacanthus abditus Brandegee, Zoe 3:348. 1893. TYPE: MEXICO. Sonora: Las Durasnillas, 14 May 1892, T. Brandegee s.n. (holotype: UC!; isotypes: F!, GH!). Phenology. Flowering: November—May; fruiting: November—May. Distribution and habitats. Western Mexico (Chihuahua, Durango, Michoacan, Nayarit, Sinaloa, and Sonora); Sonoran plants occur on slopes and along watercourses in mesquite scrub, tropical deciduous forests, and oak-pine woodlands at elevations from 230-1360 m. Illustrations. The Botany of the Voyage of H.M.S. Herald, t. 68. 1856; Systematic Botany Monographs 12:115. 1986. Local names. “Rama del toro” (fide Yetman and Van Devender 2002); “viipsumar u’us” (Pima Bajo, Rea 1069). Daniel (1986) provided a description of this species and discussed (2004b) its distribution and distinctions from the morphologically similar T rubrum Happ. Tetramerium abditum attains the 794 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 northern extent of its distribution in east-cen- tral Sonora (Fig. 39). The location of the type locality is discussed above under Henrya. SONORAN COLLECTIONS: San __ Javier, microondas en el Cerro El Durazno, 28°37'N, 109°45'W, A. Burquez 94-019[a] (MEXU); Cerro El Verde, San Javier, 28°33-34'N, 109°43-44'W, 4. Burquez et al. 96-066 (MEXU), 96-085 (MEXU); Mpio. Yécora, Santa Rosa, 28°28'N, 109°06'W, A. Burquez M. 96-228 (CAS); between Hwy. 16 E of Tonichi and Onavas, 0.5 mi S jet. Hwy. 16, 7. Daniel 3347 (ASU, CAS, MEXU); between Rosario de Tezopaco and Nuri, 7.1 mi S of turnoff to Nuri, 7: Daniel 3364 (ASU, CAS, ENCB, MEXU, MICH, MO, NY, US); Rio Cuchujaqui, 7.4 mi SE of Alamos toward Guiricoba, 7) Daniel 3378 (CAS), T. Daniel @ Tetramerium abditum 3998 (CAS); Mpio. Yécora, between San Nicolas pe and Nuri, ca. 6.4 km SW ject. Hwy. 16, ca. 28°24'N, 109°14'W, T. Daniel et al. 8570 (BR, CAS, K, L_, i MEXU, MO); Sierra de Alamos, Arroyo Uvalama FIGURE 39. Sonoran distribution of Tetramerium abdi- (Igualamas), 27°O1'N, 108°59'W, M. Fishbein et al. — tum. 1095 (ARIZ, CAS); Arroyo Guajaray, upstream from village of Guajaray, 27°37'N, 108°57'W, M. Fishbein et al. 1553 (ARIZ, CAS); 21.5 mi E of Tecoripa, P. Gallagher et al. 287 (ASU); Distr. Alamos, Agua Caliente, H. Gentry 898 (ARIZ, DS, MICH); Caramechi, Rio Mayo, H. Gentry 1178 (A, ARIZ, F, K, MEXU, MICH, MO, NY, UC, US); vicinity of Alamos, H. Gentry 4801 (ARIZ, DS, GH, MEXU, MICH, MO, NY, UC); 5 mi W of El Novillo Dam, D. Goldberg 76-66 (ARIZ); 10 mi W of Nuri, D. Goldberg s.n. (ARIZ); W of Piedras Verdes near Mocuzari Reservoir, 15 km NW of Alamos, P. Jenkins 89-127-A (ARIZ); Naranjo, 27°14’N, 108°43’W, P. Jenkins 89-316 (ARIZ); Mpio. Alamos, Arroyo Verde, 27°06'N, 108°43'W, P. Jenkins 90-313 (ARIZ); Hwy. 16 E of Onavas turn-off, 28°35'N, 109°30'W, E. Joyal 1674 (CAS, MEXU); Rio Cuchujaqui, cave next to Mesa Redonda, 27°05’N, 108°45'W, P. Martin & G. Ferguson s.n. (ARIZ, CAS); Mpio. Alamos, 3.5 mi SW of Rancho Santa Barbara, 27°06'N, 108°45'W, P. Martin & G. Ferguson s.n. (ARIZ); 6 mi NNW of Alamos, P. Martin & M. O'Rourke s.n. (ARIZ); cajon of Cuchujaqui, 27°03’, 108°44°W, P. Martin & M. O’Rourke s.n. (ARIZ); above Las Animas, 27°31.5'N, 108°42'W, P. Martin et al. s.n. (ARIZ); El Reparo near Santa Ana de Yécora, G. Monson s.n. (ARIZ); near Onavas, 15—20 km NE along Hwy. 16, 4. Rea 1069 (ARIZ); 4 km W of Agua Amarilla, 18.9 km NNW of Tepoca on Mex. 16, 28°29'N, 109°22’'W, A. Reina G. & T. Van Devender 97-371 (ARIZ, CAS, MEXU, TEX); Arroyo El Mentidero at El Chinal road, 11.3 km S of Alamos, 26°55’N, 108°55’W, A. Reina G. et al. 98-24 (ARIZ); Mpio. Onavas, 4.3 km SE of Onavas junction on Mex. 16, 28°31'N, 109°28'W, A. Reina G et al. 2001-155 (ARIZ, MO); Hwy. 16, ca. 8 km E of Tecoripa, Rancho San Juanico, R. Rondeau &9- S82 (ARIZ): Cerro Redondo, 27°05'N, 108°45'W, R. Rondeau & G. Rodda s.n. (ARIZ); ca. 5 mi below Minas Nuevas, J. Rose et al. 12669 (US): vicinity of Alamos, J. Rose et al. s.n. (F, GH); Mpio. Mazatan, Sierra de Mazatan, Canada El Carrizo, N de Rancho El Carrizo, 29°03'N, 110°12'W, J. Sanchez E. et al. 04-045 (CAS); Mpio. Alamos, Cafion Algoroba, Sierra de Alamos, ca. 27°00'N, 108°57'W, A. Sanders & L. Lubinsky 13452 (TEX); lower crossing of Rio Cuchujaqui, 8 mi SE of Alamos, ca. 26°56'N, 108°53'W, A. Sanders et al. 2636 (CAS, TEX); 2.2 mi E of San Nicolas, ca. 8 mi NE of Curea, A. Sanders et al. 2679 (ASU); vicinity of Alamos, E. Smith 3960 (CAS, MEXU, TEX); 18.5 mi SE of Rio Yaqui crossing at Mex. 16, G Starr & C. Starr 154 (ARIZ), Starr et al. 44 (ARIZ): Mpio. Alamos, Sierra de Alamos, Arroyo El Huirotal, ca. 26°57.5'N, 108°57.5'W, VK Steinmann 95-04 (ARIZ, MEXU); above El Novillo, 83 mi E of Hermosillo, L. Toolin & K. Roever 576 (ARIZ); 4 mi S of Mazocahui, R. Turner et al. 59-72 (ARIZ, CAS, UC); 12 mi S of Soyopa, R. Turner et al. 71-89 (ARIZ, UC); Arroyo Hondo, 10 mi N of Nuri, R. Turner et al. 75-33 (ARIZ, SD, UC); Rio Cuchujaqui at Rancho El Conejo, 12.4 km S of Alamos, 26°55'N, 108°55’W, T: Van Devender 92-563 (ARIZ); crossing of Rio Cuchujaqui on road to Guirocoba, ca. 8 mi SE of Alamos, 7. Van Devender et al. 84-144 DANIEL: SONORAN ACANTHACEAE 795 (ARIZ); SE slopes of Sierra de Alamos, Arroyo El Mentidero, Rancho Uvalama, 11 km WSW of Alamos, 26°S5'N, 108°55'W, T. Van Devender et al. 92-53 (CAS); El Guayabo crossing of Rio Cuchujaqui, 1.1 mi NE of Sabinito Sur, 14 km ESE of Alamos, 27°00’N, 108°48'W, T. Van Devender et al. 92-228 (ARIZ, CAS); Rio Cuchujaqui at Rancho El Conejo, 13.4 km S of Alamos, 26°54'N, 108°55'W, T. Van Devender et al. 92-615 (ARIZ): Arroyo El Mentidero at crossing of El Chinal road, 11.5 km S of Alamos, 26°55'N, 108°55'W, T. Van Devender et al. 93-73 (ARIZ, CAS); below Guajaray on Arroyo Guajaray, 6.5 km WNW of jet. with Rio Mayo, 27°36'N, 108°56'’W, 7. Van Devender et al. 93-464 (ARIZ); canyon N of Los Aguaros on Arroyo Guajaray, 27°38.5'N, 108°58'W, T. Van Devender et al. 93-490 (ARIZ, in part); Mpio. Yécora, Agua Amarilla (Los Pinitos), 15 km WNW of Tepoca, 24.7 km WNW of San Nicolas on Mex. 16, ca. 28°08’N, 109°20'W, 7: Van Devender et al. 96-25 (ARIZ, CAS); San Javier, Arroyo San Javier, 1 km S del poblado y 1.5 km N del Cerro El Verde, 28°35’N, 109°44'W, L. Varela E. 96-51 (MEXU); N side of Sierra de Alamos, S. Walker 73H34 (DES): Rancho La Sierrita, E slopes of Sierra de Alamos, ca. 6 km SSW of Alamos, 26°58'N, 108°57'W, J. Weins et al. 93-027 (ARIZ). Tetramerium fruticosum Brandegee, Zoe 5:171. 1903. TYPE: MEXICO. Baja California Sur: San José del Cabo, Jan-Mar 1901, C. Purpus 489 (holotype: UC!; isotypes: CAS!, GH!, MO!, NY!, US!). Phenology. Flowering: October; fruiting: October. Distribution and habitats. Northwestern Mexico (Baja California Sur, Sonora); Sonoran plants occur in Sonoran desertscrub (Central Gulf Coast) at an elevation between 500 and 600 m. Illustration. Proceedings of the California Academy of Sciences 49:399. 1997. Daniel (1986, 1997) provided descriptions of this species. Tetramerium fruticosum is wide- spread in Baja California Sur, but known from the Mexican mainland only by a single collection (Fig. 37) from the Central Gulf Coast subdivision of the Sonoran Desert near Puerto Libertad (1.e., the northern extent of its geographic range). Daniel (1986) discussed its distribution in some detail. SONORAN COLLECTION: Tinaja Picu in Picu Mts., 2 mi N of concrete monument set on N side of Libertad Road, 1. Wiggins 6055 (DS). Tetramerium glandulosum Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1854:171. 1854. TYPE: MEXICO. Oaxaca: Rio de las Vueltas, Dec 1842, Liebmann 10751 (holo- type: C!). Tetramerium aureum Rose, Contr. U.S. Natl. Herb. 1:349. 1895. Justicia aurea (Rose) Lindau, Bull. Herb. Boissier 5:675. 1897. TYPE: MEXICO. Colima: without specific locality, 9 Jan-6 Feb 1891, E. Palmer 1302 (holotype: US!; isotypes: GH!, K!, NY!, US!). Cardiacanthus fragrans Lindau, Bull. Herb. Boissier 7:577. 1899. TYPE: MEXICO. Oaxaca: Distr. Nochixtlan, El Parian, 15 Dec 1895, C. Seler & E. Seler 1565 (holotype: B, destroyed; lec- totype, designated by Daniel, 1986: K!; isolectotypes: A!, CAS!, GH!, MEXU!, NY!, POM!). Anisacanthus stramineus Barneby, Rhodora 72:70. 1970. TYPE: MEXICO. Puebla: near Coxcatlan, 18 Nov 1966, H. Ripley 14731 (holotype: NY!; isotype: US!). Phenology. Flowering: April, December; fruiting: April, December. Distribution and habitats. Western and southern Mexico (Chihuahua, Colima, Durango, Guerrero, Jalisco, Michoacan, Morelos, Oaxaca, Puebla, Sinaloa, and Sonora); Sonoran plants occur along watercourses in tropical deciduous forests at elevations from 350 to 400 m. Illustrations. Figure 9; Systematic Botany Monographs 12:64. 1986. Daniel (1986) provided a description of this species, which attains the northern and western extents of its distribution in southern Sonora (Fig. 37). SONORAN COLLECTIONS: La Calera, 5.5 km N of Alamos Hwy. at El Carrizal, 27°07'N, 109°01.5'W, 796 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 P. Martin & M. O'Rourke s.n. (ARIZ); 23 km E of Navojoa, R. Moran 4033 (DS, SD, US); Mpio. Alamos, Arroyo Las Rastras, SW edge of Sierra de Alamos, 26°57'N, 109°03'W, T. Van Devender et al. 93-1457 (ARIZ, CAS); Cerro Huacharibi, 4 km SW of Basiroa, 26°41'N, 108°55’W, 7: Van Devender et al. 94-946 (ARIZ, CAS). Tetramerium nervosum Nees in G. Bentham, Bot. Voy. Su/phur, 148. 1846. TYPE: ECUA- DOR. Guayas: Puna, 1836-1839, A. Sinclair s.n. (holotype: K!). Tetramerium hispidum Nees in A. de Candolle, Prodr. 11:468. 1847. TYPE: MEXICO. Hidalgo: prope las Ajuntas ad flumen Montezuma, Jan, C. Ehrenberg 1072 (lectotype, designated by Happ, 1937: B, destroyed; isolectotypes: GH!, PR!; probable isolectotype: P!). Tetramerium nervosum vat. hispidum Torr. in W.H. Emory, Rep. U.S. Mex. Bound. 2(1):125. 1859. TYPE: MEXICO. Sonora: Santa Cruz, Sep 1851, G Thurber 918 (lectotype, designated by Daniel, 1986: NY!; isolectotypes: F!, GH!, NY!). Dianthera sonorae S. Watson, Proc. Amer. Acad. Arts 24:67. 1889. TYPE: MEXICO. Sonora: vicinity of Guaymas, Sep 1887, E. Palmer 240 (holotype: GH!; isotypes: C!, K!, NY, US!). Tetramerium scabrum Happ, Ann. Missouri Bot. Gard. 24:532. 1937. TYPE: MEXICO. Sonora: Fronteras, Jun 1851, G Thurber 432 (holotype: NY!; isotypes: CAS!, GH!). See Daniel (1995a) for a complete listing of synonyms for this species. Phenology. Flowering: throughout the year, with peaks in March and October (Fig. 1); fruit- ing: throughout the year. Distribution and habitats. Southwestern United States (Arizona, New Mexico, Texas), Mexico (Baja California Sur, Campeche, Chiapas, Chihuahua, Coahuila, Colima, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, México, Michoacan, Morelos, Nayarit, Nuevo Leon, Oaxaca, Puebla, Querétaro, San Luis Potosi, Sinaloa, Sonora, Tamaulipas, Veracruz, Yucatan, Zacatecas), Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, and Peru; Sonoran plants occur on slopes, along watercourses, and in disturbed areas in Sonoran desertscrub (Arizona Upland, Central Gulf Coast), palm oases, thornscrub, tropical decid- uous forests, grasslands, oak woodlands, and oak-pine woodlands at elevations from 20 to 1450 m. Illustrations. Botany of the Voyage of H.M.S. Sulphur, t. 48. 1845; Acta Botanica Venezuelica 8:165. 1973; Fieldiana, Botany 24 (10):455. 1974: Flora of Baja California, 196. 1980; Systematic Botany Monographs \2:58. 1986; Flora of Chiapas 4:146. 1995; Flora del Valle de Tehuacan- Cuicatlan 23:88. 1999. Local names. “Abahaca del monte” (Reina G. et al. 2001-653); “cola de vibora” (Yetman 2002); “tapa chorro” (Mexican, Rea 1260): “saya huehuasira” (Guarijio, Yetman 2002); “tapachor- ro” (Varela E. 96-309, 96-456); “toskonig” (Pima Bajo, Rea 1260). Uses. Remedy for diarrhea (Rea 1260; Varela E. 96-309, 96-456); treatment for stomach ache (Reina G. et al. 2001-653). Descriptions of this widespread and morphologically variable species were provided by Daniel (1986, 1995a, 1997, 1999a) and Daniel and Acosta (2003). The morphological diversity through- out its range was discussed in detail by Daniel (1986). Sonoran plants exhibit both white and yel- low corollas and have 4-calyx lobes (see Daniel 1986 for discussion of the variation in calyx lobe number in 7: nervosum). The species is widespread throughout the eastern half of Sonora (Fig. 40), but is largely absent from the drier western half. Indeed, its occurrences in the Sonoran Desert are very rare and may be restricted to riparian sites. In northern Sonora, the species 1s especially com- mon in desert grasslands and in southern Sonora it is abundant in both thornscrub and tropical deciduous forests. One collection from Sonora (Van Devender & Reina G. 2001-184) that resembles 7: nervosum DANIEL: SONORAN ACANTHACEAE Ty by its relatively long (to 1.5 mm) trichomes on the bracteal margins and lack of glandular tri- chomes in the inflorescence differs from that species by its five calyx lobes, + antrorse (vs. straight) trichomes on the bracteal margins, and by the generally flat and straight apical portions of the bracts. Whether it represents a hybrid (e.g., between T. nervosum and T. tenuissi- mum), an undetected species, or an unusual form of 7. nervosum remains undetermined. Tetramerium nervosum and T. tenuissimum sometimes grow within several centimeters of one another (e.g., Daniel et al. 8636 and 8637), but no evidence of hybridization was noted within this or other Sonoran populations of the species. Daniel (1986) discussed the results of attempted artificial crosses between these and between these and other species of the genus. SONORAN COLLECTIONS: near Guaymas, L. Bailey 267 (F); Agua Maria, along Hwy. 16 at KM 200 W of Tepoca, ca. 28°17.5'N, 109°20'W, G Boyer et al. s.n. (ARIZ); 8 km NW of Trigo on road from Yécora to Suarahipo, D. Breedlove 61074 (CAS, MEXU, MO); near Magdalena, C. Carter s.n. (MICH); San Pedro Bay, 7’ Craig 680 (POM); between Cananea and Arizpe, 27.7 mi N crossing of Rio Bacanuchi, ca. 30°50'N, 110°20'W, T. Daniel 92] (CAS, ENCB, MEXU, MICH); between Cananea and Arizpe, 16.1 mi S of Rio Bacanuchi, 7 Daniel 927 (MICH); between Hermosillo and Sahuaripa, 3 mi W of Mazatan, 7. Daniel 953 (MICH); between Navojoa and Alamos, 24.6 mi E jet. Hwy. 15 in Navojoa, 7: Daniel 1000 (CAS, MICH); Nacapuli Canyon, ca. 4.4 mi N of San Carlos, 7: Daniel 1968 (ASU); along Hwy. 16 between Tecoripa and Tonichi, 13.4 mi W of Rio Yaqui, 7. Daniel 3343 (CAS); between Hwy. 16 E of Tonichi and Onavas, 0.5 mi S jet. Hwy. 16, 7; Daniel 3349 (ASU, CAS); between Rosario de Tezopaco and Nuri, 8.1 mi S of turn to Nuri, 7 Daniel 3362 (ASU); Sierra @ Tetramerium nervosum FIGURE 40. Sonoran distribution of Tetramerium nervo- sum. N of hwy. between Navojoa and Alamos, 2.6 mi W of Alamos, 7. Daniel 3375 (ASU), 3385 (ASU, CAS, ENCB, MICH, NY); Rio Cuchujaqui, 7.4 mi SE of Alamos toward Guiricoba, 7; Daniel 3380 (ASU, CAS, MEXU, NY, US), 3989 (CAS); 0.2 mi N of Navajoa—Alamos hwy. toward El Tezal, 2.6 mi W of Alamos, T. Daniel 3385 (CAS, MEXU); southeastern border of Alamos (road to mirador), ca. lat. 27°01'N, long. 108°56'W, T. Daniel 9772 (CAS); Rio Cuchujaqui, ca. 8 mi SE of Alamos, 7: Daniel & M. Butterwick 3193 (CAS, MEXU): Mpio. Yécora, between San Nicolas and Nuri, just S of Curea, ca. 24 km SW jet. Hwy. 16, ca. 28°19’N, 109°17'W, T: Daniel et al. 8573 (CAS); Mpio. Yécora, Arroyo La Quema, 0.3 km SE of Tepoca on Hwy. 16, ca. 28°26’N, 109°15'W, T. Daniel et al. 8636 (BR, CAS, K, MEXU, MICH); mountain W of Alamos, F- Drouet & D. Richards 3956 (F); 20 mi NE of Ures, F. Drouet et al. 3598 (F); ca. 5 km W of Pilares de Nacozari, F. Drouet et al. 3672 (F); 21 mi SE of Ures, F) Drouet et al. 3729 (F); Rancho la Brisca, 3.5 mi N of Agua Fria, 30°24’N, 110°33’W, J. Duek s.n. (ARIZ); 35.6 mi NE of Nuri, D. Dunn 9064 (RSA); Hwy. 2, Imuris, J. Dwyer 14131 (CAS, MO); ca. 1 km N of Bahia San Carlos on old road to Bahia Algadones, R. Felger & R. Devine 85-330C (ARIZ), R. Felger et al. 84-193 (ARIZ, MEXU); Arroyo Nacapules, ca. 0.5—0.8 km E of mouth of Cafion Nacapules, 6 km N of Bahia San Carlos Road, 28°00’/N, 111°02'W, R. Felger & F- Reichenbacher 85-1178 (ARIZ, CAS, TEX); Cafion Nacapuli, 4.7 mi N of San Carlos Road, 28°00'N, 111°02-03’W, R. Felger & F- Reichenbacher 85-1222 (ARIZ, CAS, MO, TEX), R. Felger & R. Thomas 11862 (ARIZ, CAS, MEXU); ca. 1 mi N of Bahia San Carlos toward Bahia Algadones, R. Felger & A. Russell 9598 (ARIZ, CAS, MEXU); El Baviso, ca. 2.5 km NE of Bahia San Carlos, R. Felger & D. Valdez Z. 84-517 (ARIZ); Sierra el Aguaje, Cafion las Barajitas, ca. 18 km NW of San Carlos, ca. 28°03'N, 111°10'W, R. Felger 798 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 & M. Wilson 95-237 (ARIZ, CAS, MEXU); Mpio. Guaymas, Cafion La Balandrona N side of Sierra El Aguaje, 28°06’N, 111°04'W, R. Felger et al. 01-719 (CAS); 4.1 mi E of Alamos, J. Fish 158 (UC); Las Camotes, 6 mi NE of Los Tanques, 27°17'N, 108°50'W, M. Fishbein et al. 80 (ARIZ); Conejos, Rio Mayo, H. Gentry 1121A (F); Alamos, Rio Fuerte, H. Gentry 2926 (ARIZ, GH, MEXU, MO, S); KM 75 on Hwy. 15, N of Guaymas, 4. Gibson & L. Gibson 2028 (RSA); 4.3 km SW of Santa Ana de Yécora on road to Nuri, 28°23'N, 109°19'W, D. Goldberg & S. McLaughlin 77-183 (ARIZ); Cohuto, C. Hartman 76 (F, GH, PH, UC, US); 4 mi N of Colonia Oaxaca, J. Hastings & R. Turner 65-28 (ARIZ, DS, SD); 16 mi E of Moctezuma, J. Hastings & R. Turner 65-101 (ARIZ, DS, SD); Rio El Naranjo crossing N of Taymuco, 27°15'N, 108°43'W, P. Jenkins & P. Martin 88-174 (ARIZ); 20 km W of Alamos, 27°07'N, 109°10'W, P. Jenkins et al. 88-65 (ARIZ); San Carlos Bay, /. Johnston 4362 (CAS, GH, UC); Las Cabras, ESE of Alamos, 26°50’N, 108°55'W, E. Joyal 1422A (CAS); Rancho El Aguilar Noria, N of Ures, 29°33'N, 110°25—26'W, E. Joyal 1817 (MEXU, TEX); 72 mi E of Hermosillo, E. Lehto et al. L19299 (ASU); El Guayabo along Rio Cuchujaqui, 27°00'N, 108°47'W, P. Martin s.n. (ARIZ); 8 km E of Alamos on road to Sabinito Sur, P. Martin s.n. (ARIZ); 16 km SE of Alamos, P. Martin s.n. (ARIZ); Rio Santo Domingo (2 km N of Rancho La Brisca), 30°25.5'N, 110°33'W, P. Martin s.n. (ARIZ); 1 km NE of Bachoco, 26°44'N, 109°21'W, P. Martin & M. O’Rourke s.n. (ARIZ); Las Chinacas and vicinity of road to Chinipas, 27°14'N, 108°39'W, P. Martin et al. s.n. (ARIZ); “Agua Amarilla,” ca, 28°27'N, 109°20'W, P. Martin et al. s.n. (TEX); Canyon de las Palmas, Sierra Baviso, 16 mi SE of Magdalena, NV. McCarten 2300 (ENCB); Rte. 15, 18.7 mi N of Imuris, L. McGill & D. Pinkava 6595 (ASU); Mpio. Guaymas, 3 km NE del Aguaje de Robinson, Canon Los Anegados, 28°02'N, 111°10'W, L. Moreno et al. 16 (ARIZ); Mpio. Nacore Chico, El Rio Bonito near La Nopalera, C. Muller 3683 (GH, LL, UC, US); Magdalena, C. Orcutt 1922:1352 (US); Nacopuli Canyon, N of San Carlos, B. Perrill 5658 (ARIZ); Valle de Teras (just E of La Angostura), E. Phillips 761 (MICH); Horconcitas, E. Phillips 837 (MICH); between Horconcitos and Bacerac, E. Phillips 852 (GH, MICH); near Onavas, 4. Rea 49/7 (ARIZ); Onavas graphite mine, ca. 28°28'N, 109°32'W, A. Rea 1260 (ARIZ); northern Sierra Aconchi, 7 mi W of San Felipe, 29°52'N, 110°21'W, F) Reichenbacher 1172 (ARIZ); Mpio. Santa Ana, Arroyo El Papago, 13 km ENE of Benjamin Hill, 30°16'N, 110°52’W, 4. Reina G. & E. Reina G. 97-1609 (CAS); Mpio. Yécora, base of Cerro E] Pilar, above Arroyo Santa Ana, 5.5 km N of Curea on Son. 12, 28°20'N, 109°15'W, A. Reina G et al. 99- 803 (CAS); Mpio. Guaymas, old road to Algodones, San Carlos, 27°58'N, 111°04’W, A. Reina G. & T: Van Devender 2000-915 (CAS); Mpio. Yécora, 1.5 km SW of Santa Ana on road to Guadalupe Tayopa, 28°23’N, 109°09.5'W, A. Reina G et al. 96-521] (ARIZ); Mpio. Imuris, Agua Caliente, 19 km N of Imuris, 30°57'N, 110°51'W, A. Reina G. et al. 2001-653 (CAS); ca. 5 mi below Minas Nuevas, J/. Rose et al. 12672 (NY, US); vicinity of Alamos, J. Rose et al. 12741 (NY, US); vicinity of Puerta de Mano, 26°59'N. 108°39'W, J. Salmon s.n. (ARIZ); 5-7 mi E of Alamos toward upper crossing of Rio Cuchujaqui, ca. 26°59'N, 108°50'W, A. Sanders et al. 8833 (CAS); Mpio. Villa Pesqueira, Sierra Pinta, ca. 33 mi S of Moctezuma, ca. 29°35'N, 110°01'W, J. Shortman & M. Wilson 96-65 (ARIZ); Distr. of Altar, near Los Molinos, F: Shreve 6014 (F); 10 mi SE of Magdalena, F) Shreve 6628 (ARIZ, F, GH, MICH, MO); Cerro las Campas, 2.6 mi ESE of Alamos on road to Rio Cuchujaqui crossing near Sabinito Sur, 27°01'N, 108°54’W, G Starr & D. Palzkill 329 (ARIZ); 7.5 mi W of Alamos, road to Navojoa, G Starr & C. Starr 242 (ARIZ); Mpio. Imuris, microondas 14 km S de Imuris, 30°52'N, 110°S0'W, P. Tenorio L. & C. Romero de T. 13495 (CAS); Mpio. Yécora, La Concepcion, 29°19'N, 109°02'W, W. Trauba s.n. (CAS): Guirocoba crossing of Rio Cuchujaqui, 12.3 km SSE of Alamos, 26°56'N, 108°53'W, T. Van Devender 92-532 (ARIZ); Rio Cuchujaqui at Rancho el Conejo, 12.4 km S of Alamos, 26°55'N, 108°55'W, 7. Van Devender 92-564 (ARIZ): 59 km ESE of Hermosillo on Sonora 16, 7: Van Devender & V. Markgraf s.n. (ARIZ); Mpio. Soyopa, Loma Maderista, 3.5 km S of Tonichi, ca. 28°34’N, 109°33'W, T. Van Devender & A. Reina G. 2001-184 (ARIZ, CAS); El Rancheria crossing of Rio Cuchujaqui, near Tojibampo, 19.5 km NNW of San Bernardo toward Mesa Colorada, 27°02’'N, 108°47'W, T: Van Devender et al. 93-368 (ARIZ, CAS); Mpio. Navojoa, Teachive de Masiaca, 26°47'N, 109°14'W, T. Van Devender et al. 95-105 (ARIZ); 1.5 km SW of Santa Ana on road to Guadallupe Tayopa, 28°23'N, 109°09.5’W, T: Van Devender et al. 97-215 (MEXU); Mpio. Nogales, Sierra Las Avispas, 14.6 km W of MEX 15 on SON 43 (to Saric), 31°07'N, 111°04°E, T. Van Devender et al. 2002-745 (CAS); Palm Canyon, 17.7 mi SE of Magdalena in Cerro Cinta de Plata, 77 Van Devender et al. s.n. (ARIZ); San Javier, cafidn entre los cerros El Potrerito y Los Amoles, 7 km S del poblado, 28°32'N, 109°45'W, L. Varela E. 96-309 (MEXU); San Javier, cerro El DANIEL: SONORAN ACANTHACEAE 799 Verde, 3 km de la carr. 16, 28°34'N, 109°44'W, L. Varela E. 96-456 (MEXU); Puerto de Huépari, NW of Aribabi, S. White 2773 (GH, MICH, US); Cafion de la Petaquilla (just W of Bavispe), S. White 3333 (GH, MICH); Canon del Agua Amarga (E of Oputo), S. White 3612 (MICH); Horconcitos, Rio Huachinera, S. White 3727 (ARIZ, MICH, GH); Colonia Morelos, S. White 4420 (MICH), 4455 (ARIZ, GH, MICH, NY, US); west- ern foothills of Sierra de la Cabellera, ca. 10 mi E of Colonia Morelos, S. White 4470 (ARIZ, GH, MICH, US); 4 mi N of Nuri, 28°10'N, 109°19'W, 4. Whittemore et al. 83-085 (CAS, TEX); 5 mi S of San Rafael, /. Wiggins 3942 (DS, US); 2 mi E of Rancho San Carlos toward Norio, /. Wiggins 6147 (DS, US); 1 mi W of jet. Cananea and Remedios roads on upper ford of Rio Magdalena, /. Wiggins 7096 (DS, MICH, US); Rio Sasabe, 10 mi E of Magdalena, /. Wiggins 7115 (DS, F, LL, MICH, UC, US); ca. 14 mi S of Divisaderos, /. Wiggins 7466 (DS, MICH, US). Tetramerium tenuissimum Rose, Contr. U.S. Natl. Herb 1:349. 1895. TYPE: MEXICO. Colima: vicinity of Colima, 27-28 Feb 1891, E. Palmer 1297 (holotype: US!; isotypes: GH!, K!, NY!, US!). Tetramerium leptocaule Happ, Ann. Missouri Bot. Gard. 24:516. 1937. TYPE: MEXICO. Michoacan: Distr. Huetamo, Tacupa, 17 Jan 1934, G Hinton et al. 5494 (holotype: K!; isotypes: DES!, GH!, MO!, NY!, US!). Phenology. Flowering: September—March; fruiting: September—March. Le Distribution and habitats. Mexico (Cam- De ly peche, Chiapas, Chihuahua, Colima, Guana- juato, Guerrero, Jalisco, Michoacan, Morelos, (3! Nayarit, Sinaloa, Sonora, Veracruz, Yucatan); Sonoran plants occur on slopes, along streams, and in disturbed areas in thornscrub and trop1- cal deciduous forests at elevations from 200 to 920 m. 29° Illustration. None found. Local names. “Rama del toro” (fide Van Devender et al. 2000); “rama toro” (Gentry 1084): “tau-i-we toro” (Guarijio, Gentry 1084). a‘ a @ Tetramerium tenuissimum Daniel (1986, 1995a) and Daniel and [27° 6 Tetramerium yaquianum Acosta (2003) provided descriptions of this species. Until recently, 7etramerium tenuissi- 114° 112° mum was thought to be quite rare. It is now : being found in regions of dry forest throughout Mexico and Central America. It reaches the northern and western extents of its distribution in north-central Sonora (Fig. 41). Sonoran plants of T. tenuissimum, which often grow in close proximity to 7; nervosum and which are often mistaken for it, can be distinguished from that species by the characters in the following couplet: “100 km” FiGURE 41. Sonoran distribution of TYetramerium tenuissimum and known distribution of 7) yaquianum. 1. Calyx 4-lobed; margin of bracts conspicuously ciliate with at least some (usually most) tri- chomes 14 mm long; bracteal apex usually twisted....................... T. nervosum 1. Calyx 5-lobed; margin of bracts + inconspicuously ciliate with trichomes 0.05—0.5 mm long; Bide tec napemmlOrstWASte ees pay. 2 wren apt ee oes ce ce oiecllens mae crepe we lege) eee T. tenuissimum SONORAN COLLECTIONS: along road to El Tezal, 2.6 mi W of Alamos, 7: Daniel 3374 (CAS, ENCB, MEXU); 3384 (ASU, NY, US); Mpio. Yécora, along Hwy. 16, 2.6 km NW of Tepoca, ca. 28°27’N, 109°16'W, 800 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 T. Daniel et al. 8565 (CAS); Mpio. Yécora, between San Nicolas and Nuri, just S of Curea, ca. 24 km SW jet. Hwy. 16, ca. 28°19'W, 109°17'W, T: Daniel et al. 8575 (CAS); Mpio. Yécora, Arroyo La Quema, 0.3 km SE of Tepoca on Hwy. 16, ca. 28°26'N, 109°15'W, T. Daniel et al. 8637 (CAS, MEXU); Canyon Sapopa, Rio Mayo, H. Gentry 1084 (ARIZ, F, GH, MO); Rio El Naranjo crossing N of Taymuco, 27°15'N, 108°43’W, P Jenkins et al. 88-320 (ARIZ, CAS); Mpio. Arizpe, Puente Caliente, Rancho Agua Caliente, ca. 3 km S of Arizpe on Son. 89, 30°20'N, 110°12'W, 4A. Reina G & T. Van Devender 2000-738 (ARIZ, CAS); Mpio. Yécora, Curea, 28°19'N, 109°17'W, A. Reina G. et al. 2001-79 (CAS, MEXU, MO), 2001-80 (CAS); Mpio. Yécora, Arroyo Santa Ana at SON 12 (Tepoca-Cd. Obregon Hwy.), 2.3 km SW of La Quema turnoff, 8.5 km (air) W of Guadalupe Tayopa, 28°21'N, 109°16'W, A. Reina G. et al. 2001-98 (CAS); Mpio. Alamos, Arroyo Mentidero at crossing of Alamos-E] Chinal road and down arroyo to Rio Cuchujaqui, ca. 12 km S of Alamos, ca. 26°55'N, 108°55'W, A. Sanders et al. 12617 (CAS); Mpio. Yécora, Arroyo Santa Ana at Son. 12, 2.3 km SW of turn to La Quema, 28°21'N, 109°16'W, 7: Van Devender & A. Reina G. 97-1060 (ARIZ, CAS, MEXU); Mpio. Soyopa, Loma Maderista, 3.5 km S of Tonichi, ca. 28°34’N, 109°33'W, 7: Van Devender & A. Reina G. 2001-199 (CAS, MO); ca. 8 mi SE of Alamos near Rio Cuchujaqui, 7: Van Devender & R. Van Devender 53-142 (ARIZ, CAS), 54-142 (ASU): T. Van Devender et al. 84-153 (ASU); ca. 13.5 mi S of Alamos, 7. Van Devender et al. 84-191 (ASU); El Guayabo crossing of Rio Cuchujaqui, 14 km ESE of Alamos, 27°00'N, 108°47'W, T. Van Devender et al. 92-257 (ARIZ, CAS), 92-1291 (ARIZ, CAS, TEX); El Rancheria crossing of Rio Cuchujaqui, ca. 22.5 km S of Alamos toward El Chinal, 26°51'N, 108°55'W, 7: Van Devender et al. 92-1192 (ARIZ, CAS, MO, TEX): Cerro las Tatemas below Microondas La Luna, 13.8 km NW of Alamos, 27°07'N, 109°02'W, T: Van Devender et al. 93-338 (ARIZ, CAS); Sierra de Alamos, El Rincon Viejo, Arroyo El] Aguaje, ca. 4 km N of Alamos, 27°04'N, 108°56'W, 7: Van Devender et al. 93-1507 (ARIZ, CAS); Mpio. Soyopa, NE side of Rio Yaqui bridge on Mex. 16, just S of Tonichi, ca. 28°34'N, 109°33'W, T: Van Devender et al. 97-1471 (CAS); Mpio. Bacanora, jct. of Cafiada Jiropa on road from Bacanora to Tonichi, 5.4 km (air) SSW of Bacanora, 28°56'N, 109°25'W, T. Van Devender et al. 99-861 (CAS, TEX); Mpio. Arivechi, Arroyo Agua Caliente, 10.2 km (air) WNW of Arivechi, 28°57'N, 109°17'W, T. Van Devender et al. 99-886 (ARIZ, CAS); Mpio. Yécora, 1.9 km W of Tepoca on Mex. 16, 28°27'N, 109°16'W, 7: Van Devender et al. 2000-61 (CAS); Mpio. Yécora, Curea, 28°19'N, 109°17'W, 7: Van Devender et al. 2001-100 (CAS). Tetramerium yaquianum T.F. Daniel, Syst. Bot. Monogr. 12:102. 1986. TYPE: MEXICO. Sonora: along rd. to Micro. Cerro Prieto, between Navojoa and Alamos, 9.7 mi E of jet. Hwy. 15 in Navojoa, 13 Mar 1984, 7: Daniel 3372 (holotype: CAS!; isotypes: ASU!, MEXU!, MICH!, NY!, US!). Phenology. Flowering: July-December, March; fruiting: September—December, March. Distribution and habitats. Endemic to Sonora where plants occur on slopes and along streams in thornscrub and tropical deciduous forests at elevations from 110—400 m. Illustration. Systematic Botany Monographs 12:76. 1986. Daniel (1986) provided a description of 7etramerium yaquianum. It is the only species of Acanthaceae known to be restricted in distribution to Sonora (Fig. 41), where it can be distin- guished from the morphologically similar 7) nervosum by the characters in the following couplet: 1. Calyx 5-lobed; bracts widely spreading (usually + horizontal), glandular pubescent with glan- dular trichomes 0.2—0.3 mm long, (acute to) rounded to truncate to emarginate at apex, the api- calinortion\ usually flatand not twisted... 0.25.0. 2% ae eee eee eee T. yaquianum 1. Calyx 4-lobed; bracts mostly erect (+ appressed to rachis), not glandular pubescent (or if glan- dular trichomes, if present, these up to 0.1 mm long), acute to acuminate at apex, the apical portionjusually 3: recurved and tasted. ......<- . ~ a. was Saas eee T. nervosum SONORAN COLLECTIONS: between Ures and Hermosillo, 17.1 mi SW of Ures, 7. Daniel 940 (MICH); between Esperanza and Rosario de Tezopaco, 33.3 mi NE jet. Hwy. 15, 7. Daniel 3361 (ASU, CAS, MICH, MEXU, NY, US); western base of Sierra Bojihuacame, 1.2-1.1 mi SE of Bachoco, 7: Daniel 3365 (ASU, CAS, DANIEL: SONORAN ACANTHACEAE 801 ENCB, K, MEXU, MICH, NY, UC, US); El Novillo (E-central Sonora), R. Felger et al. 84-298 (ARIZ): Sierra Bajihuacame, SE of Cd. Obregon, H. Gentry 14530 (ARIZ, LL); summit of Cerro Prieto, 15 km E of Navojoa toward Alamos, ca. 27°15'N, 109°17'W, A. Sanders et al. 9279 (CAS); 37 mi NE of Cajeme on road to Tesopaco, F Shreve 6158 (ARIZ, F); 1.5 mi downstream from La Estrella on Rio Yaqui, T. Van Devender et al. s.n. (ARIZ, ASU); Mesa Masiaca along road to microwave tower, 6.5 km WNW of San José de Masiaca, 26°46'N, 109°18'W, 7. Van Devender et al. 92-1109 (ARIZ, CAS, MO, TEX); Mpio. Alamos, Cerro El Chorro, 4 km NW of Yocogigua, 26°49'N, 109°03'W, T. Van Devender et al. 93-1445 (ARIZ, CAS). ACKNOWLEDGMENTS Many individuals and organizations have assisted me over the past 27 years with my studies of Sonoran Acanthaceae. Foremost among them is Tom Van Devender of the Arizona-Sonora Desert Museum in Tucson. He has sent me duplicates of his many excellent collections of Acanthaceae, specifically searched for (and found!) several critical taxa in the field, allowed me to accompany him to the beautiful highland region near Yécora, shared his knowledge of Sonora and its plant life in correspondence and through his numerous publications, and prodded me to prepare this regional account of Acanthaceae. Sonora is fortunate to have such a devoted admirer and inves- tigator of its natural and cultural resources. The illustrations represent the artistry of Nadia Strasser, Zina Deretsky, Jenny Speckles, and Alan Chou; I am most grateful to each of them. Others who generously provided assistance include: Marc Baker, Mary Butterwick, Richard Felger, Howard Gentry, Phil Jenkins, Elizabeth Moore, Ana Lila Reina G., Jesus Sanchez, Andy Sanders, Victor Steinmann, Alush Ton, and William Trauba. I am most grateful for funding from the National Science Foundation (DEB 78-09321, BSR-8304790, BSR 8609852) and the American Philosophical Society (1572 Johnson Fund) that made several field trips to Sonora possible. Direct or indirect support from Arizona State University, the California Academy of Sciences, and the Arizona-Sonora Desert Museum also helped to make some of the field work possible. Loans or other courtesies were extended from the following herbaria: A, ARIZ, ASU, BM, BR, BRIT, C, @AS. CHARA, DAV, DES, DS; DUKE, ENCB, F, G, GH, EB, K, L,. LA, LE, LL, MA, MEXU, MICH, MIN, MO, MSC, NSW, NY, P, PH, POM, PR, PRC, RM, RSA, S, SD, SMU, TEX, UC, UCR, UNM, US, and WIS. To these organizations and individuals, and to the friendly folks of Sonora, I offer this treatment of Acanthaceae with the hope that it leads to a greater understanding of the plant life in that state. It is my further hope that such understanding will assist efforts to pre- serve the state’s precious natural resources. LITERATURE CITED AMERICAN GEOGRAPHICAL SOCIETY. 1937. Sonora [1:1,000,000 Map]. American Geographical Society, New York. AXxeLRop, D.I. 1979. Age and origin of Sonoran Desert vegetation. Occasional Papers of the California Academy of Sciences (132):1—74. BENTHAM, G., AND J.D. HOOKER. 1876. Genera Plantarum, vol. |. Reeve and Co., London. BRANDEGEE, T.S. 1893. Notes concerning the flora of Sonora. Zoe 3:344—349. Brown, D.E., ed. 1982a. Biotic communities of the American Southwest-United States and Mexico. Desert Plants 4:\-342. [A reprinting of this, with minor corrections, was published by the University of Utah Press in 1994 under the title, Biotic Communities: Southwestern United States and Northwestern Mexico. | Brown, D.E. 1982b. 134.3 Sinaloan thornscrub. Desert Plants 4:101—105. Brown, D.E. 1982c. 143.1 Plains and Great Basin grasslands. Desert Plants 4:115—121. Brown, D.E. 1982d. 143.1 Semidesert grassland. Desert Plants 4:123—131. Brown, D.E. 1982e. 123.3 Madrean evergreen woodland. Desert Plants 4:59-65. Brown, D.E., AND C.H. Lowe. 1980. Biotic Communities of the Southwest [Map]. General Technical Report 802 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 35 RM-78, Rocky Mountain Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture, Tempe, Arizona. DANIEL, T.F. 1982. Anisacanthus andersonii (Acanthaceae), a new species from northwestern Mexico. Bulletin of the Torrey Botanical Club 109:148—151. DANIEL, T.F. 1983a. Carlowrightia (Acanthaceae). 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FILE NAMING PROTOCOLS: To facilitate the handling of digital files submitted by authors, the following file-naming conven- tions are to be followed: text files should bear the author’s last name (in the case of multiple authors, only the first author’s name) followed by a space and a date in the format mmyy (e.g., 0603 for June 2003) to yield a file name such as Gosliner 0603.doc or Williams 0603.rtf. [f an author has submitted two or more manuscripts and must distinguish between them, then the naming should include an additional numeral: Gosliner1 0603.doc for the first manuscript, Gosliner2 0603.doc (or .rtf) for the second. Figures should follow similar conventions, as follows: Gosliner F1 0603.tif, Gosliner F2 0603.tif, for figures in the first manu- script and, if more than one manuscript, then Gosliner1 F1 0603.tif etc. for the figures associated with the first manuscript and Gosliner2 F1 0603.tif etc. for those with the second. Following these conventions will insure that figures submitted by one author are always maintained distinct from those submitted by another. Tables submitted as Excel files should follow the same naming conventions except the file type designation will be “.xls”: e.g., Gosliner T1 0603.xls. Please note that extraneous periods are omitted in file names. BIBLIOGRAPHY FORMAT: Three bibliographic styles are accommodated in the Academy’s scientific publications, one com- monly used in scientific journals publishing papers in systematic and evolutionary biology, a second used mainly in the geologi- cal literature, and lastly, the format most commonly used in the humanities by historians of science. On request, the author will be sent a style sheet that includes samples of the three formats. Authors are also encouraged to examine a copy of the latest pub- lished Proceedings. In all instances, however, authors should not abbreviate journal names but spell them out completely. For books, the reference must include the publisher and city of publication. It is recommended that the total number of pages in the book also be given. SUBSCRIPTIONS ; ; The Proceedings series of the California Academy of Sciences is available by exchange or subscription. For information on exchanges, please contact the Academy Librarian via regular mail addressed to the Librarian, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, U.S.A. or via email addressed to amalley@calacademy.org. Subscription requests, including information on rates, should be addressed to Scientific Publications, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, U.S.A. or via email to the Editor at scipubs@calacademy.org. Subscription price for 2005: $65 (US) plus $10 for mailing to U.S. and Canadian addresses and $18 to all others. The Occasional Papers and Memoirs are not available by subscription. Each volume is priced separately. Occasional Papers, Memoirs, and individual issues of the Proceedings are available for purchase through the Academy Stores (http://store.yahoo.com/casstore/bookstore.html). COMMENTS The California Academy of Sciences also publishes, on an irregular basis, Occasional Papers and Memoirs. Prices of these publications, which are not available by subscription, are available on request. Address editorial correspondence or requests for pricing information to the Editor, Scientific Publications Office, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, U.S.A. or via email to the Editor, Scientific Publications, at scipubs@calacademy.org. 1A wii 88 TABLE OF CONTENTS ANGEL VALDES AND YOLANDA E. CAMACHO-GaRCiA: “Cephalaspidean” Heterobranchs (Gastropoda) from the Pacific Coast of Costa Rica DAvID W. GREENFIELD AND JOHN E. RANDALL: The Marine Gobies of the Hawaiian Islands. . 498 ALICIA HERMOSILLO AND ANGEL VALDES: Two New Species of Dorid Nudibranchs (Mollusca, Opisthobranchia) from Bahia de Banderas, Mexico DaAvID W. GREENFIELD AND JOHN E. RANDALL: Two New Cardinalfish Species of the Genus Apogon from Easter Island Davip W. GREENFIELD AND WM. LEO SMITH: Allenbatrachus meridionalis, a New Toadfish (Batrachoididae) from Madagascar and Reunion R.C. DREWES AND R.E. STOELTING: The California Academy of Sciences Gulf of Guinea Expedition (2001) II. Additions and Corrections to Our Knowledge of the Endemic Island Amphibians of S40 Tomé and Principe TERRENCE M. GOSLINER, JESUS ORTEA, AND ANGEL VALDES: New Data on Tropical Eastern Pacific Chromodorididae (Nudibranchia: Doridina) with Description of a New Species of Mexichromis Bertsch, 1977 GUILHERME MURICY AND JOHN S. PEARSE: A New Species of Oscarella (Demospongiae: Plakinidae) from California, with a Discussion of Fixation Methods SHIREEN J. FAHEY AND TERRENCE M. GOSLINER: A Phylogenetic Analysis of the Aegiridae Fischer, 1883 (Mollusca, Nudibranchia, Phanerobranchia) with Descriptions of Eight New Species and a Reassessment of Phanerobranch Relationships THOMAS H. DANIEL: Sonoran Acanthaceae of Sonora: Taxonomy and Phytogeography