PROCEEDINGS of the California Academy of Sciences (Series 4) Copyright © 2016 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 Publisher: Shannon Bennett, Ph.D. Chief, Institute for Biodiversity Science & Sustainibility California Academy of Sciences Editorial Board A lan E. Leviton, Ph.D., Editor Katherine Piatek, M.A., Managing Editor Michael T. Ghiselin, Ph.D., Associate Editor Tomio Iwamoto, Ph.D., Associate Editor Gary C. Williams, Ph.D., Associate Editor & Website Coordinator Michele L. Aldrich, Ph.D., Consulting Editor Cover image Farallon Islands See G. C. Williams & O. Breedy, New species of Whip-Like Gorgonian Coral in the Genus Swiftia from the Gulf of the Farallones, California Photo by Gary C. Williams Cover Design Gary C. Williams & Alan E. Leviton California Academy of Sciences ISSN 0068-547X “Pnoceedbifi t£c faUfemte Amdemy S cien c e s is an international journal that accepts manuscripts for publication in the Natural Sciences and selected areas in the Earth Sciences, such as bios¬ tratigraphy, 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 institutions of natural his¬ tory, to individuals, and to activities, such as expeditions and explorations, in the natural sciences. All manuscripts submitted for publication in any of the Academy's scientific publication series {Pro¬ ceedings, Occasional Papers, Memoirs ) are subject to peer review. Peer review includes both internal and external review, internal review by at least one Academy scientist whose interests parallel those of the sub¬ mission, and external review, ordinarily by two individuals who are recognized scholars in the field. Manuscripts accepted for publication are subject to page charges; charges may be waived on a case- by-case basis. Published by the California Academy of Sciences 55 Music Concourse Drive, Golden Gate Park, San Francisco, California 94118 U.S.A. Printed in the United States of America by Allen Press Inc., Lawrence, Kansas 66044 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 1, pp. 1-13, 9 figs. April 29, 2016 A New Species of Whip-Like Gorgonian Coral in the Genus Swiftia from the Gulf of the Farallones in Central California, with a Key to Eastern Pacific Species in California (Cnidaria, Octocorallia, Plexauridae) Gary C. Williams 13 , Odalisca Breedy 2 1 Department of Invertebrate Zoology and Geology, California Academy of Sciences, Golden Gate Park, 55 Music Concourse Drive, San Francisco, California 94118, USA. 2 Centro de lnvestigacion en Estruc- turas Microscopicas, Centro de lnvestigacion en Ciencias del Mary Limnologia, Escuela de Biologia, Universidad de Costa Rica. P.O. Boxll501-2060, Universidad de Costa Rica, San Jose, Costa Rica. Smithsonian Tropical Research Institute, P.O. Box 0843-03092, Panama, Republic of Panama. 3 Corresponding Author: Gary C. Williams ( gwilliams@calacademy.org ) A new species of plexaurid gorgonian coral in the genus Swiftia , collected by Remote¬ ly Operated underwater Vehicle, is described from the Greater Farallones National Marine Sanctuary near the Farallon Islands off the coast of central California. The species is unlike other species of Swiftia in that it is whip-like and unbranched or Y- shaped with two terminal branches. The genus Swiftia has a wide geographic distri¬ bution - not associated with coral reefs — it is known from the eastern Pacific and the western and eastern Atlantic, as well as parts of the Indo-West Pacific, from mesophotic depths and in deeper water. Keywords: Alcyonacea, Swiftia, plexaurid gorgonian, central California, Farallon Islands, outer continental shelf, taxonomic key to local species. A previously undescribed species of the gorgonian genus Swiftia was recently encountered during an exploratory survey by Remotely Operated underwater Vehicle (ROV) on board the National Oceanic and Atmospheric Administration (NOAA) ship R/V Fulmar in the then proposed expansion area of the Gulf of the Farallones National Marine Sanctuary (now known as the Greater Farallones National Marine Sanctuary) off the Sonoma County coast of northern California, USA. A single whole colony was collected by ROV, which is here designated as the holotype specimen. The gorgoniid genus Swiftia Duchassaing & Michelotti, 1864 is comprised of 15 currently rec¬ ognized species, widely-distributed in the eastern Pacific — the western Atlantic, the northeastern Atlantic, the northern Pacific, and the Indo-Pacific (Ofwegen 2015; Breedy et al. 2015). The new species described here makes a total of 16 species currently recognized as valid that comprise the genus. Most species of Swiftia vary in depth from approximately 70-732 m with a few species record¬ ed from as shallow as 18-30 m (Breedy et al. 2015) and as deep as at least 1829 m (Goldberg 2001; and CASIZ collections data base). 1 2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 1 Materials and Methods The material examined in this study is housed in the collections of the Department of Inverte¬ brate Zoology and Geology at the California Academy of Sciences (CASIZ), San Francisco. In situ photographs were taken by Peter Etnoyer during the NOAA ROV cruise on board RV Fulmar in the Greater Farallones NMS (National Oceanic and Atmospheric Administration 2014), approxi¬ mately 70 miles NW of the Farallon Islands (Fig. 1), on 6 September 2014. Photographs of pre¬ served material were taken by the authors at the California Academy of Sciences in September of 2015. The holotype was collected by ROV at 182 m in depth. Abbreviations used in the text are: CASIZ (California Academy of Sciences, Department of Invertebrate Zoology), GFNMS (Gulf of the Farallones National Marine Sanctuary), NOAA (National Oceanic and Atmospheric Adminis¬ tration), and ROV (Remote Operational Vehicle). Definitions recognized in this paper for depth zones of benthic faunal communities are as fol¬ lows: shallow-water (< 40 m), mesophotic (40-150 m), and deep-water (> 150 m). Figure 1. The Farallon Islands, approximately 40 miles west of San Francisco, California, and 70 miles southeast of the type locality of Swiftia farallonesica sp. nov., in the Greater Farallones National Marine Sanctuary. WILLIAMS & BREEDY: NEW SPECIES OF GORGONIAN CORAL 3 Systematic Account Subclass Octocorallia Order Alcyonacea Lamouroux, 1812 Family Plexauridae Gray, 1859 Swiftia Duchassaing and Michelotti, 1864 Swiftia Duchassaing and Michelotti, 1864:13; Kiikenthal, 1924:236; Deichmann, 1936:185-186; Bayer, 1956:F206; Bayer, 1981:945; Harden, 1979:109-110; Breedy, et ah, 2015:329. Stenogorgia Verrill, 1883:29 (see Kiikenthal, 1924:347 for Stenogorgia synonymy). Platycaulos Wright and Studer, 1889:61: Bayer, 1981:945. Callistephanus Wright and Studer, 1889:62: Bayer, 1981:945. Allogorgia Verill, 1928:7; Bayer, 1981:945. Generic diagnosis. — Colonies unbranched, or sparsely to copiously branched. Branches mostly free or with some anastomoses. Polyp mounds conical, prominent, or slightly raised, scat¬ tered or crowded, usually biserial or present on all sides of polyp-bearing branches. Anthocodiae with points arrangements of straight to curved bar-like rods, or frequently elongate tuberculated spindles. Type species. — Swiftia exserta (Ellis and Solander, 1786) Swiftia farallonesica Williams and Breedy, sp. nov. Figures 1-9. Leptogorgia, National Oceanic and Atmospheric Administration, 2014:1-2. Holotype.— CASIZ 196930. Type locality (Figs. 8-9).— Deep-sea rocky substrata at Football Shoal, Greater Farallones National Marine Sanctuary, off Bodega Bay, California, USA (38.00°26.33'N 123.00°34.19W); 182 m depth; 06 September 2014; Gary Williams and Peter Etnoyer; one entire specimen collect¬ ed by ROV. Habitat and distribution.— Found on hard, rugose, horizontal substrata composed of rela¬ tively dense congregations of detritus-covered rounded boulders between 181 and 190 m depth, at the type locality and vicinity (Figs. 8-9). Pink and white ophiuroids (species unidentified) were observed (on the holotype and by underwater video and still imagery) attached along portions of the lengths of several colonies (Figs. 2A & D, 3A & E). Approximately 15-20 colonies were observed in total by underwater video and still photography. Etymology.— The specific epithet is derived from the Spanish farallon (steep rock, cliff, headland, outcrop), and the Latin -icus (suffix meaning belonging to); referring to the region of the discovery of the new species — the Greater Farallones National Marine Sanctuary. Description of the Holotype External morphology (Figs. 2-3).— The proximal-most surface of the holdfast that adheres to the substratum is circular, approximately 10 mm in diameter and 2 mm in height. The colony is unbranched and whip-like, 378 mm in length by 3-4 mm in width. The external surface of the colony is pustular in appearance with retracted polyps forming low mounds on all sides of the branches, from the holdfast to the apex of the colony. Each protuberance is approximately 2.0 4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 1 mm in diameter at their bases and 0.8-1.0 mm in height. There are approximately 25-30 polyps per every 10 mm along the length of the colony. Many of the polyps throughout the colony are entirely retracted, while others are preserved in varying states of partial retraction. Sclerites (Figs. 4-7).— The sclerites of the polyp mounds and coenenchyme are predomi¬ nantly double discs and disc spindles 0.05-0.08 mm in length (Fig. 4), eight-radiates 0.04-0.12 mm long (Fig. 5), and spindles and girdled spindles (0.08-0.20 mm in length). Some girdled spindles are wider toward one end compared to the opposite end, and thus appear somewhat club-shaped in overall appearance (Fig. 6). Sclerites of the anthocodiae are mostly elongated rods with parallel sides and are variably tuberculated, 0.09-0.21 mm long (Fig. 7). All sclerites in the colony are col¬ orless. Color (Figs. 2-3).— The color of the holotype is white throughout, both in life and wet-pre¬ served. Remarks (Figs. 2-3).— All colonies observed are whip-like and unbranched, except for one colony that was recorded by underwater still imagery and is Y-shaped with a single lateral branch (Fig. 3C). All colonies that were observed by underwater video or still imagery are also uniformly white in color (Fig 3A-E). Discussion and Conclusion Key to Swiftia species in California la. Colonies multiply-branched.2 lb. Colonies unbranched (flagelliform, whip-like),or Y-shaped with a single branch; with low polyp mounds distributed on all sides of branches; colonies pure white in color. . Swiftia farallonesica Williams and Breedy, sp. nov. 2a.Colonies mostly dichotomously branched, somewhat fan-shaped; with low polyp mounds dis¬ tributed on all sides of branches; colonies pink to bright coral red in color. . Swiftia spauldingi (Nutting, 1909) 2b.Colonies densely branched with prominent polyp mounds distributed biserially along branches; colonies red with deep red polyp mounds. Swiftia kofoidi (Nutting, 1909) Taxonomic Assessment Swiftia is one of 43 currently recognized genera in the gorgonian family Plexauridae (Williams and Cairns 2015). Plexaurids are characterized by a darkly colored axis (black or brown), which is composed of a cortex of proteinaceous material (gorgonin) with spaces (loculi) that are sometimes filled with lighter calcified material. The central core of the axis is hollow and cross-chambered (Fabricius and Alderslade 2001:59; Williams 2005:54, Fig. 1). The genus Swiftia is distinguished by the following combined set of characters: coenenchymal sclerites are largely capstans, many of which are modified to a lesser or greater degree as discs; polyp mounds low-rounded to cylindrical or conical, slightly raised to prominent; anthocodiae con¬ tain relatively large, conspicuously tuberculated rods (based on Bayer 1981:932). WILLIAMS & BREEDY: NEW SPECIES OF GORGONIAN CORAL 5 Figure 2. Swiftiafarallonesica sp. nov. Wet-preserved holotype. A, D. Entire colony, scale bars = 10 mm. B. Distal apex region, scale bar = 5 mm. C. Middle portion of colony, scale bar =10 mm. 6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 1 Figure 3. Swiftia farallonesica sp. nov. Underwater photographs taken in situ by ROV, between 181 and 190 m in depth, 6 September 2014. WILLIAMS & BREEDY: NEW SPECIES OF GORGONIAN CORAL 7 Figure 4. Swiftia farallonesica sp. nov. Scanning electron micrographs of coenenchymal sclerites — double discs and disc-spindles. Scale bar = 0.03 mm. 8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 1 Figure 5. Swiftia farallonesica sp. nov. Scanning electron micrographs of coenenchymal sclerites — eight radiates. Scale bar = 0.03 mm. WILLIAMS & BREEDY: NEW SPECIES OF GORGONIAN CORAL 9 Figure 6. Swiftia farallonesica sp. nov. Scanning electron micrographs of coenenchymal sclerites — spindles. Scale bar = 0.03 mm. 10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 1 Figure 7. Swiftia farallonesica sp. nov. Scanning electron micrographs of anthocodial sclerites. Scale bar = 0.03 mm. WILLIAMS & BREEDY: NEW SPECIES OF GORGONIAN CORAL 11 Figure 8. Map of the Greater Farallones National Marine Sanctuary (central California, USA), showing type locality of Swiftia farallonesica sp. nov. (represented by red triangle). Map modified from National Oceanic and Atmospheric Admin¬ istration (2014). 12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 1 Figure 9. Map showing the known distribution of the genus Swiftia in North America and northern South America. Arrow shows type locality of Swiftia farallonesica sp. nov. Acknowledgments We are indebted to the staff members of the Greater Farallones National Marine Sanctuary and the National Oceanic and Atmospheric Administration, for the opportunity to participate in ROV cruises on board R/V Fulmar in the GFNMS during 2012 and 2014. In particular, we thank Maria Brown, Danielle Lipski, Jan Roletto, Peter Etnoyer, and Dave Minard for their support. We are grateful to Jei-Ying Chen, California Academy of Sciences, for making many of the scanning elec¬ tron micrographs, and Christina Piotrowski for curation of invertebrate material collected during the cruises. Literature Cited Bayer, F.M. 1956. Octocorallia. Page 206 in R.C. Moore, ed., Treatise on Invertebrate Paleontology Part F. Coelenterata. The University of Kansas Press and the Geological Society of America and University of Kansas Press, Lawrence, Kansas, USA. 498 pp. Bayer, F.M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata, Anthozoa), with diagnoses of new taxa. Proceedings of the Biological Society of Washington 94(3):902-947. Breedy, O., S.D. Cairns, and V. HAussermann. 2015. A new alcyonacean octocoral (Cnidaria, Anthozoa, Octocorallia) from Chiliean fjords. Zootaxa 3919 (2):327-334. WILLIAMS & BREEDY: NEW SPECIES OF GORGONIAN CORAL 13 Deichmann, E. 1936. The Alcyonaria of the western part of the Atlantic Ocean. Memoirs of the Museum of Comparative Zoology> at Harvard College 53:1-317. Duchassaing, P., and H. Michelotti. 1864. Supplement au memoire sur les coralliaires des Antilles. Mem- orie della Accademia della Scienze di Torino (2) 23:97-206. Ellis, J., and D. Solander. 1786. The Natural History ; of Many Curious and Uncommon Zoophytes, Col¬ lected by the Late John Ellis and Systematically Arranged and Described by the Late Daniel Solander. Benjamin White and Son, London, England. 208 pp. Fabricius, D., and P. Alderslade. 2001. Soft corals and sea fans — a comprehensive guide to the tropical shallow water genera of the central-west Pacific, the Indian Ocean and the Red Sea. Australian Institute of Marine Science, Townsville, Australia. 264 pp. Goldberg, W.M. 2001. The sclerites and geographic distribution of the gorgonian Swiftia exserta (Coelen- terata: Octoorallia: Holaxonia). Bulletin of the Biological Society of Washington 10:100-109. Harden, D.G. 1979. Intuitive and numerical classification of east Pacific Gorgonacea (Octocorallia). Unpub¬ lished Ph.D. dissertation, Illinois State University. 214 pp. Kukenthal, W. 1924. Gorgonaria. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer ‘Valdivia’, 1898-1899, 13(2): 1-946. National Oceanic and Atmospheric Administration. 2014. Scientists on NOAA-led mission discover new coral species off California, http://www.noaanews.noaa.gov/stories2014/ 20141105_deepseacoral.html Last updated November 05, 2014. Nutting, C.C. 1909. Alcyonaria of the Californian coast. Proceedings of the U.S. National Museum 35( 1658): 681-727. Ofwegen, L. Van. 2015. Swiftia Duchassaing & Michelotti, 1864. Accessed through: World Register of Marine Species at on 2016-02- 17. Verrill, A.E. 1928. Hawaiian shallow water Anthozoa. Bernice P. Bishop Museum Bulletin 49:1-30. Verrill, A.E. 1883. Report on the Anthozoa, and on some additional species dredged by the “Blake” in 1877- 1879, and by the U.S. Fish Commission steamer “Fish Hawk” in 1880-82. Bulletin of the Museum of Comparative Zoology Harvard 11:1-72. Williams, G.C. 2005. New taxa of octocorals (Anthozoa: Octocorallia) from the northeastern Pacific Ocean. Proceedings of the California Academy of Sciences, ser. 4, 56(5):53-65. Williams, G.C., and S.D. Cairns. 2015. Systematic list of valid octocoral genera. . Last updated December 16, 2015. Wright, E.P., and T. Studer. 1889. Report on the Alcyonaria collected by H.M.S. Challenger during the years 1873-1876. Report of the Scientific Results of the Voyage of H.M.S. Challenger 1873-1876, Zoology, 31:1-314. 14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63 Page intentionally left blank PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2, pp. 15-61, 19 figs., Appendix April 29, 2016 Herpetological Survey of Iona National Park and Namibe Regional Natural Park, with a Synoptic List of the Amphibians and Reptiles of Namibe Province, Southwestern Angola Luis M. P. Ceriaco f 2 ’ 8 , Sango dos Anjos Carlos de Sa 3 , Suzana Bandeira 3 , Hilaria Valerio 3 , Edward L. Stanley 2 , Arianna L. Kuhn 4 ’ 5 , Mariana P. Marques f Jens V. Vindum 6 , David C. Blackburn 2 , and Aaron M. Bauer 7 1 Museu National de Histdria Natural e da Ciencia, Universidade de Lisboa, Rua da Escola Politecnica, 58, 1269-102 Lisbon, Portugal. 2 Department of Natural History, Florida Museum of Natural History, Gainesville, FL, 32611, USA. 3 Instituto National da Biodiversidade e Areas de Conservaqao, Ministerio do Ambiente de Angola, Centralidade do Kilamba, Rua 26 de Fevereiro, quarteirao Nimi ya Lukemi, edlficio Qll, 3° andar, Angola. 4 American Museum of Natural Histoiy, Central Park West at 79th Street, New York, New York 10024, USA. 5 City University of New York, Graduate Center, 365 5th A ve., New York, New York, 10016, USA. 6 Department ofHerpetology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118, USA. 1 Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085-1699, USA. 8 Corresponding author: Luis M. P. Ceriaco, Email address: luisceriaco@netcabo.pt Namibe Province is the southernmost province of Angola and, as the result of sever¬ al expeditions from the nineteenth century to the present, it is one of the most her- petofaunally well-known areas of the country. The Province harbors a high diversi¬ ty of amphibians and reptiles, including roughly one-third of the reptile taxa report¬ ed for Angola as a whole. In this paper we present the results of a joint herpetologi¬ cal expedition to Namibe Province in 2013 by the California Academy of Sciences r and the Instituto Nacionalda Biodiversidade e Areas de Conservaqao, as well as a synoptic list of all the herpetological bibliographic records for the taxa known from the Province. A total of 37 herpetological taxa was collected, including at least three (then) undescribed species, two new country records, and new records for rarely cited taxa in Angola. These taxa belong to four amphibian genera and 15 reptile gen¬ era. Species accounts are provided for each of the species collected. We also highlight biogeographic patterns, conservation issues, and possible future paths for the explo¬ ration and knowledge of the herpetofauna of Namibe. A provincia do Namibie situa-se no sudoeste de Angola e e uma das mais conhecidas relativamente a sua herpetofauna. Este conhecimento e resultado de varias expe¬ dites realizadas desde o seculo XIX ate aos dias de hoje. A provincia alberga uma espetacular diversidade de anfibios e repteis, que para estes ultimos representa aproximadamente a um terqo dos taxa que ocorrem no pais. Neste artigo apresenta- mos os resultados da expediqao herpetologica levada a cabo pela California Acade¬ my of Sciences e o Instituto Nacional da Biodiversidade e Areas de Conservaqao em 2013, bem como uma lista sinoptica de todos os registos bibliograficos para os taxa conhecidos na provincia. Um total de 37 taxa de anfibios e repteis foram colectados, incluindo pelo menos tres especies novas (uma ja descrita e as outras em processo de descriqao), dois novos registos de especies para o pais, bem como o registo de espe- 15 16 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 cies raramente citadas para o pais. Estes taxa pertencem a 4 generos de anfibios e a 15 generos de repteis. Todos os resultados sao apresentados em fichas taxonomicas. Sao ainda apresentados algums comentarios relativos a padroes biogeograficos e questoes ligadas a conserva^ao, e futuros caminhos para a explora^ao e conheci- mento da herpetofauna da provincia do Namibe. Keywords: amphibians, Angola, biogeography, conservation, geographic distribution, Namibe Province, reptiles. The current knowledge of Angola’s herpetofauna is incom¬ plete in contrast to neighboring countries such as Namibia (Her¬ rmann and Branch 2013, Mar¬ ques 2015). Namibe is Angola’s southwesternmost province and is one of the better explored provinces in terms of herpetolog- ical diversity (Branch et al. 2014). Namibe Province occu¬ pies an area of 57,097 km 2 and is bordered by Huila Province to the northeast, Cunene Province to the southeast, Benguela Province to the north, Namibia to the south, and the Atlantic Ocean to the west. The province is geo¬ graphically separated from Huila by the great escarpment of Serra da Leba and Chela, which sepa¬ rates the lower elevation areas of the Namib Desert from the Huila Plateau. Topographically, the majority of the province has an elevation lower than 500 m, ris¬ ing to 1500 m at the escarpment in the east. The highest elevation is at the Serra da Neve inselberg (2403 m) in north of the province, almost at the border with Benguela. The province has two main conservation areas - Iona National Park (INP), the largest conservation area in the country with an area 15,150 km 2 , and the smaller Namibe Region¬ al Natural Park (NNP), with an area of 4,450 km 2 . Namibe lies within the African southwest arid biome, mainly comprising Kaokoveld desert, Namibian savanna, miombo woodlands, and mopane forest. The Kaokoveld desert, which extends along the coastal regions from southern Benguela Province to the Skeleton Coast in Namibia, is mostly dominated by sandy dunes and the occasion¬ al presence of Odyssea paucinervis, Sporobolus spicatus, and Acanthosicyos horridus dominated Figure 1. Map of the main sampled localities: A) Morro do Soba; B) Omauha Lodge; C) Rio Curoca; D) Entrada do Iona; E) Campo das espin- heiras; F) Iona; G) Ford car Wreck; El) Pediva; I) Tambor; J) Serra da Leba; K) Pico do Azevedo; L) Caraculo; M) Praia do navio. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 17 vegetation. The Namibian savanna woodlands in the central areas of the province are dominated by herbaceous plants of the genera Aristida and Eragrostis, dispersed shrubs of Acacia, Com¬ miphora and Combretum, and, towards the southwest, extensive populations of Welwitschia mirabilis. The eastern portions of the province support smaller areas of Angolan mopane wood¬ lands, which are dominated by the deciduous tree Colophospermum mopane, and Angolan miom- bo woodlands, which are dominated by Brachystegia trees, but also typified by Isobertlinia angolensis, Julbernardia paniculata and Baikiaea plurijuga (Grandvaux-Barbosa 1970). The province is bounded in the south by the Cunene (Kunene) River, and crossed by the Curoca and Giraul Rivers. Geologicaly, the south of the province is mostly dominated by schists, sometimes interspersed with granites, while the north of the province is mainly comprises granites (Anony¬ mous 1963). Approximately 16 species of amphibians and 95 species of reptiles are known from Namibe Province (Table 1). In separate works we are preparing a complete review of the diversity and dis¬ tribution (including an annotated checklist) of the Angolan herpetofauna based published biblio¬ graphic records prior to 2014 (Marques et al., in prep.), and an annotated checklist of the herpeto¬ fauna of Namibe Province, based on bibliographic material, unpublished museum records, and recent collections (Branch et al., in prep). This paper presents the results of an expedition con¬ ducted by a team from the California Academy of Sciences (CAS), San Francisco (USA), Villano- r va University (VU), Villanova (USA), and the Instituto Nacional da Biodiversidade e Areas de Conserva^ao (INBAC), Kilamba-Kiaxi (Angola). A total of 37 herpetological taxa were collected, including at least three new species, one of which has recently been described (Stanley et al. 2016), two new country records and new records for taxa rarely cited for the country. We then provide a brief discussion of the present status and future prospects for the study of the herpetofauna of Namibe. History of the Herpetological Exploration of the Province Namibe Province was explored by several well documented expeditions in the nineteenth and twentieth centuries. The first herpetological surveys conducted were those of the Portuguese explorer Jose de Anchieta (1832-1897), who visited the region in the late nineteenth century. Anchieta’s specimens were deposited in the Natural History Museum of Lisbon and largely stud¬ ied by the Portuguese zoologist Jose Vicente Barbosa du Bocage (1823-1907), who published sev¬ eral papers on this material (e.g., Bocage 1867, 1873, 1896). Based on material from Anchieta, as well as others, Bocage described several herpetological taxa from Namibe Province, including Anchieta’s Tree Frog, Leptopelis anchietae (Bocage, 1873), the Double-scaled Chameleon, Chamaeleo anchietae Bocage, 1872, Anchieta’s Ground Agama, Agama anchietae Bocage, 1896, Anchieta’s Shovel-snout Lizard, Meroles anchietae (Bocage, 1867), the Reticulate Sand Lizard, Meroles reticulatus (Bocage, 1867), the Speckled Sand Skink, Trachylepis punctulata (Bocage, 1872), the Speckled Western Burrowing Skink, Typhlacontias punctatissimus (Bocage, 1873), Anchieta’s Worm-Lizard, Monopeltis anchietae (Bocage, 1873), a Skaapsteker, Psammophylax occelatus Bocage, 1873 (currently a synonym of the Spotted Skaapsteker, Psammophylax rhombeatus (Linnaeus, 1758)), a new variety of Striped Sand Snake, Psammophis sibilans var. stenocephalus (Bocage, 1877), and a new species of Giant Blind Snake, Onychocephalus petersii Bocage, 1873 that is currently considered a synonym of Afrotyphlops schlegelii (Bianconi, 1847). At the dawn of the twentieth century, the Portuguese explorer Francisco Newton (1864-1909) collected herpetological specimens in the province for the Natural History Museum of the Poly¬ technic University of Porto, during a three year mission from 1903 to 1905, exploring the provinces 18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 of Kwanza-Norte, Kwanza-Sul and Namibe. The initial collections made by Newton in the two first provinces were studied and published upon by the Portuguese zoologist Jose Julio Bettencourt Fer¬ reira (1866-1948) on two different occasions (Ferreira 1906, 1906), but the Namibe Province mate¬ rial had remained unstudied until today (Ceriaco et al., in prep.). In 1925, the Vemay Angola Expe¬ dition explored central and southwestern Angola and collected specimens destined for the Ameri¬ can Museum of Natural History. The herpetological results of this expedition were partly published upon by the American herpetologist Charles M. Bogert (1908-1992), in a paper dealing with the snakes (Bogert 1940). A second paper detailing the rest of the herpetological material collected on the expedition was never published. Some years later, two Swiss scientific expeditions to Angola, 1928-1929 and 1932-1933, led by the Swiss naturalist Albert Monard (1886-1952) also explored several locations in Namibe Province. The herpetological results of these expeditions were pub¬ lished in four different papers (Monard 1931, 1937a, b, 1938). In between the two Swiss expedi¬ tions, the Pulitzer-Camegie Museum Expedition to Angola in 1930, led by influential American publisher Ralph Pulitzer (1879-1939) and conducted by Wilfrid Rudyerd Boulton (1901-1983) and his wife Laura Crayton Boulton (1899-1980), became one of the most important expeditions in terms of herpetological results. The material was studied and published upon by Karl Patterson Schmidt (1890-1957) in two papers — one dedicated to the reptiles (Schmidt 1933) and other to the amphibians (Schmidt 1936). These works resulted in the description of Pulitzer’s Thick-toed Gecko, Chondrodactylus pulitzerae (Schmidt, 1933), Boulton’s Namib Day Gecko, Rhoptropus boultoni Schmidt, 1933, and the Angolan endemic subspecies of White-Throated Monitor, Varanus albigularis angolensis Schmidt, 1933. More recently the Belgian herpetologist Raymond Laurent (1917-2005) published on a collection of amphibians and reptiles from Namibe (Laurent 1964) sent to him by the Portuguese entomologist and director of the former Museu do Dundo (northeast Angola), Antonio Barros de Machado (1912-2002). This contribution was of uttmost importance for the knowledge of the southwestern Angolan herpetofauna. In addition to the several new taxa added to the list of the provincial herpetofauna, he described four new taxa endemic to the south¬ west of the country — Bogert’s Speckled Western Burrowing Skink, Typhlacontias bogerti Lau¬ rent, 1964, Hellmich’s Wolf Snake, Lycophidion hellmichi Laurent, 1964, and two Namib Day Geckos, Rhoptropus taeniostictus Laurent, 1964 and R. boultoni montanus Laurent, 1964. Wulf Haacke conducted the last systematic field surveys in the Portuguese colonial period in 1971 and 1974 and deposited his collections in the Ditsong National Museum of Natural History (TM) in Pretoria, South Africa, though this material has not been fully published upon. After independence in 1975, Angola entered a long period of civil war, which ended only in 2002. This prevented fur¬ ther field surveys and studies. In the past decade, several field surveys have been conducted, including in Namibe. Teams from the Porth Elizabeth Museum — Bayworld (PEM) prospected the province in three different expeditions so far, and a team from CAS, INBAC and VU conducted the survey reported in this paper. These expeditions have increased our knowledge of the southern Angolan herpetofauna, and since 2008 five new taxa from southwestern Angola (2008-2013) have been described — the Chela Mountain Reed Frog, Hyperolius chelaensis Conradie, Branch, Measey and Tolley, 2012, the Namib Spiny Tailed Gecko, Afrogecko plumicaudus Haacke, 2008, subsequently made the type species of the monotypic genus Kolekanos Heinicke, Daza, Green- baum, Jackman and Bauer, 2014, Haacke’s Sand Lizard, Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012, Huntley’s Sand Lizard, Pedioplanis huntleyi Conradie, Measey, Branch and Tolley, 2012, and the Kaokoveld Girdled Lizard, Cordylus namakuiyus Stanley, Ceriaco, Ban- deira, Valerio, Bates and Branch, 2016 — all but the first endemic to Namibe Province. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 19 Material and Methods We conducted herpetological surveys in Namibe Province from 28 November to 11 December 2013, including both Iona National Park and Namibe Regional Natural Park. A total of 13 areas were surveyed (Fig. 1). In each area, we attempted to sample a combination of habitat types. Over¬ all conditions during this fieldwork were hot and dry as this was an unusually dry year. We cap¬ tured specimens using long-nooses, rubber bands, or by hand during both diurnal and nocturnal visual surveys. All specimens were euthanized following an approved IACUC protocol (#2014-2), preserved in 10% buffered formalin in the field, and then transferred to 70% ethanol for storage. Liver tissues were preserved in 95% ethanol and RNALateR. Voucher specimens and tissue sam¬ ples are deposited in the herpetological collection of the California Academy of Sciences, with a subset of specimens deposited in the reference collection of INBAC. In some cases, we further con¬ firmed species identifications by sequencing the mitochondrial 16S ribosomal RNA gene. As noted above, a complete list of all amphibians and reptile species reported from Namibe Province was assembled (Table 1). This list, including localities and associated bibliographic references was based on the ongoing project for the first atlas of the Angolan amphibians and reptiles (Marques 2015; Marques et al., in prep.). We do not include in the list unpublished museum records such as the large series in the Ditsong National Museum of Natural History collected by Wulf Haacke in the 1970s or the recent collections made by William R. Branch and Werner Conradie from the Port Elizabeth Museum (Bayworld). These specimens will be a part of a forthcoming publication (W.R. Branch, pers. comm.). However, museum material representing taxa vouchered on our expedition are noted when relevant in the species accounts. Results A total of 411 specimens were collected during the expedition, representing four amphibian genera and 15 reptile genera. In the following species accounts, we provide information on CAS voucher specimens, localities, and natural history. Latitude, longitude and elevation (in meters) of the collection site are provided in each species account. In addition, when appropriate, we provide brief taxonomic or geographic notes. Species Accounts Amphibia Anura Bufonidae Hallo well’s Toad Sclerophrys maculata (Hallowell, 1854) Material.— Leba Pass, between river and highway, 5 December 2013, 15°04'13.2"S, 13°14'37.7"E, 1676 m (CAS 254877-254878). Comments. — In Angola this species mostly occurs in the southwestern provinces of Namibe, Benguela, Bie, and Huila (Marques 2015). The nearest records are in “Cainde” and “16 km W of Vila Nova” (Poynton and Haacke 1993; Ruas 1996, 2002). It is widespread in arreas to the south, including much of northern Namibia (du Preez and Carruthers 2009). Ohler and Dubois (2016) recently presented evidence identifying the type species of Sclerophrys Tschudi, 1838 as referable to Bufo rangeri Hewitt, 1935, thus making Sclerophrys the oldest available name for the clade of African toads recently referred to as Amietophrynus Frost et al., 2006. 20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Pyxicephalidae Angola River Frog Amietia angolensis (Bocage, 1866) Material.— Leba Pass, between river and highway, 5 December 2013, 15°4T2.2"S, 13°14'38.9"E, 1676 m (CAS 254876). Comments. — This specimen represents the first record for the species for the province, although there are several records from the province of Huila, in Boca de Humpata (Laurent 1964a; Ruas 1996; Channing and Baptista 2013), and Huila (Bocage 1895; Themido 1941; Perret 1976; Ruas 1996, 2002) less than 20 km east of Leba Pass. The species is widespread across the rest of the country (Marques 2015) as well as in much of the more mesic areas of southern Africa. In Namibia it occurs only where there are permanent rivers (du Preez and Carruthers 2009). Damaraland Sand Frog Tomopterna damarensis Dawood and Channing, 2002 Material.— Pediva Hot Springs, 2 December 2013, 16°17'4.62"S, 12°33'47.86"E, 241m (CAS 254855). Comments. — The specimen was collected by locals on the border of the largest pond at Pedi¬ va Springs. The lower jaw is broken, but the specimen is otherwise in good condition. A dark pig¬ mentation is visible along the jaw-line, which identifies the specimen as a male, as is common in the genus. Comparing our specimen to the recently described Tomopterna damarensis Dawood and Channing, 2002, from Damaraland, northwestern Namibia, it agrees with the smooth dorsum and most important morphological characters. Comparison of 16S mtDNA sequence to the type speci¬ men confirms the identification of this specimen as T. damarensis (GenBank KU662310; p-dis- tance from GenBank AY255091.1, the holotype of T. damarensis, is 0.7 %). Additional details on the distribution of the species in Angola and Namibia are being prepared for publication (M. Heinicke et al., in prep.). This is the first record of the species for the country, extending the range of the species considerably northwards from the type locality at Khorixas, Namibia (Dawood and Channing 2002). Microhylidae Marbled Rubber Frog - Fig. 2 Phrynomantis annectens Werner, 1910 Material.— Omauha Lodge, 3 December 2013, 16°11'55.4"S, 12°24'0.3"E, 338 m (CAS 255056). Comments. — Both this specimen and another now in the INBAC collections were collected inside a toilet water tank, one of few available sources of standing water in the area. These speci¬ mens represent the fourth record for the species in the country. The species reaches its northern dis¬ tribution in Angola, in Novo Redondo/Gabela, Kwanza Sul Province (Poynton and Haacke 1993). Phrynomantis annectens has previously been recorded from the Mutiambo River and Caraculo in Namibe Province (Poynton and Haacke 1993). The current records represent the southernmost known distribution of the species in Angola, however it is common in Namibia and South Africa (Channing 2001; du Preez and Carruthers 2009) and is likely to occur throughout the province wherever water is available. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 21 Figure 2. Marbled Rubber Frog, Phrynomantis annectens Werner, 1910, from Palmwag, Kunene Region, Namibia. Photo courtesy of Randall Babb. Reptilia Squamata Agamidae Anchieta’s Ground Agama - Fig. 3 Agama anchietae Bocage, 1896 Material.— INP, 29 November 2013, 16°39'27.12"S, 12°26T7.04"E, 459 m (CAS 254778); Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12°29'31.rE, 359 m (CAS 254942); NNP, 28 November 2013, 15°46'27.4"S. 12°19'59.2"E, 264 m (CAS 254956). Comments.— These specimens have distinctive black-tipped spines on the palmar scales, which distinguish A. anchietae from the morphologically similar A. aculeata Merrem, 1820 that occurs sympatrically in Southern Angola and Namibia (Branch 1993). Agama anchietae was described from Angola by Bocage based on specimens from Catumbela, Benguela and Dombe (all in Benguela Province), and Mogamedes (currently Namibe, Namibe Province). In Angola, the species is known to occur in Namibe Province (Bocage 1896, 1897; Laurent 1964a), Benguela Province (Bocage 1863, 1896, 1897), and Bie Province (Schmidt 1933). Our specimens represent the southernmost localities of the species in Angola, although it is certainly distributed continu¬ ously across the entire province. Namib Rock Agama - Fig. 4 Agama planiceps Peters, 1862 Material.— ‘Lion Cave’ at 3.4 km SW of Espinheira camp, 30 November 2013, 22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Figure 3. Anchieta’s Ground Agama, Agama anchietae Bocage, 1866, from Sesfontein, Kunene Region, Namibia. Photo courtesy of Johan Marais. 16°48'45.0"S, 12°20'22.9"E, 463 m (CAS 254753); Omauha Lodge camp, 2 December 2013, 16°11'55.4"S, 12°24'0.3"E, 335 m (CAS 254832); INP, north of Tambor, 4 December 2013, 15°59'46.0"S, 12°24'24"E, 307 m (CAS 254839); INP, south side of Curoca River crossing, 29 November 2013, 16°18T5.6"S, 12°25'1.56"E, 209 m (CAS 254845), 1 December 2013, 16°18'14.8"S, 12°24'2159.8"E, 210 m (CAS 254848); Pediva Hot Springs, 2 December 2013, 16°7'19.7"S, 12°33'40.0"E, 244 m (CAS 254859); Namibe- Lubango road, road marker 59, 1.8 km west by road from Carac- ulo, north side of the road, 6 December 2013, 15°0'57.6"S, 12°38'36.8"E, 500 m (CAS 254900, CAS 254910); Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12°29'31.1"E, 359 m (CAS 254941). Figure 4. Adult male Namib Rock Agama, Agama planiceps Peters, 1862, from Pico Azevedo. Photo by Luis Ceriaco. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 23 Comments.— The species has been cited throughout Angola, although many previous records remain doubtful. Agama planiceps appears to be restricted to arid savannas and Namibe Province, or perhaps Beguela, is most probably its northernmost limit. Mertens (1938) described the sub¬ species Agama planiceps shackii from Cubal (Benguela Province), and it is likely that the central and northern Angolan records of planiceps are in fact shackii. The status of this form remains in question, but preliminary examination of topotypical material suggests that it is specifically dis¬ tinct. Our specimens fit the morphological description and current known distribution of nomino- typical planiceps. The species is cited for several localities in Namibe Province, including Biballa (Bocage 1895), Fazenda Bumbo (Laurent 1964a), and Pico Azevedo (Schmidt 1933). Some of our records represent a southern range extension of the species in the Province, although a record from the Kwito region, in Cunene Province (Angel 1923) is the southernmost Angolan record. The species likely also occurs in rocky areas extending south to the Namibian border. The majority of these specimens were seen basking on the top of rocks. Males and females present a striking sex¬ ual dimorphism, with the males being considerably larger than the females, and having an intense¬ ly red head and a dark blue body (the posterior half of the tail is usually also red). Females are typ¬ ically dark-grey on the dorsum with yellow marking on the head and dorsum. Gekkonidae Fitzsimons’ Thick-Toed Gecko - Fig. 5 Chondrodactylus fitzsimonsi (Loveridge, 1947) Material.— Espinheira, 30 November 2013, 16°47T4.3"S, 12°21'29.4"E, 457 m (CAS 254814); INP, north of Tambor, 4 December 2013, 15°59'46.9"S, 12°24'24"E, 300 m (CAS 254841). Comments.— Only four records of this species are known for the country: Ongueria, 55 km from Sa da Bandeira (presently Lubango), in Huila Province (Laurent 1964a), Praia das Conchas (Laurent 1964a), “around Mogamedes in the road to Sa da Bandeira” (Laurent 1964a) and Pico Azevedo (Schmidt 1933), both in Namibe Province. Our records represent southern range exten¬ sions for the species in the country, although the species extends into west-central Namibia (Bauer et al. 1993). In both Namibia and Angola it can occur sympatrically with other, similarly sized con¬ geners, but is generally more restricted to rocky habitats than either C. pulitzerae or C. turneri. Pulitzer’s Thick-Toed Gecko - Fig. 6 Chondrodactylus pulitzerae (Schmidt, 1933) Material.— INP, 9.65 km (by air) west-south-west of Espinheira, 30 November 2013, 16°48'43.19"S, 12°16T6.55"E, 488 m (CAS 254790-CAS 254792); Espinheira, 29 November 2013, 16°47T1.01"S, 12°21'28.77"E, 457 m (CAS 254796-254798), 16°47'8.1"S, 12°21T6.44"E, 457 m (CAS 254804, 30 November, 16°47'14.3"S, 12°21'29.4"E, 457 m (CAS 254814-254815), 16°47'15.1"S, 12°21'23.8"E, 457 m (CAS 254816), 16°47T4.0"S, 12°21'29.6"E, 457 m (CAS 254817), 16°47T7.3"S, 12°2E25.1"E, 457 m (CAS 254818), 16°47T1.1"S, 12°21'30.2"E, 457 m (CAS 254819); Omauha Lodge, 28 November 2013, 16°11'55.01"S, 12°24'3.12"E, 338 m (CAS 254830), 2 December 2013, 16°1E55.4"S, 12°24'0.3"E, 338 m (CAS 254833), 28 November 2013, 16°11'54.19"S, 12°24'2.45"E, 338 m (CAS 254843); INP, Rio Curoca in Pediva Hot Springs area, 3 December 2013, 16°17'0.93"S, 12°33'39.81"E, 247 m (CAS 254854); Namibe-Lubango road, road marker 59, 1.8 km west (by road) of Caraculo, on the north side of the road, 6 December 2013, 24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Figure 5. Adult male FitzSimons’Thick-toed Gecko, Chondrodactylus fitzsimonsi (Loveridge, 1947), from northern Kaokoveld, Kunene Region, Namibia. Photo courtesy of Johan Marais. Figure 6. Subdult male Pulitzer’sThick-toed Gecko, Chondrodactylus pulitzerae. (Schmidt, 1933), from Chimalavera, Benguela Province, Angola. Photo by Luis Ceriaco. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 25 15°0'57.3"S, 12°38'32.6"E, 497 m (CAS 254915); Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12°29'31.1"E, 359 m (CAS 254920, CAS 254943). Comments.— This species occurs from the southern regions of Angola in Namibe (Bocage 1867, 1887, 1895; Laurent 1964a; Schmidt 1933) and Cunene (Monard 1937) provinces to Malange, where it reaches its northern distribution in Capanda (Ceriaco et al. 2014). There are records from the far northwest of Namibia as well, although these have been consistently treated as either C. turneri or C. laevigatus. This species was originally described as a subspecies of C. bibronii (now regarded as limited to South Africa, southern Namibia and adjacent areas; Benyr 1995; Bauer and Lamb 2005) based on material from Pico Azevedo (Schmidt 1933). Heinz (2011) provided evidence for the specific distinctness of C. pulitzerae; see Ceriaco et al. (2014) for brief discussion ofthe nomenclatural and taxonomic history of this taxon. One specimen (CAS 254920) represents topotypical material. The species was common and found hiding on shaded areas of rocky crevices, houses, and other structures. Large-Scaled Thick-Toed Gecko Pachydactylus scutatus Hewitt, 1927 Material.— Espinheira, 30 November 2013, 16°47'51.8"S, 12°21T5.2"E, 457 m (CAS 254826). Comments. — This specimen is the first published record of Pachydactylus scutatus for Angola. Pachydatylus scutatus angolensis is now recognized as a distinct species (see below; Bauer et al. 2002). Five additional specimens, all from Iona, are present in the Ditsong National Museum of Natural History (TM 40615-18 from Espinheira, TM 40751 from 16°54'S, 12°35'E). Angolan Thick-Toed Gecko - Fig. 7 Pachydactylus angolensis (Loveridge, 1944) Material.— Namibe-Lubango road, 2 km E (by road) of Mangueiras, south side of the road, 5 December 2013, 15°2'37"S, 13°9'36"E, 625 m (CAS 254887). Comments.— This poorly known taxon was described from Benguela Province (Loveridge Figure 7. Juvenile Angolan Thick-toed Gecko, Pachydactylus angolensis (Loveridge, 1944), from Chimalavera, Benguela Province, Angola. Photo by Luis Ceriaco. 26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 1944a). Laurent (1964a) subsequently reported additional specimens from the “environs de Mogamedes” (now Namibe, Namibe Province). More recently, Wulf Haacke collected 16 speci¬ mens from both Bengela and Namibe Provinces. The Namibe localities include Lungo, Lucira, San Nicolau, and Saco de Giraul. There is a single record from extreme northern Namibia (J. Boone, pers. comm.), suggesting that the species actually has a fairly broad range from south of the Kunene to Hanha in Benguela (TM 46558). Speckled Thick-Toed Gecko Pachydactylus punctatus Peters, 1854 Material.— INP, 29 November 2013, 16°39'24.1"S, 12 0 26'12.2"E, 460 m (CAS 254781- 254784); Espinheira, 29 November 2013, 16 o 47T1.01"S, 12°21'28.77"E, 457 m (CAS 254799- 254800), 16°47'7.02"S, 12°21T6.86"E, 457 m (CAS 254806); 16°47'4.2"S, 12°21T6.62"E, 457 m (CAS 254809-254810), 16°47T2.71"S, 12°21'28.67"E, 457 m (CAS 254812), 16°47'20.3"S, 12°2E27.6"E, 457 m (CAS 254960). Comments.— The species is known from southwestern Angola in the provinces of Benguela (Bocage 1867, 1895; Boulenger 1885; Hellmich 1957b; Laurent 1954), Huila(Monard 1931, 1937, Laurent 1964a), Cunene (Laurent 1964a), and Namibe (Schmidt 1933, Laurent 1964a). Members of the P. punctatus complex have been confused with other southern African Pachydactylus, including P occellatus and P. geitje (Bocage 1867, 1885, 1895; Boulenger 1905; Frade 1963). The previous report of P serval in Angola (Monard 1931) likely corresponds to P punctatus. Aphylo- geographic study of P. punctatus is being currently undertaken (Heinz 2011) and it appears that this taxon comprises multiple unnamed cryptic species. At least two, possibly three, taxa in this com¬ plex occur in southwestern Angola. Pachydactylus punctatus sensu lato is the most common ter¬ restrial gecko in most of northern Namibia as well as southern Angola. Kaokoland Rock Gecko Pachydactylus cf. oreophilus McLachlan and Spence, 1967 Material.— Omauha Lodge, 28 November 2013, 16°1L55.01"S, 12°24'3.12"E, 338 m (CAS 254829). Comments. — Pachydactylus oreophilus was described from near Sesfontein in northwestern Namibia. Specimens assigned to this species extend northwards at least as far as the southern low¬ land portions of Benguela Province, Angola. Preliminary molecular phylogenetic data suggest that northern populations, including all of those in Angola and possibly those along the Kunene River in Namibia, are not conspecific with the nominotypic form. The specimen was collected at night, basking near a lamp 2.5 m off the ground, in Omauha Lodge. Barnard’s Namib Day Gecko - Fig. 8 Rhoptropus barnardi Hewitt, 1926 Material.— Approximately 7.35 km north-west (by road) of Pico Azevedo, 7 December 2013, 15°28'30.7"S, 12°27'47.5"E, 420 m (CAS 254759, CAS 254761); Omauha Lodge, 4 Decem¬ ber 2013, 16°12T.2"S, 12 o 24'0.1"E, 343 m (CAS 254837); INP, Rio Curoca crossing, North side of the river, 1 December 2013, 16°18'6.8"S, 12°25T3.0"E, 206 m (CAS 254844); INP, Rio Curoca crossing, south side of the river, 1 December 2013, 16°18T4.7"S, 12°25'0.0"E, 210 m (CAS 254846-254847); INP, Rio Curoca in the Pediva Hot Springs area, 2 December 2013, CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 27 Figure 8. Adult Barnard’s Namib Day Gecko, Rhoptropus barnardi Hewitt, 1926, from Kamanjab, Kunene Region, Namibia. Photo courtesy of Johan Marais. 16°17'0.93"S, 12°33'39.81"E, 247 m (CAS 254852), 16°17T4.3"S, 12°33'35.9"E, 238 m (CAS 254856), 16°17'24.0rS, 12°33'43.9"E, 270 m (CAS 254863), Namibe-Lubango road, 2 km east (by road) of Mangueiras, south side of the road, 5 December 2013, 15°2'40.8"S, 13°9'32.6"E, 664 m (CAS 254890); NNP, 28 November 2013, 15°46'23.4"S, 12°19'58.9"E, 264 m (CAS 254954). Comments.— Until now, this species has been known only from one published locality in Angola. Laurent (1964a) cites the specimen from a locality “60 km on the road from Mogamcdes [presently Namibe] to Sa da Bandeira [presently Lubango]”, the same locality from which he described R. taeniostictus, which we also collected (see account below). This species is widely dis¬ tributed in northwestern Namibia, occurring as far inland as the Otavi-Grootfontein region, due south of western Cuando Cubango Province. It is rupicolous and can be found on small rocky piles and ridges, as well as on larger boulders. The extent of its distribution in Angola is poorly known, in part becase many records from Namibe and Huila are assignable to a morphologically similar, but undescribed congener (see Rhoptropus sp. below). Two-Pored Namib Day Gecko - Fig. 9 Rhoptropus biporosus Fitzsimons, 1957 Material.— INP, 29 November 2013, 16°32'0.48"S, 12°26'44.16"E, 378 m (CAS 254779, 16°39'26.04"S, 12°26'13.5"E, 460 m (CAS 254780); INP, 20 km south-south-west (by air) of Espi- nheira, 30 November 2013, 16°55'54.1"S, 12°14'42.0"E, 631m (CAS 254786-254788), PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 28 Figure 9. Close-up of head of adult specimen of Two-Pored Namib Day Gecko, Rhoptropus biporosus FitzSimons, 1957 from northwest of Palmwag, Kunene Region, Namibia. Photo courtesy of Johan Marais. 16°48'43.19"S, 12°16'16.55"E, 485 m (CAS 254959), Espinheira, 29 November 2013, 16°47T9.9"S, 12°21'27.4"E, 457 m (CAS 254794), 16°47'7.08"S, 12°21'16.02"E, 457 m (CAS 254802-254803), 16°47'7.02"S, 12°21'16.86"E, 457 m (CAS 254805), 16°47'4.26"S, 12°21T6.62"E, 457 m (CAS 254811), 16°47T2.71"S, 12°21'28.67"E, 457 m (CAS 254813), 16°47'20.2"S, 12°21'27.9"E, 457 m (CAS 254958), 30 November 2013, 16°47'14.3"S, 12°21 '29.4"E (CAS 254820), 16°47'8.7"S, 12°21'30.3"E, 457 m (CAS 254821), 16°47T8.1"S, 12°21'26.2"E, 457 m (CAS 254822), 16°47'33.6'S, 12°21'19.0"E, 457 m (CAS 254823), 16°47'41.5"S, 12°21T7.3"E, 457 m (CAS 254824), 16°47'45.3"S, 12°21'15.9"E, 457 m (CAS 254825); NNP, 28 November 2013, 15°46'27.4"S, 12°19'59.2"E, 264 m (CAS 254957-254958). Comments.— The species occurs in the rocky outcrops in arid habitats inland of the northern Namib dune fields in the vicinity of Orupembe, in the Kaokoveld and across the Cunene River to Angola (Bauer and Good 1996). The only published record of this species for Angola is from the Pico Azevedo region (Bauer and Good 1996), although Wulf Haacke collected numerous speci¬ mens from localities across southern Namibe, as well as from near Otchinjau, Cunene Province (specimens in Ditsong National Museum of Natural History). Boulton’s Namib Day Gecko - Fig. 10 Rhoptropus boultoni boultoni Schmidt, 1933 Material.— INP, 3.4 km southwest (by air) of Espinheira, vicinity of “Lion Cave”, 30 November 2013, 16°48'73.5"S, 12°20'23.2"E, 463 m (CAS 254752); Approximately 7.35 km north-west (byroad) of Pico Azevedo, 7 December 2013, 15°28'33.2"S, 12°27'45.7"E, 421 m (CAS 254757-254758); Espinheira, 16°47'29.4"S, 12°21'6.06"E, 457 m (CAS 254795); Omauha Lodge, 28 November 2013, 16°11'52.5"S, 12°23'59.3"E, 335 m (CAS 254828, 2 December 2013, CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 29 Figure 10. Adult Boulton’s Namibe Day Gecko, Rhoptropus boultoni boultoni Schmidt, 1933, from east of Kaman- jab, Kunene Region, Namibia. Photo courtesy of Johan Marais. 16°12T.2"S, 12°24'0.1"E, 343 m (CAS 254834); INP, Rio Curoca crossing, south side of river, 1 December 2013, 16°18T4.7"S, 12°25'0.0"E, 210 m (CAS 254849-254850), 2 n d December 2013, 16°17T9.7"S, 12°33'40.0"E, 244 m (CAS 254857-254858, CAS 254861-254862), 29 November 2013, 16°18T5.6"S, 12°25'1.56"E, 209 m (CAS 254865); Leba Pass, between river and highway, 5 December 2013, 15°4'13.2"S, 13°14'37.7"E, 1676 m (CAS 254880); Namibe-Lubango road, 2.0 km east (by road) of Mangueiras, south side of the road, 5 December 2013, 15°2'40.8"S, 13°9'32.6"E, 664 m (CAS 254892), 15°2'40.7"S, 13°9'31"E, 640 m (CAS 254894, 15°0'55.1"S, 12°38'32.8"E, 497 m (CAS 254902); Namibe-Lubango road, road marker 59, 1.8 km west by road of Caraculo, 6 December 2013, 15°00'55.1"S, 12°38'32.8"E, 497 m (CAS 254903); Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12°29'31.1"E, 359 m (CAS 254921-254926), 15°32'5.8"S. 12°29'29.5"E, 366 m (CAS 254946-254947, CAS 254949-254950); Espinheira, 29 November 2013, 16°47'20.2"S, 12°21'27.9"E, 457 m (CAS 254958). Comments.— This taxon is widespread from northwestern Namibia north at least to northern Namibe Province. Montane Namib Day Gecko Rhoptropus boultoni montanus Laurent, 1964 Material.— Leba Pass overlook, 5 December 2013, 15°4'37.2"S, 13°13'58.5"E, 1682 m (CAS 254866, CAS 254867; 15°4'38.3"S, 13°13'57.0"E, 1682 m (CAS 254868); 15°4'36.0"S, 13°14T.6"E, 1682 m (CAS 254869-254872); Leba Pass, between river and highway, 15°04'13.2" S, 13°14'37.3"E, 1676 m (CAS 254882). Comments.— The subspecies was described from the Leba Escarpment (“60 km on the road to Mogamedes [now Namibe, Namibe Province] from Sa da Bandeira [now Lubango, Huila 30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Province]”, Laurent 1964a). A large series of specimens in the Ditsong National Museum of Nat¬ ural History are derived from localities near Lubango, in Huila. Our specimens are topotypical and were collected on the Namibe side of the provincial boundary. Specimens were found basking on high elevation granite rocks covered with bryophytes. Molecular phylogenetic studies (A. Kuhn, unpublished) reveal that this taxon is specifically distinct from R. boultoni. Its formal elevation to specific status will be justified in detail elsewhere. Angolan Namib Day Gecko Rhoptropus taeniostictus Laurent, 1964 Material.— Namibe-Lubango road, 2 km east (by road) of Mangueiras, south side of the road, 5 December 2013, 15°2'40.8"S, 13°9'32.6"E, 664 m (CAS 254889); Namibe-Lubango road, road marker 59, 1.8 lan west (by road) of Caraculo, on the north side of the road, 6 December 2013, 15°0'58.0"S, 12°38'37.3"E, 490 m (CAS 254895);15°0'57.9"S, 12°38'42.3"E, 472 m (CAS 254897- 254898), 15°0'57.6"S, 12°38'36.8"E, 500 m (CAS 254901), 15°1'0.7"S, 12°38'31.9"E, 492 m (CAS 254904-254905), 15°0'58.8"S, 12°38'33.8"E, 491 m (CAS 254908), 15°0'58.9"S, 12°38'32.4"E, 497 m (CAS 254911), 15°0'57.3"S, 12°38'32.6"E, 497 m (CAS 254916), 15°E0.1"S, 12°38'31.9"E, 497 m (CAS 254917-254918), 15°1'0.9"S, 12°38'30.4"E, 503 m (CAS 254919). Comments. — The Angolan endemic R. taeniostictus was described from a single specimen from “60 km on the road from Mogamcdes [presently Namibe] to Sa da Bandeira [presently Luban¬ go]”. Laurent (1964a) also considered the populations of R. barnardi from Mucungo cited by Schmidt (1933) as referable to R. taeniostictus. The species appears restricted to Namibe Province and is represented by many specimens in our collection as well as more widespread Namibe local¬ ities in the Ditsong National Museum of Natural History. Rhoptropus sp. Material.— Espinheira, 29 November 2013, 16°47'32.7"S, 12°21T4.4"E, 562 m (CAS 254801); Omauha Lodge, 4 December 2013, 16°12T.2"S, 12°24'0.r'E, 343 m (CAS 254836, CAS 254955); INP, north of Tambor, 4 December 2013, 15°59'46.9"S, 12°24'24.0"E, 300 m (CAS 254842, CAS 254762, CAS 254766), 15°28'31.7"S, 12°27'43.9"E, 408 m (CAS 254765, CAS 254760); Leba Pass, between river and highway, 5 December 2013, 15°4T2.1"S, 13°14'36.5"E, 1680 m (CAS 254873), 15°4'13.2"S, 13°14'37.7"E, 1676 m (CAS 254879, CAS 254881, CAS 254883); Namibe-Lubango road, 2 m east (by road) of Mangueiras, south side of the road, 5 December 2013, 15°2'40.8"S, 13°9'32.6"E, 664 m (CAS 254890-254891), 15°2'40.7"S, 13°9'31.0"E, 640 m (CAS 254893, CAS 254894). Comments. — This undescribed species is morphologically similar to both R. barnardi and R. biporosus, but appears to be endemic to southern Angola. Populations from the Escarpment are both morphologically and genetically different from those below the Escarpment.This taxon is cur¬ rently under study as part of a phylogenetic analysis and revision of the genus as a whole (A. Kuhn and A. Bauer, in prep.). Scincidae Bogert’s Speckled Western Burrowing Skink - Fig. 11 Typhlacontias punctatissimus bogerti Laurent, 1964 Material.— Espinheira, 29 November 2013, 16°47'7.02"S, 12°21T6.86"E, 457 m (CAS CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 31 254807); Pico Azevedo, 7 De¬ cember 2013, 15°32'2.4"S, 12°29'31.1"E, 359 m (CAS 254932-254938), 15°32'5.8"S, 12°29'29.5"E, 366 m (CAS 254944-254945). Comments.— Haacke (1997) reviewed the taxonomic and nomenclatural history of Typhlacontias punctatissimus and its subspecies and recog¬ nized two sympatric subspecies in southern Angola — T. puncta¬ tissimus punctatissimus Bocage, 1873, and the Angolan endemic T. punctatissimus bogerti Lau¬ rent, 1964. In all of our speci¬ mens, the second and third upper labials are in contact with the eye and there is a second supraoc¬ ular. Both characters fit the description presented by Haacke (1997) as diagnostic for T. punctatissimus bogerti. The species is known to be viviparous and one female specimen (CAS 254945) contains an almost fully developed neonate. Variable Skink Trachylepis varia (Peters, 1867) Material.— Leba Pass, 5 December 2013, 15°4T2.1"S, 13°14'36.2"E, 1680 m (CAS 254874), 15°4'13.2"S, 13°14'37.7"E, 1676 m (CAS 254884). Comments.— The species occurs throughout Angola, with many records in the provinces of Benguela (Parker 1936; Bocage 1895, 1896; Monard 1937; Hellmich 1957a; Mertens 1938; Boulenger 1905) and Huila (Bocage 1895; Monard 1937). Although other records from Namibe are 12 km W of Humbia (TM 40128-29) and Chapeau Armado turnoff (TM 41131). This species is typically associated with relatively mesic microclimates and is, therefore, excluded from the hyper- arid areas of Namibe. This skink has a broad distribution across much of sub-Saharan Africa and includes several cryptic species, two of which are present in Angola. There are records from across Angola and from all bordering countries as well. The phylogeography of the T. varia complex is presently under study (J. Weinell, pers. comm.). Speckled Sand Skink -Fig. 12 Trachylepis punctulata (Bocage, 1872) Material.— Espinheira, 29 November 2013, 16°47'20.6"S, 12°21'27.2"E, 457 m (CAS 254793); Namibe-Lubango road, road marker 59, 1.8 km west (by road) of Caraculo, on the north side of the road, 6 December 2013, 15°0'55.1"S, 12°38'32.8"E, 497 m (CAS 254903); Praia do Navio coastal dunes, ca 124 km SSW of Namibe, 8 December 2013, 16°16'20.4"S, 11°49'53.9"E, 8 m (CAS 254769-254771), 16°16'39.3"S, 11°49'20.5"E, 8 m (CAS 254775). Comments.— The species was originally described by Bocage based on material from “Rio Coroca, sur le littoral de Mossamedes, Angola” (Bocage 1872). The type locality is presumably the Figure 11 . Adult Bogert’s Speckled Western Burrowing Skink Typhla¬ contias punctatissimus bogerti Laurent, 1964, from Pico Azevedo. Photo by Edward Stanley. 32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 region near the mouth of Curoca River, from the vicinity of Tombwa (formerly Porto Alexandre). Our specimens were collected among plants between dunes along the coast south of Tombwa, and these agree morphologically with the original description for the species. A comparison with the type material was impossible due to its destruction in the fire that destroyed the Lisbon Museum in 1978. Several uncatalogued specimens in the Museu Nacional de Historia Natural do Porto col¬ lected in 1905 by the Portuguese explorer Francisco Newton are congruent with the specimens col¬ lected by us. Newton’s specimens are still in their original jar and are labeled “Mossamedes” (pre¬ sumably refering to the province as a whole, not the city of Mossamedes = Namibe). They are part of a collection of vertebrates that the explorer made in the region. The herpetological specimens were only partly studied and published upon (Ferreira 1904, 1906; Ceriaco et al. 2014), in contrast to the bird and mammals collections (Seabra 1906a, 1906b, 1906c, 1906d, 1907). It is probable that these specimens are from Tombwa, as this was the main place where Newton collected while in the province (see bird records - Seabra 1906a). This is a common species in much of Namibia, Botswana, and central South Africa, as well as portions of Zambia, Zimbabwe, and Mozambique (Portik and Bauer 2012). Figure 12. Adult Speckled Sand Skink, Trachylepis punctulata (Bocage, 1872), from Kamanjab, Kunene Region, Namibia. Photo courtesy of Johan Marais. Hoesch’s Skink-F ig. 13 Trachylepis hoeschi (Mertens, 1954) Material.— Rio Curoca in the Pediva Hot Springs area, 2 December 2013, 16°17'0.93"S, 12°33'39.81"E, 247 m (CAS 254851); NNP, 15°46'25.9"S, 12°19'59.0"E, 247 m (CAS 254952). Comments.— The only published Angolan record is from Laurent (1964a), from “Plage das Conchas,” Namibe Province. Our specimens are, respectively, 75 and 135 km SE of Laurent’s site, the Ditsong National Natural History Museum holds a small series of this species (TM 40733-37) CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 33 Figure 13. Adult Hoesch’s Skink, Trachylepis hoeschi (Mertens, 1954), from Kamanjab, Kunene Region, Namibia. Photo courtesy of Johan Marais from Iona National Park. The Angolan records extend the core distribution of its range in north¬ western Namibia (Branch 1998). Angolan Blue-Tailed Skink Trachylepis laevis (Boulenger, 1907) Material.— INP, north of Tambor, 4 December 2013, 15°59'47.1"S. 12°24'25.6"E, 314 m (CAS 254838). Comments.— The species was described by Boulenger from Maconjo, in northern Namibe Province (Boulenger 1907b). Laurent (1964a) recorded this species in Namibe Province from “Munhino 50 km west of Sa da Bandeira.” Hellmich (1957a) cited the species for Piri-Dembos, Kwanza Norte Province, but this record is dubious. The Ditsong National Natural History Muse¬ um houses numerous specimens from localities in Namibe and southern Benguela below the Escarpment. The species occurs also in the Kamanjab area and Damaraland in northwestern Namibia (Bauer et al. 1993). This lizard is extremely dorsoventrally depressed in association with its crevice dwelling habits and was, for a time, placed in a monotypic genus, Oelofisa , in recogni¬ tion of its highly autapomorphic morphology (Steyn and Mitchell 1965). Western Three-Striped Skink Trachylepis occidentalis (Peters, 1867) Material.— Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12 0 29'31.1"E, 359 m (CAS 254931). 34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Comments.— The species has been cited from 35 km south of the city of Namibe (Laurent 1964a) and from Curoca River (Bocage 1895). Three specimens in the Ditsong Natural National History Museum originate from the Rio Curoca mouth and from Namibe. In addition to our spec¬ imen from Pico Azevedo several individuals of the species were observed near Espinheira camp (specimens not collected). In Angola this skink takes refuge in holes it digs in the sand at the base of spiny shrubs of the genus Blepharis. It is widely distributed in western South Africa and much of central and western Namibia (Branch 1988). Wedged-Snouted Skink -Fig. 14 Trachylepis acutilabris (Peters, 1862) Material.— INP, 3.4 km south-west (by air) of Espinheira, vicinities of “Lion Cave”, 30 November 2013, 16°48'54.4"S, 12°20T3.7"E, 450 m (CAS 254751); INP, car wreck 20 km south- south-west (by air) of Espinheira, 30 November 2013, 16°55'53.81"S, 12°14'45.42"E, 616 m (CAS 254789); Namibe-Lubango road, road marker 59, 1.8 km (by road) of Caraculo, north side of the road, 6 December 2013, 15°0'59.3"S, 12°38'33.6"E, 488 m (CAS 254899), 15°0'58.8"S, 12°38'33.8"E, 491 m (CAS 254907); Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12°29'3U"E, 359 m (CAS 254927-254931). Comments.— This species is similar to lacertid lizards in morphology and diet (Castanzo and Bauer 1992). Its elongate toes and countersunk lower jaw are consistent with its burrowing habits. It typically occupies burrows at the base of vegetation in sandy soils from Namibia through west¬ ern Angola to the Democratic Republic of Congo and Cabinda (Branch 1998). Western Rock Skink Trachylepis sulcata (Peters, 1867) Material.— INP, Rio Curoca in Pediva Hot Springs area, 3 December 2013, 16°17'0.93"S, 12°33'39.81"E, 247 m (CAS 254853); Leba Pass, 5 December 2013, 15°4T2.1"S, 13°14'36.2"E, 1680 m (CAS 254875); Namibe-Lubango road, 2 km east (by road) of Mangueiras, south side of the road, 5 December 2013, 15°2'40.7"S, 13°9'31"E, 625 m (CAS 254886), 15°2'40.8"S, 13°9'32.6"E, 664 m (CAS 254888). Comments. — Trachylepis sulcata is a rupicolous skink ranging from the Western Cape Province of South Africa north to southern Angola. Trachylepis sulcata ansorgii (Boulenger 1907b) was described from southern Angola to accommodate specimens with bright throat and infralabial coloration. Laurent (1964a) and Mertens (1971) considered it valid and the latter iden¬ tified some Namibian specimens as intergrades between T. s. ansorgii and T. s. sulcata, whereas Haacke (1972) considered specimens on the Namibian side of the Kunene river to be referable to T. s. ansorgii. Some specimens from west of the Great Escarpment in northwestern Namibia exhib¬ it the diagnostic coloration of ansorgii, but the two subspecies seem to have no fixed differences in scalation (Bauer et al. 1993). Although Portik et al. (2011) did not include typical T. s. ansorgii in their molecular sampling, preliminary integration of samples from our collection into their data set reveals no significant difference from putative T. s. ansorgii from Namibe and the nominotyp- ical form. We therefore treat T. sulcata as a monotypic species. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 35 Figure 14. Adult Wedged-Snouted Skink, Trachylepis acutilabris (Peters, 1862), from Kamanjab, Kunene Region, Namibia. Photo courtesy of Johan Marais. 36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Lacertidae Anchieta’s Shovel-Snout Lizard Meroles anchietae (Bocage, 1867) Material.— Praia do Navio coastal dunes, ca 124 km SSW of Namibe, 8 December 2013, 16°16'29.1"S, 11°49'05.0"E, 8 m (CAS 254773). Comments.— Bocage (1867) described this species from “Mossamedes” (Bocage 1867). Sur¬ prisingly, this remains the only published locality for this species in Angola (Bocage, 1867, 1895), despite it being common and widely distributed in barchan dunes from the Klinghardt Mountains north through the Namib of western Namibia. In the Newton collections in Porto, there are sever¬ al uncatalogued specimens corresponding to this species from “Mossamedes” and the Ditsong National Natural History Museum has material collected by Wulf Haacke from Porto Alexandre and Foz de Cunene. Reticulate Sand Lizard - Fig. 15. Meroles reticulatus (Bocage, 1867) Material.— Praia do Navio coastal dunes, ca 124 km SSW of Namibe, 8 December 2013, 16°16'39.3"S, 11°49'20.5"E, 8 m (CAS 254776). Comments.— Bocage (1867) described this species from “Mossamedes.” Bocage (1895) sub¬ sequently clarified that the types had come from the littoral zone at Rio Coroca [= Rio Curoca, southern Namibe Province, Angola]. The range of this species extends towards Namibia to the area of Conception Bay on the central coast. Although it is well documented within its Namibian range, this specimen is only the third published locality for Angola. The species is, however, well repre¬ sented from numerous localities in Namibe by specimens in the Ditsong National Museum of Nat¬ ural History. Figure 15. Adult specimen of Anchieta’s Shovel-snout Lizard, Meroles anchietae, (Bocage, 1867) from gravel plains north of Henties Bay, Erongo Region, Namibia. Photo courtesy of Johan Marais. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 37 Haacke’s Sand Lizard Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012 Material.— 5 km NW (by road) of Pico Azevedo, 7 December 2013, 15°28'33.6"S, 12°27'41.4"E, 399 m (CAS 254767), 15°28'31.7"S, 12°27'43.9"E, 408 m (CAS 254763-253764); RNN, 28 November 2013, 15°46'22.5"S, 12°19'57.7"E, 262 m (CAS 254951), 15°46'25"S, 12°19'54.9"E, 262 m (CAS 254953); Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12°29'31.1"E, 359 m (CAS 254939); Pediva Hot Springs, south side of the river, 2 December 2013, 16°17'37.7"S, 13°33'37.2"E, 235 m (CAS 254860), 16°17'24.01"S, 12°33'43.9"E, 270 m (CAS 254864); INP, north of Tambor, 4 December 2013, 15°59'43.4"S, 12°24'23.3"E, 306 m (CAS 254840); Omauha Lodge, 4 December 2013, 16°12T.2"S, 12 o 24'00.1"E, 343 m (CAS 254835). Comments.— Pedioplanis haackei is one of the latest additions to the herpetofauna of Ango¬ la, and is endemic to southern Angola. Each of our specimens has 10 to 12 longitudinal rows of ventral scales, a semi-transparent lower eyelid with a brille formed of two large scales, five to six supralabials anterior to the subocular, two rows of granules separating supraoculars from supracil- iaries, and the typical coloration with dots on the flanks, concordant with the diagnostic characters presented by Conradie et al. (2012) in the species description. Genetically, our specimens (Gen- Bank accession numbers KU662311-KU662318) have an average 16S p-distance of 1% from those of Conradie et al. (2012) (GenBank accession numbers HE794000, HE793999.1; HE793998.1; HE793997.1; HE793996.1; HE793995.1; HE793994.1; HE793994.3). Benguela Sand Lizard Pedioplanis benguellensis (Bocage, 1867) Material.— Namibe-Lubango road, road marker 59, 1.8 km W (by road) from Caraculo, north side of the road, 6 December 2013, 15°00'57.8"S, 13°38'41.4"E, 476 m (CAS 254909), 15°00'58.8"S, 13°38'33.8"E, 491m (CAS 254906), 15°00'57.5"S, 12°38'38.3"E, 482 m (CAS 254896). Comments.— All of these specimens have ten to eleven longitudinal series of ventral plates and a single transparent scale in the lower eyelid. However, they also have two rows of granules separating supraoculars from supraciliaries, a character given by Conradie et al. (2012) as synapo- morphic for P haackei. The number of upper labials in front of the subocular is variable: CAS 254896 has three, CAS 254906 has five, and CAS 254909 has four. Molecular comparisons with specimens of P. benguellensis from Conradie et al. (2012) show these specimens to be conspecif- ic (average uncorrected 16S p-distance of 1% from our specimens [GenBank accession numbers KU662319 to KU662321] from those of Conradie et al. [2012] [GenBank accession numbers HE794014.1, HE794012.1, HE794011.1, HE794010.1 and HE794013.1]). Bushveld Lizard Heliobolus lugubris (Smith, 1838) Material.— Namibe-Lubango road, 2 km E of Mangueiras, south side of the road, 5 Decem¬ ber 2013, 15°02'37.0"S, 13°09'36.0"E, 625 m (CAS 254885). Comments.— The species is widespread over much of southern Africa, particularly on sandy substrates (Bauer et al. 1993). In Angola, the majority of both bibliographic (Bocage 1867, 1895; Monard 1937; Mertens 1938; Boulenger 1921) and museum records (e.g., TM 46525) are from Benguela. However, there are also records of the species for Namibe Province from “Maconjo” 38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 (Bocage 1895), “Capangombe” (Bocage 1895), and “Konondoto” (Boulenger 1921). The species’ distribution in the country extends to the southeast in Huila and Cunene Provinces. Our specimen is a sub-adult (55.5 mm SVL). Cordylidae Kaokoveld Girdled Lizard Cordylus namakuiyus Stanley, Ceriaco, Bandeira, Valerio, Bates and Branch, 2016 Material. — 7.5 km NW (by road) of Pico Azevedo, 7 December 2013, 15°28'33.2"S, 12°27'45.7"E, 421 m (CAS 254754-254755, CAS 256530-256531); Namibe-Lubango road, road marker 59, 1.8 km W (by road) from Caraculo, north side of the road, 6 December 2013, 15°00'59.4"S, 12°38'31.3"E, 503 m (CAS 254912-254914); 15°00'57.3"S, 12°38'32.6"E, 509 m (CAS 256529). Comments. — Based on a combination of morphological and molercular data, Stanley et al. (2016) described this new species endemic to the arid lowlands west of the southern Angolan escarpment. The majority of the type material of C. namakuiyus was collected during this trip. The new species is morphologically and genetically distinct from its sister taxon Cordylus machadoi Laurent, 1964, which occurs in the highlands of Huila, not far from Namibe. In addition to an aver¬ age 7.1% uncorrected p-distance for the mitochondrial marker ND2, the two species differ in the extent of osteodermal armament. The more complete body armor of C. namakuiyus may be an adaptation to the semi-arid and refuge-scarce habitat where the species occurs. Within the new species there is also a degree of internal genetic structure, with specimens from Iona (PEM R18005) and Pico Azevedo (CAS 254754, 254755, 256530, 256531) more closely related to one another than to the two specimens from Caraculo. Specimens in the American Museum of Natural Histiry identified as Cordylus cordylus and collected in 1925 during the Vernay Expedition in Angola are assignable to C. namakuiyus. Although the Vernay specimens lack specific locality information, the expedition field notes mention that significant numbers of unidentified lizards were collected at “Pico Azevedo” and “100 km east of Mogamedes” the same areas where we col¬ lected eight specimens of C. namakuiyus. Dwarf Plated Lizard - Fig. 16 Cordylosaurus subtessellatus (Smith, 1844) Material. — INP, 20 km SSW of Espinheira, 30 Novem¬ ber 2013, 16°55'54.1"S, 12°14'42.0"E, 631m (CAS 254785). Comments. — The species is known from the coastal areas of Benguela Province (Bocage 1867, 1895; Boulenger 1887) and from the Curoca River in Namibe Province (Bocage 1895). This specimen represents the southernmost record for Angola. FlGURE 16 Adult Dwarf Pla,ed Lizard ’ Cordylosaurus subtessellatus . (Smith, 1844), from near Espinheira. Photo by Edward Stanley. The specimen was found hiding CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 39 in a granite boulder crevice. Extralimitally, the species is widely distributed from the Little Karoo in the Western Cape of South Africa, north through the entire length of Namibia (Branch 1998). Gerrhosauridae Desert Plated Lizard - Fig. 17 Gerrhosaurus skoogi (Andersson, 1916) Material.— Praia do Navio coastal dunes, ca 124 km SSW of Namibe, 8 December 2013, 16°16'20.4"S, 11°49'53.9"E, 8 m (CAS 254772), 16°16'29.1"S, 11°49'50.0"E, 8 m (CAS 254774), 16 0 16'42.1"S, 11°49'21.7"E, 8 m (CAS 254777). Comments.— This species was encoun¬ tered basking at the sun in the coastal dunes SSW of Namibe, especially in dune valleys areas. When approached, these lizards dive into the sand, disappearing rapidly. The species is easily identified by its unique morphology and peculiar ecology. Sand trails resulting from the specimens walking in the dunes were noted. A total of five specimens were collected, three of which are at CAS and two at INBAC. Several additional animals were observed in the area but not collected. Males have a distinct black throat and venter, and are considerably larger than the females. The only published Angolan records are from the type locality, Porto Alexan¬ dre, between Mossamedes and the mouth of the Cunene River (Andersson 1916; Fitzsimons 1953), approximately in the same area as our material. Although the species is the most dis¬ tinctive of all Gerrhosaurus (Nance 2007), it is unambiguously nested among more typical taxa, implying its unique features are relatively r Lamb and Bauer 2013). Figure 17. Adult female Desert Plated Lizard, Ger- rhosaurus skoogi Andersson, 1916, from the coastal dunes, near Praia do Navio. Photo by Arianna Kuhn derived autapomorphies (Lamb et al. 2003; Varanidae Angolan Rock Monitor Varanus albigularis angolensis Schmidt, 1933 Material.— 7.5 km NW (by road) of Pico Azevedo, adult, found in a rock crevice, 7 Decem¬ ber 2013, 15°28'33.6"S, 12°27'41.4"E, 399 m (CAS 254768). Comments.— The subspecies was described from Gauca, Bihe (Bie) Province (Schmidt 1933). According to the original description, the subspecies differed from the nominotypic form by having larger scales everywhere on the body so that the scales around the body are about 125 40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 instead of 150, and the transverse rows of scales from the collar to thighs are 75 instead of 100 (Schmidt 1933). Laurent (1964b) noted that the morphological differences between the nomino- typic form and angolensis are quite subtle and that angolensis may, in fact, be a synonym of albigu- laris. However, Bayless (2002) considered the material from nearby localities, such as Bibala and Caraculo as V a. angolensis. Given the currently accepted distribution of angolensis and the lower number of scales around the midbody and between the collar and the thighs, we tentatively identi¬ fy our specimen as the Angolan subspecies. The subspecies appears to be the prevalent form in Angola (and possibly extending to some neighboring regions of the DRC, even if the nominotyp- ic form occurs sympatrically, especially in the southern regions of the country (Bayless 2002). Pythonidae Southern African Rock Python - Fig. 18 Python natalensis Smith, 1840 Material.— Beginning of the forested areas, at the start of the climb to Leba Pass (by road), near Bruco village, 5 December 2013, 15°07'15.82"S, 13°11'11.56"E. Individual observed but not collected. Comments.— A local at a site near Bruco village was sell¬ ing a single live individual of Python natalensis , presumably collected nearby. In the province, this species is known from Maconjo (Bocage 1895; Broadley 1984) and from Giraul River (Bocage 1896; Broadley 1984). Python natalensis was for many years considered as a sub¬ species of Python sebae (Gmelin, 1789) (Broadley 1984), but was elevated to specific status by Broadley (1999) based on mor¬ phological differences as well the evidence of the overlapping dis¬ tributions (Broadley and Cotterill 2004). Although the current taxo¬ nomic arrangement appears appropriate, molecular analyses are needed to conclusively resolve the relationship between P natalensis and P sebae (Alexander 2007). Spawls and Branch (1995) and Bellosa et al. (2007) provided maps with the distribution ranges for both species, with P natalensis occurring in central and southern Angola, as far north as the Kwanza River, overlaping in Luanda Province with P sebae , which occurs in northern regions of Angola, including Cabinda. According to the local selling this individual, this species is sometimes collected for food or sold to tourists as pets. Figure 18. Adult Southern Rock Python, Python natalensis (Gmelin, 1788), being sold by a local near Bruco village. Photo by Luis Ceriaco. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 41 Lamprophiidae Karoo Sand Snake - Fig. 19 Psammophis notostictus Peters, 1867 Material.— Espinheira, 30 November 2013, 16°47'13.8"S, 12°21'27.5"E, 457 m (CAS 254827); Pico Azevedo, 7 December 2013, 15°32'2.4"S, 12°29'31.1"E, 359 m (CAS 254940). Comments.— Psammophis notostictus is easily recognizable from all other southern African Psammophis by its single cloacal shield and the presence of two preoculars (Broadley 1975b, 1977, 2002). These two specimens have both of these diagnostic characters. The species is known for Angola, but only from Namibe Province. The closest published records of the species are in Rio Sao Nicolau (Loveridge 1940; Broadley 1975b, 2002), Mo^amedes [Namibe city] (Bocage 1887; Loveridge 1940), and Curoca River (Loveridge 1940; Broadley 2002). Our records expand the known distribution of the species further south in the country, although it is continuous southwards throughout much of western southern Africa (Branch 1998). Figure 19 - Adult Karoo Sand Snake, Psammophis notostictus Peters, 1867, from Espinheira. Photo by Luis Ceriaco. Discussion Namibe Province hosts a high diversity of reptile taxa, with approximately one-third of all the reptile species known for Angola (see Table 1). Not surprisingly for an arid region, the diversity of amphibians is considerably lower in this province. However, the anuran species Tomopterna damarensis, adapted to drier climates, is reported here for the first time. The lizard families Scin- cidae and Gekkonidae are the most species-rich groups for the province with 21 and 20 species/subspecies known, respectively, and for the snakes the family Lamprophiidae has the high¬ est diversity of taxa, with 13 species known for the province. These numbers, however, are under¬ estimates. We did not take into account unpublished voucher specimens; as noted above, a more complete synopsis of the Namibe Province taxa will be provided elsewhere (W.R. Branch, pers. comm.). There are several examples of taxa not previously recorded for Namibe, but which are found both north and south of Namibe Province and can be expected to be found here in the 42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 future.The presence of conspicuous species, such as Anchieta’s Dwarf Python, Python anchietae Bocage, 1887, known from Benguela Province and from the northern regions of Namibia, is an excellent example of this pattern. Biogeographically, the province is interesting because it repre¬ sents the northern limit of several southern African habitats and species. The entry of the coastal Namib Desert from Namibia into the southern coastal areas of the province as far north as the city of Namibe, as well as the continuation of the Kaokoveld and the Namibian savanna woodlands pro¬ vides a clear dispersal path for southern African taxa. Likewise, the so-called Pro-Namib extends from northwestern Namibia, through Namibe, to the southern areas of Benguela Province. Despite these similarities, the influence of the Cunene River as a barrier should not be underestimated. There are a number of endemic Angolan species of widespread genera in southern Africa, such as Pedioplanis haackei, P huntleyi, and the undescribed species of Rhoptropus collected in our expe¬ dition, as well as the endemic genus Kolekanos. Given the concentration of reptile diversity in Namibe Province, conservation in this region is of special concern. Of the 95 species of reptiles that occur in the province (Table 1), several have small distributions within Namibe and ten are endemic, including the Slender Feather-tailed Gecko, Kolekanos plumicaudus and the recently described Kaokoveld Girdled-Lizard, Cordylus namakuiyus. As noted by Marques (2015), the majority of the amphibians and reptiles of Angola (1) have not been accessed by the IUCN or are listed as Data Deficient, and (2) are known from fewer than five published records for the country since the first studies published in early 1860s. Roughly one third of Namibe Province, 19,600 km 2 out of 57,091 km 2 , is protected as either national park or nature reserve. In contrast to more populated provinces, human activities in Namibe that present significant threats to the herpetofauna are limited. The majority of the human population is concentrated around Namibe and Tombwa, and the main economic activities are fish¬ eries and traditional pastoralism. While there are no studies of the impact of livestock on the her¬ petofauna within Namibe Province, negative impacts are known worldwide in other regions, including South Africa (Bauer and Branch 2003; Fabricius et al. 2003; Smart et al. 2005). Yet the low densities of livestock and the nomadic nature of the populations practicing pastoralism suggest that this is not a major threat to the amphibians and reptiles of Namibe. In contrast to neighboring Namibia (Herrmann and Branch 2013), there are no major mining activities in the province, even if these activites show recent signs of increase, which may in the future threathen some species. Other threats, such as climate change, are believed to have negative consequences on the distribu¬ tion and abundance of southern African lizards (Erasmus et al. 2002; Bates et al. 2014). Extreme climatic events can facilitate wildfires, which unambiguously affect the availability of important habitats for reptiles (Meik et al. 2002). Lastly, for several taxa (e.g., pythons, chameleons, varanids), the impact of human harvesting for food, traditional medicine and the pet trade should not be overlooked. This is known from other African countries (Weldon et al. 2007; Alves et al. 2008; Segniagbeto et al. 2013), and we did encounter one instance of a python being sold during our brief survey. Additional faunal surveys are clearly still needed for Namibe Province. Even if this is one of the most herpetofaunally well known provinces in Angola, the new species recently described sig¬ nal that diversity likely remains underestimated, especially for groups of species that are similar in external appearance. In addition to surveying new areas, it is also important to sample type locali¬ ties of previously described species. Because many of the original topotypes were lost or destroyed, especially material described by Bocage and originally housed in the Lisbon Museum (Ceriaco 2014), new topotypic material with associated genetic resources will help to address many taxo¬ nomic issues, including for groups containing undescribed cryptic diversity. Angola sits at a cros¬ sroads of southern and central Africa and is important to understanding phylogenetic and biogeo- CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 43 graphic patterns across sub-Saharan Africa. The specimens reported here were collected in the first of an ongoing series of joint American-Angolan herpetofaunal expeditions that will build local capacity within Angola and provide accessible data resources to the scientific and conservation communities through georeferenced biodiversity informatics databases (e.g., GBIF.org, Vertnet.org), molecular databases (GenBank), and digitized morphological resources (high-resolu¬ tion x-ray CT-scans). Combining data from new field surveys with information from both museum specimens and literature records (Marques et al., in prep.) will provide the first detailed picture of Angola’s herpetofaunal diversity. Acknowledgments We thank the Angolan Ministry of Environment and INBAC for providing the institutional support and permits for this work. Sold Kuedikuenda, former director of INBAC provided the nec¬ essary collecting and exporting permits and gave institutional support to the expedition. The Sec¬ retary of State for Biodiversity, Paula Francisco, provided institutional support and guidance to the expedition since its beginning. We also thank provincial and regional offices of Biodiversity Affairs for all their support and cooperation. Alvaro Baptista and his family provided logistical help dur¬ ing field work in Namibe. We also want to thank to the Methodist University of Angola, especial¬ ly Teresa Silva Neto and Luis Sebastiao for their support during our stay in Angola. Two anony¬ mous reviewers provided thoughtful criticisms that gave us pause and that we believe led to an improvement of the presentation. However, we accept full responsibility of all acts of commission and/or omission that persist herein. This work was partly funded by a US National Science Foundation grant to DCB and AMB (DEB 1202609 and 1019443) and a grant to DCB and AMB from the JRS Biodiversity Founda¬ tion. Literature Cited Adolphs, K. 2006. Bibliotheca Cordyliformium. 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Dr. Karl Jordan’s Expedition to South West Africa and Angola: Herpetological collection. Novitates Zoologicae 40:115-146. Pasteur, G. 1964. Recherches sur revolution des lygodactyles, lezards Afro-Malgaches actuels. Travaux de I’lnstitut Scientifique Cherifien, Serie Zoologie 9:1-132. Perret, J.-L. 1976. Revision des amphibiens africains et principalement des types conserves au Musee Boca¬ ge de Lisbonne. Arquivos do Museu Bocage, Segunda Serie 6(2): 15-34. Poynton, J.C., and W.D. Haacke. 1993. On a collection of amphibians from Angola including a new species of Bufo Laurenti. Annals of the Transvaal Museum 36(2):9-16. Ruas, C. 1996. Contribuigao para o conhecimento da fauna de batraquios de Angola. Parte I: Familias Pipi- dae, Bufonidae, Microhylidae, Ranidae, Hemisidae e Arthroleptidae. Garcia da Orta Serie de Zoologia 21 (1): 19—41. Ruas, C. 2002. Batraquios de Angola em colecgao no Centro de Zoologia. Garcia da Orta Serie de Zoologia 24(1-2): 139-146. Schmidt, K.P. 1936. The amphibians of the Pulitzer-Angola Expedition. Annals of the Carnegie Museum 25:127-133. Schmidt, K.P. 1933. The reptiles of the Pulitzer-Angola Expedition. Annals of the Carnegie Museum 22: 1-15. Seabra, A. 1906a. Aves de Porto Alexandre. Jornal de Sciencias Mathematicas, Physicas e Naturaes, Segun¬ da Serie 7(27): 143-148. Seabra, A. 1906b. A proposito de algumas especies de Microchiropteros d’Angola. Annaes de Sciencias Naturaes 10:81-82. Seabra, A. 1906c. Ribeirinhas e palmipedes das margens do Rio Cunene. Annaes de Sciencias Naturaes 10:83-90. Seabra, A. 1906d. Aves da exploracao de Fr. Newton em Angola - Subsidios para o conhecimento da destri- CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 49 buigao geographica das aves d’Africa occidental. Annaes de Sciencias Naturaes do Porto 10:153-159. Seabra, A. 1907. Sur quelques oiseaux d’Angola envoyes par Francisco Newton. Contribution a l’etude de la distribution geographique des oiseaux de l’Afrique occidentale. Bulletin de la Societe Portugaise des Sciences Naturelles 1:41-45. Segniagbeto, G.H., F. Petrozzi, A. Aidam, and L. Luiselli. 2013. Reptiles traded in the fetish market of Lome, Togo (West Africa). Herpetological Conservation and Biology’ 8(2):400-408. Smart, R., M. J. Whiting, and W. Twine. 2005. Lizards and landscapes: integrating field surveys and inter¬ views assess the impact of human disturbance on lizard assemblages and selected reptiles in a savanna in South Africa. Biological Conservation 112(2005):23—31. Stanley, E.L., L.M.P. Ceriaco, S. Bandeira, H. Valerio, M.F. Bates, and W.R. Branch. 2016. A review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the description of a new species from the Pro-Namib desert. Zootaxa 4061:201-226. Themido, A.A. 1941. Repteis e batraquios das colonias Portuguesas (Catalogo das colecgdes do Museu Zoo- logico de Coimbra). Memorias e Estudos do Museu Zoologico da Universidade de Coimbra 119:1-28. Tilbury, C. 2010. 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African Journal of Herpetology 56(1): 77-83. 50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 51 Appendix Table 1 52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Notes 1 Because of the morphological similarity among many Tomopterna species, some older records might be referable to this taxon. 2 Records attributed in the literature to Tomopterna cryptotis , e.g., 25 km W of Virei (Poynton and Haacke 1993; Ruas 1996, 2001) and Miranda (Boulenger 1907a; Poynton and Haacke 1993; Ruas 1996, 2001) are likely referable to T. tandyi, but this requires reexamination of the relevant material. 3 Some records may be referable to L. bradfieldi Hewitt, 1932. 4 Lygodactylus lawrencei Hewitt, 1926 is a rocky, dry savanna species from the Kaokoveld regions in northern Namib¬ ia, extending into southern Angola (Pasteur 1964:70 [Fig. 18]; Branch 1998:247; Uetz and Hosek 2014). New surveys and aquisition of new fresh material is absolutely required to clarify the extention range of L. lawrencei in Angola, given the lack of literature records for this species in the country. 5 Certainly in error, see Greer (1967). 6 It is plausible that these records correspond to the subspecies angolensis. However, due to the destruction of the specimens used by Bocage in the Lisbon Museum fire and the impossibility of confimring their identity, we opt to main¬ tain the original identification. 7 The taxonomy of this species is in flux and a revisionary work on the nigrolineatus complex is undeiway (D.G. Broadley and M. Bates, pers. comm.). 8 The species was described by Broadley and Schatti (1997: 172), from Namibia, near the Cunene River at Ruacana, western Ovamboland. Bauer et al. (2001:75-76, 79) suggested that this species should be expected to occur in southern Angola, due to the continuity of the mopaneveld habitat of the species on either side of the Cunene River. 9 The species has an irregular distribution from eastern Zimbabwe and the Okavango Swampa, to Angola and Lake Malawi, through Cameroon (Hughes 1985:519 [Fig. 11]; Branch 1998:94; Chirio and LeBreton 2007:518; Wallach et al. 2014:546). CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 53 Table 1 - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References AMPHIBIANS Anura Pipidae Genus Xenopus Wagler, 1827 Xenopus petersii Bocage, 1895 Mossamedes [= Namibe] (Bocage 1867; Ruas 1996, 2002). Bufonidae Genus Sclerophrys Tschudi, 1838 Sclerophrys gutturalis (Power, 1927) Mossamedes [= Namibe] (Bocage 1867); Cima [= Giraul de Cima] (Poynton and Haacke 1993; Ruas 1996, 2002); Saco do Giraul (Poynton and Haacke 1993; Ruas 1996, 2002); Curoca (Poynton and Haacke 1993; Ruas 1996, 2002). Sclerophrys maculata (Hallowell, 1854) Cainde (Poynton and Haacke 1993; Ruas 1996, 2002); 16 km W of Vila Nova (Poynton and Haacke 1993, Ruas 1996, 2002); this study. Sclerophrys garmani (Meek, 1897) Mossamedes [= Namibe] (Bocage 1895; Ruas 1996, 2001). Genus Poyntonophrynus Frost, Grant, Falvovich, Bain, Haas, Haddad, de Sa, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Biotto, Moler, Drewes, Nussbum, Lynch, Green and Wheeler, 2006 Poyntonophrynus dombensis (Bocage 1895) Assungao (Poynton and Haacke 1993; Ruas 1996). Poyntonophrynus grandisonae (Poyn- ton and Haacke 1993) 5 km E of Assungao (Poynton and Haacke 1993; Ruas 1996); Carac- ulo (Poynton and Haacke 1993; Ruas 1996); Salona [= Saiona] River, 2 km N of Cainde (Poynton and Haacke 1993; Ruas 1996); 20 km W of Virei (Poynton and Haacke 1993; Ruas 1996). Microhylidae Genus Phrynomantis Peters, 1867 Phrymomantis annectens Wemer, 1910 Mutiambo River (Poynton and Haacke 1993; Ruas 1996); Caraculo (Poynton and Haacke 1993; Ruas 1996); this study. Brevicipitidae Genus Breviceps Merrem, 1820 Breviceps adspersus Peters, 1882 Biballa [= Bibala] (Bocage 1895; Ruas 1996, 2002); Mossamedes [= Namibe] (Bocage 1873b). Hemisotidae Genus Hemisus Wagler, 1827 Hemisus guineensis Cope, 1865 Mossamedes [= Namibe] (Bocage 1887b, 1895; Ruas 1996). Arthroleptidae Genus Leptopetlis Gunther, 1859 Leptopelis anchietae (Bocage, 1873) Mossamedes [= Namibe] (Bocage 1873b; Boulenger 1882; Loveridge 1957). Phrynobatrachidae Genus Phrynobatrachus Gunther, 1862 Phrynobatrachus natalensis (Smith 1849) Cunene mouth (Poynton and Haacke 1993; Ruas 1996, 2002). 54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References Pyxicephalidae Genus Amietia Dubois, 1987 Amietia angolensis (Bocage, 1866) This study. Genus Tomopterna Dumeril and Bibron, 1841 Tomopterna damarensis Dawood and Channing, 1999 This study 1 . Tomopterna krugerensis Passmore and Carruthers, 1975 Namibe (Channing 2001). Tomopterna tandyi Channing and Bogart, 1996 Namibe (Channing 2001) 2 . Tomopterna tuberculosa (Boulenger, 1882) Biballa [= Bibala] (Bocage 1895; Ruas 1996); bottom of Leba Pass (Poynton and Haacke 1993; Ruas 1996). REPTILES Testudines Pelomedusidae Genus Pelomedusa Wagler, 1830 Pelomedusa subrufa (Bonnaterre, 1789) Mossamedes [= Namibe] (Bocage 1887, 1895; Loveridge 1941); Mucungo (Shmidt 1933; Loveridge 1941); Maconjo (Bocage 1895; Loveridge 1941); Capangombe (Bocage 1887, 1895; Loveridge 1941). Testudinidae Genus Kinixys Bell, 1827 Kinixys belliana Gray, 1830 Capangombe (Bocage 1895; Loveridge and Williams 1957). Genus Stigmochelys Gray, 1873 Stigmochelys pardalis (Bell, 1828) Mossamedes [= Namibe] (Bocage 1895); Capangombe (Loveridge and Williams, 1957). Squamata Gekkonidae Genus Afrogecko Bauer, Good and Branch, 1997 Afrogecko ansorgii (Boulenger, 1907). Maconjo (Boulenger 1907b; Bauer et al. 1997). Genus Chondrodactylus Peters, 1870 Chondrodactylus fitzsimonsi (Loveridge, 1947) Praia das Conchas (Laurent 1964a); around Mo 9 amedes [= Namibe] in the road to Sa da Bandeira [= Lubango] (Laurent 1964a); this study. Chondrodactylus pulitzerae (Schmidt, 1933) Mossamedes [= Namibe] (Bocage 1867, 1895; Loveridge 1947; Laurent 1964a); Pico Azevedo (Schmidt 1933; Barbour and Loveridge 1946, 1947; Marx 1959); Curoca River (Bocage 1887; Loveridge 1947); this study. Genus Hemidactylus Oken, 1817 Hemidactylus longicephalus Bocage, 1873. Capangombe (Bocage 1873b, 1895, 1897; Loveridge 1947); Curoca Rriver (Bocage 1895, 1897; Loveridge 1947). CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 55 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References Genus Kolekanos Heinicke, Daza, Greenbaum, Jackman and Bauer, 2014 Kolekanos plumicaudus (Haacke, 2008). Tambor (Haacke 2008; Mashinini and Mahlangu 2013); Curoca River (Haacke 2008); 11 km NE of Iona along track towards Oncocua (Haacke 2008; Heinicke et al. 2014). Genus Lygodactylus Gray, 1864 Lygodactylus capensis (Smith, 1849) Mucungu (Schmidt 1933; Loveridge 1947); Capangombe (Bocage 1895) 3 . Lygodactylus lawrencei Hewitt, 1926 4 Genus Pachydactylus Wiegamnn, 1834 Pachydactylus angolensis Loveridge, 1944 Mogamedes [= Namibe] to S a da Bandeira [= Lubango] at Praia das Conchas (Laurent 1964a); Lucira (Bauer 1999); San Nicolau [= Sao Nicolau] (Bauer 1999); Lungo (Bauer 1999); Saco de Giraul (Bauer 1999); this study. Pachydactylus caraculicus FitzSimons, 1959 Lungo (FitzSimons 1959); Caraculo (FitzSimons 1959; Haacke 1970; Mashinini and Mahlangu 2013); Giraul de Cima, river (FitzSimons 1959); 36 mi. northwest of Mocamedes [= Namibe] (Bauer 1999). Pachydactylus cf. oreophilus McLach- land and Spence, 1967 Assuncao (= Assuncao) (Bauer 1999); Caraculo (Bauer 1999); 20 km W Virei (Bauer 1999); 6 km S of Coroca River towards Iona (Bauer 1999); Saiona River, 25 km NW Cainde (Bauer 1999); Mutiambo River on road to Lucira (Bauer 1999); Tambor (Bauer 1999); 7 km from Iona towards Oncocau, Iona Reserve (Bauer 1999); Furnas (Bauer 1999). Pachydactylus punctatus Peters, 1854 60 km of the road of Mogamedes [= Namibe] to Sa da Bandeira (Laurent 1964a); Mogamedes [= Namibe] (Laurent 1964a); 35 km south of Mogamedes [= Namibe] (Laurent 1964a); Pico Azevedo (Schmidt 1933; Loveridge 1947); 11 mi NE of Mocamedes [= Namibe] (Bauer and Branch 1995). Pachydactylus rangei (Andersson, 1908) Mossamedes [= Namibe] (Haacke 1976b); Curoca River (Haacke 1976b); Porto Alexandre (Haacke 1976b); Cunene mouth (Haacke 1976b); Lacrau (Haacke 1976b); Namib Desert (Mertens 1937). Pachydactylus vanzyli (Steyn and Haacke, 1966) Espinheira (Haacke 1976a); Kakolo windmill (Haacke 1976a). Genus Rhoptropus Peters, 1869 Rhoptropus afer Peters, 1869 Maconjo (Bocage 1873b); Capangombe (Bocage 1873b, 1895, 1897b); Mogamedes [= Namibe] (Boulenger 1885); Curoca River (Bocage 1887b, 1895, 1897b). Rhoptropus barnardi Laurent 1964a 60 km from Mogamedes [= Namibe] to Sa da Bandeira [= Luban¬ go] (Laurent 1964a); this study. Rhoptropus biporosus Fitzsimons, 1957 Pico Azevedo (Bauer and Good 1996); this study. Rhoptropus boultoni boultoni Schmidt, 1933 60 km from Mogamedes [= Namibe] to S a da Bandeira [= Luban¬ go] (Laurent 1964a); Pico do Azevedo (Schmidt 1933; Mertens 1938; Barbour and Loveridge 1946; Marx 1959; McCoy and Rich¬ mond 1966); this study. 56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & Referencess Rhoptropus boultoni montanus Lau¬ rent, 1964 This study. Rhoptropus taeniostictus Laurent 1964a Mucungu (Schmidt 1933); “60 km from Mogamedes [= Namibe] to Sa da Bandeira [= Lubango] (Laurent 1964a); this study. Lacertidae Genus Heliobolus Fitzinger, 1843 Heliobolus lugubris (A. Smith, 1838) Maconjo (Bocage 1895); Capangombe (Bocage 1895); Konondoto (Boulenger 1921); this study. Genus Meroles Gray, 1838 Meroles anchietae (Bocage, 1867) Mogamedes (= Namibe) (Bocage 1867, 1895, 1897; Boulenger 1887; Loveridge 1936); this study. Meroles reticulatus (Bocage, 1867) Mogamedes (= Namibe) (Bocage 1867; Boulenger 1887; Loveridge 1936); Coroca River (Bocage 1895, 1897; Boulenger 1887; Bauer and Gunther 1995; this study. Genus Nucras Gray, 1838 Nucras tessellata (Smith 1838) Maconjo (Bocage 1895; Broadley 1972); 34 km from Mogamedes [= Namibe] to Sa da Bandeira [= Lubango] (Laurent 1964; Broadley 1972). Genus Pedioplanis Fitzinger, 1843 Pedioplanis benguellensis (Bocage, 1867) Maconjo (Boulenger 1921; Bauer and Gunther 1995; Conradie et al. 2012); Capangombe (Bocage 1895; Monard 1937); Mossamedes [= Namibe] (Bocage 1887, 1895; Monard 1937); this study. Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012 Red Canyon at Lake Arco (Conradie et al. 2012); 10 km south of Lake Arco (Conradie et al. 2012); Road to Tambor at giant Welwitchia (Con¬ radie et al. 2012); Road from Lake Arco to Espinheira (Conradie et al. 2012); Omauha Lodge (Conradie et al. 2012); Road to Tambor (Con¬ radie et al. 2012); 20 km north of Omauha Lodge (Conradie et al. 2012); this study. Pedioplanis huntleyi Conradie, Measey, Branch and Tolley, 2012 Omauha Lodge (Conradie et al. 2012). 14 km west of Moimba (Con¬ radie et al. 2012); 23 km west of Moimba (Conradie et al. 2012); 26 km east of Iona (Conradie et al. 2012); 16 km east of Iona (Conradie et al. 2012); 8 km northeast of Iona (Conradie et al. 2012); Road to Onocua 7 km NE from Iona (Conradie et al. 2012); 26 km SE of Onocua (Conradie et al. 2012). Scincidae Genus Eumecia Bocage, 1870 Eumecia anchietae Bocage, 1870 Mogamedes [= Namibe] (Boulenger 1887) 5 . Genus Mochlus Gunther, 1864 Mochlus sundevallii (Smith, 1849) Campangombe (Bocage 1895); Mogamedes (= Namibe) (Bocage 1867; 1895); Curoca River (Bocage 1895); lOmls E of Caracul [= Caraculo] (Haacke 1965; Broadley 1966). CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 57 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References Genus Panaspis Cope, 1868 Panaspis cabindae (Bocage, 1866) Capangombe (Bocage 1895, 1897). Genus Sepsina Bocage, 1866 Sepsina angolensis Bocage, 1866 Capangombe (Bocage 1895; Monard 1937). Sepsina copei Bocage, 1873 Biballa [= Bibala] (Bocage 1895, 1897). Genus Trachylepis Fitzinger, 1843 Trachylepis acutilabris (Peters, 1862) Cahinde-Ongueira (Hellmich 1957a); Mossamedes [= Namibe] desert, 35 km south from the city (Laurent 1964a); Curoca River (Bocage 1895); this study. Trachylepis bayonii bayonii (Bocage, 1872) Mogamedes [= Namibe] (Boulenger 1887). Capangombe (Bocage 1895); 50 km Mogamedes [= Namibe] to Sa Trachylepis binotata (Bocage, 1867) da Bandeira [= Lubango]" (Laurent 1964a); Maconjo (Bauer et al. 2003). Trachylepis chimbana (Boulenger, 1887) Chimba River (Boulenger 1887; Bocage 1872, 1895, 1897; Broadley 1975a); Assungao (Broadley 1975a); Maconjo (Bocage 1895, 1897; Broadley 1975a), Mogamedes [= Namibe] (Boulenger 1887); Lucira (Broadley 1975a); Chapeau Armando [= Chapeu Armando] turnoff, Mossamedes [= Namibe] (Broadley 1975a); Caraculo (Broadley 1975a); 14 km NE Caraculo (Broadley 1975a); Capangombe (Bocage 1895, 1897; Broadley 1975a); Mossamedes [= Namibe] (Boulenger 1887); Saiona River, NW of Cainde (Broadley 1975a); Cainde (Broadley 1975a); Coroca River (Broadley 1975a). Trachylepis hoeschi (Mertens, 1954) Irachylepis lacertijormis (Peters, 1864) Camde (Broadley 19/6); 14 km NL ol Caraculo (Broadley 19/6a). Trachylepis laevis (Boulenger, 1907) Maconjo (Boulenger 1907b); Munhino (Laurent 1964a); this study. Trachylepis occidentalis (Peters, 1867) Curoca River (Bocage 1895), 35 km south of the city of Mogamedes [= Namibe] (Laurent 1964a); this study. Inamango River on Lucira road (Broadley 1975a); Mucungu (Schmidt 1933; Broadley 1975a); Sao Nicolau [= Sao Nicolau] (Broadley 1975a); 17 km N of Sao Nicolau [= Sao Nicolau] (Broadley 1975a); Caraculo (Broadley 1975a); 15 km W of Carac- ulo (Broadley 1975a); Cima [= Giraul de Cima] (Broadley 1975a); Trachylepis punctulata (Bocage, 1872) Pico Azevedo (Broadley 1975a); 23 km W of Virei (Broadley 1975a); Coroca River (Bocage 1872, 1895, 1897; Boulenger 1887; Broadley 1975a, 2000); 6 km S of Rio Coroca on Iona road (Broadley 1975a); Porto Alexandre [= Tombua] (Broadley 1975a); 30 km N of Tambor (Broadley 1975a); Octchinfengo River on Onocua road, Iona Reserve (Broadley 1975a); Cunene mouth (Broadley 1975a); this study. Trachylepis sulcata (Peters, 1867) Capangombe (Bocage 1895); Munhino (Laurent 1964a); this study. 58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References Trachylepis varia (Peters, 1867) Mogamedes [= Namibe] (Bocage 1867); Biballa [= Bibala] (Bocage 1872); this study. Genus Typhlacontias Bocage, 1873 Typhlacontias johnsonii Anderson, 1916 Curoca River (Bocage 1895; Haacke 1997); Porto Alexandre (= Tombwa) [= Tombua] (Haacke 1997); Lacrau (Haacke 1997). Typhlacontias punctatissimus punctatis¬ simus Bocage, 1873 Coroca River (Bocage 1873, 1887 1895, 1897); Mossamedes [= Namibe] (Boulenger 1887); Porto Alexandre [= Tombua] (Haacke 1997). Typhlacontias punctatissimus bogerti Laurent, 1964 Mossamedes [= Namibe] desert, 35 km south from the city (Lau¬ rent 1964a); Curoca River (Haacke 1997); Mogamedes [= Namibe] (Haacke 1997); 10 km S of Mogamedes [= Namibe] (Haacke 1997); 34 km S of Mogamedes [= Namibe] (Haacke 1997); 8 km SE of Pico Azevedo (Haacke 1970); Kakolo windmill (Haacke 1997); this study. Typhlacontias rudebecki Haacke, 1997 Sao Nicolau (Haacke 1997). Varanidae Genus Varanus Merrem, 1820 Varanus albigularis angolensis Schmidt, 1933 This study. Varanus albigularis albigularis (Daudin, 1802) Chimba River (Bocage 1895); Biballa [= Bibala] (Bocage 1895) 6 . Chamaeleonidae Genus Chamaeleo Laurenti, 1768 Chamaeleo anchietae Bocage, 1872 Namibe (Tilbury 2010: 451). Chamaeleo dilepis Leach, 1819 Mogamedes [= Namibe] (Bocage 1867b; 1887; 1895); Chimba (Hellmich 1957a). Chamaeleo namaquensis Smith, 1831 Mogamedes [= Namibe] (Gunther 1865; Bocage 1867, 1872, 1895; Boulenger 1887). Agamidae Genus Agama Daudin, 1802 Agama aculeata Merrem, 1820 Biballa [= Bibala] (Bocage 1895); Mogamedes [= Namibe] (Bocage 1887b, 1895); Virei-Cahinde (Hellmich 1957a); Molundo (Monard 1937); Chimporo (Monard 1937). Agama anchietae Bocage, 1896 Mogamedes [= Namibe] (Bocage 1896, 1897); 100 km southeast of Mogamedes [= Namibe] (Laurent 1964); Maconjo (Boulenger and Power 1921); this study. Agama planiceps planiceps Peters, 1862 Biballa [= Bibala] (Bocage 1895); Pico Azevedo (Schimdt 1933); this study. Cordylidae Genus Cordylus Laurenti, 1768 Cordylus namakuiyus Stanley, Cerlaco, Bandeira, Valerio, Bates and Branch, 2015 Caraculo, on the road from Lubango and Namibe (Stanley et al. 2016); Pico Azevedo (Stanley et al. 2016); road between Namibe and Omauha Lodge (Stanley et al. 2016); this study. CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 59 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References Gerrhosauridae Genus Cordylosaurus Gray, 1865 [1866] Cordylosaurus subtessellatus (Smith, 1844) Curoca River (Bocage 1895); this study. Genus Gerrhosaurus Wiegmann, 1828 Gerrhosaurus nigrolineatus Hallowed, 1857 Capangombe (Bocage 1895; Loveridge 1942) 7 . Gerrhosaurus skoogi Andersson, 1916 Porto Alexandre [= Tomboa] (FitzSimons 1953); this study. Genus Matobosaurus Bates and Tol¬ ley, 2013 Matobosaurus maltzahni (De Grys, 1938) Chimba River (Bocage 1895); Mogamedes [= Namibe] (Bocage 1895); Tambor (Bates et al. 2013); Omauha Lodge (Bates et al. 2013). Serpentes Typhlopidae Genus Afrotyphlops Broadley and Wallach, 2009 Afrotyphlops anomalus (Bocage, 1873) Biballa [= Bibala] (Bocage 1895; Broadley and Wallach 2009); Mogamedes [= Namibe] (Bocage 1873). Afrotyphlops schlegelii (Bianconi, 1847) Biballa [= Bibala] (Bocage 1873, 1886, 1895, 1897a; Loveridge 1933). Leptotyp hlopid ae Genus Leptotyphlops Fitzinger, 1843 Leptotyphlops scutifrons (Peters, 1854) Biballa [= Bibala] (Bocage 1873, 1895), Capangombe (Bocage 1895). Pythonidae Genus Python Daudin, 1803 Python natalensis Smith, 1840 Maconjo (Bocage 1895; Broadley 1984); Giraul River (Bocage 1896; Broadley 1984); this study. Viperidae Genus Bids Gray, 1842 Bids arietans (Merrem, 1820) Mogamedes (= Namibe) (Gunther 1865; Managas 1981). Bids caudalis (Smith, 1839) Capangombe (Bocage 1895, Managas 1981); Mogamedes [= Namibe] (Gunther 1865; Monard 1937; Managas 1981); Mossamedes [= Namibe] Desert, 35 km south from the city (Lau¬ rent 1964a); Curoca River (Bocage 1895; Monard 1937; Managas 1981). Genus Causus Wagler, 1830 Causus resimus (Peters, 1862) Chimba River (Bocage 1895; Managas 1981); Biballa [= Bibala] (Bocage 1895; Managas 1981); Maconjo (Bocage 1895); Macujo [= Maconjo] (Managas 1981). Causus rhombeatus (Lichtenstein, 1823) Cuce River (Ferreira 1897); Mossamedes [= Namibe] (Bocage 1887a, 1895). 60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 2 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References Lamprophiidae Genus Aparallactus Smith, 1849 Aparallactus capensis capensis Smith, 1849 Biballa [= Bibala] (Bocage 1895, 1897a; Loveridge 1944b; de Witte and Laurent 1947). Genus Boaedon Dumeril, Bibron and Dumeril, 1854 Boaedon fuliginosus complex Biballa [= Bibala] (Bocage 1895); Capangombe (Bocage 1895). Genus Lycophidion Fitzinger, 1843 Lycophidion hellmichi Laurent, 1964 Capolopopo (Laurent 1964a; Broadley 1991, 1996). Lycophidion multimaculatum Boettger, 1888 Mo 9 amedes [= Namibe] (Bocage 1895; Laurent 1968; Broadley 1996). Genus Hemirhagerrhis Boettger, 1896 Hemirhagerrhis viperina (Bocage, 1873) Cuce River (Ferreira 1897); Munhino (Bogert 1940; Broadley 1995b; Broadley and Hughes 2000); Maconjo (Bocage 1895; Broadley 1995b; Broadley and Hughes 2000); Capangombe (Bocage 1895; Broadley 1995b; Broadley and Hughes 2000); Huxe (Broadley 1997b; Broadley and Hughes 2000); Lungo (Broadley 1997b; Broadley and Hughes 2000); Caraculo (Broadley 1997b; Broadley and Hughes 2000). Genus Psammophis Boie, 1825 Psammophis leopardinus Bocage, 1887 Capangombe (Bocage 1887; Broadley 2002); Mogamedes [= Namibe] (Bocage 1887, 1895; Loveridge 1957); Iona (Broadley 2002; Hughes and Wade 2002). Psammophis mossambicus Peters, 1882 Mossamedes [= Namibe] (Loveridge 1940; Broadley 2002); Coroca River (Bocage 1895); Porto Alexandre [= Tomboa] (Loveridge 1940). Psammophis namibensis Broadley, 1975 Mossamedes [= Namibe] (Broadley 1975b, 2002b); Pico Azevedo (Broadley 2002); Coroca River (Broadley 1975b, 2002); Cunene mouth (Broadley 1975b, 2002b); Cunene Forde, 15 km NE, Iona Res. (Broadley 2002). Psammophis notostictus Peters, 1867 Sao Nicolao River (Boulenger 1896; Loveridge 1940; Broadley 1975b; 2002); Coroca River (Bocage 1887, 1895; Monard 1937; Loveridge 1940; Broadley 1975b, 2002); this study. Psammophis subtaeniatus Peters, 1882 Biballa [= Bibala] (Bocage 1895, 1897a; Loveridge 1940); Macon¬ jo (1895, 1897a; Loveridge 1940; Broadley 2002); Chao de Chella (Broadley 2002). Psammophis trigrammus Gunther, 1865 Sao Nicolao [= Sao Nicolau] River (Bocage 1887; Loveridge 1940; Broadley 1977, 2002; Wallach et al. 2014); Catara River (Broadley 2002); Coroca River (Broadley 2002b); Iona Reserve, 7 km to Oncocua (Broadley 2002). Genus Prosymna Gray, 1849 Prosymna frontalis (Peters, 1867) Mo 9 amedes [= Namibe] (Boulenger 1893). Prosymna visseri Fitzsimons, 1959 near Caracul [= Caraculo], S. Angola (Fitzsimons 1959; Bauer et al. 2001); Balabaia (Broadley 1980); 5 km S of Chibemba (Broadley 1980). CERIACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 61 Table 1 (continued) - Only published records are included. Additional species known from the province based on unpublished museum records are not included unless also supported by published records. Taxon Occurrences & References Elapidae Genus Afronaja Wallach, Wiister and Broadley 2009 Afronaja mossambica (Peters, 1854) Maconjo (Mananas 1981; Broadley 1974). Afronaja nigricincta (Bogert, 1940) Munhino (Bogert 1940; Managas 1981); Maconjo (Managas 1981); Capangombe (Bocage 1895); Cunene mouth (Managas 1981). Afronaja nigricollis (Reinhardt, 1843) Capangombe (Ferreira 1900b; Managas 1981). Genus Aspidelaps A. Smith, 1849 Aspidelaps lubricus cowlesi Bogert, 1940 Munhino (Bogert 1940; Managas 1981; Broadley and Baldwin 2003). Genus Boulengerina Dollo, 1886 Boulengerina melanoleuca Hallowed, 1857 Capangombe (Ferreira 1900b; Managas 1981). Genus Elapsoidea Bocage, 1866 Elapsoidea semiannulata semiannulata Bocage, 1882 Maconjo (Bocage 1895, 1897c; Loveridge 1944b; Broadley 1971, 1998b; Managas 1981). Colubridae Genus Coluber Linnaeus, 1758 Coluber zebrinus Broadley and Schatti, 1997 8 Genus Crotaphopeltis Fitzinger, 1843 Crotaphopeltis hotamboeia (Laurenti, 1768) Biballa [= Bibala] (Bocage 1895). Genus Philothamnus Smith, 1840 Philothamnus angolensis Bocage 1882 Capangombe (Bocage 1882a, 1897a; Loveridge 1951, 1957; Chirio and LeBreton 2007; Wallach et al. 2014). Philothamnus irregularis (Leach, 1819) Capangombe (Bocage 1882a, 1887, 1895); Mossamedes [= Namibe] (Bocage 1887, 1895). Philothamnus ornatus Bocage, 1872 9 Philothamnus semivariegatus (Smith, 1840) Maconjo (Bocage 1882a); Capangombe (Bocage 1895; Haacke 1985). Genus Telescopus Wagler, 1830 Telescopus finkeldeyi Haacke, 2013 5 km north Namibe [= Namibe] (Haacke 2013). Genus Thelotornis A. Smith, 1849 Thelotornis capensis oatesi (Gunther, 1881) Biballa [= Bibala] (Bocage 1895; Loveridge 1944b). 62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63 Page intentionally left blank PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3, pp. 63-157, 8 figs., 10 tables, Appendix April 29, 2016 Exotic Terrestrial Macro-Invertebrate Invaders in California from 1700 to 2015: An Analysis of Records Robert V. Dowell', Raymond J. Gill 2 , Daniel R. Jeske 3 , and Mark S. Hoddle 4 1 California Department of Food and Agriculture 3288 Meadowview Road, Sacramento, California, USA 95832; Email', bdowell396@gmail.com. 2 California Department of Food and Agriculture 3288 Meadowview Road, Sacramento, California, USA 95832; Email', raymond.gill@cdfa.ca.gov. 3 Department of Statistics, University of California, Riverside, California, USA; Email: Daniel.Jeske@ucr.edu. 4 Department of Entomology and the Center for Invasive Species Research, University of California, Riverside, California, USA; Email: mark.hoddle@ucr.edu. We document that 1,686 exotic terrestrial macro-invertebrates from 278 families have developed self-sustaining populations in California, USA over the period 1700 to 2015. The majority of exotic arthropods are insects (84.4%) followed by mites (7.5%) and spiders (3.2%). The insect Order Hemiptera, Auchenorrhyncha and Sternorrhyncha combined, has the greatest insect representation at 27.6%. The Aphididae is the most successful group of invaders with 183 species established in California. Small, less mobile plant feeders (e.g., plant feeders like thrips, scales, mealybugs, whiteflies and aphids) have successfully invaded more often than large mobile invertebrates (e.g., bees and wasps, butterflies and moths, grasshoppers and katydids, or beetles). The greatest number of invaders (43.6%) most likely came to California from invasion bridgeheads established elsewhere in North America (USA and Canada). The date of first detection is known for approximately 1,116 (66%) invertebrate species and the temporal invasion rate has not remained constant over time. Fewer invaders were detected over 1970-1989 at a rate of ~ 6 species per year. This rate increased to ~ 9.7 new species detected per year from 1990 to 2010, an increase of ~ 62% since 1989. Invasion rates into California have not increased sig¬ nificantly post 11 September 2001 when the Department of Homeland Security assumed responsibility from United States Department of Agriculture-Animal Plant Health Inspection Service (USDA APHIS) for border protection. It is anticipated that invasion rates for invertebrates will continue to increase and more exotic organ¬ isms will establish in California because of increasing trade and tourism and a cor¬ responding lack of increased resources for pest surveillance, detection, and post¬ incursion management. Keywords: invasion rates, invasive species, change point analyses, impacts of invasive species Over the last 25 years there has been growing awareness that exotic invasive organisms can greatly modify our urban, agricultural, and native habitats (Mattson et al. 1994, Barthell et al. 1998, Pimentel et al. 2000, Jass and Klausmeier 2001, Beatty 2002, Huber et al. 2002, Hoddle et al. 2004, Sierwald et al. 2005, Footitt et al. 2006, Haack 2006, Bolger et al. 2000, Kenis et al. 2009, Auke- ma et al. 2010, Aukema et al. 2011, Engelkes and Mills 2011, Roy et al. 201 la,b). The literature is rich in examples, and generally includes instances of invasive species that adversely affect native 63 64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 flora and fauna, reduce agricultural productivity, and impact urban areas. Some invasive species affect human, animal, and plant health, while others degrade and deny citizens the use of parklands and wilderness areas (Dowell 1988; Horsfall 1983, Dowell and Gill 1989, Knight and Rust 1990, Dowell and Krass 1992, Daehler and Strong 1995, Holway 1998, Suarez and Case 2002). How¬ ever, biological invasions are complex processes whose individual components are not easily understood (Kolar and Lodge 2001, Crooks 2005, Barney and Whitlow 2008). The environmental and economic impacts that result from invasion events are the reason that regulatory agencies like the California Department of Food and Agriculture (CDFA), United States Department of Agriculture Animal Plant Health Inspection Service (USDA-APHIS), and the Department of Homeland Security (DHS) are charged to detect and prevent the establishment of non-native species within their jurisdictions. Following the passage of the 2002 Homeland Securi¬ ty Act, USDA-APHIS personnel were transferred into Customs and Border Protection (CBP) under the auspecies of DHS in 2003 and this resulted in the incorporation of the protection of US Agri¬ culture into CBP’s anti-terrorism mission (GAO 2006). US agriculture is a $1 trillion per year enterprise, and this significant change in focus from pest and disease exclusion to detecting terror¬ ist activity raised concerns that US agriculture may experience increased vulnerability to invasive pests and diseases affecting crops and lifestock (GAO 2006). For invasive species managers, researchers, and regulatory agencies, a first step in managing the invasion process is the development of lists of successful invaders and their likely area of ori¬ gin (Dowell and Gill 1989). This information may help identify likely future invaders and their- probable sources, and assist with resource allocation decisions for monitoring, detection, and man¬ agement programs (Crooks et al. 1983, Dowell and Gill 1989, Work et al. 2005, Koch et al. 2011). Additionally, these lists can be used to look for temporal, geographic origin, and species compos- tion changes in invasion rates. The suggestion that the transfer of pest detection and exclusion responsibilities from USD A-APHIS to CBP could have increased the rate of unwanted pest intro¬ ductions (GAO 2006) is a hypothesis that can be examined statistically with a chronologically con¬ structed species detection list. Dowell and Gill (1989) published a list of exotic invaders in California and their likely origins over the period 1955-1988. Although this study (Dowell and Gill 1989) has been cited widely, it encompassed a limited timeframe and utilized one data source. Using Dowell and Gill (1989) as a reference point, the goals of this paper are: 1) to expand the timeframe of analyses to include exot¬ ic terrestrial macro-invertebrates that successfully invaded California from 1700 to 2015; 2) to uti¬ lize a wider range of information resources that document breeding populations of invasive macro¬ invertebrates in California; and 3) use these expanded datasets to examine the composition and ori¬ gins of invading species and how they have changed over time. In particular, this dataset provided an opportunity to determine whether or not invasion rates into California increased after CBP assumed responsibility for managing unwanted pest and disease incursions. Here we refer to exot¬ ic invaders in the broadest sense, non-native species that developed reproducing populations in California. These organisms may or may not cause economic or environmental problems, but the commonality underlying this resident community is their non-native status. Additionally, some species that developed reproducing populations in California may have done so temporarily before being successfully eradicated. In the absence of eradication efforts, these species would likely still be in California, and are therefore considered species that successfully developed breeding popu¬ lations before being deliberately extirpated. DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 65 Methods Construction of an Invasive Terrestrial Invertebrate Species List for Califor¬ nia.— To develop a list of exotic terrestrial macro-inverterbrate invaders (Appendix), we exam¬ ined print and electronic sources for information on breeding populations within the state of Cali¬ fornia, USA. The following criteria were used to develop a documented evidence list (McGeoch et al. 2012): 1) One or more authorities noted that the organism was found in California, and 2) these sources noted that the organism was not native to California; 3) There was evidence of a breeding population of the organism outside of a quarantine situation initiated to contain a possible incipi¬ ent infestation. For example, the discovery of a single individual, such as the male Queensland fruit fly, Bactrocera tryoni (Diptera: Tephritidae) found in California in 1991, was not considered evi¬ dence of a breeding population (Centre for Agriculture and Biosciences International 2016). Organ¬ isms discovered in ports, or at nurseries importing stock from outside California were considered to be contaminants and were within the State of California’s quarantine system and therefore were not included in the list; 4) For analysis of electronic sources, like BugGuide, we included species for which we could find another source to independently confirm the presence of breeding popu¬ lations or when there were one or more pictures posted on web sites documenting the invader of interest from locations outside of quarantine situations within California; 5) If an organism had invaded California several times, and reproducing populations were detected (e.g., oriental fruit fly, Bactrocera dorsalis ) (Stephens et al. 2007), only the first successful reproducing population was considered; 6) Arthropod biological control organisms (i.e., predators and parasitoids) that were intentionally released in California were not included in the list. Neither was the deliberately intro¬ duced European honey bee, Apis mellifera. The point sources of pests arriving in California may not necessarily correspond with the pest’s native home range. Therefore, if a newly detected invasive species was found in California and populations of this pest existed previously elsewhere within North America (i.e., the US or Cana¬ da), the parsimonious decision was made that assumed in the absence of contrary evidence (e.g., molecular-based data if it existed) that the population of interest originated from populations already established in North America and did not represent a novel introduction directly from the native range of the invader. This decision was based partly on the volume of traffic entering Cali¬ fornia each year. In 2010 (most recent data available) some 27.3 million private vehicles entered California along with 7.14 million commercial trucks (California Department of Food and Agri¬ culture 2011). For example, the red imported fire ant, Solenopsis invicta (Hymenoptera: Formici- dae), almost certainly entered California from long-established populations in the eastern United States and it did not invade California directly from its native South America. We have identified in the Appendix invertebrates that are classified not native to California and in some instances are classified as pests. For plant, home and garden, and the environment, we identify those invertebrates listed as pests in the University of California IPM online (UC IPM 2014), sheep and goat pests are from Pirelli (2013), dairy cattle pests are those identified by Geden et al. (2010), horse insect pests are from Meyer et al.(1991), swine pests are from Loftin (2013), and chicken pests are from Stafford III (2011). These sources tend to deal with animals that are established in California or are frequently detected there. We have also identified some pests not established in California (IPPC 2007) but which have been detected here and which are generally considered pests in those areas where they occur. These include the gypsy moth ( Lymantria dispar) (Liebhold 2003), Japanese beetle (Popillia japonica) (Anon. 1997), Mexican bean beetle ( Epilach- na varivestis) (Sanchez-Arroyo 2015), oriental fruit fly (Bactrocera dorsalis), Mexican fruit fly (Anastrepha ludens), Caribbean fruit fly ( Anastrepha suspensa), peach fruit fly (Bactrocera zona- 66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 ta ), Mediterranean fruit fly ( Ceratitis capitata ), melon fly ( Bactrocera cucurbitae), white striped fruit fly ( Bactrocera albistrigata ), guava fruit fly ( Bactrocera correcta) (White and Elson-Harris 1992), boll weevil ( Anthonomus grandis ) (Lange et al. 2008), varroa mite ( Varroa jacobsoni ) (Bessin 2001), red palm weevil {Rhynchophorus vulneratus ) (Rugman-Jones et al. 2013), Diaprepes root weevil {Diaprepes abbreviatus ) (Grafton-Cardwell et al. 2004), Asian longhorn beetle ( Anoplophora glabripennis ) (USDA APHIS 2015), and cereal leaf beetle (Walentra 2014). The year the organism was identified from California is listed unless there is published infor¬ mation supporting the use of an earlier date. Organisms listed in overview works (e.g., Essig 1915, 1926) are assumed to have entered the state in the decade prior to the work’s publication date. For -34% of recorded exotic invertebrates, the date of introduction was either unknown or was uncer¬ tain and therefore not included in analyses. Statistical Analyses. — Change point detection analyses aim to identify when a sequence of time ordered observations changes with respect to the underlying model that generates data. A common type of change point analysis is to look for a change in either the mean or the variance of the data generating model. Change point analyses were used to determine if the model for the annu¬ al number of exotic macroarthropods incurred a change point over the period 1935 to 2010. The time period 2011-2015 was not included because of the time lag in the publication of non-native animals discovered in California therein. Although the data source available for analyses extends to periods earlier than 1935, data quality was perceived to be a potential issue that could influence analyses, hence a conservative approach was taken and this 75 yr period was selected. The change point detection method used was CUSUM (cumulative sum) that detects shifts in the mean in the data series under analysis. The software package Change Analyzer, (2013), was used to identify change points using the CUSUM method. Succinctly, the CUSUM analysis was computed as fol¬ lows: first, the overall average number of exotic arthropds was calculated for the time interval of interest; second, the cumulative sum for the time interval of interest was initialized at zero; and third, cumulative sums were calculated by sequentially adding deviations of current values from the average to the previous cumulative sum. A CUSUM chart, a plot of the cumulative sums ver¬ sus time, was used to identify transitions or change points in the sequence. Potential change points were evaluated using bootstrap procedures to determine their statistical significance. When a sig¬ nificant change point was detected, its location was determined by identifying the last point before the change and the first point after the change. This sequence was divided at the change point and the CUSUM procedure was repeated on each of the new segments to determine if additional sig¬ nificant change points existed. Of particular interest in change point analyses performed on this data set was that of testing the hypothesis that incursions of invasive arthropods into California increased after 11 September 2001 (GAO 2006). We used chi-square analysis (Little and Hills 1978) to compare the composition of species established in California to the worldwide composition of the taxa they represent, and to determine if the composition of the invaders by area of origin differs from global composition. All analyzes herein include the exotic organisms listed in an earlier work (Dowell and Gill 1989) and organisms are grouped by family rather than by order as done in Dowell and Gill (1989). Family-level clas¬ sification provides finer taxonomic resolution and accommodates recent taxonomic nomenclatural changes. We also compared the composition by taxa of invaders from each region to their over-all composition. For this analysis, the expected values were generated by multiplying the percentage of each invading taxa over-all by the total number of invaders from each region. For example the percentage of Coleoptera in the over-all composition of invaders was multiplied by the number of invaders recorded from North America. DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 67 Results and Disussion Composition of Invaders.— We identified 1,686 species of exotic terrestrial macro-inver¬ tebrates that have developed breeding infestations within California over the time period 1700 to 2015 (Appendix). These organisms include earthworms (Annelida: Oligochaeta), mites and spiders (both Arthropoda: Arachnida), centipedes (Arthropoda: Chilopoda), millipedes (Arthropoda: Diplopoda), slugs and snails (both Mollusca: Gastropoda), woodlice (Arthropoda: Isopoda), insects (Arthropoda: Insecta), and one amphipod (Arthropoda: Amphipoda) from 278 different families. The majority of these exotic organisms are insects (84.4%) followed by mites (7.4%) and spiders (3.2%) (Table 1) (Figure 1). A number of organisms on the list (see Appendix) devel¬ oped breeding populations in California that were or are being subjected to Official Action (quarantine and eradica¬ tion) in accordance with inter¬ nationally accepted guidelines (IPPC 2007) by the California Department of Food and Agri¬ culture often in collaboration with the United States Depart¬ ment of Agriculture. Organisms subject to official action are not considered to be established in California. These include the Mediterranean fruit fly, Cerati- tis capitata; Mexican fruit fly, Anastrepha ludens; peach fruit fly, Bactrocera zonata; guava fruit fly, Bactrocera correcta; gypsy moth, Lymantria dispar, boll weevil, Anthonomus gran- dis; Japanese beetle, Popillia japonic a’, Caribbean fruit fly, Anastrepha suspensa; melon fly, Bactrocera cucurbitae; Hall scale, Nilotaspis halli; red palm weevil, Rhynchophorus vulneratus; white striped fruit fly, Bactrocera albistrigata; European grapevine moth, Lobesia botrana\ Mexican Bean Beetle, Epilachna varivestis; Asian gypsy moth, Lymantria dispar asiatica; and Asian longhorn beetle, Anoplophora glabripennis. Table 1. Composition by taxon of exotic terrestrial macro-mverte- brates found breeding in California 1700-2010. Taxon 3 Number of organisms Percent of total organisms Hemiptera: Auchenorrhyncha + Stemorrhyncha 467 27.6 Coleoptera 344 20.4 Lepidoptera 172 10.2 Acari 125 7.4 Diptera 122 7.2 Hymenoptera 96 5.6 Thysanoptera 63 3.7 Hemiptera- Heteroptera 50 2.9 Araneae 53 3.2 Mollusca 43 2.5 Phthiraptera 34 2.0 Dicytoptera 19 1.11 Opisthopora 16 1 Psocoptera 16 1 Isopoda 11 0.6 Orthoptera 9 0.5 Dermaptera 9 0.5 Diploda 9 0.5 Siphonaptera 9 0.5 Thysanura 5 0.3 Collembola 4 0.24 Chilopoda 2 0.12 Embiidina 2 0.12 Amphipoda 1 0.06 Isoptera 1 0.06 Phasmida 1 0.06 Symphyla 1 0.06 Total 1686 a See Appendix for list of organisms. 68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 69 The Hemiptera, Auchenorrhyncha, and Stemorrhyncha combined, has the greatest representa¬ tion (Table 1) with 27.6% of all organisms. Next speciose are the Coleoptera (20.4%), Lepidoptera (10.2%), Acari (7.4%), Diptera (7.2%) and Hymenoptera (5.6%). Remaining taxons have no more than 3.6% of recorded organisms (Table 1). This differs somewhat from a list of non-native species for all of North America that had the Coleoptera ranked first followed by the Hymenoptera, Hemiptera, Diptera and Lepidoptera (Yamanaka et al. 2015). Yamanaka et al. (2015) included releases of biological control agents and excluded species that were eradicated, the opposite of this study. As seen in other studies (Lawton et al. 1986, Gaston et al. 2001, Karatayev et al. 2009), the composition of invading animals deviates significantly from random (x 2 = 257.497; df=25; p < 0.001) (Table 2) based on their worldwide numbers (Borror et al. 1989, Brusca 1997, Shelley 1999, Halliday et al. 2000, Triplehorn and Johnson 2005, Chapman 2009, Platnick 2011, Ramel 2011). Hemiptera (Auchenorryncha and Stemorrhyncha), Acari, Thysanoptera, Psocoptera, Siphonaptera, Dermaptera, Phthiraptera, Dictyoptera, Isopoda, and Thysanura invaded California more often than expected. Conversely, Coleoptera, Lepidoptera, Diptera, Hymenoptera, Araneae, Collembola, Chilopoda and Orthoptera invaded less often than expected. In general, small, less mobile plant feeders (e.g., whiteflies, thrips, mites, aphids and scales) invaded California more often than expected and larger more mobile insects (e.g., bees and wasps, butterflies and moths, grasshoppers, tme bugs, spiders, and beetles) less often than expected. Similar invasion patterns have been seen in other studies (Lawton et al. 1986, Gaston et al. (2001). The most successful group of invaders was the Aphididae (183 species [10.9% of the total]) (Table 3). After the aphids the most numerous invaders were, in descending order, the Curculion- idae (83 species [4.9% of invasive species]), Staphylinidae (74 species [4.4%]), Eriophyidae (72 species [4.2%]), Diaspididae (71 species [4.2%]),), and Pseudococcidae (59 species [3.5%]). Other highly successful invaders were in the Cicadellidae (Hemiptera), Thripidae (Thysanoptera), Coc- cidae (Hemiptera), Psyllidae (Hemiptera), Aleyrodidae (Hemiptera), Formicidae (Hymenoptera), Chrysomelidae (Coleoptera), Tetranychidae (Acari), Miridae (Hemiptera: Heteroptera), Tenebrion- idae (Coleoptera), and four families of moths (Lepidoptera: Gelechiidae, Noctuidae, Pyralidae and Tortricidae) (Table 3). Successful invaders included many well known invertebrates including the codling moth {Cydia pomonella), silverfish {Lepisma saccharina ), European earwig ( Forficula auricularia ), pill- bug ( Armadillidium vulgare), cabbage butterfly ( Pieris rapae ), American ( Periplaneta americana) and German (Blattella germanica) cockroaches, brown garden snail ( Cornu aspersum), house fly (Musca domestica ), praying mantis {Mantis religiosus ), and Argentine ant (Linepithema humile) (Appendix). As discussed below, many successful invaders are important pests of agricultural, hor¬ ticultural and floricultural crops, wilderness areas, livestock, urban gardens, houses, pets, and peo¬ ple. Origins of Invaders. — The origins of 980 of listed invaders are known or suspected (Table 4). The greatest number of invaders came to California from elsewhere in North America (i.e., USA Figure 1 (left). Examples of non-native macro-terrestrial arthropods that have been detected in California. A. Avocado thrips, Scirtothrips perseae Nakahara (Thysanoptera: Thripidae). (Detected 1992). B. Brown widow spider, Latrodectus geometricus Koch (Araneae: Theridiidae) (detected 2003). C. Glassy-winged sharpshooter, Homalodisca vitripennis (Ger- mar) (Hemiptera: Cicadellidae) (detected 1994). D. Brown marmorated stink bug, Ha/yomorpha halys Stal (Hemiptera: Pentatomidae) (detected 2005). E. Citrus leafminer, Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae) (detected 1999). F. Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae) (detected 2008). G. Palm weevil, Rhyn- chophorus vulneratus (Panzer) (Coleoptera: Curculionidae) (detected 2010) (eradicated 2015). H. Gold spotted oak borer, Agrilus auroguttatus Waterhouse (Coleoptera: Buprestidae) (detected 2004). 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Table 2. Expected versus observed number of exotic invertrbrate invaders based on their worldwide numbers. Taxon Expected number of invading species 11 Observed number of invading species 1 Hemiptera: Auchenorrhyncha plus Sternorrhyncha 3 83 468 Coleoptera 3 496 312 Lepidoptera 3 248 167 Acari b 70 125 Diptera 3 248 109 Hymenoptera 3 190 94 Thysanoptera 3 7 59 Hemiptera- Heteroptera 3 58 55 Araneae c 80 52 Mollusca d 39 43 Phthiraptera 3 5 34 Psocoptera 3 5 16 Dictyoptera 3 10 19 Isopoda e 8 11 Diplopoda b 16 9 Orthoptera 3 33 9 Dermaptera 3 3 9 Siphonaptera 3 4 8 Thysanura 3 1 5 Collembola 3 10 4 ChilopodaS 5 2 Embiidina 3 1 2 Isoptera 3 4 1 Phasmida 3 4 1 Symphyla® 1 1 3 Based on numbers in Borrer et al. 1989 and Triplehom and Johnson 2005. b Based on numbers in Halliday et al. 2000. c Based on numbers in Platnick 2011. cl Based on numbers in Chapman 2009. e Based on numbers in rusca 1997. 1 Based on numbers in Ramel 2011. § Based on numbers in Shelley 1999. b rounded to the nearest whole number or to one if less than 0.50. 1 See Appendix for list of organisms, y} = 2,688.37; df = 25; p < 0.001. DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 71 Table 3. The twenty families with the largest Table 4. Number of exotic terrestrial macro¬ number of exotic terrestrial macro-invertebrates invertebrates by origin, found breeding in California. and Canada) (422 species; 43.6%). These invaders were comprised of species native to the USA (e.g. the glassy-winged sharpshooter, Homalodisca vitripennis, which is native to the southeastern USA) (e.g., indigenous exotics (Dodds et al. [2010]) and species exotic to the USA but which likely invaded California from invasion bridgeheads within the USA (e.g., red imported fire ant and European gypsy moth). Europe was the second largest source of exotic invaders (24.8%) followed by Asia (including Japan) (10.2%), Tropical America (Central America and South America) (7.2%), Australia (4.9%), Africa (4.7%), Hawaii (2.2%), Mexico (2.0%), and the Pacific Region (areas bordering the Pacif¬ ic Ocean including New Zealand but excluding Hawaii and Japan) (0.8%) (Table 4). Mexico and Hawaii are listed separately because of their relatively close proximity to Cali¬ fornia, high reciprocal volumes of trade and tourist traffic, and the belief that numerous invaders enter California from these two areas (Dowell and Gill 1989). Organisms from the Mediterranean Region were assigned to either Africa or Europe as appropriate. The distribution of recorded invaders was not uniform (% 2 = 1,386.31; df = 8; p < 0.001) with more species than expected infil¬ trating California from North America and Europe and fewer invaders than expected originating from Tropical America, Australia, Africa, Hawaii, Mexico and the Pacific Region (Table 4). Not all taxa invaded California from all regions (Table 5). Only the Acari, Diptera, Hemiptera Auchenorrhyncha and Stemorrhyncha, and Lepidoptera have representatives from every region. No region has invaders from every taxa (Table 5). Europe has representatives from 20 of 26 taxa followed by North America (15), Africa (15), Tropical America (11), Asia and Australia (10), Hawaii (9), and Mexico (6). Chi-square analysis found the composition of invaders from North America (% 2 = 48.04, Order Family Number of Species 3 Hemiptera Aphididae 183 Coleoptera Curculionidae 83 Coleoptera Staphylinidae 764 Acari Eriophyidae 72 Hemiptera Diaspididae 71 Hemiptera Pseudococcidae 59 Thysanoptera Thripidae 49 Hemiptera Cicadellidae 47 Hemiptera Psyllidae 37 Hemiptera Coccidae 35 Coleoptera Chrysomelidae 30 Lepidoptera Pyralidae 26 Hymenoptera Formicidae 25 Hemiptera Miridae 24 Acari Tetranychidae 23 Lepidoptera Tortricidae 21 Hemiptera Aleyrodidae 20 Lepidoptera Gelechiidae 20 Coleoptera Tenebrionidae 19 Lepidoptera Noctuidae 16 3 See Appendix for list of organisms. Origin Number of organisms 3 Percent of total North America 422 43.6 Europe 243 24.8 Asia 100 10.2 Tropical America 71 7.2 Australia 48 4.9 Africa 46 4.7 Hawaii 2227 2.2 Mexico 20 2.0 Pacific Region 8 0.8 Total 980 a See Appendix 1 for list of organisms, y} = 1,386.31; df=8; p< 0.001. Assumes an average of 108.8 invaders from each region. 72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 df = 10, p< 0.001), Europe (f = 4.975, df= 11, P< 0.001), Africa (x 2 = 13.67, df = 2, p < 0.01), Asia (x 2 = 43.89, df = 5, p < 0.001), Tropical America (x 2 = 11.86, df = 3, p < 0.01), and Australia (X 2 = 20.0, df = 3, p < 0.001) deviated significantly from that seen in Table 1 (Table 5). There were too few species recorded from Hawaii, Mexico and the Pacific region to analyze (Little and Hills 1978). Invasion Frequencies into California Post September 11 , 2001 — Change Point Analyses.— Count data for the number of recorded invasive species by year over the period 1935-2010 (Fig. 2) and the CUSUM plot for this 75-year data set are shown in Fig. 3. Three sig¬ nificant change points were identified, 1941, 1971, and 1983 (Table 6). The mean number of exot- Table 5. Likely origins of the exotic terrestrial macro-invertebrates found breeding in California by taxon. Taxon 3 Mexico North America Europe Africa Asia Hawaii Pacific Region Tropical America Australia Acari 3 24 18 2 4 2 1 4 1 Amphipoda 0 0 0 0 0 0 0 0 1 Araneae 0 4 8 1 0 0 0 4 0 Chilopoda 0 0 2 0 0 0 0 0 0 Coleoptera 1 78 50 4 10 1 0 7 6 Collembola 0 0 3 0 0 0 0 0 0 Dermaptera 0 1 1 1 0 0 0 0 0 Dictyoptera 0 4 2 4 2 0 0 0 0 Diplopoda 0 1 6 0 2 0 0 0 1 Diptera 1 32 17 2 7 4 2 7 5 Embiidina 0 0 2 0 0 0 0 0 0 Hemiptera- Heteroptera 0 17 14 1 43 1 0 3 1 Hemiptera: Auchenorrhyncha + Sternorrhyncha 9 141 58 23 58 8 3 32 21 Hymenoptera 0 33 6 5 5 3 0 3 9 Isopoda 0 0 8 1 0 0 0 0 0 Isoptera 0 0 0 0 0 1 0 0 0 Lepidoptera 5 66 19 1 5 1 1 7 1 Mollusca 0 10 22 1 0 1 0 0 0 Orthoptera 0 5 3 0 0 0 0 0 0 Phasmida 0 1 0 0 0 0 0 0 0 Phthiraptera 0 0 0 0 0 0 0 0 1 Psocoptera 0 0 4 0 0 0 0 0 0 Siphonaptera 0 0 1 0 0 0 0 1 0 Symphyla 0 0 0 0 0 0 0 0 0 Thysanoptera 1 5 9 0 3 0 1 3 1 a See Appendix for list of organisms. DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 73 Year Figure 2. Total number of invertebrates detected in Cal¬ ifornia 1935-2010. Year Figure 3. Plot of CUSUM values for counts of exotic arthropods recorded in California over the period 1935 to 2010 . ic arthropods before and after the change was detected with 95% confidence intervals for the year of the detected change, and thus the degree of confidence was interpreted as indicating that declar¬ ing that the detected change was real. The mean estimates of the numbers of detected exotic arthro¬ pods before and after the detected changes are presented in Table 6. There is no statistical evidence to support the hypothesis that the number of invasive species recorded in California increased after 2001. A truncated data set spanning 1991-2010 (20 years) was further analyzed to assess the effect of 2001 on incursion rates in California. Again no significant change points were detected (results not shown) supporting the results of the analyses using the larger 1935-2010 data set. Table 6. Significant change point results indicating year of detected change, 95% confidence interval (in years), level of confidence that the detected change point is real, and the observed change from the mean num¬ ber of exotic arthopods detected before the change point to the new mean after the change occurred. Year change was detected 95% confidence interval Confidence level (%) Mean number of species before change was detected Mean number of species after change was detected 1941 1936-1949 96 13.83 8.6 1971 1949-1976 95 8.6 4.92 1983 1978-1996 94 4.92 8.89 The change point analysis detected three transitional points underlying exotic arthropod species records for California over the period 1935-2010, 1941, 1971, 1983. The causes underly¬ ing these changes are undetermined and beyond the scope of this study, which sought to determine if a significant change in arthropod numbers occurred after 2001. There are several important caveats to consider here when interpreting the significance of these results. The first concerns intensity and consistency of sampling efforts, resources invested in surveying for invasive pests, and technical capability, especially with regard to recognition and identification of new species. It is highly unlikely that these factors were consistent each year over 1935-2010. Short duration high intensity surveys focusing on invasive ants, spiders, or scale pests, as part of a research project for example, could greatly increase the number of records for new species in a particular taxon. This type of survey work, while valuable, is not uniform through time or across taxa and high intensity 74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 effects with narrow focus probably affects the quality of data sets for analysis. Additionally, data were not adjusted for other covariates that could influence invasion rates and detection and identification of new resident species. These additional factors include increasing rates of tourism, economic development, and population growth, all of which have been correlated with invasion likelihood. Political policies such as the construction and operation of additional border inspection stations, or subsequent closures and reduced hours for stations during economic recessions may also influence the rate at which exotic arthropods enter and establish in Califor¬ nia. Although no significant change in the num¬ ber of exotic arthropods in California was detect¬ ed after 2001, this does not necessarily mean that the numbers of invasive pests entering and estab¬ lishing has not changed. There may be a signifi¬ cant lag period before detection and confirmation of new species occurs, and subsequent analyses of data sets with a longer time series post-2001 may be needed to confirm or refute the result pre¬ sented here. Temporal Patterns of Invasion.— Detection times for 1079 exotic terrestrial macro¬ invertebrates are presented in Table 7. The greatest number (419; 38.8%) of these organisms was detected in California prior to 1930 (Table 7). The temporal invasion rate has not remained constant over time (% 2 = 61.37; df = 7; p < 0.01) with fewer than expected invaders being recorded during the period 1970-1989. The rate of inva¬ sion since 1989 has increased to levels similar to those recorded from 1950 to 1969 (Table 7). Other studies concluded that the rate of introduction of exotic pests was increasing (Aukema et al. 2010, Kirkendall and Faccoli 2010, Roy et al. 2011). The origins of invaders have not remained constant over the time period of this study. There has been a marked increased in the number of invaders from Australia between 1980 and 2010, Asia between 2000 and 2010, Hawaii between 1980 and 1999, and Europe between 1990 and 2010 (Table 8). These trends may reflect increased trade and tourism, which are proxies for propagule pressure. Similarly, the composition of invaders has not remained constant over time. There were marked increases in the number of Coleoptera (11 versus 45), Diptera (9 versus 33) and Hemiptera (Heteroptera) (5 versus 15) detected between 1990 and 2015 and Hymenoptera (11 versus 25) between 1990 and 1999 compared to the preceeding decades (1970-1989) (Table 9). Examining the origin and composition data over time in the Appendix reveals interesting trends. Eighteen invasive wood boring Coleoptera (Buprestidae, Cerambycidae and Curculionidae) were discovered in California between 2000 and 2015 with five species from Asia, eight from North America, four species from Europe, and one from Africa. There were 10 Hymenoptera from Australia discovered between 1980 and 2010: eight Eulophids and two Encyritids. Australia con- Table 7. Number of exotic terrestrial macro¬ invertebrates discovered breeding in California by time period in which they were discovered. Time Period 3 Number Exotic Invaders Average number per year < 1900 112 — 1901-1929 312 108 1930-1939 99 9.9 1940-1949 76 7.6 1950-1959 84 8.4 1960-1969 95 9.5 1970-1979 53 5.9 1980-1989 68 6.8 1990-1999 99 9.96 2000-2010 118 7.9 Total 1116 a See Appendix for data. The time periods <1900 and 1901-1929 were excluded from the analysis because we cannot assign the dates of arrival to specific decades. j} = 61.37; df = 7; p<0.01 using an average number of invaders of 121.8 per decade. Data from 2011-2015 excluded due to lag in publication of invasive species information. DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 75 Table 8. Number of exotic terrestrial macro-invertebrates discovered breeding in California by time and likely origin. Region 3 <1901 1901- 1929 1930- 1939 1940- 1949 1950- 1959 1960- 1969 1970- 1979 1980- 1989 1990- 1999 2000- 2015 Australia 5 2 1 2 3 0 2 4 4 15 Pacific Region 0 0 1 0 1 4 0 0 0 1 Africa 4 9 6 2 0 2 3 4 3 7 Asia 4 21 8 10 5 6 2 4 4 16 Tropical America 1 10 7 2 1 7 5 7 7 6 Hawaii 0 2 1 0 2 7 4 7 8 6 Mexico 2 1 1 0 1 21 0 1 5 1 Europe 26 53 17 117 9 10 2 10 12 14 North America 32 73 24 23 42 40 21 30 31 36 3 See Appendix for list of organisms. tributed nine Psyllidae, two Aphalaridae and one Triozidae to the California fauna between 1980 and 2010 (Appendix). These insects, especially the Hymenoptera and Psyllidae must have escaped quarantine detections and were moved to California on living plant material. Comparison with the earlier study by Dowell and Gill (1989). — Many of the trends reported by Dowell and Gill (1989) were repeated here. The Hemiptera, Coleoptera, Lepidoptera, Acari, Diptera, and Hymenoptera had the greatest number of invaders but the Acari were second after the Hemiptera. North America was still the origin of the greatest number of invaders entering California but the order after that was the Pacific Region (including Hawaii), Europe, Asia, Trop¬ ical America (including Mexico), Africa and Australia. Dowell and Gill (1989) reported that Hawaii contributed more invaders than any the other region except North America and Europe. In this larger analysis, Hawaii is far less important as a point of origin although this US state still con¬ tributes more exotic species to California than the rest of the Pacific Region (Table 4). Factors that allect the list. — We have adopted a conservative approach to listing organ¬ isms with the requirement of evidence of a breeding population within California. This criterion differs from other groups, such as the Lepidopterists Society, that develop regional lists of organ¬ isms. In many of these other lists the occurrence of a single individual constitutes a new regional record that in some instances may over-estimate the number of species successfully established in a given area (Rickard and Grishin 2010, Bordelon and Knudson 2011, Rickard and Pasko 2011). This conservative approach has been influenced by regulatory concerns because listing the pres¬ ence of an exotic organism as opposed to a breeding population may trigger official actions (IPPC 2007) by the CDFA and/or United States Department of Agriculture including the imposition of exterior/interior quarantines or eradication programs (Dowell 1988). As noted elsewhere (Kirkendall and Faccoli 2010), the taxonomy of many arthropod organ¬ isms is fluid with changes occurring from the order level (e.g., loss of the order Homoptera and the resurrection of Hemiptera that includes Auchenrrhyncha and Stemorrchyncha), to the family and species level (e.g., Crambidae versus Pyralidae or Solenopsis invicta versus Solenopsis wagneri). We have attempted to use the most current taxonomy for species listed here realizing there may be professional disagreement about the use of particular species names and classifications. There are often differences of opinion as to whether an organism is native or exotic to Cali¬ fornia. This is especially true for those organisms that invaded before or around 1900 (Powell et al. 76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Table 9. Taxons of exotic terrestrial macro-invertebrates found breeding in California by time period in which they were detected. Taxon 3 <1900 1901- 1929 1930- 1939 1940- 1949 1950- 1959 1960- 1969 1970- 1979 1980- 1989 1990- 1999 2000- 2015 Acari 5 13 26 14 8 8 7 10 4 Amphipoda 1 Araneae 1 4 5 Chilopoda Coleoptera 8 59 13 15 10 15 6 5 13 32 Collembola 2 1 1 Dermaptera 2 5 1 Dictyoptera 1 4 1 2 2 1 2 1 3 Diplopoda Diptera 6 32 1 5 3 11 3 6 10 23 Embiidina 1 1 Hemiptera- Heteroptera 3 7 1 1 4 3 2 7 8 Hemiptera: Auchenor- rhyncha + Stemorrhyncha 46 112 42 20 39 26 19 30 30 25 Hymenoptera 5 11 2 4 5 7 3 8 18 7 Isopoda 2 Isoptera 1 Lepidoptera 16 21 7 5 12 21 78 4 5 9 Mollusca 7 2 1 4 3 5 1 1 Orthoptera 1 6 1 1 2 Phasmida 1 Phthiraptera 1 11 1 1 2 1 3 5 2 Psocoptera 3 1 1 Siphonaptera Symphyla Thysanoptera 1 Thysanura a See Appendix for list of organisms. 2000). We have listed the authorities that note that each organism is not native to California rec¬ ognizing that new information (molecular or phylogenetic analyzes) may change this. For exam¬ ple, the gopher beetle, Ceratophyus gopherinus, has long been considered an invasive species in California as all other known species in the genus are found in Europe (Cartwright 1966). Howev¬ er, C. gopherinus does not occur in Europe thus casting doubt on the exotic origins of this beetle. Current thinking is that C. gopherinus is a native species of limited distribution in California (Arnett et al. 2002). California does not maintain a statewide survey for invasive invertebrates except for a limited DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 77 number of crop pests (Gilbert et al. 2013). The discovery of exotic macro-invertebrates not target¬ ed by CDFA depends on individuals noting and reporting damage to crops or ornamental plants, capturing unidentified organisms in local faunal surveys (e.g., light brown apple moth, Epiphyas postvittana) (Brown et al. 2010), or periodic CDFA surveys that target specific groups of organ¬ isms (e.g., wood boring beetles, which resulted in the detection of Scolytus schevyrewi). One under- appreciated avenue to the detection of new organisms that have invaded California are citizens who find species they do not recognize and then send specimens to experts for identification. One citi¬ zen scientist has discovered five exotic ant species and one exotic spider species in southern Cali¬ fornia (Martinez 1992, 1993, 1996, 1997, 2006; Martinez et al. 2011, 2014). Pathways into California.— There are numerous pathways by which the organisms listed in the Appendix were introduced into California. Human alterations to the landscape (i.e., irriga¬ tion of desert areas and propagation of exotic plants) probably facilitated the self-introduction of the giant swallowtail butterfly, Papilio cresphontes. Others, especially the wood boring beetles (e.g., Scolytus schevyrewi and the Asian longhorn beetle) probably arrived in wooden pallets of commercial goods or in firewood transported across state lines. For example, the gold spotted oak borer (Agrilus auroguttatus ) was likely introduced in cut oak firewood from the Dragoon Moun¬ tains in southern Arizona (Lopez et al. 2014). Large numbers of non-native species are thought to have arrived on living plants either shipped into the state commercially, arriving in vehicles (cars, trucks, and trains) moving here from other states, or via smuggling from overseas (e.g., red palm weevil [Hoddle 2015]). Several pestiferous fruit flies (Tephritidae) (e.g., Mediterranean fruit fly, Ceratitis capitata, and the oriental fruit fly, Bactrocera dorsalis ) probably arrived as larvae in smuggled fruit. During the 1700’s to early 1900’s soil, as well as and construction debris, was used as ballast in cargo ships and the soil was dumped at the destination port (Lindroth 1957). This route was likely the avenue through which many of the Collembola, Isopoda, Staphylinidae, Dermaptera and Embiidina, among the soil inhabiting invertebrates, were introduced into California. Some introduction pathways are unusual. Fire ants, S. wagneri, arrived in California in the soil of potted plants but also with beehives stacked on palletts originating in fire-ant infested areas of the US brought into California to pollinate crops, especially almonds (CDFA unpublished data). Slate, a fine grain metamorphic rock used for construction, represents another pathway for exotic organisms to enter California. Slate is stored as thin slabs on pallets with gaps between the slabs. These gaps provide dry, cool sites for many organisms, especially slugs, and lygaeid bugs, to over¬ winter. The pallets, with overwintering organisms included, are shipped to California in unheated boat holds making it more likely that the hitch-hiking organisms will survive and disembark in Cal¬ ifornia alive (California Department of Food and Agriculture, unpublished data) Rumors of Bioterrorism.— Speculation exists that pestiferous organisms are being delib¬ erately introduced into California (Paine et al. 2010). For example, Paine et al. (2010) suggested that eucalyptus-specific insects have been deliberately introduced to kill Eucalyptus spp., non¬ native trees that are viewed by some as noxious weeds (Paine et al. 2010). It has been similarly suggested that Mediterranean fruit flies are punitively introduced into California to challenge the State’s eradication programs. There are no data to support these suggestions. However, the large number of eucalyptus feeding insects discovered in California since 1990 is considered suspicious because there were essentially no introductions in the preceding 100+ years during which live eucalyptus were imported into the state (Paine et al. 2010). Impacts of the exotic species. — Dowell and Gill (1989) noted that exotic pests accounted for 62% of all reported crop losses in 1978; the last year for which there are data. The economic losses to invasive pests in California were estimated at $3 billion (US) dollars in 1995 (Metcalf 1995). 78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Although 314 (19.2%) Table 10. Percentage of exotic terrestrial macro-invertebrates pests of the 1,686 exotic listed in University of California on-line IPM Guides. invaders are condsidered as pests in California (see Appendix), exotic invaders account for a dispropor¬ tionately large number of pest species mentioned in University of California on line IPM guides and else¬ where (Table 10). Over 42% of the pest inverte¬ brates mentioned in the University of California Integrated Pest Manage¬ ment Guidelines for agricultural, ornamental, and native plants are exotic in origin (Table 10; see also Appendix). This overview analysis treats all listed pests equally, which is not the case. For example, cottony cushion scale, now a minor pest of citrus, is given the same weight as California red scale, Aonidiella aurantii, which is a major citrus pest even though both have been subjected to classical biological control programs. These observations suggest that exotic pests caused a more disproportionate level of crop losses than would be anticipated by their percent composition in the total pest complex. Exotic invaders represent some of the most common and troublesome urban pests (cockroaches and Agrentine ants) supporting an urban pest control industry with annual rev¬ enues estimated at $1 to 2 billion (US) per year (Rust, pers. comm.) that exists to manage these pests. Non-native species can have a devastating impact on our natural environment. The gold spot¬ ted oak borer has killed over 20,000 oaks in southern California since 2008 (Coleman and Seybold 2010). Its continued spread in infested firewood, an unregulated conduit for invasive wood pests, that poses a significant threat to ecosystems dominated by oak trees in California. The Argentine ant has displaced the native ants in many coastal areas of California, and that in turn has adverse¬ ly affected native horned lizards that feed on native ants, which has contributed to the loss of the horned lizards in some cases (Suarez and Case 2002). The cause of range reduction of the native veined white, Pier is marginalise, is thought to have occurred from competition for cruciferous host plants with the pestiferous exotic cabbage butterfly, Pieris rapae (Pyle 1981). Conclusion Terrestrial macro-invertebrates have been invading California since the advent of European traders and settlers. The invasion process is complex with significant changes in the composition and origins of the invaders over time. The one common thread is human activity, especially trans¬ portation networks that facilitate rapid and long distance trade and tourism and human-mediated habitat alteration (i.e., gardenification and urbanization). The impacts of increasing numbers of ter¬ restrial macro-invertebrate species on the agriculture, wilderness, and urban areas of California will almost certainly continue to increase. Acknowledgements Over the years a number of colleagues have examined the list of invaders and we thank them: Situation Percentage of listed pests that are not native to California 3 Agricultural crops 55.2 Urban vegetables 70.3 Urban fruit trees and vines 67.7 Urban flowers 64.5 Urban homes, stmctures, people and pets 53.8 Urban ornamental trees and shmbs 51.4 Natural environment 42.9 a Available at www.ipm.ucdavis.edu. Accessed 15 December 2014. DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 79 F. Andrews, R Amaud, J. Beardsley, L. Bezark, D. Burdick, T. Eichlin, E. Fisher, A. Gilbert, R. Goeden, D. Headrick, T. Henry, K. Kim, D. Kissinger, W. Laudenslayer, F. Legner, R. Penrose, J. Powell, M. Rust, T. Seeno, W. Sheppard, R. Somerby, J. Sorensen, P. Ward, D. Weissman, and A. Wheeler. 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Cavey, and R. Komsa. 2005. Arrival rate of nonindigenous insect species into the United States through foreign trade. Biological Invasions 7:323-332. Yamanaka, T., N. Moriomoto, G.M. Nishida, K. Kiritani, S.Moriya, and A.M. Liebhold. 2015. Comari- son of insect invasions in North America, Japan and their islands. Biological Invasions 10:3049-3061. 84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix with Addendum Footnotes to Appendix table a an organism that was or is the object of Official Action (quarantine and eradication) by the California Department of Food and Agriculture and is not established in California. b We have identified those animals in Appendix 1 that are considered pests in or by California. For plant, home and garden and the environment we identify those animals listed as pests in the University of California IPM online (UC IPM 2013), sheep and goat pests are from Pirelli (2013), Dairy cattle pests are those identified by Geden et al. (2010), Horse insect pests are from Meyer et al.(1991), swine pests are from Loft in (2013) and chicken pests are from Stafford 111 (2011). These sources tend to deal with animals that are established in California or are frequently detected there. We have also identified some pests not established in California but which have been detected here and which are generally considered pests in those areas where they occur including the gypsy moth ( Lymantria dispar) (Liebhold 2003), Japanese beetle ( Popil- lia japonica) (Anon. 1997), Mexican bean beetle ( Epilachna varivestis ) (Sanchez-Arroyo 2015), oriental fruit fly ( Bactro- cera dorsalis ), Mexican fruit fly ( Anastrepha ludens), Caribbean fruit fly ( Anastrepha suspensa), peach fruit fly ( Bactro- cera zonata ), Mediterranean fruit fly ( Ceratitis capitata), melon fly ( Bactrocera cucurbitae), white striped fruit fly ( Bactro- cera albistrigata), and guava fruit fly ( Bactrocera correcta) (White & Elson-Harris 1992), boll weevil ( Anthonomus gran- dis ) (Lange et al. 2008), varroa mite, ( Varroa jacobsoni) (Bessen 2010), red palm weevil ( Rhynchophorus vulneratus ) (Rug- man-Jones et al. 2013), Diaprepes root weevil (Diaprepes abbreviatus ) (Grafton-Cardwell et al. 2004), Asian longhorn Bee¬ tle (Anoplophora glabripennis) (United States Department of Agriculture, Animal and Plant Health Inspection Service 2015). DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 85 Appendix. Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest b Acari-Acaridae Acarus siro (Linnaeus 1758) Essig 1958 Aleuroglvphus ovatus (Troupeau 1879) 1984 Dowell & Gill 1989 Rhizoglyphus robini Claparede 1869 <1915 CDFA 1914 3:539 yes Tyrophagus neiswanderi Johnston & Bruce 1965 1982 Europe Dowell & Gill 1989 yes Tyrophagus putrescentiae (Schrank 1781) Hughes 1976 yes Acari-Astigmata Sancassania berlesi (Michael 1903) Hughes 1976 Acari-Cheyletidae Chelyletiella yasguri (Smiley 1965) 1973 North America Dowell & Gill 1989 Acari-Dermanyssidae Ornithonyssus bacoti (Hirst 1913) <1930 CDFA 1941 30:352 Acari-Eriophyidae Abacarus hystrix (Nalepa 1896) 1940 CDFA 1940 29:242 Acalitus calycophthirus (Nalepa 1891) 1984 North America Dowell & Gill 1989 Acaphylla steinwedeni Keifer 1943 1942 Asia CDFA 1945 34:174 Acarapis near dorsalis Morganthaler 1934 1959 Dowell & Gill 1989 Acarapis woodi (Rennie 1921) 1986 North America Dowell & Gill 1989 Acaricalus hederae (Keifer 1939) 1939 CDFA 1939 28:490 Aceria ajugae (Nalepa 1892) 1959 CDFA 1959 48:127 Aceria aloinis Keifer 1941 1940 CDFA 1940 29:242, Keifer 1952 Aceria cynodontis Wilson 1959 Keifer et al. 1982 Aceria diospyri (Keifer 1944) 1943 Keifer 1952 yes Aceria ficus (Cotte 1920) 1922 Europe CDFA 1939 28:266 yes Aceria fraxinivorus (Nalepa 1909) 1944 Keifer et al. 1982, Keifer 1952 Aceria genistae (Nalepa 1892) 1994 Europe Chan & Turner 1998 Aceria lycopersici (Wolffenstein 1879) 1940 Tropical America Jeppson et al. 1975 Aceria mori (Keifer 1939) 1939 Keifer 1952 Aceria neocynodonis Keifer 1960 1960 North America CDFA 1960 49:200 Aceria paradianthi Keifer 1952 1952 CDFA 1952 41:65 Aceria parthenii Keifer 1952 1952 CDFA 1952 41:147 Aceria peucedani (Canestrini 1891) Jeppson et al. 1975 Aceria sheldoni (Ewing 1937) 1937 CDFA 1937 26:433 Aceria tulipae (Keifer 1938) 1938 Europe CDFA 1938 27:185 Aculops fuchsiae Keifer 1972 1982 Tropical America Dowell & Gill 1989 yes Aculops lycopersici (Massee 1937) 1940 Australia Keifer et al. 1982 yes Aculus cornutus (Banks 1905) 1914 North America Jeppson et al. 1975 yes Aculus ligustri (Keifer 1943) 1938 CDFA 1938 27:190 86 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest b Aculus schlectendali (Nalepa 1890) 1938 CDFA 1939 28:234 Aculus teucrii (Nalepa 1892) 1986 North America Dowell & Gill 1989 Anthocoptes punctidorsa Keifer 1943 1943 Keifer 1952 Calacarus adornatus (Keifer 1940) 1939 Keifer 1952 Calacarus carinatus (Green 1890) Keifer et al. 1982 Calepitrimerus muesebecki Keifer 1940 1954 Mexico CDFA 1940 29:113 Calepitrimerus vitis (Nalepa 1905) Europe Keifer 1952 Cecidophyes collegiatus Keifer 1961 Jeppson et al. 1975 Cecidophyes malpighianus (Canestrini & Massalongo 1893) 1939 Keifer 1952 Cecidophyopsis grossulariae (Collinge 1907) 1991 CPPDR 1991 10:15 Colomerus vitis (Pagenstecher 1857) <1915 Amrine & Stasney 1994 Coptophylla lamimani (Keifer 1939) 1939 Keifer 1952 Cosetacus camelliae (Keifer 1945) 1940 Asia CDFA 1945 34:137 Diptacus gigantorhynchus (Nalepa 1892) 1938 Europe Keifer 1952 yrs Epitrimerus pyri (Nalepa 1870) 1913 North America CDFA 1915 4:12 yes Eriophyes ajugae (Nalepa 1892) 1958 Dowell & Gill 1989 Eriophyes canestrinii (Nalepa 1891) 1950 Keifer 1952 Eriophyes car ids Keifer 1944 1943 Europe Keifer 1952 Eriophyes celtis Kendall 1929 1961 North America Dowell & Gill 1989 Eriophyes convolvens (Nalepa 1892) Europe Keifer 1952 Eriophyes cynodoniensis Sayed 1946 Jeppson et al. 1975 yes Eriophyes eriobotiyae (Keifer 1938) 1938 CDFA 1938 27:188 Eriophyes granati (Canestrini & Massalongo 1893) Keifer et al. 1982 yes Eriophyes pyri (Pagenstecher 1857) 1894 North America CDFA 1915 4:6 yes Eriophyes spermaphaga (Keifer 1979) 1979 Dowell & Gill 1989 Eriophyes tiliae (Nalepa 1890) 1932 Keifer 1952 Eriophyes vaga (Keifer 1962) 1979 Tropical America Dowell & Gill 1989 Eriophyes wisteriae Keifer 1939 1939 CDFA 1939 28:329 Gammaphytoptus camphorae Keifer 1939 1938 Asia CDFA 1939 28:148 Mackiella phoenicis Keifer 1939 1938 Africa Keifer 1952 Oxypleurites maxwelli (Keifer 1939) 1939 Europe CDFA 1914 31:175 Phyllocoptes bougainvilleae Keifer 1959 1996 Tropical America CPPDR 1997 16:10 Phyllocoptes calulmi Keifer 1940 1939 CDFA 1940 29:113 Phyllocoptes destructor Keifer 1940 1940 CDFA 1940 29:161 Phyllocoptes gracilis (Nalepa 1890) Jeppson et al. 1975 Phyllocoptruta oleivora (Ashmead 1879) 1889 North America CDFA 1934 23:201 yes Phytoptus avellanae Nalepa 1889 1939 North America Keifer 1952 Phytoptus hedericola Keifer 1943 1942 Keifer 1952 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 87 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Rhynacaphytoptus fwifoliae Keifer 1939 1939 Africa CDFA 1939 28:273 Shevtchenkella depressus (Nalepa 1894) Keifer 1952 Tegolophus myersi (Keifer 1939) 1938 Keifer 1952 Tegolopkus zizyphagus (Keifer 1939) 1939 Keifer 1952 Tegonotus carinatus (Nalepa 1891) 1964 Europe Dowell & Gill 1989 Tetra concava (Keifer 1944) 1939 Europe Keifer 1952 Trisetacus pseudotsugae Keifer 1965 1969 North America Dowell & Gill 1989 Vasates immigrans (Keifer 1940) 1939 Keifer 1952 Vasates magnolivora (Keifer 1952) 1939 North America Keifer 1952 Acari-Glycyphagidae Glycyphagus destructor (Schrank 1781) Hughes 1976 Glycyphagus domesticus (De Geer 1778) Hughes 1976 Acari-Ixodidae Argas persicus (Oken 1881) <1925 Furman & Loomis 1984, Essig 1926a Boophdus annulatus (Say 1905) <1925 Furman & Loomis 1984, Essig 1926a Rhipicephalus sanguineus (Latreille 1806) Furman & Loomis 1984 Acari-Laelapidae Melittiphis alvearius (Berlese 1895) 1988 Europe Dowell & Gill 1989 Acari-Penthaleidae Penthaleus major (Duges 1834) Europe CDFA 1940 29:507 yes Acari-Phytoptidae Phantacrus lobatus Keifer 1965 1978 North America Dowell & Gill 1989 Acari-Psoroptidae Psxoroptes equi (Raspail 1834) <1925 Essig 1926a, 1958 Psoroptes ovis (Hering 1838) <1925 Essig 1926a, 1958 Acari-Pyemotidae Siteroptes graminum (Reuter 1907) 1965 Hawaii Dowell & Gill 1989 Acari-Rhyncaphytoptidae Rhyncaphytoptus ulmivagrans Keifer 1939 1939 Europe CDFA 1939 28:420 Trimeroptes aleyrodiformis (Keifer 1951) North America Jeppson et al. 1975 Acari-Tarsonemidae Steneotarsonemus ananas (Tiyon 1898) 1980 Hawaii Dowell & Gill 1989 Steneotarsonemus pallidus (Banks 1899) <1925 Jeppson et al. 1975, Essig 1926a yes Tarsonemus setifer Ewing 1939 Europe Jeppson et al. 1975 Acari-Tenuipalpidae Brevipalpus lewisi McGregor 1949 1942 Pritchard & Baker 1958 yes Brevipalpus obovatus Donnadieu 1875 Jeppson et al. 1975, Pritchard & Baker 1958 Brevipalpus phoenicis (Geijskes 1936) Jeppson et al. 1975, Pritchard & Baker 1958 88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 1 Acari-Tetranychidae Biyobia praetiosa Koch 1835 1879 Jeppson et al. 1975 Eotetranychus sexmaculatus (Riley 1890) <1915 CDFA 1915 4:16 yes Eotetranychus uncatus Gaiman 1952 North America Jeppson et al. 1975 Eurytetranychus buxi (Gamian 1935) Jeppson et al. 1975 Eutetranychus banksi (McGregor 1914) <1915 CDFA 1915 4:13 yes Mononychellus caribbeanae (McGregor 1950) 1978 CPPDR 1996 15:79 Mononychellus erythrinae Tuttle, Baker & Abbatiello 1976 1978 Dowell & Gill 1989 Oligonychus aceris (Shinier 1869) 1959 North America Dowell & Gill 1989 Oligonychus ilicis (McGregor 1917) 1935 North America CDFA 1939 28:240 Oligonychus perseae Tuttle, Baker & Abbatiello 1976 1992 Mexico CPPDR 1992 11:46 Oligonychus pratensis (Banks 1912) Baker & Tuttle 1994 yes Oligonychus punicae (Hirst 1926) Mexico Baker & Tuttle 1994 Panonychus citri (McGregor 1916) 1885 North America Baker & Tuttle 1994 yes Panonychus ulmi (Koch 1836) <1925 North America Essig 1926a yes Paraplonobia myops (Pritchard & Baker 1955) Jeppson et al. 1975 Petrobia cf. apicalis (Banks 1917) 1978 North America Dowell & Gill 1989 Schizonobia sp Womisley 1940 1965 Dowell & Gill 1989 Schizotetranychus celarius (Banks 1917) Asia Jeppson et al. 1975 Tetranychus evansi Baker & Pritchard 1960 1965 Pacific Region Jeppson et al. 1975 Tetranychus merganser Boudreaux 1954 1963 North America Dowell & Gill 1989 Tetranychus tumidus Banks 1900 North America Jeppson et al. 1975 Tetranychus urticae (Koch 1836) 1876 Europe Baker & Tuttle 1994 yes Tetranycopsis horridus (Canestrini & Fanzago 1876) Europe Jeppson et al. 1975 Acari-Uropodidae Fuscuropoda marginata (Koch 1839) 1964 Dowell & Gill 1989 Acari-Tydeidae Lorryna formosa Cooreman 1958 1984 Europe Dowell & Gill 1989 Acari-Varroidae Varroa jacobsoni Oudemans 1904 1989 North America CPPDR 1989 8:5 yes Amphipoda-Talitridae Talitrus sylvaticus (Haswell 1880) 1918 Australia Mallis 1942 Araneae-Agelenidae Agriope trifasciata (Forskal 1775) Sierwald et al. 2005, Lew 2010 Gea heptagon (Hentz 1850) Sierwald et al. 2005, Lew 2010 Metazygia zilloides Banks 1989 2009 North America Martinez et al. 2014 Tegeneria domestica (Clerck 1757) Europe Herbert et all987 Tegeneria pagana (Koch 1840) Herbert et al. 1987 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 89 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Araneae-Amphinectidae Metaltella simoni (Keyserling 1878) 1994 Tropical America Vetter & Visscher 1994 Araneae-Araneidae Araneus diadematus Clerck 1758 BugGuide 2011, Lew 2010 Larinoides sclopetarius (Clerek 1757) 2004 North America Martinez 2006 Zygiella x-notata Clerck 1757 Lindroth 1957, Lew 2010 Neoscona crucifer a Lucas 1839 1991 North America Levi 1992 Araneae-Desidae Badumna longinqua (Koch 1867) Bryant et al. 2010, Lew 2010 Araneae-Dictynidae Dictyna major Menge 1869 Lindroth 1957, Lew 2010 Araneae -Dysderidae Dysdera crocata (Koch 1838) Europe Prentice et al. 2001, Bolger et al. 2008 Araneae -Gnaphosidae Haplodrassus signifer Koch 1839 Lindroth 1957, Berrian 2005 Scotophaeus blackwalli (Thorell 1871) Herbert 1987, Lew 2010 Trachyzelotes barbatus (Koch 1866) Crespo & Mendes 2010, Prentice et al. 1998 Trachyzelotes jaxartensis (Kroneberg 1875) Crespo & Mendes 2010, Lew 2010 Trachyzelotes lyonneti (Audouin 1826) Bolger et al. 2008, Crespo & Mendes 2010 Urozelotes rusticus (Koch 1872) Beatty 2002, Prentice et al. 1998 Zelotes nilicola (Cambridge 1874) Prentice et al. 1998, Bolger et al. 2008 Araneae-Linyphidae Pityohyphantes phiygianus (Koch 1836) Jennings & Graham 2007, Lew 2010 Araneae-Mimetidae Ero furcata (Villers 1789) Lew 2010, Bolger et al. 2008 Araneae-Miturgidae Cheircanthium mildei Koch 1864 Lew 2010, BugGuide 2011, Hogg et al. 2010 Araneae -Oecobiidae Oecobius annulipes Lucas 1849 Prentice et al. 1998 Oecobius navus Blackwall 1859 Lew 2010, Bolger et al. 2008 Oecobius putus (Cambridge 1876) Lew 2010, Bolger et al. 2008 Araneae-Philodromidae Thanatus vulgaris Simon 1870 Donale & Redner 1978, Lew 2010 Araneae -Pholcidae Artema atlanta Walckenaer 1837 BugGuide 2011, Lew 2010 Holocnemus pluchei (Scopoli 1763) Europe Prentice et al. 2001, Bolger et al. 2008 Pholcus phalangioides (Fuesslin 1775) Europe Lew 2010, Bolger et al. 2008 Spermophora senoculata (Duges 1836) Beatty 2002, Lew 2010 90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Araneae-Salticidae Menemerus bivittatus (Dufour 1831) Prentice et al. 2001 Menemerus semilimbatus (Hahn 1827) 1990’s Europe Manolis & Carmichael 2010 Phlegm fas data (Hahn 1826) Sicrwald et al. 2005, Lew 2010 Salticus scenicus (Clerck 1757) Jennings & Graham 2007, Lew 2010 Araneae-Scytodidae Scytodes thoracica (Latreille 1802) Beatty 2002, Lew 2010 Araneae-Sicariidae Loxosceles laeta (Nicolet, 1849) 1936 Tropical America Robinson 2005 Loxosceles rufescens (Dufour 1820) Europe Lew 2010, Yigit et al. 2008 Araneae -Sparassidae Heteropoda venatoria (Linnaeus 1767) Sherriffs 1936, Lew 2010 Olios abnormis (Blackwall 1866) Platnick 2010, Lew 2010 Araneae-Theridiidae Achaearanea acoreensis (Berland 1932) Lew 2010, Draney 2001 Achaearanea ohlerti (Thorell 1870) Lindroth 1957, Lew 2010 Achaearanea tepediorum (Koch 1841) Tropical America Lew 2010, Campbell 2010 Dipoena prona Menge 1868 Lindroth 1957, Berrian 2005 Enoplognatha ovata (Clereck 1757) Oxford & Reillo 1994 Latrodectus geometricus (Koch 1841) 2003 North America Vincent et al. 2008 yes Steatoda albomaculata (De Geer 1778) Sierwald 2005, Lew 2010 Steatoda grossa (Koch 1838) Europe Draney 2001, Lew 2010 yes Steatoda triangulossa (Walckenaer 1802) Draney 2001, Prentice et al. 2001 Theridion hemerobium Simon 1914 Europe Lindroth 1957, Lew 2010 Theridion melanumm (Hahn 1820) Lindroth 1957, Lew 2010 Theridion murarium (Emerton 1882) Herbert et al.1987, Lindroth 1957, Lew 2010 Araneae-Zoropsidae Zoropsis spinimana (Dufour 1820) 1996 Africa Griswold & Ubick 2001 yes Chilopoda-Scolopendromorpha-Cryptopidae Cryptops hortensis (Donovan 1810) Europe Shelley 2002 yes Chilopoda-Scutigeromorpha-Scutigeridae Scutigera coleoptrata Linnaeus 1758 <1915 Europe Lewis 1981, Essig 1915 yes Coleoptera-Aderidae Aderus populneus (Creutzer in Panzer 1796) Majka 2011 Coleoptera-Anobiidae Anobium punctatum (De Geer 1774) Europe Philips 2002, Evans & Hogue 2006 Ernobius mollis (Linnaeus 1758) 1990 North America Seybold & Tupy 1993 \Gibbium psylloides (Czempinski 1778) Philips 2002 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 91 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Lasioderma haemorrhoidale (Illiger 1807) 1981 White 1990 Lasioderma serricorne (Fabricius 1792) <1925 Campbell et al. 1989, Essig 1926a yes Mezium affine Boieldieu 1856 Philips 2002 Mezium americanum Laporte 1840 Philips 2002 Ptinus fur (Linneaus 1758) Bousquet 1990, California Beetle Project 2006 Niptus holoeucus (Faldermann 1836) Philips 2002 Stegobium paniceum (Linnaeus 1758) <1925 White 1962, Essig 1926a yes Tricorynus herbarius (Gorham 1883) White 1965 Trigonogenius globulus Sober 1849 Arnett et al. 2002 Coleoptera-Anthicidae Anthicus floralis (Linnaeus 1758) Campbell et al. 1989 Stricticomus tobias (Marseul 1879) Peck & Thomas 1998 Coleoptera-Anthribidae Araecerus fasciculatus (De Geer 1775) 1994 CPPDR 1996 15:74 Coleoptera-Bostrichidae Dinoderus minutus (Fabricius 1775) Campbell et al. 1989 Heterobostrychus brunneus (Murray 1867) Ivie 2002a Lyctus brunneus (Stephens 1830) <1925 Europe Campbell et al. 1989 yes Lyctus linearis (Goeze 1777) <1925 Europe Campbell et al. 1989 Micrapate sp. Casey 1898 2003 Penrose 2004 Rhyzopertha dominica (Fabcicius 1792) <1930 Ebling 1975 yes Coleoptera-Brentidae Cylas formicarius (Fabricius 1798) 1993 North America CPPDR 1993 12:3 yes Trichapion simile Kirby 1811 Lindroth 1957 Coleoptera-Buprestidae Agrilus anxius Gory, 1841 1992 North America Dreistadt et al. 2004 yes Agrilus auroguttatus Schaeffer 1905 2004 North America Westcott 2005 yes Agrilus cyanescans (Ratzeburg 1837) 1949 Tropical America Chamberlin 1949 Chrysobothris costifrons costifrons Waterhouse 1887 2005 North America Basham et al. 2015 Xenorhipis osborni Knull 1936 1947 North America Thatcher 1948 Coleoptera-Carabidae Amara anthobia Villa & Villa 1833 North America Lindroth 1957 Agonum mulleri (Herbst 1784) Ball & Bousquet 2001 Carabus nemoralis Muller 1764 1919 North America Brown 1940 Laemostenus complanatus (Dejean 1828) North America Lindroth 1957 Perigona nigriceps (Dejean 1831) North America Bousquet 2012 Trechus obtusus Erichson 1837 North America Lindroth 1957, Kavanaugh & Eiwin 1985 92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest*’ Coleoptera-Cerambycidae a AnopIophorus glabripennis (Motachulsky 1853 2005 Asia CPPDR 2006 22:4 yes Arhopalus syriacus (Reitter 1895) 2002 Europe Penrose 2003 Callidellum rufipenne (Motschulsky 1860) Aukema et al. 2010, California Beetle Project 2006 Icosium tomentosum Lucas 1854 2006 Europe CPPDR 2007 23:5, Bohne 2006 Nathrius brevipennis (Mulsant 1839) 1932 Europe Linsley 1933 Neoclytus acuminatus (Fabricius 1775) 2002 North America Penrose 2004 Phoracantha recurva Newman 1840 1995 Australia CPPDR 1997 16:10 yes Phoracantha semipunctata Fabricius 1775 1984 Australia Dowell & Gill 1989 yes Phymatodes testaceus (Linnaeus 1758) North America Lindroth 1957 Saperda populneus (Linnaeus 1758) Europe Fumiss & Carolin 1977 Coleoptera-Chrysomelidae Acanthoscelides obtectus (Say 1831) <1915 North America CDFA 1920 9:344 yes Altica carduorum (Guerin-Memeville 1858) Riley et al. 2003 Amblycerus robiniae (Fabricius 1781) 1970 North America Dowell & Gill 1989 Bruchus brachialis Fahraeus 1839 1949 North America Davidson & Lyon 1987, Evans & Flogue 2006 Bruchus pisorum (Linnaeus 1758) <1915 CDFA 1915 4:287 Bruchus rufimanus Boheman 1833 <1915 Europe CDFA 1915 4:289, Essig 1926a Callosobruchus maculatus (Fabricius 1775) <1915 Campbell et al. 1989 Cassida nebulosa Linnaeus 1758 <1894 Europe Barber 1916 Chaetocnema sp. Stephens 1831 1948 CDFA 1948 37:211 yes Chelymorpha cassidea (Fabricius 1775) 1909 North America Essig 1958 Chiysolina bankii (Fabricius 1775 2006 Europe CPPDR 2007 23:5-6 Chrysophthartam-fuscum Boheman 1859 2003 Australia CPPDR 2006 22:25 yes Crioceris asparagi (Linnaeus 1758) 1904 Europe CDFA 1915 4:267 Crioceris duodecimpunctata (Linnaeus 1758) 1975 Dowell & Gill 1989 Diabrotica balteata LeConte 1865 1924 Tropical America Metcalf 1995 Diorhabda elongate (Brulle 1832) Riley et al. 2003 Epitrix fasciata Bratchley 1918 <1915 CDFA 1915 4:262 yes Epitrix tuberis Gentner 1944 1968 North America Dowell & Gill 1989 yes Galerucella nymphaeae (Linnaeus 1758) 1924 Lindroth 1957 Longitarsus jacobaeae Waterhouse 1858 LeSage 1988 Mimosetes nubigens (Motschulsky 1874) 1991 North America Johnson & Seeno 1993 Phyllotreta cruciferae (Goeze 1777) Riley et al. 2003 yes Phyllotreta striolata (Fabricius 1803) Campbell et al. 1989, Riley et al. 2003 yes Specularius impressithorax (Pic 1913) 2003 Gulmahamad 2006 Trachymela sloanei (Blackburn 1897) 1997 Australia CPPDR 1998 17:4-6 yes Xanthogaleruca luteola (Muller 1766) 1924 Europe Metcalf 1995, Riley et al. 2003 yes DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 93 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Zabrotes subfasciatus Boheman 1833 1965 Tropical America Dowell & Gill 1989 Zeugophora scutellaris Suffrian 1840 Riley et al. 2003, California Beetle Project 2009 Coleoptera-Cleridae Necrobia ruficoUis (Fabricius 1785) <1925 Campbell et al. 1989, Essig 1926a Necrobia rufipes (De Geer 1775) <1925 Campbell et al. 1989, Essig 1926a Necrobia violacea (Linnaeus 1758) <1925 Campbell et al. 1989, Essig 1926a Paratillus earns (Newman 1840) Europe Grace & Wood 1985 Tarsostenus univittatus (Rossi 1792) Arnett 1983 Thaneroclerus buquet (Lefebvre 1835) California Beetle Project 2006 Coleoptera-Coccinellidae b Epilachna varivestis Mulsant 1850 1946 North America CDFA 1946 35:198 yes Coleoptera-Colydiidae Microprius rufulus (Motschulsky 1863) Europe Ivie 2002b Coleoptera-Cryptophagidae Cryptophagus laticollis Lucas 1849 Majka & Langor 2010 Ciyptophagus pilosus Gyllenhal 1827 Campbell et al. 1989 Coleoptera-Curculionidae Agasphaerops nigra Horn 1876 1876 O’Brien & Wibmer 1982 Anialus haemorrhous (Herbst 1795) Lindroth 1957 Anthonomus eugenii Cano 1894 1923 Tropical America CDFA 1923 12:360 yes a Anthonomus grandis (Boheman 1843) 1984 North America Dowell & Gill 1989 yes Apion longirostre Olivier 1807 1964 North America Dowell & Gill 1989 Araecerus coffeae (Fabricius 1801) 2005 North America CDFA 2005 Araptus schwarzi (Blackman 1942) 2010 Mexico Takahashi 2011 Atrichonotus taeniatulus (Berg 1881) 1997 North America CPPDR 1997 16:49 Barypeithes pellucidus (Boheman 1843) 1947 North America Van Dyke 1947 Campylirhynchus castor (Fabricius 1792) O’Brien & Wibmer 1982 Caulophilus oryzae (Gyllenhal 1838) <1915 CDFA 1915 4:307 Ceutorhynchus assimilis (Paykull 1792) 1945 Europe CDFA 1953 42:204 Ceutorhynchus rapae Gyllenhal 1837 1920 North America Lindroth 1957 Coccotrvpes advena Blandford 1894 2006 North America Penrose 2007, Atkinson 2015 Coccotrvpes dactyliperda (Fabricius 1801) 1931 Africa Wood 1982 Coniatus splendidulus Fabricius 1781 2010 North America Eckberg & Foster 2011 Cosmobaris americana Casey 1920 1943 North America O’Brien & Wibmer 1982 Cosmobaris scolopacea Germar 1818 1939 Europe Van Dyke 1950 Cnptorhynchus lapathi (Linnaeus 1758) Europe Fumiss & Carolin 1977 Dactylotnpes longicollis (Wollaston 1864) 2009 Africa LeBonte & Takahashi 2012 Diaprepes abbreviatus (Linnaeus 1758) 2005 North America Grafton-Cardwell et al. 2004 yes 94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Euwallacea fornicatus (Eichhoff 1868) 2004 Asia Haack 2006, Atkinson 2015 yes Gonipterus scutellatus Gyllenhal 1833 1994 Australia CPPDR 1994 13:4 yes Gymnetron antirrhini (Paykull 1800) North America Findroth 1957 Gymnetronpascuorum (Gyllenhal 1813) North America Findroth 1957 Gymnetron tetrum (Fabricius 1792) 1942 Europe Findroth 1957, CDFA 1942 31:177 Hylastinus obscurus (Marsham 1802) 1939 North America Davidson & Fyon 1987, Wood 1982 Hylurgus ligniperda (Fabricius 1787) 2002 Asia Penrose 2004, Atkinson 2015 yes Hypera brunnipennis (Boheman 1834) 1939 North America Metcalf 1995 yes Hypera meles (Fabricius 1792) North America Davidson & Fyon 1987 Hypera nigrirostris (Fabricius 1775) 1945 North America CDFA 1945 34:175 Hypera postica (Gyllenhal 1813) 1923 North America CDFA 1923 12:359 yes Hypera punctata (Fabricius 1775) 1908 North America Davidson & Fyon 1987, Evans & Hogue 2006 yes Hypothenemus californicus Hopkins 1915 Wood 1982 Hypothenemus eruditus Westwood 1836 Tropical America Wood 1982 Hypurus bertrandi (Perris 1852) Anderson 2002 Ips cadigraphus calligraphic (Germar 1824) North America Connor and Wilkson 1983 Lissorhoptrus oryzophilus Kuschel 1952 1969 North America Dowell & Gill 1989 yes Listroderes costirostris Schonherr 1826 1926 Tropical America CDFA 1926 15:116, Evans & Hogue 2006 Listronotus hyperodes (Dietz 1889) 1959 North America Dowell & Gill 1989 Macrancylus littoralis (Broun 1880) 1949 O’Brien & Wibmer 1982 Magdalis barbicornis (Fatreille 1804) 1975 North America Anderson & Cline 2011 Mecinus pvraster (Herbst 1795) Aukema et al. 2010, California Beetle Project 2006 Naupactus leucoloma (Boheman 1840) 1988 North America CPPDR 1988 7:8 Orthotomicus erosus (Wollaston 1857) 2004 Europe Haack 2004, CPPDR 2006 22:5 yes Otiorhynchus cribricollis Gyllenhal 1834 1927 Europe Findroth 1957, CDFA 1929 18:567, Evans & Hogue 2006 yes Otiorhynchus meridionalis Gyllenhal 1834 1931 Europe CDFA 1931 20:470, Evans & Hogue 2006 Otiorhynchus ovatus (Finnaeus 1758) 1923 Europe Findroth 1957, CDFA 1923 12:360, Evans & Hogue 2006 Otiorhynchus rugostriatus (Goeze 1777) 1920 Europe Findroth 1957, CDFA 1930 19:17, Evans & Hogue 2006 Otiorhynchus sulcatus (Fabricius 1775) 1925 Europe Findroth 1957, CDFA 1927 16:654, Evans & Hogue 2006 yes Pagiocerus frontalis (Fabricius 1801) 2010 North America Feathers 2015 Pantomorus cervinus (Boheman 1840) 1879 Europe CDFA 1930 19:17 yes Phloeosinus armatus Reitter 1887 1989 Africa Wood 1992 Phloeotribus liminaris Harris 1852 2002 North America Penrose 2004 Piss odes strobi (Peck 1817) 1972 North America Dowell & Gill 1989 Proctorus decipiens (FeConte 1876) 1966 North America Dowell & Gill 1989 Pselactus spadix (Herbst 1795) 1966 North America O’Brien 1970 Rhynchaenus solids (Finnaeus 1758) Findroth 1957 a Rhynchophorus vulneratus (Panzer 1789) 2010 Asia Rugman-Jones, et al. 2013 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 95 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Scolytus multistriatus (Marsham 1802) 1951 North America Powell & Hogue 1979, Wood 1982 yes Scolytus rugulosus (Ratzburg 1837) 1913 North America Powell & Hogue 1979, Wood 1982 yes Scolytus schevyrewi Semenov 1902 2002 Asia Penrose 2004, Atkinson 2015 yes Sitona crinitus (Herbst 1795) O’Brien & Wibmer 1982 Sitona cylindricollis Fahraeus 1840 1957 North America Dowell & Gill 1989 Sitona /lavescens (Marsham 1802) 1907 North America CDFA 1948 37:211, Bright 1994 Sitona hispidula (Fabricius 1776) 1917 North America Metcalf 1995, Bright 1994 Sitona lepidus Gyllenhal 1834 1907 North America Campbell et al. 1989, CDFA 1948 37:211 Sitona lineata (Linnaeus 1758) 1966 North America Dowell & Gill 1989 Sitona lineellus (Bonsdorff 1785) North America Bright 1994 Sitophilus granarius (Linnaeus 1758) 1769 Europe Metcalf 1995 Sitophilus oryzae (Linnaeus 1763) 1769 Europe Metcalf 1995 Sitophilus zeamais (Motschulsky 1855) Campbell et al. 1989 Sphenophorus venatus Chittenden 1904 1968 North America Dowell & Gill 1989 Tranes internatus Pascoe 1870 O’Brien & Wibmer 1982 Trichobaris trinotata (Say 1831) 1914 North America CDFA 1915 4:303 Tychius picirostris (Fabricius 1787) 1950 Europe CDFA 1950 39:183 Xyleborinus saxeseni (Ratzeburg 1837) < 1915 Essig 1915, Wood 1982, Ivie 2002a Xyleborus californicus Wood 1975 1944 Ivie 2002a Xyleborus dispar (Fabricius 1792) <1926 North America Essig 1926a Xyleborus pfeili (Ratzeburg 1837) 2002 Europe Penrose 2004, Atkinson 2015 Xyleborus xylographus (Say 1826) 1990 North America Wood 1982, Ivie 2002a Xylosandrus germanus Blandford 1894 2002 Asia Penrose 2004, Atkinson 2015 Coleoptera-Dermestidae Anthrenus coloratus Reitter 1881 1969 Asia Dowell & Gill 1989 Anthrenus scrophulariae Linnaeus 1758 Campbell et al. 1989, CDFA 1958 47:238, 163 Anthrenus verbasci Linnaeus 1767 California Beetle Project 2006 Attagenus fasciatus (Thunberg 1795) 1974 Dowell & Gill 1989 Attagenus unicolor (Brahrn 1791) California Beetle Project 2006 Dermestes ater (De Geer 1774) Haines 1981 Dermestes lardarius Linnaeus 1758 <1925 Campbell et al. 1989, Essig 1926a Dermestes maculatus De Geer 1774 Powell & Hogue 1979 yes Dermestes marmoratus Say 1823 <1925 Powell & Hogue 1979, Essig 1926a Dermestes sardous Kuster 1846 1967 Europe Beal 1994 Trogoderma granarium Everts 1898 1953 Asia CDFA 1953 42:202 Coleoptera: Dytisicidae Copelatus glyphicus (Say 1823) 1963 Leech 1970 96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest^ Coleoptera-Elateridae Conoderus bellus (Say 1824) 1924 North America Stone 1975 Conoderus exsul Sharp 1877 1937 Hawaii Stone 1975 Conoderus falli (Lane 1956) 1963 North America Stone 1975 Heteroderes amplicollis (Gyllenhal 1817) 1938 North America Stone 1975 Coleoptera-Endomychidae Mvcetaea subterranea (Fabricus 1801) Belov 2009 Symbiotes gibberosus (Lucas 1849) BugGuide 2011 Coleoptera-Erotylidae Megalodacne fasciata (Fabricius 1777) 1983 North America Kitayama 1986, Evans & Hogue 2006 Coleoptera-Histeridae Margarinotus merdarius (Floffman 1803) Caterino 2010 Coleoptera-Hydrophilidae Sphaeridium lunatum Fabricius 1792 1947 North America Bixby 1948 Sphaeridium scarabaeoides (Linnaeus 1758) 1920 Europe Blackwelder 1931, Evans & Hogue 2006 Coleoptera-Laemophloeidae Laemophloeus ferrugineus (Stephans 1828) <1925 Campbell et al. 1989, Essig 1926a Laemophloeus pusillus (Schonherr 1817) Campbell et al. 1989 Coleoptera-Languriidae Languria mozardi Latreille 1807 Essig 1958 Pharaxonotha kirschi Reitter 1875 1956 North America Dowell & Gill 1989 Coleoptera-Latridiidae Aridius nodifer (Westwood 1839) Campbell et al. 1989 Cartodere constricta (Gyllenhal 1827) Majka et al. 2009 Corticaria serrata (Paykull 1798) Majka et al. 2009 Cortinicara gibbosa (Herbst 1793) Majka et al. 2009 Dienerella argus (Reitter 1884) Majka et al. 2009 Dienerella jilum (Aube 1850) Majka et al. 2009 Latridius minutus (Linnaeus 1767) Majka et al. 2009 Coleoptera-Mycteridae Hemipeplus marginipennis (LeConte 1854) 1958 North America CDFA 1959 48:126 Coleoptera-Mycetophagidae Mycetophagus quadriguttatus Muller 1821 Campbell et al. 1989 Typhaea stercorea (Linnaeus 1758) Campbell et al. 1989 Coleoptera-Nitidulidae Aethina tumida Murray 1867 2005 North America Mussen 2010 Carpophilus dimidiatus (Fabricius 1792) <1925 Lindroth 1957, Essig 1926a Carpophilus hemipterus (Linnaeus 1758) <1925 Campbell et al. 1989, Essig 1926a yes DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 97 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Carpophilus lugubris Murray 1864 1974 Dowell & Gill 1989 Nitidula carnaria (Schall 1783) Dodge 1937 Nitidula flavomaculata Rossi 1790 <1933 Dodge 1937, Simms & Fothergill 2014 Stelidota geminate (Say 1825) 1957 North America Dowell & Gill 1989 Coleoptera-Oedemeridae Nacerdes melanura (Linnaeus 1758) Fumiss & Carolin 1977 Coleoptera-Ptinidae Ptinus fur {Linnaeus 1758) CDFA 1958 47:243 Coleoptera-Scarabaeidae Aphodius fimetarius Linnaeus 1758 1923 Europe Evans & Flogue 2006, Gordon & Skelley 2007 Aphodius fossor Linnaeus 1758 Europe Evans & Flogue 2006 Aphodius granarius (Linnaeus 1767) Europe Campbell et al. 1989 Aphodius hamatus Say 1824 Europe Evans & Hogue 2006 Aphodius lividus (Olivier 1789) Europe Evans & Hogue 2006 Aphodius pardalis LeConte 1857 Europe Evans & Hogue 2006 Aphodius pseudolividus Balthasar 1941 Europe Evans & Hogue 2006 Aphodius vittatus Say 1825 Europe Evans & Hogue 2006 Australaphodius frenchi (Blackburn 1892) 2008 Africa BugGuide 2011 Cotinis mutabilis Goiy & Percheron 1883 1925 Tropical America CDFA 1935 24:417, Essig 1926a yes Onthophagus taurus (Schreber 1759) 1993 North America Hoebeke & Beucke 1997 Pleurophonts caesus (Creutzer 1796) North America Hatch 1946 a Popillia japonica Newman 1841 1961 North America Dowell & Gill 1989 yes Coleoptera-Silvanidae Ahasverus advena (Waltl 1834) <1925 Metcalf 1995, Campbell et al. 1989, Essig 1926a Oryzaephi/us mercator (Fauvel 1889) Thomas 2002 yes Oryzaephilus surinamensis (Linnaeus 1758) <1921 CDFA 1921 10:142 yes Silvanus bidentatus Fabricius 1792 California Beetle Project 2006 Coleoptera: Staphylinidae Aleochara bimaculata Gravenhorst 1802 Hatch 1957 Aleochara fumata Gravenhorst 1802 Klimaszewski 1984 Aleochara lanuginosa Gravenhorst 1802 Hatch 1957 Aleochara lata Gravenhorst 1802 Hatch 1957 Aleochara puberula Klug 1834 Klimaszewski 1984 Aleochara tristis Gravenhorst 1806 California Beetle Project 2006 Aleochara villosa Mannerheim 1830 Hatch 1957 Aloconota cambrica (Wallaston 1855) California Beetle Project 2006 Amischa analis (Gravenhorst 1802) Muona 1984 Anotylus rugosus (Fabricius 1775) Moore & Orth 1977 98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Anthobium atrocephalum (Gyllenha 1827) Moore & Legner 1975 Atheta amicula (Stephens 1832) Muona 1984 Atheta coriaria Kraatz 1856 Muona 1984 Atheta dilutipennis (Motschulsky 1858) Muona 1984 Atheta fungi (Gravenhorst 1806) Muona 1984 Atheta harwoodi Williams 1930 Muona 1984 Atheta iongicornis (Gravenhorst 1802) Muona 1984 Atheta nigricornis (Thomson 1852) Muona 1984 Atheta pallidicornis (Thomson 1856) Muona 1984 Autalia puncticollis Sharp 1864 1947 North America Moore & Legner 1975 Autalia rivularis (Gravenhorst 1802) Moore & Orth 1977 Bisnius sordidus (Gravenhorst 1802) Hatch 1957, Smetana 1995 Carpelimus pusillus (Gravenhorst 1802) Moore & Legner 1975 Cilea silphoides (Linnaeus 1767) Hatch 1957 Cordalia obscura (Gravenhorst 1802) Muona 1984 Dinaraea angustula (Gyllenhal 1810) Muona 1984 Falagria concinnus Erichson 1839 Hoebeke 1985 Gauropterus fulgidus (Fabricius 1787) 1931 Hatch 1957 Gyrohypnus angustatus (Stephens 1833) 1931 Klimaszewski 1984 Gyrohypnus fracticornis (Muller 1776) Klimaszewski 1984 Habrocerus capillaricornis Gravenhorst 1806 California Beetle Project 2006 Homalota plana (Gyllenhal 1810) California Beetle Project 2006 Leptacinus intermedins Donisthorpe 1935 1927 Klimaszewski 1984 Leptacinus pusillus (Stephens 1833) 1894 Europe Klimaszewski 1984 Mannerheimia sp. Newton et al. 2001 Meotica apicalis Benick 1953 Muona 1984 Ocypus ater (Gravenhorst 1802) 1936 North America Evans & Hogue 2006 Ocypus olens (Muller 1764) 1920 Europe Van Dyke 1945 Oligota parva Kraatz 1858 Moore & Legner 1975 Olisthaerus megacephalus (Zetterstedt 1828) Moore & Legner 1975 Omalium caesum Gravenhorst 1806 Moore & Legner 1975 Omalium rivulare (Paykull 1789) Moore & Legner 1975 Oxypoda opaca (Gravenhorst 1802) Moore & Legner 1975 Oxytelus sculptus Gravenhorst 1806 Hatch 1957 Paraphloeostiba gayndahensis (MacLeay 1871) 1995 Australia Newton et al. 2001 Phacophallus parumpunctatus (Gyllenhal 1827) 1891 Europe Klimaszewski 1984 Phacophallus tricolor (Kraatz 1859) 1904 Asia Klimaszewski 1984 Philonthus cognatus Stephens 1832 California Beetle Project 2006 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 99 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest^ Philonthus debilis Gravenhorst 1802 California Beetle Project 2006 Philonthus discoideus Gravenhorst 1802 1902 Smetana 1995 Philonthus jurgans Tottenham 1937 California Beetle Project 2006 Philonthus longicornis Stephens 1832 1890 Smetana 1995 Philonthus politus (Linnaeus 1758) Smetana 1995 Philonthus rectangulus Sharp 1874 California Beetle Project 2006 Philonthus ventralis Gravenhorst 1802 California Beetle Project 2006 Placusa complanata Erichson 1839 Moore & Fegner 1975 Placusa tachyporoides (Waltl 1838) Moore & Fegner 1975 Porrhodites fenestralis (Zetterstedt 1828) Campbell 1984 Quedius mesomelinus (Marsham 1802) Hatch 1957 Stenus morio Gravenhorst 1806 Moore & Fegner 1975 Tachyporus abdominalis (Fabricius 1781) Campbell 1979 Tachyporus nitidulus (Fabricius 1781) Campbell 1979 Xantholinus linearis (Olivier 1794) Hatch 1957 Xantliolinus longiventris Heer 1839 Newton et al. 2001 Coleoptera-Tenebrionidae Alphitobius diaperinus (Panzer 1797) Campbell et al. 1989 yes Alphitophagus bifasciata (Say 1824) Campbell et al. 1989 Cvnaeus angustus (FeConte 1851) Campbell et al. 1989 Eleodes suturalis (Say 1824) 1963 North America Dowell & Gill 1989 Gnathocerus cornutus (Fabricius 1798) <1925 CDFA 1958 47:245, Essig 1926a Gnathocerus maxillosus (Fabricius 1801) CDFA 1958 47:245 Opatroides punctulatus Brulle 1832 2003 Europe Aalbu et al. 2009 Palorus ratzburgi (Wissmann 1848) 1954 North America Campbell et al. 1989 Palorus subdepressus (Wollaston 1864) 1942 North America Campbell et al. 1989 Sitophagus hololeptoides (Faporte 1840) Peck & Thomas 1998 Tenebrio molitor Linnaeus 1758 <1925 Campbell et al. 1989 Tenebrio obscurus Fabricius 1792 <1925 Campbell et al. 1989, Essig 1926a Tribolium audax Triplehorn 1968 Sokoloff 1972 Tribolium castaneum (Herbst 1797) <1921 CDFA 1921 10:142 yes Tribolium confusum duVal 1868 <1925 Campbell et al. 1989, Essig 1926a yes Tribolium destructor Uyttcnboogaart 1933 <1925 Sokoloff 1972, Essig 1926a Tribolium ferrugineum (Fabricius 1781) <1925 Essig 1926a Coleoptera-Trogidae Trox scabra (Finnaeus 1767) Findroth 1957 Coleoptera-Trogossitidae Lophocateres pusillus Klug 1833 1954 North America Arnett 1985 100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Tenebroides mauritanicus (Linnaeus 1758) <1925 Flatch 1961, Essig 1926a Coleoptera-Zopheridae Microprius rufulus (Motschulsky 1863) Lord et al. 2011 Collembola-Entomobryidae Entomobrya unostrigata Stach 1930 1938 Christiansen & Bellinger 1980 Seira domestica (Nicolet 1841) <1925 Europe Essig 1926a Willowsia platani (Nicolet 1841) 2009 Europe BugGuide 2011 Collembola-Hypogastruridae Xenylla martima Tullberg 1869 <1925 Europe Essig 1926a Dermaptera-Anisolabididae Anisolabis maritima (Bonelli 1832) 1921 Langston & Powell 1975 Euborellia annulipes (Lucas 1847) 1885 Langston & Powell 1975 Euborellia cincticollis (Gerstacker 1883) 1946 Africa CDFA 1950 39:183 Dermaptera-Chelisochidae Chelisoches mono (Fabricius 1775) 1905 Langston & Powell 1975 Dermaptera-F orficulidae Doru lineare (Eschscholtz 1822) 1865 Langston & Powell 1975 Forficula auricularia Linnaeus 1758 1923 Europe Langston & Powell 1975 yes Dermaptera-Labiduridae Labidura riparia Pallas 1773 1952 North America CDFA 1953 42:40 yes Dermaptera-Labiidae Labia minor (Linnaeus 1758) 1909 Langston & Powell 1975 Marava arachidis (Yersin 1860) 1910 Langston & Powell 1975 Dictyoptera-Blaberidae Pycnoscelus surinamensis (Linnaeus 1758) 1988 North America Dowell & Gill 1989 Dictyoptera-Blattellidae Blattella germanica (Linnaeus 1767) <1925 Powell & Hogue 1979, Atkinson et al. 1991, Essig 1926a yes Biattella vaga Hebard 1935 1942 Asia CDFA 1942 31:176 yes Phyllodromica trivittata (Serville 1839) 2004 Europe CPPDR 2011 25:7 Supella longipalpa (Fabricius 1798) 1940 North America CDFA 1952 41:50 yes Dictyoptera-Blattidae Blatta lateralis Walker 1868 1978 Africa Dowell & Gill 1989 yes Blatta orientalis Linnaeus 1758 Powell & Hogue 1979 yes Neostylopyga rhombifolia (Stoll 1813) <1925 Asia Arnett 1985, Essig 1926a Periplaneta americana (Linnaeus 1758) <1925 Africa Powell & Hogue 1979, Essig 1926a yes Periplaneta australasiae (Fabricius 1775) <1925 Africa Powell & Hogue 1979, Essig 1926a Periplaneta brunnea Burmeister 1838 1970 North America Dowell & Gill 1989 yes Periplaneta fuliginosa (Serville 1839) 1966 North America Dowell & Gill 1989 yes DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 101 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Dictyoptera-Ectobidae Luridiblatta trivattata (Servile 1839) 2009 Africa Lee 2010,Bugguide 2015 Dictyoptera-Mantidae Iris oratorius (Linnaeus 1758) 2007 Europe Arnett 1985 Mantis religiosus (Linnaeus 1758) 1959 Arnett 1985 Stagmomantis Carolina Johannson 1763 Arnett 1985, Essig 1926a Stagmomantis limbata (Hahn 1835) 1864 Buckett 1966 Tenodera angustipennis Saussure 1869 1952 Arnett 1985 Tenodera aridifolia Stoll 1813 1932 Arnett 1985 Diploda-J ulida-J ulidae Brachyiulus lusitanus Verhoeff 1898 North America Hoffman 1999, Shelley 2002, Reeves 2000 Cylindroiulus caeruleocinctus (Wood 1864) Europe Shelley 2002 Cylindroiulus truncomm (Silvestri 1896) Europe Shelley 2002 Diploiulus latistriatus (Curtis, 1845) Europe Lindroth 1957, Verhoeff 1944 Ophyiulus pilosus (Newport 1843) Europe Shelley 2002 Opiona fisheri Gardner & Shelley 1989 Europe Shelley 2002 Diploda-Polydesmida-Paradoxosomatidae Akamptogonus novarae (Humbert & DeSaussure 1869) Australia Shelley 2002 Oxidus gracilis (Koch 1847) Asia Shelley 2002 Diploda-Polydesmida-Polydesmidae Oxidus gracilis (Koch 1847) Asia BugGuide 2011 Diptera-Agromyzidae Chromatomyia syngenesiae Hardy 1849 1914 Spencer 1981 Liriomyza brassicae (Riley 1885) Spencer 1981 Liriomyza sativae Blanchard 1938 Spencer 1981 yes Liriomyza langei Frick 1951 Tropical America Spencer 1981 yes Melanagromyza splendida Frick 1953 1967 Hawaii Dowell & Gill 1989 Ophiomvia simplex (Loew 1869) <1915 Spencer 1981 Phytomyza crassiseta Zetterstedt 1860 1962 Pacific Region Dowell & Gill 1989 Phytomyza ilicicola Loew 1872 <1915 Davidson & Lyon 1987 Phytomyza ranunculi (Schrank 1803) 1965 Pacific Region Dowell & Gill 1989 Phytomyza rufipes Meigen 1830 2006 Europe Scheffer & Winkler 2008 Diptera-Anthomyiidae Delia antiqua (Meigen 1826) <1925 North America Davidson & Lyon 1987, Essig 1926a Delia platura (Meigen 1826) <1915 North America Spencer 1981, CDFA 1915 4:334 Delia radicum Linnaeus 1758 North America Griffiths 1991 Pegomya hyoscyami (Panzer 1809) <1925 Essig 1926a Pegomya betae (Curtis 1847) 1893 CDFA 1915 4:331 102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Pegomya wygodzinskyn Albuquerque 1954 1948 Australia CDFA 1948 37:204 Diptera-Bibionidae Dilophus sayi (Hardy 1966) 1950 North America Powell & Hogue 1979 Diptera-Braulidae Braula coeca Nitzsch 1818 Amett 1985 Diptera-Calliphoridae Calliphora vicina Robineau-Desvoidy 1830 Powell & Hogue 1979, James 1955 Calliphora vomitoria (Linnaeus 1758) James 1955 Chrysomya megacephala (Fabricius 1794) 1988 Tropical America Dowell & Gill 1989 Chrysoma ntfifacies (MacQuart 1843) 1987 North America Dowell & Gill 1989 Cochliomyia hominivorax (Coquerel 1858) James 1955 Lucilia illustris (Meigen 1826) 1769 Europe Metcalf 1995 Lucilia silvamm (Meigen 1826) James 1955 Lucilia sericata (Meigen 1826) 1769 Metcalf 1995, James 1955 Phormia regina (Meigen 1826) <1925 James 1955, Essig 1926a Pollenia rudis (Fabricius 1794) <1925 Cole 1969, James 1955, Essig 1926a Diptera-Cecidomyiidae Cecidomyia balsamicola (Linter 1888) 1963 North America Dowell & Gill 1989 Contarinia sorghicola (Coquillett 1899) 1960 Asia Dowell & Gill 1989 Dasineura affinis (Kieffer 1886) 1992 Europe CPPDR 1992 11:6 Dasineura balsamicola (Lintner 1888) 1964 CDFA 1965 54:81 Dasineura gleditchiae (Osten Sacken 1866) 1978 North America Dowell & Gill 1989 yes Dasineura leguminicola (Lintner 1879) <1925 North America Pritchard 1953, Essig 1926a Dasineura rhodophaga (Coquillett 1900) 1996 North America CPPDR 1996 15:77 Mayetiola destructor (Say 1817) 1879 North America Pritchard 1953 Mayetiola violicola (Coquillett 1900) 1967 North America Dowell & Gill 1989 Monarthropalpus flava (Schrank 1776) 1915 Europe Cole 1969 yes Rhopalomyia chrysanthemi (Ahlberg 1939) <1925 Cole 1969, Essig 1926a Sitodiplosis mosellana (Gehin 1857) North America Davidson & Lyon 1987 Diptera-Chaoboridae Mochlonyx cinctipes (Coquillett, 1903) 2003-2008 North America Woodward et al. 2010 Diptera: Chironomidae Goeldichironomus amazonicus (Fittkau 1968) 1990 Tropical America Sublette & Mulla 1991 Diptera-Chloropidae Meromyza americana Fitch 1856 <1925 North America Davidson & Lyon 1987, Essig 1926a yes Oscinella frit (Linnaeus 1758) <1925 Davidson & Lyon 1987, Essig 1926a Diptera-Culicidae Aedes aegypti (Linnaeus 1762) 1979 North America Dowell & Gill 1989 yes DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 103 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Culex erraticus (Dyar & Knab 1906) 1994 North America Lothrop et al. 1995 Culex pipiens Linnaeus 1758 < 1920 North America Essig 1926a Diptera-Drosophilidae Dettopsomyia nigrovittata (Malloch 1924) 1949 Australia Wheeler 1951 Drosophila suzukii Matsumura 1931 2008 Asia Walsh et al. 2011 yes Drosophila virilis Sturtevant 1916 Cole 1969 Drosophila bifurca Patterson & Wheeler, 1942 2010 Castrezana 2010 Zaprionus indianus Gupta 1970 2010 Castrezana et al. 2010 Diptera-Ephydridae Hydrellia griseola (Fallen 1813) 1922 North America Metcalf 1995 Ochthera mantis (De Geer 1776) <1925 Europe Essig 1926a Diptera-F annidae Fannia canicular is (Linnaeus 1761) <1925 Huckett 1975, Essig 1926a yes Fannia incisurata (Zetterstedt 1838) Cole 1969 Fannia manicata (Meigen 1826) Cole 1969 Fannia scalar is (Fabricius 1794) <1925 Cole 1969, Essig 1926a Diptera-Hippoboscidae Melophagus ovinus Linnaeus 1758 <1925 North America Bequeart 1953-1954, Essig 1926a Pseudolynchia canariensis (Macquart 1839) <1945 Europe Cole 1969 Diptera-Lonchaeidae Dasiops alveofrons Me Alpine 1961 1945 North America Cole 1969 Diptera-Muscidae Coenosia attenuate Stein 1903 2002 Hoebeke et al. 2003 Haematobia irritans (Linnaeus 1758) <1900 North America Davidson & Lyon 1987, Essig 1926a yes Musca autumnalis De Geer 1776 1968 North America Dowell & Gill 1989 yes Musca domestica Linnaeus 1758 1769 Europe Metcalf 1995 yes Stomoxys calcitrans (Linnaeus 1758) <1932 Cole 1969 Diptera-Oestridae Gasterophilus haemorrhoidalis (Linnaeus 1758) <1925 Cole 1969, Essig 1926a yes Gasterophilus intestinalis (De Geer 1776) <1925 Cole 1969, Essig 1926a yes Gasterophilus nasal is (Linnaeus 1758) <1925 Cole 1969, Essig 1926a yes Hypoderma bovis (Linnaeus 1758) <1925 North America Cole 1969, Essig 1926a yes Hypoderma lineatus (De Villers 1789) <1925 North America Cole 1969, Essig 1926a yes Oestrus ovis Linnaeus 1758 <1925 Europe Cole 1969, Essig 1926a Diptera-Otitidae Euxesta stigmatias Loew 1868 1996 Tropical America CPPDR 1996 15:154 Diptera-Phoridae Chonocephalus bentacaisei (Santos Abreu 1921) 2008 Africa Disney & Brown 2009 104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest^ Megaselia scutellaris (Wood 1909) 2008 Europe Disney & Brown 2009 Megaselia seticauda (Malloch 1914) 1996 Mexico CPPDR 1996 15:153 Diptera-Piophilidae Piophila casei (Linnaeus 1758) <1925 North America Cole 1969, Essig 1926a Diptera: Platystomatidae Pogonortalis doclea (Walker 1849) Australia Me Alpine 2001 Diptera-Psilidae Psila rosae (Fabricius 1794) 1963 Europe Dowell & Gill 1989 Diptera-Sciaridae Pnyxia scabiei (Hopkins 1895) Cole 1969 Diptera-Sepsidae Decachaetophora aeneipes (Meijere 1913) Cole 1969 Diptera-Stratiomyidae Inopus rubriceps (Macquart 1847) 1948 Australia Kessel 1948 yes Neoexaireta spinigera (Weidemann 1830) 1966 Hawaii Dowell & Gill 1989 Diptera-Syrphidae Eristalinus taeniops (Wiedemann 1818) 2006 Africa Hauser 2006 Eristalis arbustorum (Linnaeus 1758) Nayar 1968 Eristalis tenax (Linnaeus 1758) <1925 Europe Powell & Hogue 1979, Essig 1926a Eumerus narcissi Smith 1928 1927 CDFA 1930 19:760, Davidson & Lyon 1987 yes Eumerus strigatus (Fallen 1817) <1925 CDFA 1930 19:575, Essig 1926a yes Eumerus tuberculatus Rondani 1857 CDFA 1933 22:142 Merodon equestris (Fabricius 1794) 1914 Europe Metcalf 1995 yes Myatropha florae (Linneaus 1758) 2006 Europe BugGuide 2011 Syritta flaviventris Macquart 1842 2006 Europe Hauser 2007 Syrphus ribesi (Linnaeus 1758) <1925 Europe Essig 1926a Diptera-Tephritidae a Anastrepha ludens (Loew 1873) 1954 North America CDFA 1954 43:168 yes a Anastrepha suspensa (Loew 1862) 1984 North America Dowell & Gill 1989 yes a Bactrocera albistrigata (Meijere 1911) 2009 Asia CPPDR 2011 25:9 yes a Bactrocera cor re eta (Bezzi 1916) 1986 Asia Dowell & Gill 1989 yes a Bactrocera cucurbitae (Coquillett 1849) 1956 Hawaii Dowell & Gill 1989 yes a Bactrocera dorsalis (Hendel 1912) 1960 Hawaii Dowell & Gill 1989 yes b Bactrocera zonata (Saunders 1842) 1985 Asia Dowell & Gill 1989 yes Bactrocera oleae (Gmelin 1790) 1998 Europe CPPDR 1998 17:66 yes a Bactrocera scutellata (Hendel 1912) 2009 Asia CPPDR 2011 25:9 a Ceratitis capitata (Weidemann 1824) 1975 Tropical America Dowell & Gill 1989 yes Chaetorellia succinea (Costa 1844) 1996 North America Pitcairn et al. 2000 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 105 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 1 Rhagoletis completa Cresson 1929 1925 North America CDFA 1930 19:249 yes Rhagoletis pomonella (Walsh 1867) 1983 North America Dowell & Gill 1989 yes Terellia fuscicornis (Loew 1844) 1994 Europe CPPDR 1994 13:57 Diptera-Tipulidae Tipula paludosa Meigan 1830 1999 North America Umble & Rao 2005 yes Tipula oleracea Linnaeus 1758 2003 North America Umble & Rao 2005, CPPDR 2006 22:7 yes Embiidina-Oligotomidae Haploembia solieri (Rambur 1842) <1900 Europe Powell & Hogue 1979, Ross 1957 Oligotoma nigra Hagen 1866 1890 Europe Powell & Hogue 1979, Ross 1957 Hemiptera-Aclerdidae Aclerda tokionis (Cockerell 1896) 1900 Asia Gill 1993 Hemiptera-Adeligidae Adelges abietis (Linnaeus 1758) North America Martineau 1984 Adelges nordmannianae (Eckstein 1890) North America Fumiss & Carolin 1977 Adelges piceae (Ratzeburg 1844) 1928 North America Blackman & Eastop 1994, Rageno- vich & Mitchell. 2006 Hemiptera-Aleyrodidae Aleurodicus dugesii Cockerell 1896 1992 Mexico CPPDR 1992 11:78 yes Aleuroplatus berbericolus Quaintance & Baker 1917 1935 North America Mound & Halsey 1978 Aleurothrixus floccosus (Maskell 1896) 1966 Tropical America Dowell & Gill 1989 yes Aleurotuba jelinekii (Frauenfeld 1867) 1963 Europe Dowell & Gill 1989 Aleurotuberculatus aucubae (Kuwana 1932) 2002 Asia CPPDR 2006 22:15 Aleurotuberculatus jasmini (Takahashi 1932) 1997 Hawaii CPPDR 1996 15:156 Aleyrodes proletella (Linnaeus 1758) 2001 Europe Penrose 2001 Bemisia argentifolii Bellows & Perring 1994 1991 North America Bellows et al. 1994 yes Bemisia tabaci (Gennadius 1889) North America Mound & Halsey 1978 yes Dialeurodes citri (Ashmead 1885) 1906 North America Metcalf 1995, yes Dialeurodes citrifolii (Morgan 1893) 1985 North America Dowell & Gill 1989 Parabemisia myricae (Kuwana 1927) 1978 Asia Dowell & Gill 1989 Paraleyrodes minei Iaccarino 1990@ 1983 Mexico Dowell & Gill 1989 Siphoninus phillyreae (Haliday 1835) 1988 Europe Dowell & Gill 1989 yes Tetraleurodes acaciae (Quaintance 1914) 1900 Mexico Mound & Halsey 1978 Tetraleurodes sp Cockerell 1902 1983 Tropical America Dowell & Gill 1989 Trialeurodespackardi (Morrill 1903) North America Mound & Halsey 1978 Trialeurodes ruborum (Cockerell 1897) 1953 North America CDFA 1953 42:203 Trialeurodes vaporariorum (Westwood 1856) <1915 North America CDFA 1915 4:197 yes Trialeurodes variabilis (Quaintance 1900) 2002 North America CPPDR 2003 21:10 Hemiptera-Anthocoridae Anthocoris nemoralis (Fabricius 1794) 1990 North America Hagen & Dreistadt 1990; Lattin 2007 106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Buchananiella continua (White 1880) 1934 Europe Lattin 2001 Dufouriellus ater (Dufour 1833) Eleniy & Froeschner 1988; Lattin 2007 Macrotrachelia nigronitens (Stal 1860) 2001 Tropical America Lewis et al. 2005; Lattin 2007 Xvlocoris galactinus (Fieber 1837) Lattin 2007 Xvlocoris sordid us (Reuter 1871) Lattin 2007 Hemiptera-Aphalaridae Ctenary>taina eucalypti (Maskell 1890) 1991 Australia CPPDR 1991 10:5 yes Ctenarytaina spatulata Taylor 1977 1991 Australia CPPDR 1991 10:42 Psyllopsis fraxinicola (Foerster 1848) 1920 Europe Hodkinson 1988 Hemiptera-Aphididae Acyrthosiphon festucae (Theobald 1917) 1970 Blackman & Eastop 1994 Acyrthosiphon kondoi Shinji 1938 1975 Asia Dowell & Gill 1989 Acyrthosiphon lactucae (Passerini 1860) Blackman & Eastop 1984, Foottit et al. 2006 Acyrthosiphon loti (Theobald 1913) 1975 Blackman & Eastop 1984 Acyrthosiphon malvae (Mosley 1841) Smith & Parron 1978 Acyrthosiphon pisum (Flams 1776) <1915 North America Davidson & Lyon 1987, Foottit et al. 2006 yes Aloephagus myersi Essig 1950 1939 Africa Blackman & Eastop 1994, Foottit et al. 2006 Aphis arundinis (Fabricius 1775) 1881 North America Blackman & Eastop 1994, Essig 1917 Aphis citricola Van der Goot 1912 Mexico Blackman & Eastop 1994 yes Aphis craccivora Koch 1854 <1915 Tropical America Blackman & Eastop 1994, Foottit et al. 2006 yes Aphis cytisorum Flartig 1841 Smith & PaiTon 1978, Foottit et al. 2006 Aphis fabae Scopoli 1763 <1915 North America CDFA 1915 4:104, Foottit et al. 2006 yes Aphis gossypii Glover 1877 <1915 CDFA 1915 4:89, Foottit et al. 2006 yes Aphis hederae Kaltenbach 1843 1910 Blackman & Eastop 1994, Essig 1917, Foottit et al. 2006 yes Aphis middletonii (Thomas 1879) North America Smith & Parron 1978 Aphis nasturtii Kaltenbach 1843 Blackman & Eastop 1984, Foottit et al. 2006 Aphis nerii Fonscolombe 1841 <1915 CDFA 1915 4:91, Foottit et al. 2006 Aphis pomi De Geer 1773 <1915 North America CDFA 1915 4:92, Foottit et al. 2006 Asiphonella cynodonti (Das 1918) 1958 North America CDFA 1959 48:127 Aulacorthum circumflexum (Buckton 1876) Blackman & Eastop 1984, Foottit et al. 2006 Aidacorthum solani (Kaltenbach 1843) Blackman & Eastop 1994, Foottit et al. 2006 yes Betulaphis brevipilosa Bomer 1940 Blackman & Eastop 1994, Foottit et al. 2006 Brachycaudus cardui (Linnaeus 1758) <1916 Blackman & Eastop 1994, Essig 1917 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 107 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Brachycaudus persicae (Passerini 1860) 1910 North America Blackman & Eastop 1984, Foottit et al. 2006 yes Brachycaudus rumexicolens (Patch 1917) 1984 Europe Dowell & Gill 1989 Brachycaudus schwartzi (Bomer 1931) 1975 Europe Blackman & Eastop 1994, Foottit et al. 2006 Brachycaudus tragopogonis (Kaltenbach 1843) 1975 Europe Dowell & Gill 1989 Brachycolus asparagi Mordvilko 1928 1984 North America Dowell & Gill 1989 Brevicoryne brassicae (Linnaeus 1758) 1876 North America Davidson & Lyon 1987, Foottit et al. 2006 yes Calaphis betulicola (Kaltenbach 1843) Blackman & Eastop 1994, Foottit et al. 2006 Cal aphis flava Mordvilko 1928 Blackman & Eastop 1994 Calaphis juglandis Goeze 1778 1952 Asia CDFA 1952 41:180 yes CaUipterinella calliptera (Hartig 1841) Blackman & Eastop 1994, Foottit et al. 2006 Callipterinella minutissima (Stroyan 1953) Blackman & Eastop 1994, Foottit et al. 2006 Capitophorus hippophaes (Walker 1852) Blackman & Eastop 1994, Foottit et al. 2006 Carolinaia howardi (Wilson 1911) 1916 Essig 1915 Cavariella pastinacae (Linnaeus 1758) Blackman & Eastop 1994 Cerataphis lataniae (Boisduval 1867) <1916 Blackman & Eastop 1984, Essig 1917, Foottit et al. 2006 yes Cerataphis orchidearum Westwood 1879 Blackman & Eastop 1994, Foottit et al. 2006 Chaetosiphon tetrarhodum (Walker 1849) Blackman & Eastop 1984, Foottit et al. 2006 Chaitophoms leucomelas Koch 1854 1966 Europe Blackman & Eastop 1994, Foottit et al. 2006 yes Chromaphis juglandicola (Kaltenbach 1843) 1896 Europe CDFA 1915 4:82, Foottit et al. 2006 Cinara atlantica (Wilson 1919) Blackman & Eastop 1994 Cinara cupressi (Buckton 1881) 1916 Europe Foottit et al. 2006 yes Cinara pilicornis (Hartig 1841) Blackman & Eastop 1994, Foottit et al. 2006 Cinara tujafdina (del Guercio 1909) Fumiss & Carolin 1977, Foottit et al. 2006 Colopha ulmicola (Fitch 1859) Blackman & Eastop 1994 Coloradoa rufomaculata (Wilson 1908) Blackman & Eastop 1984 Cryptomyzus ribis (Linnaeus 1758) Blackman & Eastop 1984, Foottit et al. 2006 Diuraphis noxia (Mordvilko 1913) 1988 North America Dowell & Gill 1989 yes Drepanaphis acerifoliae Thomas 1878 <1916 CDFA 1914 3:85 yes Drepanosiphum oregonensis Granovsky 1939 Blackman & Eastop 1994, Foottit et al. 2006 108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Drepanosiphum platanoides (Schrank 1801) 1908 Europe Foottit et al. 2006 Dysaphis apiifolia (Theobald 1923) Blackman & Eastop 1984, Foottit et al. 2006 yes Dysaphis crataegi (Kaltenbach 1843) Blackman & Eastop 1984, Foottit et al. 2006 yes Dysaphis foeniculus (Theobald 1922) Blackman & Eastop 1984, Foottit et al. 2006 yes Dysaphis plantaginae (Passerini 1860) Blackman & Eastop 1984, Foottit et al. 2006 Dysaphis tulipae (Fonscolombe 1841) <1925 Europe Blackman & Eastop 1984, Essig 1926a, Foottit et al. 2006 yes Elatobium abietina (Walker 1849) 1915 Europe Fumiss & Carolin 1977, Foottit et al. 2006 Eriosoma americana (Riley 1879) <1916 Blackman & Eastop 1984 yes Eriosoma Ianigera (Hausmann 1802) 1870 North America Metcalf 1995 Eriosoma pyricola Baker & Davidson 1916 <1925 North America Blackman & Eastop 1994, Essig 1926a, Foottit et al. 2006 Eriosoma rileyi Thomas 1878 1913 North America Blackman & Eastop 1984 Eucallipterus tiliae (Linnaeus 1758) 1909 North America Davidson 1909, Foottit et al. 2006 Eucarazzia elegans (Ferrari 1872) 1984 Africa Dowell & Gill 1989 Euceraphis betulae Koch 1855 Blackman & Eastop 1994, Foottit et al. 2006 Euceraphis punctipennis (Zetterstedt 1828) Fumiss & Carolin 1977, Foottit et al. 2006 Eulachnus agilis (Kaltenbach 1843) Blackman & Eastop 1994, Foottit et al. 2006 Eulachnus rileyi (Williams 1843) Blackman & Eastop 1994, Foottit et al. 2006 Euthoracaphis umbellulariae (Essig 1932) 1929 Asia CDFA 1941 30:353, Blackman & Eastop 1994, Foottit et al. 2006 yes Forda formicaria Heyden 1837 Blackman & Eastop 1984, Foottit et al. 2006 Forda marginata Koch 1857 Blackman & Eastop 1984, Foottit et al. 2006 Hayhurstia atriplicis (Linnaeus 1761) Foottit et al. 2006 Flyadaphis coriandri (Das 1918) 1999 Europe CPPDR 1999 18:19, Foottit et al. 2006 Hyadaphis foeniculi (Passerini 1860) Blackman & Eastop 1984, Foottit et al. 2006 yes Hyalopterus pruni (Geoffroy 1762) 1881 North America Metcalf 1995, Foottit et al. 2006 yes Hyperomyzus lactucae (Linnaeus 1758) <1916 Blackman & Eastop 1984, Foottit et al. 2006 Hysteroneura setariae (Thomas 1878) 1955 North America CDFA 1966 55:93, Dowell & Gill 1989 \ldiopterus nephrelepidis Davis 1909 <1915 CDFA 1915 4:101, Foottit et al. 2006 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 109 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Illinoia azaleae (Mason 1925) 1954 CDFA 1954 43:76 Illinoia liriodendri (Monell 1879) 1974 North America Blackman & Eastop 1994 yes Lipaphis erysimi (Kaltenbach 1843) Blackman & Eastop 1984 yes Macrosiphoniella sanborni (Gillette 1908) Blackman & Eastop 1994, Foottit et al. 2006 Macrosiphum eurphorbiae (Thomas 1878) Blackman & Eastop 1994 yes Macrosiphum avenae (Fabricius 1775) 1916 Essig 1915, Davidson & Lyon 1987, CDFA 1915 4:86 yes Macrosiphum rosae (Linnaeus 1758) <1915 CDFA 1915 4:105, Foottit et al. 2006 Macrosiphum tiliae (Monell 1879) Blackman & Eastop 1994 Melanaphis sacchari (Zehntner 1895) 1997 Asia CPPDR 1997 16:49, Foottit et al. 2006 yes Melanocallis caiyaefoliae (Davis 1910) 1958 North America CDFA 1959 48:127 Melaphis rhois (Fitch 1866) <1916 North America Essig 1915 Metopolophium festucae (Theobald 1917) 1970 Foottit et al. 2006 Mindarus abietinus Koch 1857 Blackman & Eastop 1994, Foottit et al. 2006 Mon ell ia cancel!a (Fitch 1855) <1932 Blackman & Eastop 1984 yes Monelliopsis caryae (Monell 1879) <1925 North America Essig 1926a yes Myzaphis rosarum (Kaltenbach 1843) <1915 Blackman & Eastop 1984, Foottit et al. 2006 Myzocallis castaneae (Fitch 1856) 1914 Essig 1915 Myzocallis castanicola Baker 1916 1914 Europe Blackman & Eastop 1984, Foottit et al. 2006 Myzocallis coiyli (Goeze 1778) <1916 North America Blackman & Eastop 1994, Foottit et al. 2006 Myzocallis punctatus (Monell 1879) North America Blackman & Eastop 1994 Myzocallis walshii (Monell 1879) North America Blackman & Eastop 1994 Myzus ascalonicus Doncaster 1946 North America Blackman & Eastop 1984, Foottit et al. 2006 Myzus cerasi (Fabricius 1775) <1915 Davidson & Lyon 1987, CDFA 1915 4:96, Foottit et al. 2006 yes Myzus certus (Walker 1849) Blackman & Eastop 1984, Foottit et al. 2006 Myzus dianthicola Hille Ris Lambers 1966 Blackman & Eastop 1984 Myzus formosanus Takahashi 1923 1958 Essig 1958 Myzus hemerocallis Takahashi 1921 1959 Asia Dowell & Gill 1989 Myzus ligustri (Mosley 1841) Blackman & Eastop 1984, Foottit et al. 2006 110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Myzus lythri (Schrank 1801) Blackman & Eastop 1994, Foottit et al. 2006 Myzus ornatus Laing 1932 1926 Europe Essig 1926b Myzus persicae (Sulzer 1776) <1915 North America Blackman & Eastop 1994, CDFA 1915 4:98 yes Myzus varians Davidson 1912 Asia Blackman & Eastop 1984 Nasonovia ribisnigri (Mosley 1841) 2001 North America CPPDR 1998 17:69, Foottit et al. 2006 yes Nearctaphis bakeri (Cowen 1895) <1915 Blackman & Eastop 1984 Neophyllaphis araucariae Takahashi 1937 Blackman & Eastop 1994, Foottit et al. 2006 Neophyllaphis podocarpi Takahashi 1920 1954 Australia Blackman & Eastop 1994, Foottit et al. 2006 yes Neotoxoptera formosana Takahashi 1921 1927 Asia Blackman & Eastop 1984, Essig 1958, Foottit et al. 2006 Neotoxoptera oliveri (Essig 1935) 1927 Asia Blackman & Eastop 1984, Foottit et al. 2006 Neotoxoptera violae Pergande 1900 <1915 Blackman & Eastop 1984, CDFA 1915 4:101 Ovatus crataegarius (Walker 1850) Smith & Parron 1978, Foottit et al. 2006 Pachypappa pseudobyrsa (Walsh 1863) 1953 North America Smith & Parron 1978 Pemphigus betae Doane 1900 <1915 CDFA 1915 4:75 yes Pemphigus bursarius (Linnaeus 1758) 1955 CDFA 1955 44:164, Foottit et al. 2006 yes Pentalonia nigronervosa Coquerel 1859 <1916 Blackman & Eastop 1994, Foottit et al. 2006 Periphyllus californiensis (Shinji 1917) Fumiss & Carolin 1977, Foottit et al. 2006 yes Periphyllus lyropictus (Kessler 1886) Fumiss & Carolin 1977, Foottit et al. 2006 yes Periphyllus testudinaceus (Femie 1852) Fumiss & Carolin 1977, Foottit et al. 2006 yes Phorodon humuli (Schrank 1801) 1890 North America Blackman & Eastop 1994, Foottit et al. 2006 Phyllaphis fagi (Linnaeus 1767) Europe Blackman & Eastop 1994, Foottit et al. 2006 yesS Pleotrichophorus chrysanthemi (Theobald 1920) Foottit et al. 2006 Pterocallis alni (De Geer 1773) Europe Blackman & Eastop 1994, Foottit et al. 2006 Pterocomma populea (Kaltenbach 1843) Blackman & Eastop 1994, Foottit et al. 2006 Pterocomma salicis (Linnaeus 1758) Blackman & Eastop 1994, Foottit et al. 2006 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 111 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest** Rhopalomyzus poae (Gillette 1908) 1955 North America Dowell & Gill 1989 Rhopalosiphoninus latysiphon (Davidson 1912) Foottit et al. 2006 Rhopalosiphoninus staphyleae (Koch 1854) Blackman & Eastop 1984, Foottit et al. 2006 Rhopalosiphum insertum (Walker 1849) 1955 North America Smith & Parron 1978 Rhopalosiphum maidis (Fitch 1856) <1916 Blackman & Eastop 1994, Essig 1917, Foottit et al. 2006 yes Rhopalosiphum nymphaeae (Linnaeus 1761) <1916 CDFA 1917 6:65, Foottit et al. 2006 Rhopalosiphum padi (Linnaeus 1758) Blackman & Eastop 1994 yes Rhopalosiphum rufiabdominalis (Sasaki 1899) Blackman & Eastop 1984, Foottit et al. 2006 Sarucallis kahawaluokalani (Kirkaldy 1907) 1937 North America Blackman & Eastop 1994 yes Schizaphis graminum (Rondani 1852) <1925 North America Metcalf 1995, Essig 1926a, Foottit et al. 2006 yes Schizolachnus pineti (Fabricius 1781) Blackman & Eastop 1984, Foottit et al. 2006 Shivaphis celti Das 1918 2002 North America CPPDR 2006 22:8-9, Foottit et al. 2006 yes Sinomegoura citricola (van der Goot 1917) 2009 Asia CPPDR 2011 25:16 Sipha flava (Forbes 1884) Smith & Parron 1978 yes Sipha maydis Passerini 1860 2007 Europe CPPDR 2011 24:9-11 Sitobion fragariae (Walker 1848) Foottit et al. 2006 Smynthurodes betae Westwood 1849 Blackman & Eastop 1994, Foottit et al. 2006 Takecallis arundicolens (Clarke 1903) 1903 Asia Blackman & Eastop 1994, CDFA 1915 4:84, Foottit et al. 2006 yes Takecallis arundinariae (Essig 1917) 1911 Asia Blackman & Eastop 1994, Foottit et al. 2006 Takecallis taiwanus (Takahashi 1926) Asia Blackman & Eastop 1994, Foottit et al. 2006 Tetraneura nigriabdominalis (Saski 1899) Blackman & Eastop 1994 Tetraneura ulmi (Linnaeus 1758) Blackman & Eastop 1994, Foottit et al. 2006 Thecabius affmis (Kaltenbach 1843) Blackman & Eastop 1994 Therioaphis riehmi (Bomer 1949) 1956 Dowell & Gill 1989 Therioaphis trifolii (Monell 1882) 1954 Europe Metcalf 1995, Foottit et al. 2006 Tinocallis platani (Kaltenbach 1842) 1935 Europe Blackman & Eastop 1994, Foottit et al. 2006 Tinocallis ulmifolii (Monell 1879) 1914 North America Blackman & Eastop 1994 112 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Tinocallis uImipar\>ifoIiae Matsumura 1998 2001 Asia CPPDR 1998 17:10 Tinocallis zelkowae (Takahashi 1919) 1983 Asia Dowell & Gill 1989 Toxoptera aurantii (Fonscolombe 1841) 1907 Asia CDFA 1915 4:78, Foottit et al. 2006 yes Tuberculatus annulatus (Hartig 1841) Blackman & Eastop 1994, Foottit et al. 2006 Uroleucon achilleae Koch 1855 1966 Europe Blackman & Eastop 1984, Foottit et al. 2006 Vesiculaphis caricis (Fullaway 1910) 1962 Dowell & Gill 1989 Hemiptera-Artheneidae Holcocranum saturejae (Kolenati 1845) 1997 Europe Wheeler & Stoops 1999 Hemiptera-Asterolecaniidae Asterolecanium arabidis (Signoret 1876) 1940 North America Gill 1993 Asterolecanium bambusae (Boisduval 1869) Asia Gill 1993 Asterolecanium grandiculum Russell 1941 Tropical America Gill 1993 Asterolecanium minus Russell 1941 North America Gill 1993 Asterolecanium quercicola (Bouche 1851) North America Gill 1993 yes Asterolecanium variolosum (Ratzeburg 1970) 1913 Europe Gill 1993 Planchonia stentae (Brain 1920) 1980 Africa Miller et al. 2005 Pollinia pollini (Costa 1828) 1892 Europe Gill 1993 Hemiptera-Cercopidae Philaenus spumarius (Linneaus 1758 ) BugGuide 2011, Karban & Strauss 2004 Hemiptera-Cicadelldidae Agalliota barretti (Ball 1900) 1972 North America Dowell & Gill 1989 Allygus mixtus Fabricius 1794 2007 North America CPPDR 2011 24:6 Anoscopus flavostriatus (Donovan 1799) North America Hamilton 1983 Anoscopus serratulae (Fabricius 1775) Europe Hamilton 1983 Aphrodes costatus (Panzer 1799) Europe Hamilton 1983 Balclutha rosea Scott 1876 1982 Tropical America Dowell & Gill 1989 Circulifer tenellus (Gillette & Baker 1895) 1899 Africa CDFA 1915 4:64, Hamilton 1983 yes Empoasca bipunctata (Oshanin 1871) 1955 Europe Hamilton 1983 Empoasca fabae (Harris 1841) <1915 North America CDFA 1915 4:62, 21:377 yes Empoasca cf. guevarai Gonzalez 1955 1996 Tropical America CPPDR 1996 15:5 Empoasca mexara Ross & Moore 1957 1964 Mexico CDFA 1965 54:81 Erythroneura comes (Say 1825) 1868 North America CDFA 1929 18:766 Eupteryx decemnotata Rey 1891 2009 Europe CPPDR 2011 25:13 Euptery>x melissae Curtis 1837 1903 Europe Hamilton 1983 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 113 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Eus cel is obsoleta (Kirschbaum 1858) 1956 North America Hamilton 1983, Bugguide 2006 Eus cel is variegausa (Kirshbaum 1858) 1947 Europe Gross 2006 Fieberiella florii (Stal 1864) 1947 North America Hamilton 1983 yes Graphocephala versuta (Say 1830) 2003 CPPDR 2006 22:43 Homalodisca vitripennis (Germar 1821) 1994 North America CPPDR 1994 13:8 yes Idiocerus decimaquartus (Schrank 1776) 1928 Europe Hamilton 1983 Idona minuenda (Moznette 1919) 1970 Tropical America Dowell & Gill 1989 Japan anus hyalinus (Osbom 1900) 1975 North America Dowell & Gill 1989 Latalus misellus (Ball 1899) 1980 North America Dowell & Gill 1989 Macropsis mendax (Fieber 1868) 1944 Europe Hamilton 1983 Macropsis ulmi (Scott 1873) 1955 Europe Dowell & Gill 1989 Negosiana dual is (DeLong 1942) 1979 North America Dowell & Gill 1989 Opsius stactogalus Fieber 1866 1914 Europe Hamilton 1983 Psammotettix emarginatus Greene 1971 1979 North America Dowell & Gill 1989 Rhytidodus decimasquartus (Schrank 1776) North America Hamilton 1983 Ribantiana tenerrima (Herrich-Schaeffer 1834) BugGuide 2011 Ribantiana ulmi (Linnaeus 1758) 1926 Europe Hamilton 1983 Sanctanus sonorus DeLong & Hershenberger 1946 1986 North America Dowell & Gill 1989 Scaphytopius nitridus (DeLong 1943) 1975 Tropical America Dowell & Gill 1989 Sophonia rufofascia Kuoh & Kuoh 1983 1995 Hawaii CPPDR 1996 15:4 yes Stirellus cf. bicolor Van Duzee 1892 1987 North America Dowell & Gill 1989 Texananus gladius DeLong 1938 1963 North America Dowell & Gill 1989 Thamnotettix zelleri (Kirschbaum 1868) 1995 Europe CPPDR 1995 14:4 Trypanalebra balli Young 1957 1983 North America Dowell & Gill 1989 Typhlocyba prunicola Edwards 1914 1949 Europe Hamilton 1983 Typhlocyba quercus (Fabricius 1777) 1955 North America Hamilton 1983 Typhlocyba rosae (Linnaeus 1758) BugGuide 2011 Hemiptera-Cimicidae Cimex lectularius Linnaeus 1758 <1900 Milne & Milne 1980 yes Hemiptera-Coccidae Ceroplastes cistudiformis Cockerell 1893 1897 Tropical America Gill 1988 yes Ceroplastes sinensis Del Guercio 1900 1887 Asia Gill 1988 yes Coccus hesperidum Linnaeus 1758 1880 Africa Gill 1988, Miller et al. 2005 Coccus longulus (Douglas 1881) 1909 Africa Gill 1988, CDFA 1915 4:140 Coccus pseudomagnoliarum (Kuwana 1914) 1900 Asia Gill 1988 yes Eucalymnatus tessellatus (Signoret 1873) 1897 Tropical America Gill 1988, CDFA 1915 4:139, Miller et al. 2005 Eulecanium caryae (Fitch 1857) 1936 North America Gill 1988 Eulecanium cerasorum (Cockerell 1899) 1909 Asia Gill 1988, Miller et al. 2005 yes 114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Eulecanium excrescens (Ferns 1920) <1925 Asia Gill 1988, Essig 1926a Eulecanium kunoense (Kuwana 1907) 1896 Asia Gill 1988 Eulecanium tiliae (Linnaeus 1758) 1984 North America Gill 1988 Neopulvinaria innumerabilis (Rathvon 1854) <1915 North America Gill 1988, CDFA 1915 4:135 Parasaissetia nigra (Nietner 1861) 1906 Asia Gill 1988 Parthenolecanium corni (Bouche 1844) 1881 North America Gill 1988 Parthenolecanium fletcheri (Cockerell 1893) 1889 North America Gill 1988, Miller et al. 2005 Parthenolecanium persicae (Fabricius 1776) 1897 Europe Gill 1988, Miller et al. 2005 Parthenolecanium pruinosum (Coquillett 1891) <1915 Tropical America Gill 1988 yes Parthenolecanium quercifex (Fitch 1859) North America Gill 1988 Physokermes hemicryphus (Dalman 1825) 1958 Europe CDFA 1959 48:127 Physokermes piceae (Schrank 1801) 1958 North America Dowell & Gill 1989 Protopulvinaria pyriformis (Cockerell 1894) 1945 North America Gill 1988 Pulvinaria citricola Kuwana 1914 1939 North America Gill 1988 Pulvinaria delottoi Gill 1979 1973 Africa Gill 1988, Miller et al. 2005 Pulvinaria floccifera (Westwood 1870) North America Gill 1988 yes Pulvinaria hydrangeae Steinweden 1946 North America Gill 1988 Pulvinaria mesembryanthemi (Vallot 1829) 1971 Africa Miller et al. 2005 Pulvinaria phaiae Lull 1899 1906 Asia Gill 1988, Miller et al. 2005 Pulvinaria psidii Masked 1893 1992 Hawaii CPPDR 1992 11:6 yes Pulvinaria rhois Ehrhom 1898 North America Gill 1988 Pulvinaria urbicola Cockerell 1893 1993 North America CPPDR 1992 11:8 Pulvinaria vitis (Linnaeus 1758) <1915 North America Gill 1988, CDFA 1915 4:135 Saissetia coffeae (Walker 1852) 1914 Africa Gill 1988, Miller et al. 2005 yes Saissetia miranda (Cockerell & Parrott 1899) Tropical America Gill 1988 Saissetia oleae (Olivier 1791) 1862 Africa Gill 1988 yes Hemiptera-Coreidae Anasa tristis (De Geer 1773) 1854 Tropical America Metcalf 1995, Essig 1926a yes Centrocoris variegatus Kolenati 1845 2009 Europe CPPDR 2011 25:13 Hemiptera-Dactylopiidae Dactylopius opuntiae (Cockerell 1896) Mexico Gill 1993 Dactylopius tomentosus (Lamarck 1801) Mexico Gill 1993 Hemiptera-Delphacidae Delphacodes fulvidorsum (Metcalf 1923) 1982 Tropical America Dowell & Gill 1989 Eurysa magnifrons (Crawford 1914) 1959 CDFA 1959 48:147 Liburniella ornata (Stal 1862) 1939 North America Wilson & Gill 1985 Nilaparvata wolcotti Muir & Giffard 1924 1975 North America Dowell & Gill 1989 Peregrinus maidis (Ashmead 1890) 1933 Tropical America CDFA 1934 23:414 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 115 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 1 Sogatella kolophon (Kirkaldy 1907) 1985 Tropical America Dowell & Gill 1989 Hemiptera-Diaspididae Abgrallaspis cyanophylli (Signoret 1869) 1931 Tropical America Gill 1997 Abgrallaspis degeneratus (Leonardi 1896) 1928 Europe Gill 1997 Aonidia lauri (Bouche 1833) 1940 Asia Gill 1997 Aonidiella aurantii (Masked 1879) 1872 Asia Gill 1997 yes Aonidiella citrina (Coquillett 1891) 1872 Asia Gill 1997 yes Aspidiotus nerii (Bouche 1833) 1882 Europe Gill 1997 yes Aspidiotus spinosus (Comstock 1883) 1932 Asia Gill 1997 Aulacaspis rosae (Bouche 1833) 1931 Europe Metcalf 1995 Carulaspis juniperi (Bouche 1851) Europe Gill 1997 Carulaspis minima Targioni-Tozzetti 1868 1931 Europe Gill 1997 yes Chionaspis americana Johnson 1896 1933 North America Gill 1997 Chionaspis corni Cooley 1899 1900 North America Gill 1997 Chionaspis etrusca Leonardi 1908 1923 Africa Gill 1997 Chionaspis wistariae Cooley 1897 <1925 Asia Gill 1997, Miller et al. 2005, Essig 1926a Chtysomphalus aonidium (Linnaeus 1758) <1925 Essig 1926a Chtysomphalus bifasciculatus Fends 1938 1930 Asia Gill 1997, Miller et al. 2005 Chtysomphalus dictyospermi (Morgan 1889) 1909 Asia Gill 1997 Clavaspis ulmi (Johnson 1896) 1967 North America Gill 1997 Comstockiella sabalis (Comstock 1883) 1917 North America McKenzie 1956 Diaspidiotus ancylus (Putnam 1878) 1931 North America Gill 1997 Diaspidiotus liquidambaris (Kotinsky 1903) 1942 North America Gill 1997 Diaspidiotus osborni (Newell & Cockerell 1898) 1904 North America McKenzie 1956 Diaspidiotus uvae (Comstock 1881) 1946 North America Gill 1997 Diaspis boisduvalii Signoret 1869 1930 Tropical America Gill 1997 Diaspis bromeliae (Kemer 1778) 1934 Tropical America Gill 1997 Diaspis cocois Lichtenstein 1882 1919 Tropical America McKenzie 1956, Miller et al. 2005 Diaspis echinocacti (Bouche 1833) 1931 Tropical America Gill 1997 Dynaspidiotus britannicus (Newstead 1898) 1939 Europe Gill 1997 Epidiaspis leperii (Signoret 1869) 1882 Europe Gill 1997 yes Fiorinia ftoriniae (Targioni-Tozzetti 1867) 1931 North America Gill 1997 Fiorinia japonica Kuwana 1902 2008 Asia CPPDR 2011 25:17-18 Furcadaspis zamiae (Morgan 1890) 1906 Europe Gill 1997, Miller et al. 2005 yes Hetniberlesia candidula (Cockerell 1900) 1957 North America Dowell & Gill 1989 Hemiberlesia lataniae (Signoret 1869) 1931 North America Gill 1997 yes Hemiberlesia rapax (Comstock 1881) 1928 Elawaii Gill 1997 yes Kuwanaspis pseudoleucaspis (Kuwana 1902) 1938 Asia Gill 1997 116 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Lepidosaphes beckii (Newman 1869) 1889 North America Gill 1997 yes Lepidosaphes chinensis Chamberlin 1925 1927 Asia Gill 1997, Miller et al. 2005 Lepidosaphes conchiformis (Gmelin 1789) 1904 Africa Gill 1997 yes Lepidosaphes gloverii (Packard 1869) <1925 Asia Gill 1997, Essig 1926a Lepidosaphes pinnaeformis (Bouche 1851) 1936 Asia Gill 1997 Lepidosaphes noxia McKenzie 1946 1944 Asia McKenzie 1956 Lepidosaphes pallida (Green 1896) 1935 Hawaii Gill 1997 Lepidosaphes sciadopitvsi McKenzie 1955 1949 Asia Gill 1997 Lepidosaphes ulmi (Linnaeus 1758) 1870 Europe Gill 1997 yes Leucaspis portaeaureae Ferris 1942 1938 Pacific Region Gill 1997 Lindingaspis rossi (Masked 1891) 1892 Australia Gill 1997 yes Melanaspis obscurus (Comstock 1881) 1897 North America Gill 1997 yes Mercetaspis harperi (McKenzie 1949) 1949 Asia Gill 1997 a Nilotaspis halli (Green 1923) 1934 Africa Gill 1997 Odonaspis penicillata (Green 1905) 1936 Asia Gill 1997 Odonaspis ruthae Kotinsky 1907 1931 North America Gill 1997 yes Parlatoreopsis chinensis (Marlatt 1908) 1948 Asia Gill 1997 Parlatoria blanchardi (Targioni-Tozzetti 1892) 1905 Africa Gill 1997 Parlatoria camelliae Comstock 1883 1886 Asia McKenzie 1956, Miller et al. 2005 Paralatoria crotonis Douglas 1881 1935 Asia Gill 1997 Parlatoria oleae (Colvee 1880) 1890 Europe CDFA 1934 23:415 yes Parlatoria pergandei Comstock 1881 1889 North America Gill 1997 Parlatoria pittospori Masked 1891 1935 Australia Gill 1997, Miller et al. 2005 Pinnaspis aspidistrae (Signoret 1896) 1896 North America Gill 1997, Miller et al. 2005 Poliaspis cycadis Comstock 1883 1993 Gill 1997 Pseudaulacaspis pentagona Targioni-Tozzetti 1855 1888 Asia Gill 1997 Quadraspidiotus forbesi (Johnson 1896) 1934 North America Gill 1997 Quadraspidiotus juglansregiae (Comstock 1881) 1913 North America Gill 1997 yes Quadraspidiotus ostraeformis (Curtis 1843) 1959 North America Dowell & Gill 1989 Quadraspidiotus perniciosus( Comstock 1881) 1870 Asia Gill 1997, Miller et al. 2005 yes Selenaspidus albus McKenzie 1953 1935 Africa Gill 1997 Selenaspidus rubidus McKenzie 1953 1936 Africa Miller et al. 2005 Toumeyella liriodendri (Gmelin 1790) 1942 North America Gill 1988 yes Unaspis euonymi (Comstock 1881) 1945 Asia Gill 1997 yes Hemiptera-Eriococcidae Acanthococcus araucariae (Masked 1897) 1880 Australia Gill 1993, Miller et al. 2005 Acanthococcus coccineus (Cockered 1894) Gill 1993 Acanthococcus insign is (Newstead 1891) Europe Gill 1993 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 117 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest^ Acanthococcus pittospori (Ferns 1955) 1954 Australia Gill 1993 Gossyparia spuria (Modeer 1778) 1893 Europe Gill 1993 yes Hemiptera-Flatidae Siphanta acuta (Walker 1851) 1983 Hawaii Dowell & Gill 1989 Hemiptera-Issidae Acanalonia conica (Say 1830) 2002 North America CPPDR 2001 20:8 Agalmatium bilobum (Fieber) 1877 1946 Europe Gnezdilov & O’Brien 2006 Asarcopus palmarum Florvath 1921 1924 Africa Essig 1926a Caliscelis bonellii (Latreille 1807) 1965 Europe O’Brien 1967 Hemiptera-Lecanodiaspididae Lecanodiaspis prosopidis (Masked 1894) North America Gill 1993 Hemiptera-Leptopodidae Patapius spinosus (Rossi 1790) 1941 Europe Slater & Baranowski 1978 Hemiptera-Lygaeidae Blissus insularis Barber 1918 1967 North America Dowell & Gill 1989 yes Blissus leucopterus (Say 1832) 1885 North America CDFA 1915 4:208 Lachnesthus singalensis (Dohrn 1860) 1965 Asia Slater & Lattin 1965 Lamprodema maura (Fabricius 1803) 1979 Dowell & Gill 1989 Megalonotus sabulicola (Thomson 1870) 1921 North America Henry & Froeschner 1988 Hemiptera-Margarodidae Icerya purchasi Masked 1876 1868 Australia Gill 1993 yes Kuwania quercus (Kuwana 1902) 1966 Asia Dowell & Gill 1989 Margarodes meridional is Morrison 1927 1954 North America CDFA 1954 43:134 Hemiptera-Membracidae Idioderma sp. 1988 North America Dowell & Gill 1989 Hemiptera-Miridae Calocoris norwegicus (Gmelin 1790) 1920 Henry & Froeschner 1988 yes Campyloneura virgida (Herrick-Schaeffer 1835) 1964 Dowell & Gill 1989 Capsus ater (Linnaeus 1758) Wheeler & Hemy 1992 Cyrtopeltis tenuis Reuter 1895 Henry & Froeschner 1988 Halticotoma valida Townsend 1892 CDFA 1943 32:279 Heterotoma planicorne (Pallas 1772) 1964 North America Dowell & Gill 1989 Lopus decolor (Fallen 1807) Wheeler & Hemy 1992 Melanotrichus concolor (Kirschbaum 1856) Wheeler & Hemy 1992 Melanotrichus virescens (Douglas & Scott 1865) Wheeler & Henry 1992 Pilophorus clavatus (Linnaeus 1767) Henry & Froeschner 1988 Plagiognathus chry>santhemi (Wolff 1804) Wheeler & Henry 1992 Polymerus unifasciatus (Fabricius 1794) Henry & Froeschner 1988 118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Psallus ancorifer (Fieber 1858) 1914 Europe Wheeler & Henry 1992 Pseudatomoscelis seriatus (Reuter 1876) North America CPPDR 26:264 Taylorilygus pallidulus (Blanchard 1852) 1917 North America Wheeler & Henry 1992 Trigonotydus tenuis Reuter 1893 Wheeler & Henry 1992 yes Trigonotylus uhleri (Reuter 1876) 1993 North America Daehler & Strong 1995 Hemiptera-Nabidae Anaptus major (Costa 1842) Henry & Froeschner 1988 Hemiptera-Ortheziidae Orthezia insignis Brown 1887 1911 Gill 1993, CDFA 1915 4:110, Miller et al. 2005 Hemiptera-Oxycarenidae Metopoplax ditomoides (Costa 1847) 2002 Europe CPPDR 2006 22:9-10 Hemiptera-Pentatomidae Agonoscelis puberula Stal 1854 1997 North America CPPDR 2007 23: 6-7 Aelia americana Dallas 1851 1980 North America Dowell & Gill 1989 Bagrada hilaris (Burmeister 1835) 2008 Africa CPPDR 2011 25:14-15 Elasmucha lateralis (Say 1831) 1969 North America Dowell & Gill 1989 Euschistus servus (Say 1832) Henry & Froeschner 1988 yes Halyomorpha halys (Stal 1855) 2005 North America CPPDR 2006 22:8-9 yes Murgantia histrionica (Hahn 1834) <1915 North America CPPDR 1983 2:630, Essig 1917 Nezara viridula (Linnaeus 1758) 1986 North America Dowell & Gill 1989 yes Hemiptera-Phoenicococcidae Phoenicococcus marlatti Cockerell 1899 1890 Africa Gill 1993 Hemiptera—Phylloxeridae Daktulosphaira vitifoliae (Fitch 1855) 1852 North America Metcalf 1995 yes Phylloxera stellata Duncan 1922 1953 North America CDFA 1953 42:172 Hemiptera-Pseudococcidae Allococcus sp. 1980 Africa Dowell & Gill 1989 Antonina crawi Cockerell 1900 1900 Asia McKenzie 1967, Miller et al. 2002 Antonina graminis (Masked 1897) 1957 North America Dowell & Gill 1989 Antonina pretiosa Ferris 1953 1915 Asia McKenzie 1967, Miller et al. 2005 Balanococcus diminutus (Leonardi 1918) 1906 Europe McKenzie 1967 Brevennia rehi (Lindinger 1943) 1967 Asia Miller et al. 2002 Cataenococcus olivaceus (Cockerell 1896) 1960 Tropical America Dowell & Gill 1989 Chorizococcus brevicruris McKenzie 1960 1965 Hawaii Dowell & Gill 1989 Chnaurococcus trifolii (Forbes 1885) 1916 North America McKenzie 1967 Chorizococcus rostellum (Lobdell 1930) 1932 North America McKenzie 1967 Crisicoccus azaleae (Tinsley 1898) 1898 Asia McKenzie 1967 Crisicoccus pini (Kuwana 1902) 1918 Asia McKenzie 1967, Miller et al. 2002 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 119 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest^ Delottococcus cf. confusus De Lotto 1977 2006 Africa CPPDR 2007 23:10 Dysmicoccus brevipes (Cockerell 1893) 1938 North America McKenzie 1967 Eriococcus pittospori (Ferns 1955) Gill 1993 Ferrisia gi/Ii Gullan 2003 1968 North America CPPDR 2006 22:16 yes Ferrisia malvastra (McDaniel 1962) 1960’s Pacific Region CPPDR 1997 16:8-9 Ferreisia virgata (Cockerell 1893) 1963 Tropical America Dowell & Gill 1989 Fleterococcus nudus (Green 1926) 1959 North America Dowell & Gill 1989 Hypogeococcus spinosus Ferris 1953 1951 North America McKenzie 1967 Maconellicoccus hirsutus (Green 1908) 1999 North America CPPDR 1999 18:69 yes Nipaecoccus aurilanatus (Maskell 1890) 1895 Australia McKenzie 1967 yes Nipaecoccus nipae (Maskell 1893) 1897 Tropical America McKenzie 1967, Miller et al. 2002 yes Palmicultor lumpurensis (Takahashi 1950) 2006 North America CPPDR 2011 24: 11-12 Phenococcus aceris (Signoret 1875) 1971 North America Dowell & Gill 1989 Phenacoccus celtisifoliae Flollinger 1917 1936 North America McKenzie 1967 Phenacoccus dearnessi King 1901 1922 North America McKenzie 1967 Phenacoccus gossypii Townsend & Cockerell 1898 McKenzie 1967 Phenacoccus graminicola Leonardi 1908 1953 Europe CDFA 1959 48:127 Phenacoccus madeirensis Green 1923 1911 North America McKenzie 1967 yes Planococcus citri (Risso 1813) 1880 Europe Metcalf 1995 yes Planococcus ficus (Signoret 1875) 1994 Africa CPPDR 1994 13:8 Planococcus kraunhiae (Kuwana 1902) 1915 Asia McKenzie 1967 Pseudantonina arundinariae McConnell 1941 1959 North America McKenzie 1967 Pseudococcus affinis (Maskell 1894) 1946 North America McKenzie 1967 Pseudococcus calceolariae (Maskell 1897) 1915 Australia Metcalf 1995, Miller et al. 2002 yes Pseudococcus comstocki (Kuwana 1902) 1967 North America Dowell & Gill 1989 yes Pseudococcus importatus McKenzie 1960 1963 Tropical America Dowell & Gill 1989 Pseudococcus longispinus (Targioni-Tozzetti 1867) 1916 Asia Metcalf 1995 yes Pseudococcus maritimus (Ehrhom 1900) 1909 North America McKenzie 1967 yes Pseudococcus microcirculus McKenzie 1960 1954 Tropical America McKenzie 1967, Miller et al. 2002 Pseudococcus nakaharai Gimpel & Miller 1996 1973 North America Miller et al. 2005 Pseudococcus sorghiellus (Forbes 1885) 1964 North America McKenzie 1967 Rhizoecus arabicus Hambleton 1976 1996 Tropical America CPPDR 1996 15:5 yes Rhizoecus associatus (Flambleton 1946) 1964 Tropical America McKenzie 1967 yes Rhizoecus cacticans (Flambleton 1946) 1948 Tropical America McKenzie 1967 yes Rhizoecus falcifer Kunckel d'Herculais 1878 1917 North America McKenzie 1967, Miller et al. 2002 Rhizoecus kondonis Kuwana 1923 1921 Asia McKenzie 1967 120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest** Rhizoecus leucosomus (Cockerell 1901) 1956 North America McKenzie 1967 Rhizoecus nakaharai Hambleton 1976 McKenzie 1967 Rhizoecus dianthi Green 1926 1954 Asia McKenzie 1967, Miller et al. 2002 Spilococcus geraniae (Rau 1938) 1966 North America Dowell & Gill 1989 Spilococcus mamillariae (Bouche 1844) 1938 Miller et al. 2002 Trionymus americanus (Cockerell 1899) 1940 North America McKenzie 1967 Trionymus caricis McConnell 1941 1961 North America McKenzie 1967 Trionymus mocus Ferris 1953 1952 McKenzie 1967 Vryburgia amaryllidis (Bouche 1873) 1925 Africa McKenzie 1967, Miller et al. 2005 Vyburgia brevicruris (McKenzie 1960) 1935 McKenzie 1967 Vrybergia trionymoides DeLotto 1961 1994 Africa CPPDR 1994 13:84 Hemiptera—Psyllidae Acizzia acaciae-baileyanae (Froggatt 1901) 1987 Australia Dowell & Gill 1989 yes Acizzia cf.jucunda (Tuthill 1952) (undescribed #22) 1992 Australia CPPDR 1992 11:7 Acizzia cf. hakeae (Tuthill 1952) 2000 Australia CPPDR 2000 19:27-28 Acizzia uncatoides (Ferris & Klyver 1932) 1954 Pacific Region CDFA 1955 44:164 yes Agonoscena pistaciae Burckhardt & Lauterer 1989 Percy 2005 Arytaina genistae (Latreille 1804) 2010 North America Percy et al. 2012 Ary’tainilla spartiophila Forster 1848 1995 North America Percy et al. 2012 Bactericera cockerelli (Sulc 1909) 1930’s North America Liu & Trumble 2007 yes Blastopsylla occidentalis Taylor 1985 1983 Australia Hodkinson 1988 Boreioglycaspis melaleucae Moore 1964 2009 North America Arakelian 2009 yes Cacopsylla buxi (Forster 1848) 1934 Europe CDFA 1931 20:694 Cacopsylla fatsiae (Jensen 1957) 1937 Asia Hodkinson 1988 Cacopsylla pyricola (Fderster 1848) 1883 North America CDFA 1954 43:42, Essig 1915 yes Cacopsylla tobirae Miyatakel964 2007 Asia Percy et al. 2012 yes Calophya schini Tuthill 1959 1984 Tropical America Dowell & Gill 1989 yes Cryptoneossa triangula Taylor 1990 1995 Australia CPPDR 1995 14:31 Ctenaiytaina eucalypti (Masked 1895) 1991 Australia CPPDR 1991 10: 5-7 yes Ctenaiytaina longicauda Taylor 1987 1983 Australia Dowell & Gill 1989 Ctenaiytaina spatulata Taylor 1997 1991 Australia Percy et al. 2012 Diaphorina citri Kuwayama 1908 2008 North America CPPDR 2011 25:19 yes Eucalyptolyma maideni Froggat 1901 2000 Australia CPPDR 2000 19:26 yes Euphyllura olivine (Costa 1877) 2007 Europe CPPDR 2011 24:7 yes Freysuila dugesi Aleman 1887 1997 Mexico Percy et al. 2012 Glycaspis brimblecombei Moore 1964 1998 Australia CPPDR 1998 17:4-10 yes Freysuila dugesii Aleman 1887 1997 Mexico CPPDR 1997 16:42 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 121 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Heteropsylla cubana Crawford 1914 1986 Hawaii Dowell & Gill 1989 Heteropsylla flexuosa Muddiman, Hodkinson & Hollis 1992 1996 Tropical America CPPDR 1996 15:5 Homotoma ficus (Linnaeus 1758) 1969 Europe Dowell & Gill 1989 Pachypsylla celtidismamma (Fletcher 1882) 1989 North America CPPDR 1989 8:9 Pachypsylla celtidisvesicu/a (Riley 1884) 1960 North America Dowell & Gill 1989 Platycorypha nigrivirga Burckhardt 1987 2008 Tropical America CPPDR 2011 25:18 Pseudacysta perseae (Heidemann 1908) 2004 Tropical America CPPDR 2006 22:10-11 yes Psylla uncatoides (Ferris & Klyver 1932) 1955 Hawaii Dowell & Gill 1989 Trioza alacris Flor 1861 1911 Europe Percy et al. 2012 Trioza eugeniae Froggatt 1901 1988 Australia Percy et al. 2012 Hemiptera-Pyrrhocoridae Scan tins aegyptius Linnaeus 1758 2009 Europe Bryant 2009 yes Hemiptera-Rhopalidae Jadera haematoloma (Herrich-Schaeffer 1847) < 1917 Tropical America Henry and Froeschner 1988, Van Duzee 1917 Liorhyssus hyalinus (Fabricius 1794) <1925 Europe Essig 1926a Rhopalus tigrinus (Schilling 1817) 1998 Asia Wheeler & Hoebeke 1999 Hemiptera-Rhyparochromidae Raglius alboacuminatus Goeze 1778 2002 Europe Henry 2004, CPPDR 2006 22:12 Xanthochilus saturnius (Rossi 1790) 1997 Europe Henry & Adamski 1998 Hemiptera-Tingidae Corythucha gossypii (Fabricius 1794) 2000 Hawaii CPPDR 2007 23: 8 Coiythucha montivaga Drake 1919 1971 Dowell & Gill 1989 Gargaphia arizonica Drake & Carvalho 1944 1970 North America Dowell & Gill 1989 Pseudacysta per sea (Heidemann 1908) 2004 North America CPPDR 2006 22:10-11 Stephanitis pyriodes (Scott 1874) Asia Gyeltshen & Hodges 2006 Teleonemia scrupulosa Stal 1873 1994 North America CPPDR 1994 13:84 Hemiptera-Triozidae Trioza alacris Flor 1861 1911 Europe CDFA 1911-12 1:86 Trioza chenopodii Reuter 1876 1997 Europe CPPDR 1997 16:41 Trioza eugeniae Froggatt 1901 1988 Australia Dowell & Gill 1989 yes Hymenoptera-Agaonidae Acophila sp. Ishii 1934 1997 Hawaii Beardsley & Rasplus 2010 Blastophaga psenes (Linnaeus 1758) 1891 Europe Arnett 1985, CDFA 1915 4:366 Eupristina verticil lata Waterston 1921 1994 CPPDR 1994 13:62 Josephiel/aMicrocarpac Beardsley & Rasplus 2001 1997 Asia Beardsley & Rasplus 2010 Odontofroggatia galili Wiebes 1980 1994 CPPDR 1994 13:62 Odontofroggatia ishii Wiebes 1980 1994 CPPDR 1994 13:62 Walkerella microcarpae Boucek 1993 1994 CPPDR 1994 13:62 122 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Hymenoptera-Anthophoridae Ceratina dallatorreana (Friese 1869) 1949 Africa Daly 1973 Hymenoptera-Apidae Apis mellifer scutellata Lepeletier 1836 1985 North America Dowell & Gill 1989 yes Hymenoptera-Argidae Schizocerella pilicornis Holmgren 1868 North America University of California 2011 Hymenoptera-Braconidae Dendrosoter protuberans (Nees 1834) 1978 North America Hajek & Dahlsten 1981 Hymenoptera-Cephidae Cephus cinctus Norton 1872 1890 Asia Ivie 2001 Hymenoptera-Ceraphronidae Ceraphron abnormis Perkins 1910 Krombein & Hurd 1979 Hymenoptera-Chrysididae Chrysis angolensis Radoszkowski 1881 1959 North America CDFA 1959 48:127 Pseudomalus auratus (Linnaeus 1758) Bohart & Kimsey 1982 Hymenoptera-Colletidae Hylaeus bisinuatus Forster 1871 Krombein & Hurd 1979 Hylaeus punctatus (Brulle 1832) 1981 Europe Snelling 1982 Hymenoptera-Cynipidae Plagiotrochus suberi Weld 1926 1926 Europe Zuparko 1996 Hymenoptera-Encyritidae PsyUaephagus parvus Riek 1962 2007 Australia Jones et al. 2011 Psyllaephagus perplexus Riek 1962 2007 Australia Jones et al. 2011 Hymenoptera-Eulophidae Aprostocetus diplosidis Crawford 1907 1972 North America Summers 1976 Aprostocetus sp. #1 Westwood 1833 1991 Australia CPPDR 1991 10:42 Aprostocetus sp. #2 Westwood 1833 1994 Australia CPPDR 1995 14:5, Headrick et al. 1995 Cirrospilus sp. #1 1991 Australia Headrick et al. 1995 Epichiysocharis bunveili Schauff 2000 1999 Australia CPPDR 1999 18:16 Oncastichus goughi Headrick & LaSalle 1995 1980’s Australia Headrick et al. 1995 QuadrastichodeUa nova Girault 1922 1957 Australia Timberlake 1957 Selitrichodes globulus La Salle & Gates 2009 2008 Australia La Salle et al. 2009 Hymenoptera-Eupelmidae Macroneura versicularis (Retzius 1783) North America Krombein & Hurd 1979 Hymenoptera-Eurytomidae Bruchophagus gibba Boheman 1836 <1915 CDFA 1915 4:368 Bruchophagus platypterus (Walker 1834) North America Davidson & Lyon 1987 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 123 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest*’ Bruchophagus roddi (Gussakovsky 1933) Davidson & Lyon 1987 Eurytoma orchidearum (Westwood 1869) 1914 Tropical America Krombein & Hurd 1979, Essig 1915 Eurytoma tumoris Bugbee 1962 1964 Krombein & Hurd 1979 Harmolita grand is (Riley 1884) 1915 North America Metcalf 1995 Elarmolita websteri (Howard 1896) <1925 Krombein & Hurd 1979, Essig 1926a Sycophila sp. Walker 1871 1994 Hawaii CPPDR 1994 13:62 Systole geniculata Foerster 1861 Europe Krombein & Hurd 1979 Tetramesa tritici (Fitch 1859) 1853 Metcalf 1995 Hymenoptera-Figitidae Melanips opacns (Hartig 1840) Europe Krombein & Hurd 1979 Hymenoptera-F ormicidae Brachymyrmex patagonicus Mayr 1868 2010 North America Martinez et al. 2011 Camponotus savi Emery 1893 1963 North America Dowell & Gill 1989 Cardiocondyla ectopia Snelling 1974 1972 Mackay 1995 Cardiocondyla nuda (Mayr 1866) 1958 Asia Dowell & Gill 1989 Hypoponera gleadowi (Forel 1895) 1999 CPPDR 1999 18:19 Hypoponera punctatissima (Roger 1859) Krombein & Hurd 1979 Linepithema humile (Mayr 1868) 1908 Tropical America Knight & Rust 1990 yes Monomorium pharaonis (Linnaeus 1758) <1925 Africa Knight & Rust 1990, Essig 1926a yes Ochetomyrmex auropunctata (Roger 1863) Tropical America Arnett 1985 Paratrechina currens (Mostchoulsky 1863) 1964 CDFA 1965 54:81 Paratrechina longicornis (Latreille 1802) 1967 North America Dowell & Gill 1989 Paratrechina vividula (Nylander 1846) Mexico Smith 1979, Ward 2005 Pheidole fer\>ens Smith 1858 1995 Hawaii Martinez 1996 Pheidole moerens Wheeler 1908 1995 North America Martinez 1997 Pheidole teneriffana Forel 1893 1989 Africa Snelling 1992, Davidson & Lyon 1987, Martinez 1992 Plagiolepis alluaudi Emery 1894 Africa Wetterer 2013 Solenopsis wagneri Santschi 1916 1997 North America CPPDR 1997 16:41 yes Strumigenys silvestri Emery 1906 Ward 2005 Tapinoma melanocephalum (Fabricius 1793) 1991 North America CPPDR 1991 10:68 yes Tetramorium bicarinatum (Nylander 1846) 1990 North America Martinez 1993 Tetramorium caespitum (Linnaeus 1758) <1925 North America Knight & Rust 1990, Essig 1926a yes Tetramorium insolens (Smith 1861) Bolton 1979 Tetramorium pacificum Mayr 1870 1944 Krombein & Hurd 1979 Tetramorium simillium (Smith 1851) Bolton 1979 Trichoscapa membranifera (Emery 1869) 1963 North America Dowell & Gill 1989 124 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest * 5 Hymenoptera-Ichneumonidae Stilpnus gagates Gravenhorst 1807 1985 North America Dowell & Gill 1989 Hymenoptera-Megachilidae Megachile apicalis Spinola 1808 1982 North America Barthell et al. 1998, Cooper 1984 Megachile argentata (Fabricius 1793) 1949 North America Hurd 1954 Megachile concinna Smith 1879 North America Barthell et al. 1998 Megachile rotundata (Fabricius 1787) 1949 North America Powell & Hogue 1979, Barthell et al. 1998 Hymenoptera-Pteromalidae Anisopteromalus calandrae (Howard 1881) Krombein & Hurd 1979 Cerocephala rufa (Walker 1833) Krombein & Hurd 1979 Cyclogastrella deplanata (Nees 1834) Krombein & Hurd 1979 Theocolax elegans (Westwood 1874) Krombein & Hurd 1979 Trichomalus perfectus (Walker 1837) Krombein & Hurd 1979 Hymenoptera-Spechidae Cemonus lethifer (Shuckard 1837) North America Krombein & Hurd 1979 Rhopalum clavipes (Linnaeus 1758) Lindroth 1957 Hymenoptera-Tenthredinidae Ardis sulcata (Cameron 1882) 1937 Europe Middlekauff 1958 Caliroa cerasi (Linnaeus 1758) 1875 North America Metcalf 1995 yes Cladius difformis (Panzer 1799) 1921 Metcalf 1995 Endelomyia aethiops (Fabricius 1781) North America Krombein & Hurd 1979 Fenusa dohrnii (Tischbein 1846) 1986 North America Dowell & Gill 1989 Nematus ribesii (Scopoli 1763) North America Arnett 1985 Pachynematus sporax Ross 1945 1951 North America CDFA 1954 43:170 Pristiphora abbreviata (Hartig 1837) 1881 North America Krombein & Hurd 1979, CDFA4:360 yes Tomostethus multicinctus (Rohwer 1909) 1950 North America CDFA 1950 39:183 Hymenoptera-Torymidae Ditropinotus aureoviridis Crawford 1907 <1915 CDFA 1915 4:370, Dowell & Gill 1989 Megastigmus pistaciae Walker 1871 1967 Asia Dowell & Gill 1989 yes Megastigmus transvaalensis (Hussey 1956) 1960 Africa Krombein & Hurd 1979, Dowell & Gill 1989 Hymenoptera-Vespidae Polistes dominulus (Christ 1791) 1997 North America CPPDR 1997 16:44 yes Polistes apachus Saussure 1857 1921 North America Bohart & Bechtel 1957 yes Vespula germanica (Fabricius 1793) 1983 North America Vetter et al. 1995 yes Isopoda-Armadillidiidae Armadillidium nasatum Budde-Lund 1885 Europe Jass & Klausmeier 2001 Armadillidium vulgare (Latreille 1804) <1925 Europe Jass & Klausmeier 2000, Essig 1926a DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 125 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Isopoda-Trichoniscidae Haplophthalmus danicus Budde-Lund 1879 Europe Jass & Klausmeier 2001 Isopoda-Ligiidae Ligia exotica Roux 1828 Jass & Klausmeier 2001 Ligidium elrodii (Packard 1873) Europe Jass & Klausmeier 2001 Isopoda-Platyarthridae Niambia capensis (Dollfus 1895) Africa Jass & Klausmeier 2001, Brusca et al. 2001 Isopoda-Oniscidae Platyarthrus aiasensis Legrand 1954 Europe Jass & Klausmeier 2001, Isopoda-Porcellioniade Porcellio dilatatus Brandt & Ratzeburg 1831 Europe Jass & Klausmeier 2001, Porcellio Iaevis Latreille 1804 <1915 Europe Jass & Klausmeier 2001, Essig 1915 Porcellio scaber Latreille 1804 Europe Jass & Klausmeier 2001 Porcellionides pruinosus (Brandt 1833) Lindroth 1957, Richardson 1905 Isoptera-Rhinotermitidae Coptotermes formosanus Shiraki 1909 1992 Hawaii CPPDR 1992 11:4 yes Lepidoptera-Argyresthiidae Argyresthia conjugella Zeller 1839 Zhang 1994, Carter 1984 Argyresthia pruniella (Clerk 1759) Carter 1984 Lepidoptera-Arctiidae Hyphantria cunea (Drury 1773) <1915 North America Lindroth 1957, CDFA 1915 4:391, Carter 1984 yes Lepidoptera-Bucculatrigidae Buccidatrix thurberiella Busck 1914 Mexico Zhang 1994 yes Bucculatrix tridenticola Braun 1963 1963 North America Dowell & Gill 1989 Lepidoptera-Choreutidae Choreutis pariana (Clerck 1759) 1944 North America CDFA 1944 33:248 Lepidoptera-Coleophoridae Coleophora mayrella (Hiibner 1813) 1952 Europe CDFA 1952 41:182 Homaledra sabalella (Chambers 1880) 2000 North America CPPDR 2000 19:29 yes Lepidoptera-Cosmopterygidae Periploca nigra Hodges 1962 1962 North America Powell & Opler 2009, Dowell & Gill 1989 Pyroderces badia Hodges 1962 <1925 North America Essig 1926a, Powell & Opler 2009 Lepidoptera-Crambidae Asciodes gordialis Guenee 1854 2005 North America CPPDR 2006 22: 21 Diaphania nitidalis (Stoll 1781) 1992 North America CPPDR 1992 11:71 Eoreuma loftini (Dyar 1917) 1945 Asia CDFA 1945 34:175 Glyphodes onychinalis (Guenee 1854) 2000 Pacific Region CPPDR 2000 19:30 Hellula rogatalis (Hulst 1886 Powell 2005 126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 1 Lineodes elcodes Dyar 1910 2003 Tropical America CPPDR 2006 22:20 Loxostege sticticalis (Linnaeus, 1761) Carter 1984 yes Ostrinia penitalis (Grote 1796) 1952 North America CDFA 1953 42:204 Parapidiasia teterrella (Zincken 1821) 1954 North America Powell 1992 Udea ferrugalis (lliibncr 1796) 1907 North America Campbell 1927 Uresiphita reversals (Guenee 1854) 1929 CDFA 1933 22:410 yes Lepidoptera-Ethmiidae Pyramidobela angel arum Keifer 1936 1934 Mexico CDFA 1942 31:177 Lepidoptera-Gelechiidae Anarsia lineatella Zeller 1839 1900 North America Metcalf 1995, Carter 1984 yes Athrips rancidella (Herrich-Schaffer 1854) 1983 Europe Dowell & Gill 1989 Chysoesthia lingulacella (Clemens 1860) North America Powell & Opler 2009 Dichomeris marginella (Fabricius 1781) 1922 Europe CDFA 1928 17:17, 47 Euscrobipalpa artemisiella (Treitschke 1833) Powell & Hsu 2004 Euscrobipalpa artiplicella (F. von Roslerstamm 1841) Powell & Hsu 2004 Euscrobipalpa instabiliella (Douglas 1846) Powell & Hsu 2004 Euscrobipalpa obsoletella (F. von Roslerstamm 1841) Powell & Hsu 2004 Keiferia lycopersicella (Walsingham 1897) 1922 Tropical America CDFA 1934 23:301 yes Metzneria lappella (Linnaeus 1758) 1958 North America Powell & Opler 2009 Mirificarma eburnella (Denis & Schiffermullcr 1775) 1969 Powell & Hsu 2004, Powell & Opler 2009 Mirificarma formosella (Hubner 1796) 1969 Europe Dowell & Gill 1989 Pectinophora gossvpiella (Saunders 1844) 1965 Tropical America Dowell & Gill 1989 yes Phthorimaea operculella (Zeller 1873) 1854 Tropical America Metcalf 1995, Carter 1984 yes Platyedra subcinerea (Haworth 1828) 1990 Europe Powell & Opler 2009 Recurvaria nanella Denis and Schiffermullcr 1775 Aukema et al. 2010, Sitotroga cerealella (Olivier 1789) <1915 Arnett 1985 Stenolechia bathyrodyas Meyrick 1935 1980 Asia Dowell & Gill 1989 Symmetrischema striatellum (Murtfeldt 1900) 1930’s Powell & Hsu 2004, Powell & Opler 2009 Symmetrischema tango lias (Geyen 1913) 1936 Tropical America McDunnough 1939 Lepidoptera-Geometridae Alsophila pometaria (Harris 1841) <1915 North America CDFA 1915 4:477 yes Disclisioprocta stellata (Guenee 1857) 2006 North America CPPDR 2007 23:12 yes Lepidoptera-Gracillariidae Caloptilia azaleella (Brants 1913) 1960’s North America Powell & Opler 2009 Caloptilia fraxinella (Ely 1915) Zhang 1994 Caloptilia negundella (Chambers 1876) Aukema et al. 2010 Caloptilia syringella (Fabricius 1794) Zhang 1994 Lithocolletis mespilella (Hubner 1805) Zhang 1994 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 127 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Phyllocnistis citrella Stainton 1856 1999 North America CPPDR 1999 18:79 yes Phyllonoiycter crataegella (Clemens 1859) <1952 North America CDFA 1952 41:181 Phyllocnistis sp. Zeller 1848 1963 North America Dowell & Gill 1989 Lepidoptera-Heliodinidae Aetole bella Chambers 1875 1890 Europe Powell & Opler 2009 Lepidoptera-Limacodidae Sibine stimulea (Clemens 1860) 1965 North America Dowell & Gill 1989 Lepidoptera-Lymantriidae Leucoma salicis (Linnaeus 1758) 1960 North America Dowell & Gill 1989 a Lymantria dispar (Linnaeus 1758) 1976 North America Dowell & Gill 1989 yes a Lvmantria dispar asiatica Vnukovskij 1926 2005 Asia CPPDR 2006 22: 19-20 Orgyia antiqua (Linnaeus 1758) <1925 North America Martineau 1984, Essig 1926a Lepidoptera-Lyonetiidae Bedellia somnulentella (Zeller 1847) Powell & Hsu 2004, Powell 2005 Lepidoptera-Noctuidae Ascalapha odorata (Linnaeus 1758) <1925 Mexico Powell & Hogue 1979, Essig 1926a Agrotis malefida Guenee 1852 1960 North America Dowell & Gill 1989 Amphipyra tragopoginis Clerk 1759 Aukema et al. 2010, Apamea indocilis Walker 1856 1970 Dowell & Gill 1989 Athetis mindara (Bames & McDunnough 1913) 1958 CDFA 1960 49:246 Discestra trifolii (Hufnagel 1766) Zhang 1994 Euxoa ochrogaster (Guenee 1852) 1970 Dowell & Gill 1989 Feltia jaculifera (Guenee 1852) Powell 2005 Macronoctua onusta Grote 1874 1963 Dowell & Gill 1989 Noctua pronuba (Linnaeus 1758) 2001 North America Powell 2002 Peridroma saucia (Hubner 1808) Aukema et al. 2010, yes Pseudaletia unipunctata (Haworth 1809) CDFA 1959 48:20, Powell 2005 Sideridis rosa Harvey 1874 1970 North America Dowell & Gill 1989 Spodoptera exigua (Hubner 1808) 1876 Tropical America Davidson & Lyon 1987, Powell & Opler 2009 Spodoptera frugiperda (J.E. Smith 1797) 1931 CDFA 1931 20:613 Trichoplusia ni (Hubner 1803) North America Powell 2005 yes Lepidoptera-Nymphalidae Dione vanillae (Linnaeus 1758) 1875 Mexico Hogue 1974 Lepidoptera-Oecophoridae Agonopterix alstroemeriana (Clerck 1759) 1983 North America Powell 1991 Agonopterix nen>osa (Haworth 1811) <1957 Europe Hodges 1974, Powell & Opler 2009 128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Batia lunaris (Haworth 1828) 1956 Powell 1964b, Powell & Opler 2009 Borkhausenia nefrax Hodges 1974 1936 Powell 1964a Depressaria daucella (Denis & Schiffemitiller 1775) 1957 Hodges 1974, Powell & Opler 2009 Depressaria pastinacella (Duponchel 1838) Zhang 1994, Carter 1984 Endrosis sarcitrella (Linnaeus 1758) 1902 Zhang 1994, Powell & Opler 2009 Esperia sulphurella (Fabricius 1775) 1966 Europe Powell 1968b Hofmannophila pseudospretella (Stainton 1849) 1871 Hodges 1974, Powell & Opler 2009 Lepidoptera-Papilionidae Papilio cresphontes Cramer 1777 1959 North America CDFA 1959 48:127 Lepidoptera-Pieridae Pieris rapae (Linnaeus 1758) 1882 North America Metcalf 1995, Shapiro and Manolis 2007 yes Lepidoptera-Plutellidae Homadaula anisocentra Meyrick 1922 1963 Dowell & Gill 1989 yes Plutella porrectella (Linnaeus 1758) <1871 North America Powell 1967, Powell & Opler 2009 Plutella xylostella (Linnaeus 1758) North America Powell & Hogue 1979 yes Lepidoptera-Psychidae Apterona helix (Siebold 1850) 1940 Asia CDFA 1944 33:250, Powell & Opler 2009 Oiketicus townsendi (Townsend 1894) 1968 North America Dowell & Gill 1989 Thyridopteryx ephemeraeformis (Haworth 1803) 1953 North America Zhang 1994 Lepidoptera-Pterophoridae Emmelina monodactylus (Linnaeus 1758) Powell & Hsu 2004 Gillmeria pallidactyla (Haworth 1811) 1871 Powell & Hsu 2004, Powell & Opler 2009 Hellinsia longifrons (Walsingham 1915) Powell & Hsu 2004 Megalorrhipida defectalis (Walker 1864) Powell & Hsu 2004 Platyptilia carduidactyla (Riley 1869) 1922 North America Metcalf 1995 Platyptilia pallidactyla (Haworth 1811) Powell 2005 Lepidoptera-Pyralidae Achroia grisella (Fabricius 1794) 1908 Europe Zhang 1994, Carter 1984 Acrohasis indigenella (Zeller 1848) 1924 North America CDFA 1959 48:206 Acrobasis nuxvorella Neunzig 1970 1924 North America Metcalf 1995 Aglossa caprealis (Hubner 1809) 1936 Powell & Opler 2009, Aglossa pinguinalis (Linnaeus 1758) 1993 Solis 2008, Amyelois transitella (Walker 1863) 1947 North America CDFA 1948 37:204 Cadra cautella (Walker 1863) 1913 Neunzig 1990, Powell & Opler 2009 Cadra figulilella (Gregson 1871) 1930 North America CDFA 1931 20:64 Duponchelia fovealis Zeller 1847 2004 Europe CPPDR 2006 22: 20-21 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 129 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Ectomyelois ceratoniae (Zeller 1839) 1983 North America Dowell & Gill 1989 yes Ephestia elutella (Hiibner 1796) 1911 Asia Neunzig 1990 Ephestia kuehniella (Zeller 1879) 1889 Zhang 1994, Powell & Opler 2009 EtieUa zinckenella (Treitschke 1832) 1885 Europe CDFA 1915 4:433, Stone 1965 Euchromius ocelleus (Haworth 1811) <1900 North America Powell & Opler 2009 Eumysia mysiella (Dyar 1905) 1975 Dowell & Gill 1989 Galleria mellonella (Linnaeus 1758) 1875 North America Zhang 1994 Hymenia perspectalis (Hiibner 1796) North America Zhang 1994 Hypsopygia costalis (Fabricius 1775) 1968 North America Pohl et al. 2005 Paralipsa gularis (Zeller 1877) <1921 CDFA 1921 10:142 Phycitodes albatella (Ragonot 1887) Zhang 1994 Plodia interpunctella (Hiibner 1813) 1881 Europe Metcalf 1995 yes Pyralis farinalis Linnaeus 1758 <1925 Zhang 1994, Essig 1926a Pyrausta laticlavia (Grote & Robinson 1867) 1880’s North America Powell & Opler 2009 Pyrausta volupialis (Grote 1877) 1991 North America Powell & Opler 2009 Pyla fusca (Haworth 1828) Lafontaine & Wood 1997 Lepidoptera-Sesiidae Podosesia syringae (Harris 1839) 1979 North America Dowell & Gill 1989 yes Svnanthedon saxifragae (Edwards 1881) 1950 Smith 1950 Synanthedon tipuliformis (Clerck 1759) 1880 North America Zhang 1994 Synanthedon exitiosa (Say 1823) 1914 North America CDFA 1915 4:421 yes Lepidoptera-Sphingidae Manduca sexta (Linnaeus 1763) 1871 North America Metcalf 1995 yes Lepidoptera-Symmocidae Oegoconia quadripuncta (Haworth 1828) North America Powell 1992 Symmoca signatella Herrich-SchaefFer 1854 Europe CDFA 1942 31:175 Lepidoptera-Tineidae Diyadaula terpsichorella (Busck 1910) 1963 Tropical America CPPDR 1999 18:81, Powell & Opler 2009 Lindera tessellatella (Blanchard 1854) Powell & Hsu 2004 Monopis crocicapitella (Clemens 1859) Powell & Hsu 2004 Monopis rusticella (Hubner 1796) Powell & Hsu 2004 Nemapogon granellus (Linnaeus 1758) 1963 Arnett 1985 Niditinea spretella (Denis & Schiffermiiller 1775) CDFA 1943 32:258 Oinophila v-flava (Haworth 1828) Europe Powell 1964b, Powell 1992, Powell & Opler 2009 Opogona omoscopa (Meyrick 1892) Hawaii Powell 1992 yes Phereoeca praecox Gozmany & Vari 1973 1966 Africa CPPDR 2000 19:29 130 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Setomorpha rutella Zeller 1852 CDFA 1958 47:248, Powell & Hsu 2004 Tinea paUescentella Stainton 1851 Powell & Hsu 2004, Powell & Opler 2009 Tinea pellionella (Linnaeus 1758) Arnett 1985, Essig 1926a yes Tineola biseUiella Hummel 1823 Arnett 1985, Essig 1926a yes Trichophaga tapetzella (Linnaeus 1758) <1940 Arnett 1985, Essig 1926a Lepidoptera-Tortricidae Acleris variegana (Denis & Schiffermuller 1775) Powell 2005, Powell & Opler 2009 Ancylis comptana (Frolich 1828) North America Essig 1926a, 1958, Powell & Opler 2009 Bactra priapeia Heinrich 1923 North America Powell 1997 Choristoneura conflictana (Walker 1863) North America Dowell & Gill 1989 Cnephasia Iongana (Haworth 1811) Europe CDFA 1948 37:204 Crocidosema plebejana Zeller 1847 North America Powell 1992 Cydia pomonella (Linnaeus 1758) North America Metcalf 1995, Powell & Opler 2009 Cydia prunivora Walsh 1868 North America Essig 1926a Ditula angustiorana (Haworth 1811) Europe CDFA 1933 22:351 Endothenia albolineana (Kearfoot 1907) North America Dowell & Gill 1989 Epinotia solandriana (Linnaeus 1758) Aukerna et al. 2010, Epiphyas postvittana (Walker 1863) 2007 Australia Powell & Opler 2009 Grapholita molesta (Busck 1916) North America CDFA 1942 31:175 ^Lobesia botrana (Denis & Schiffermuller 1775) Europe Gullian et al. 2011 yes Lorita scarificata (Meyrick 1917) Zhang 1994 Platynota stultana Walsingham 1884 North America Powell 1980 yes Rhyacionia frustrana (Scudder, in Comstock 1880) North America Dowell & Gill 1989 Spilonota ocellana (Denis & Schiffermuller 1775) North America CDFA 1939 28:539, Powell & Opler 2009 Zeiraphera canadensis Mutura & Freeman 1967 Martineau 1984, Ferguson 1978 Zeiraphera hesperiana Mutura & Freeman 1967 1968 North America Powell 1968a Zeiraphera vancouverana McDunnough, 1925 North America Dowell & Gill 1989 Lepidoptera-Yponomeutidae Zelleria cf. hepariella Stainton 1849 1977 Europe CPPDR 2000 19:31 Lepidoptera-Zygaenidae 1969 Elarrisina metallica Stretch 1885 1966 Mexico CDFA 1943 32:98 Mollusca-Agriolimacidae 1951 Deroceras panormitanum (Lessona & Pollonera 1882) Europe McDonnel et al. 2009 Deroceras reticulatum (Muller 1774) 2002 Ebeling 1975 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 131 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest^ Mollusca-Arionidae Arion ater Linnaeus 1758 1912 North America Dowell & Gill 1989 Arion distinctus Mabille 1868 Roth & Sadeghian 2003 Arion hortensis Ferussac 1819 Europe Lindroth 1957, McDonnel et al. 2009 yes Arion intermedius Normand 1852 <1925 Lindroth 1957, McDonnel et al. 2009 Arion rufus (Linnaeus 1758) <1925 Roth & Sadeghian 2003 Arion silvaticus Lohmander 1937 Roth & Sadeghian 2003 Mollusca-Carychiidae <1925 Carychium minimum Muller 1774 <1925 Europe Roth & Sadeghian 2003 Mollusca-Daudebardiidae <1925 Oxychilus alliarius Miller 1822 Europe Lindroth 1957, Hanna 1939 Oxychilus cellarius Muller 1774 Europe Lindroth 1957, Hanna 1939 Oxychilus draparnaldi (Beck 1837) Europe Lindroth 1957, Hanna 1939 Oxychilus helveticus (Blum 1881) Lindroth 1957 Mollusca-F errussaciidae Cecilioides acicula (Muller 1774) <1925 Europe Roth & Sadeghian 2003 Mollusca-Helicellidae <1925 Cernuella virgata (Mendes de Costa 1778) <1925 Europe Roth & Sadeghian 2003 Cochlicella barbara (Linnaeus 1758) Europe Roth & Sadeghian 2003 Cochlicella ventrosa (Ferrussac 1821) <1925 Dowell & Gill 1989 Mollusca-Helicidae <1925 Cepaea nemoralis (Linnaeus 1758) Europe CDFA 1940 29:241 Eobania vermiculata (Muller 1774) <1900 Roth & Sadeghian 2003 Cantareus aspertus (Bom 1778) Europe CDFA 1941 30:352 yes Cornu awspersum (Muller 1774) <1925 Europe CDFA 1939 28:302, Hanna 1939 yes Otala Iactea (Muller 1774) Europe CDFA 1940 29:241 Theba pisana (Muller 1774) Europe CDFA 1919 8:7, Hanna 1939 yes Mollusca-Helicodiscidae <1925 Lucilla singleyana (Pilsbry 1889) North America Roth & Sadeghian 2003 Mollusca-Hygromiidae HeliceUa maritima (Drapamaud 1805) Africa Dowell & Gill 1989 Xerotricha conspurcata (Drapamaud 1801) Europe CPPDR 1996 15:8 Mollusca-Limacidae <1925 Lehmannia valentiana (Ferussac 1822) Europe Roth & Sadeghian 2003 yes Umax flavus Linnaeus 1758 Europe Lindroth 1957, McDonnel et al. 2009 yes Limax maximus Linnaeus 1758 Europe Lindroth 1957, McDonnel et al. 2009 yes Mollusca-Milacidae Mil ax gagates (Drapamaud 1801) <1900 Europe Lindroth 1957, McDonnel et al. 2009 132 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Mollusca-Patulidae Discus rotundatus (Muller 1774) <1925 Europe Roth & Sadeghian 2003 Mollusca-Pristilomatidae Hawaiia minuscula (Binney 1841) <1925 Hawaii Dowell & Gill 1989 Vitrea contracta Lowe (Westerlund 1871) <1925 Europe Roth & Sadeghian 2003 Mollusca-Punctidae <1925 Paralaoma servilis (Shuttleworth 1852) Roth & Sadeghian 2003 Mollusca-Pupillidae Pupoides albilabris (C. B. Adams 1841) North America Roth & Sadeghian 2003 Mollusca-Spiraxidae Euglandina rosea (Ferussac 1818) North America Roth & Sadeghian 2003 Mollusca-Subulinidae Rumina deco data (Linnaeus 1758) North America Dowell & Gill 1989 Mollusca-Succineidae Novisuccinea ovalis (Say 1817) North America Roth & Sadeghian 2003 Succinea Iuteola Gould 1848 North America Roth & Sadeghian 2003 yes Mollusca-Testacellidae Testacella haliotidea Drapamaud 1801 Europe McDonnel et al. 2009 Mollusca-Valloniidae Vallonia costata (Muller 1774) North America Roth & Sadeghian 2003 Vallonia excentrica Sterki 1893 North America Roth & Sadeghian 2003 Vallonia pulchella (Muller 1774) North America Lindroth 1957, Hanna 1939 Opisthopora-Lumbricidae Allolophora chlorotica (Savigny 1826) Wood & James 1993 Aporrectodea caliginosa (Savigny 1826) Wood & James 1993 Aporrectodea trapezoides (Duges 1828) Wood & James 1993 Aporrectodea tuberculata (Eisen 1874) Wood & James 1993 Aporrectodea sp. Wood & James 1993 Dendrobaena rubida (Savigny 1826) Wood & James 1993 Eisenia foetida Savigny 1826 Wetzel & Reynolds 2014, Bugg 2014 Eisenia rosea (Savigny 1826) Wood & James 1993 Eiseniella tetraedra (Savigny 1826) 1942 Wood & James 1993 Lumbricus rubellus Hoffmeister 1843 Wetzel and Reynolds 2014, Bugg 2014 Lumbricus terrestris Linnaeus 1758 Wetzel & Reynolds 2014, Bugg 2014 Octolasion cyaneum (Savigny 1826) <1925 Wood & James 1993 Opisthopora-Megascolecidae Amynthas corticis (Kinberg 1867) Hendrix & Bohlen 2002, Bugg 2014 Amynthas gracilis (Kinberg 1867) <1925 Hendrix & Bohlen 2002, Bugg 2014 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 133 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Microscolex dubius (Fletcher 1887) Wood & James 1993 Microscolex sp Rosa 1887 <1925 Wood & James 1993 Orthoptera-Gryllidae <1925 Acheta domestica (Linnaeus 1758) <1925 North America Arnett 1985, Weissman & Rentz 1977 Allonemobius fasciatus (De Geer 1773) <1925 Dowell & Gill 1989 Giyllodes supplicans (Walker 1859) <1900 North America Dowell & Gill 1989 Orthoptera-Gryllotalpidae Scapteriscus borellii Giglio-Tos 1894 <1931 North America CPPDR 2011 24: 14-16 Orthoptera-Tettigoniidae Anabrus simplex Haldeman 1852 1943 North America CDFA 1943 32:201 Neoconocephalus robustus (Scudder 1862) 1995 North America Dowell & Gill 1989 Phaneroptera nana Fieber 1853 <1925 Europe Strohecker 1952 Phaneroptera quadripunctata Brunner 1878 1949 Europe Strohecker 1952, Cantrall 1972 Platycleis tesselata (Charpentier 1825) 1926 Europe CDFA 1960 49:143, Rentz 1963 Phasmida-Phasmatidae Carausius morosus (Sinety 1901) North America CPPDR 2001 20:11 yes Phthiraptera-Boopidae Heterodoxus longitarsus (Piaget 1880) 1894 Australia Essig 1926a, 1958 Heterodoxus spiniger (Enderlein 1909) 1945 Amett 1985 Phthiraptera-Gyropidae Gyropus oval is Burmeister 1838 1976 Emerson 1972, Essig 1926a Gliricola porcelli (Schrank 1781) Emerson 1972, Essig 1926a Phthiraptera-Haematopinidae 1973 Haematopinus asini (Linnaeus 1758) 1959 Kim et al. 1986, Essig 1926a yes Haematopinus eurysternus Nitzsch 1818 <1925 Kim et al. 1986, Essig 1926a yes Haematopinus suis (Linnaeus 1758) 1959 Kim et al. 1986, Essig 1926a yes Phthiraptera-Hoplopleuridae Hoplopleura captiosa Johnson 1960 2005 Kim et al. 1986 Hoplopleura pacifica Ewing 1924 1924 Kim et al. 1986 Phthiraptera-Linognathidae 1940 Linognathus africanus Kellogg & Paine 1911 Kim et al. 1986 Linognathus setosus (Olfers 1816) Kim et al. 1986, Essig 1926a Linognathus stenopsis (Burmeister 1838) Kim et al. 1986, Essig 1926a Linognathus vituli (Linnaeus 1758) Kim et al. 1986, Essig 1926a Phthiraptera-Menoponidae <1925 Menacanthus stramineus (Nitzsch 1818) Essig 1926a Menopon gallinaeLirmaeus 1758 Emerson 1972, Essig 1926a Phthiraptera-Pediculidae Pediculus humanus Linnaeus 1758 Kim et al. 1986 134 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 5 Phthiraptera-Philopteridae 1996 Campanulotes bidendatus Scopoli 1763 1968 Essig 1926a, 1958 Goniocotes chrysocephalus Giebel 1874 1925 Emerson 1972 Goniocotes creber Kellogg 1896 1909 Essig 1958 Goniocotes gigas Taschenberg 1879 <1915 Essig 1926a, 1958 Goniocotes gaUinae (De Geer 1778) 1997 Emerson 1972 Goniocotes maculatus Taschenberg 1882 <1915 Emerson 1972 Goniocotes parviceps (Piaget 1880) <1925 Emerson 1972 Goniocotes rectangulatus Nitzsch 1866 1995 Emerson 1972 Lipeurus caponis (Linnaeus 1758) 2004 Emerson 1972, Essig 1926a Lipeurus maculosus Clay 1938 Emerson 1972 Lipeurus numidae (Denny 1842) 1943 Emerson 1972 Lipeurus pavo Clay 1938 Emerson 1972 Phthiraptera-Pthiridae 1946 Pthirius pubis (Linnaeus 1758) Kim et al. 1986 Phthiraptera-Polyplacidae Polyplax spinulosa (Burmeister 1839) <1925 Kim et al. 1986, Essig 1926a Phthiraptera-Trichodectidae 1999 Bovicola bovis (Linnaeus 1758) 1907 Emerson 1972, Essig 1926a yes Trichodectes equi (Denny 1842) 1972 Emerson 1972, Essig 1926a yes Trichodectes canis (De Geer 1778) 1931 Essig 1926a, 1958 Phthiraptera-Trimenoponidae 1996 Trimenopon hispidum (Burmister 1838) 1904 Amett 1985 Pscoptera-Ectopsocidae Ctopsocus strauchi Enderlein 1906 1949 BugGuide 2011 Pscoptera-Elipsocidae 1985 Cuneopalpus cyanops (Rostock 1876) Europe Mockford 1993 Propsocus pulchripennis Perkins 1899 <1915 Mockford 1993 Psocoptera-Lachesillidae Lachesilla pedicularia (Linnaeus 1758) Mockford 1993 Psocoptera-Liposcelididae Liposcelis corrodens Heymons 1909 Mockford 1993 Liposcelis decolor (Pearman 1925) Mockford 1993 Liposcelis bostiychophilus Badonnel 1931 Powell & Hogue 1979 Liposcelis rufa Broadhead 1950 Europe Mockford 1993 Psocoptera-Peripscoidae Peripsocus reductus Badonnel 1943 <1925 Mockford 1993 Psocoptera-Psyllipsocidae <1925 Psyllipsocus ramburii Selys-Longchamps 1872 Mockford 1993 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 135 Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2010. Taxon Year detected Probable origin Reference Listed as a pest* 1 Psocoptera-Stenopsocidae Graphopsocus cruciatus (Linnaeus 1768) Mockford 1993 Psocoptera-Trichopsocidae Trichopsocus clarus (Banks 1908) BugGuide 2011 Psocoptera-Trogiidae Cerobasis annulata (Hagen 1865) Europe Mockford 1993 Lepinotus inquilinus Heyden 1850 Mockford 1993 Lepinotus patruelis Pearman 1931 Europe CDFA 1942 31:176 Lepinotus reticulatus Enderlein 1905 Mockford 1993 Siphonaptera-Ceratophyllidae Nosopsyllus fasciatus (Bose 1800) Hubbard 1968, Lewis et al. 1988, Essig 1926a Nosopsyllus londiniensis (Rothschild 1903) Lewis et al. 1988 Siphonaptera-Leptopsyllidae Leptopsylla segnis (Schonherr 1811) Hubbard 1968, Lewis et al. 1988, Essig 1926a Siphonaptera-Pulicidae Ctenocephalides canis (Curtis 1826) Hubbard 1968, Essig 1926a yes Ctenocephalides felis (Bouche 1835) Hubbard 1968, Lewis et al. 1988, Essig 1926a yes Echidnophaga gallinacea (Westwood 1875) Lewis et al. 1988, Essig 1926a yes Pul ex irritans Linnaeus 1758 Tropical America Lewis et al. 1988, Essig 1926a Xenopsylla cheopis Rothschild 1903 Europe Hubbard 1968, Lewis et al. 1988 Symphyla-Scolopendrellidae Symphylella isabellae (Grassi 1886) Europe Hilton 1931 Thysanura-Lepismatidae Ctenolepisma ciliata (Dufour 1831) Wygodzinsky 1972 Ctenolepisma lineata (Fabricius 1775) Wygodzinsky 1972 Ctenolepisma longicaudata Escherich 1905 Wygodzinsky 1972 Lepisma saccharina Linnaeus 1758 Wygodzinsky 1972, Essig 1926a yes Thermobia domestica (Packard 1873) Wygodzinsky 1972, Essig 1926a yes 136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Addendum to Appendix. Exotic terrestrial macro-invertebrates found breeding in California 1700-2016 Taxon Year detected Probable origin Reference Listed as a pest Araneae-Corinnidae Falconina gracilis (Keyserling 1891) 2013 Tropical America Valle et al. 2013 Araneae-Theridiidae Steatoda nobilis (Thorell 1875) 2012 Europe Vetter and Rust 2012 Coleoptera-Anthicidae Omonadusformicarius (Goeze 1777) North America Majka 2011, California beetle project 2016 Coleoptera-Buprestidae Chiysobothris analis LeConte 1860 2012 North America Westcott et al. 2015 Coleoptera-Carabidae Amara apricaria (Paykull 1790) Europe Sokolov and Kavanaugh 2014 Amara familiaris (Duftschmid 1812) North America Bosquet 2012 Harpalus affinis (Schrank 1781) Europe Sokolov and Kavanaugh 2014 Pterostrichus melanarius (llliger 1798) North America Bosquet 2012 Coleoptera-Chrysomelidae Microtheca ochrolma Stal 1860 2011 North America Gilbert et al. 2011 Oulema melanopus (Linneaus 1758) 2015 North America Dowell and Pickett (in prep) yes Coleoptera-Curculionidae Euwallacea n. sp. #2 2013 Anon. 2016 Coleoptera-Dermestidae Thylodrias contractus Motshulsky 1839 California beetle project 2016 Coleoptera-Erotylidae Dacne picta Crotch 1873 1990 Asia Savaiy 1995 Coleoptera-Hyborosoridae Hybosorus illigeri Reiche 1853 California beetle project 2016, Ocampo 2002. Coleoptera-Hydrophilidae Cercyon atricapillus (Marsham 1802) Smetana 1988, California beetle project 2016 Cercyon haemorrhoidales (Fabricius 1775) Smetana 1988, California beetle project 2016 Cercyon lateralis (Marsham 1802) Smetana 1988, California beetle project 2016 Cercyon pygmaeus (llliger 1801) Smetana 1988, California beetle project 2016 Cercyon quisquilus (Linneaus 1761) Smetana 1988, California beetle project 2016 Cercyon terminal is (Marsham 1802) Smetana 1988, California beetle project 2016 Coloeoptera-Latridiidae Adistema watsoni (Wollaston 1871) California beetle project 2016 Cartodere notifer (Westwood 1839) Majka et al. 2009a Corticaria ferruginea Marsham 1802 Majka et al. 2009a DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 137 Addendum to Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2016 Taxon Year detected Probable origin Reference Listed as a pest Coleoptera-Staphylinidae L inotylustetra carinatus (Block 1799) California beetle project 2016, Klimaszewski et al. 2012. Bisnius cephalotes (Gravenhorst 1802) California beetle project 2016, Majka et al. 2006 Crataraea suturalis (Mannerheim 1830) California beetle project 2016, Klimaszewski et al.2012 Lithocharis ochracea (Gravenhorst 1802) California beetle project 2016, Hamilton 1889 Myrmecocephalus concinnus (Erichson 1840) California beetle project 2016, Klimsezewski et al. 2012 Nehemitropia lividipennis (Mannerheim 1831) California beetle project 2016, Klimsezewski et al. 2012 Philonthus carbonarius (Gravenhorst 1802) California beetle project 2016. Majka et al. 2006. Philonthus varians (Paykull 1789) California beetle project 2016, Staniec 2002 Quedius fulgidus (Fabricius 1792) Majka et al. 2009b Tasgius ater (Gravenhorst 1802) California beetle project 2016, Brunlce et al. 2011 Coleoptera-Tenebrionidae Alphitobius laevigatus (Fabricius 1781) California beetle project 2016, Bosquet 1990. Latheticus oryzae (Waterhouse 1880) California beetle project 2016, Wittenberg et al. 2005 Diptera-Agromyzidae Liriomyza huidobrensis (Blanchard 1926) Tropical America Steck 2006 Napomyzae vanescens (Walker 1853) Spencer 1981 Phytomyza vomitoriae Kulp 1968 Fonsdale and Scheffer 2011 Diptera-Cecidomyiidae Monarthropalpus buxi, (Geoffroy 1764) Anon. 1985, 2014 Diptera-Culicidae Aedes albopictus (Skuse 1849) Anon. 2015 yes Aedes quinquefasciatus Say 1823 Juliano and Founibos 2005, Cornell et al. 2003 Diptera-Dolichopodidae Medetera johnthomasi Bickel & Amaud 2011 2011 Bickel and Amaud 2011 Diptera-Muscidae Athergonia orientalis (Schiner 1868) Cole 1969, Hancock et al. 2014 Athergonia reversura Villeneuve 1963 Bryant 2016, Hancock et al. 2014 Diptera-Drosophilidae Drosophila gentica Wheeler & Takada 1962. 2014 TropicalAmerica Grimaldi et al. 2015 Drosophila flavohirta Malloch 1924 2014 Australia Grimaldi et al. 2015 Drosophila melanogaster Meigen 1830 Baudy et al. 2004, Varela et al. 2015 Diptera-Tephritidae Euaresta bullans Weidemann 1830 Foote and Blanc 1963 138 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Addendum to Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2016 Taxon Year detected Probable origin Reference Listed as a pest Hemiptera-Adgelidae Pineus boerneri Annand, 1928 Lazzari and Cardoso 2011 Hemiptera-Aleyrodidae Aleurotulus nephrolepidis (Quiantance, 1900) Anon. 2006 Hemiptera-Aphididae Anoecia corni (Fabricius, 1775) Foottit et al. 2006, Swain 1919 Aphis eurphorbiae Kaltenbach, 1933 1933 Foottit et al. 2006 Aphis polygonata (Nevsky, 1929) Foottit et al. 2006, Blackman and Eastop 2007 Aphis spiraecola Patch, 1914 Foottit et al. 2006, Reuther et al. 1978 Aphis thalictri Koch, 1854 Foottit et al. 2006, Swain 1919 Asiphonella dactylonii Theobald 1923 Foottit et al. 2006, Anon. 1969 Atarsaphis agrifoliae (Ferris, 1921) 1921 Foottit et al. 2006 Brachycaudus helichiysi (Kaltenbach, 1843) Foottit et al. 2006. Bentley et al. 2009 Capitophorus elaeagni (Del Guercio, 1894) Foottit et al. 2006, Bari and Natwick 2015 Ctenocallis zelkowae (Shinji, 1924) 1983 Foottit et al. 2006 Greenidea psidii van der Goot, 1916 1998 Foottit et al. 2006 Liosomaphis berberidis (Kaltenbach, 1843) Foottit et al. 2006, Swain 1919 Metolophium dirhodum (Walker, 1849) Foottit et al. 2006, Glidow and Rochow 1983 Monaphis antennata (Kaltenbach, 1843) Foottit et al. 2006, Essig 1911 Panaphis juglandis (Geoze, 1778) 1952 Foottit et al. 2006, Olson 1974 Sitobion avenae (Fabricius, 1775) Foottit et al. 2006, Summers and Godfrey 2009 Therioaphis maculata (Buckton, 1899) Foottit et al. 2006, Smith 1959 Tnberolachnus salignus (Gmelin, 1790) Stary and Schlinger 1967 Hemiptera-Cicadellidae Alebra wahlbergi Boheman, 1845 1945 Hamiliton 2010, 2011 Aphnodes bicinctus (Schrank, 1776) Flamiliton 2011, Sherf and MacNab 1986 Dalbulus maidis DeLong, 1923 Hamiliton 2011, Summers et al. 2004 Empoascas maragdula Fallen, 1806 1916 Hamiliton 2011, van Duzee 1917 Kvboasca bipunctata (Oshanin 1871) Flock 1950 Sophonia orientalis (Matsumura, 1912) Hamiliton 2011, Bryant et al. 2016 Hemiptera-Miridae Brachynotocoris puncticornis Reuter, 1880 2013 Europe Dowell (in prep) Closterotomus norwegicus (Gmelin, 1790) 1966 Scudder and Footitt 2006 Coridromius chenopoderis Tatamic and Cassis 2008 Australia Tatamic and Cassis 2008. DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 139 Addendum to Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2016 Taxon Year detected Probable origin Reference Listed as a pest Heteropsallus planicornis (Pallas 1772) Wheeler and Flenry 1992 Lepidargyrus an cor ifer (Fieber 1858) 1912 Scudder and Footitt 2006 Megalonotus sabulicola (Thompson 1870) 1921 Scudder and Footitt 2006 Hemiptera-Psyllidae Psylla buxi Linneaus, 1758 Tuthill 1943 Psyllopsis fraxinicola Foerster 1848 Tuthill 1943 Hymenoptera-Agaonidae Eupristina altissima Saunders 1882 Boucek 1997 Hymenoptera-Apidae Xylocopa appendiculata Smith 1852 2012 Asia Dahlberg et al. 2013 Hymenoptera-F ormicidae Phiedole megacephala (Fabricius 1793) 2014 Leathers 2014 Lepidoptera-Crambidae Terastia meticulosalis Guenee & Agathodes 1854 2015 North America Hodel et al. 2016 Lepidoptera-Gracillaridae Phyllonorycter mespilella (Hubner 1805) 1936 Europe Maier 2001 Lepidoptera: Pyralidae Diaphania hyalinata Linneaus 1767 Capinera 2014, California moth database 2016 Lepidoptera-Yponomeutidae Ocnerostoma piniariella Zeller 1847 California moth database 2016, Felt 1922. Siphonaptera-Pulicidae Hectopsylla psittaci Von Frauenfeld 1860 TropicalAmerica Schwann et al. 1983 Thysanoptera-Aeolothripidae Aeolothrips collaris Priesner 1919 Europe CPPDR 16:44 Aeolothrips ericae Bagnall 1920 Europe CPPDR 14:6 Aeolothrips melaleucus (Flaliday 1852) Bailey 1957 Anaphothrips obscurus Comstock 1875 Hoddle & Mound 2012 Aptinothrips rufus Haliday 1836 Hoddle & Mound 2012 Rhipidothrips gratiosus Uzel 1895 Europe Bailey 1957 Thysanoptera-Phlaeothripidae Amynothrips andersoni O'Neill 1968 Hoddle & Mound 2012 Androthrips ramachandrai Kamy 1926 Hoddle & Mound 2012 Cartomothrips browni Stannard 1962 Hoddle & Mound 2012 Cephalothrips monilicornis O.M. Reuter 1880 Europe Cott 1956 Gynaikothrips ficorum (Marchal 1908) Mexico Clausen 1978 yes Gyaikothrips uzeli Zimmerman 1900 Hoddle & Mound 2012 Haplothrips leucanthemi (Schrank 1781) Cott 1956, Essig 1926a Plaplothrips robustus Bagnall 1918 Hoddle & Mound 2012 Haplothrips verbasci (Osborn 1897) Keifer et al. 1982 Liothrips floridensis Watson 1913 Hoddle & Mound 2012 140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 3 Addendum to Appendix (continued). Exotic terrestrial macro-invertebrates found breeding in California 1700-2016 Liothrips vaneeckei Priesner 1920 Europe CDFA 28:479 Macrophthalmothrips argus (Kamy, 1920) Hoddle & Mound 2012 Nesothrips propinquus (Bagnall, 1916) Hoddle & Mound 2012 Neurothrips magnafemoralis (Hinds 1902) CDFA 28:538 Thysanoptera-Thripidae Anaphothrips obscurus (Muller 1776) Hoddle et al. 2004 Aptinothrips rufus (Haliday) 1836 Europe Essig 1926a Aptinothrips stylifer Tryborn, 1894 Hoddle & Mound 2012 Asprothrips seminigricornis (Girault 1926) Hoddle et al. 2004 Chaetanaphothrips orchidii (Moulton 1907) Hoddle et al. 2004 Chirothrips aculeatus Bagnell 1927 Bailey 1957 Chirothrips manicatus (Haliday 1836) Hoddle et al. 2004 Danothrips trifasciatus Sakimura 1975 Pacific Region CPPDR 15:73 Dendrothrips howei Mound 1999 2011 North America Hoddle & Mound 2012 Dendrothrips ornatus (Jablonowski 1894) North America Dowell & Gill 1989 Drepanothrips reuteri Uzel 1895 Europe CDFA 17:455 Frankliniella hemerocallis Crawford 1948 North America CPPDR 16:44 Frankliniella schultzei (Trybom 1910) Hoddle & Mound 2012 Frankliniella tenuicornis (Uzel 1895) Hoddle & Mound 2012 Heliothrips haemorrhoidalis (Bouche 1833) Tropical America CDFA 17:366 yes Hercinothrips bicinctus (Bagnall 1919) Hoddle & Mound 2012 Hercinothrips femoralis (Reuter 1891) Bailey 1957, Essig 1926a Limothrips angulicornis Jablonowski 1894 Europe Bailey 1957 Limothrips cerealium (Haliday 1836) Bailey 1957 Leucothrips nigripennis Reuter 1904 Hoddle & Mound 2012 Microcephalothrips abdominalis (Crawford 1910) Hoddle et al. 2004 Monilothrips kempi Moulton 1929 Bailey 1957 Neohvdatothrips burungae (Hood 1935) Tropical America CPPDR 22:21-22 Neohydatothrips samayunkur (Kudo 1995) North America CPPDR 14:28 Odontothrips loti (Haliday 1852) Hoddle et al. 2004 Parthenothrips dracaenae (Heeger 1854) Bailey 1957, Essig 1926a Pseudanaphothrips achaetus (Bagnall) 1916 Australia CPPDR 19:25 Pseudodendrothrips mori (Niwa, 1908) Hoddle & Mound 2012 Scirtothrips citri (Moulton 1909) Asia CDFA 7:465 yes Scirtothrips inermis Priesner 1933 Europe Dowell & Gill 1989 Scirtothrips longipennis (Bagnall 1909) CDFA 20:345 Scirtothrips perseae Nakahara 1997 1996 Tropical America CPPDR 15:4 yes Taeniothrips inconsequens (Uzel 1895) Asia Davidson & Lyon 1987, Bailey 1957 yes Tenothrips frici (Uzel 1895) Hoddle et al. 2004 Thrips australis (Bagnall 1915) Australia Bailey 1957 Thrips hawaiiensis (Morgan 1913) Hawaii Dowell & Gill 1989 Thrips nigropilosus Uzel 1895 Gentile & Bailey 1968 Thrips tabaci Lindeman 1888 North America CDFA 4:56, Thrips trehernei Priesner 1927 Hoddle et al. 2004 Thrips simplex (Morison 1930) Hoddle & Mound 2012 Thrips vulgatissimus Haliday 1836 Hoddle et al. 2004 Trichromothrips cyperaceae (Bianchi 1945) Asia Hoddle et al. 2004 Trichromothrips xanthius (Williams 1917) Hoddle et al. 2004 DOWELL ET AL.: MACRO-INVERTEBRATE INVADERS IN CALIFORNIA 141 Literature Cited Appendix AAlbu, R.L.K. 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Pan-Pacific Entomologist 72:27-30. 158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63 Page intentionally left blank PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 4, pp. 159-162, 1 fig. April 29, 2016 Tetramerium vargasiae (Acanthaceae), a New Species from the Basin of the Rio Balsas in Guerrero, Mexico Thomas F. Daniel 1 and Ramiro Cruz Duran 2 1 Department of Botany, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118, U.S.A.; Email: tdaniel@calacademy.org. 2 Departamento de Biologia Comparada, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico, Apartado postal 70-181, Delegacion Coyoacan, 04510 Mexico, D.F., Mexico; Email: ramcrudur@yahoo.com Tetramerium vargasiae is newly described and illustrated from the arid portion of the basin of the Rio Balsas in Guerrero, Mexico. It is distinguished from its congeners by the combination of the following characters: shrubby habit, linear leaves, pubescent (eglandular trichomes only) and entire bracts that are 16 to 19 mm long, five-lobed calyx, pseudopapilionaceous and white (with purple markings) corollas, and glabrous capsules that are seven mm long. It is compared to the superficially similar T. tetramerioides from Oaxaca, Mexico. Keywords: Tetramerium, Guerrero, Mexico, endemic species, Acanthaceae Se describe e ilustra Tetramerium vargasiae como especie nueva de la cuenca arida del Rio Balsas en Guerrero, Mexico. Se distingue de sus congeneres por la combi- nacion de los siguientes caracteres: habito arbustivo, hojas lineares, bracteas pubes- centes (solo tricomas eglandulares) y enteras de 16 a 19 mm de largo, caliz con cinco lobulos, corolas pseudopapilionaceas y blancas (con marcas de color purpura) y cap- sulas glabras siete mm de largo. Se compara la especie nueva con T. tetramerioides , una especie superficialmente similar del estado de Oaxaca, Mexico. Tetramerium Nees consists of 29 currently recognized species (Daniel 1986, 2003) that occur from the United States to Bolivia. Most species (22) occur in Mexico, and 19 of them are endem¬ ic there. Arid and semi-arid regions in western and southern Mexico, from Jalisco to the Isthmus of Tehuantepec in Oaxaca, are especially rich in species of Tetramerium with at least 17 occurring there and 10 endemic to the region (Daniel 1986). Another distinctive species from this region was collected in the arid portion of the basin of the Rio Balsas in Guerrero. It is described below. Tetramerium vargasiae T.F. Daniel & Cruz Duran, sp. nov. Type. Mexico: Guerrero: Mpio. Huitzuco de los Figueroa, 0.7 km NE de San Francisco Ozomatlan [ca. 17°56'0.63"N, 099°19'37.41"W], 650 m, bosque tropical caducifolio, 30-III-1990 (fir, frt), A. Vargas P. 288 (holotype: FCME!). Tetramerium vargasiae differs from congeners by the combination of its shrubby habit, linear leaves, pubescent (eglandular trichomes only) and entire bracts that are 16-19 mm long, 5-lobed calyx, pseudopapilionaceous and white (with purple markings) corollas, and glabrous capsules that are 7 mm long. 159 160 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 4 Shrubs to 1 m tall; older stems with bark exfoliating in strips; young stems ± terete, ± evenly pubescent with antrorse to antrorsely appressed eglandular trichomes 0.05-0.2 mm long, nodes more densely pubescent. Leaves (immature; plants mostly leafless during anthesis) subsessile (petioles to 2 mm long), blades linear, sometimes ± conduplicate, 10-17 mm long, 0.6-0.9 mm wide, length:width = 16.7-18.9, both surfaces and margin covered with antrorsely appressed eglandular trichomes 0.05-0.2 mm long, only midvein evident. Inflorescence of terminal (some¬ times terminating short axillary branches) and congested spikes to 46 mm long, 14-19 mm across (measured flat) near midspike, rachis densely and evenly pubescent with flexuous to antrorse to antrorsely appressed eglandular trichomes to 0.5 mm long, median internodes 6 mm long. Bracts ± coriaceous, lanceolate to lance-elliptic, 16-19 mm long, 1.8-3.2 mm wide, obscurely to ± promi¬ nently and palmately 3-veined, apically straight, erect, and mucronate, mucro 0.5-0.6 mm long, abaxial surface densely pubescent with antrorse to antrorsely appressed eglandular trichomes to 0.4 mm long, margin densely ciliate with spreading-flexuous, silky eglandular trichomes to 2.5 mm long (at least some trichomes > 1 mm long). Bracteoles lance-subulate, 9-14 mm long, 0.8-1.1 mm wide, only midvein evident (sometimes obscure), abaxial surface and margin pubescent like bracts. Flowers sessile. Calyx 5-lobed, 6.5-8.5 mm long, lobes lance-subulate to subulate, 6-7 mm long, 0.4-0.5 mmm wide, abaxial surface and margin pubescent like bracteoles. Corolla pseudopapil- ionaceous, white with purple markings on upper lip, 20 mm long, externally glabrous, tube 10.5 mm long, upper lip obovate, 9.5 mm long, 2.5 mm wide, lower lip 9 mm long, lower-central lobe conduplicate, 7 mm long, 4.6 mm wide, lateral lobes obovate, 7 mm long, 3.5 mm wide. Stamens ca. 7 mm long, thecae ca. 1.2 mm long. Capsules 7 mm long, glabrous, stipe 2.5 mm long, head 4.5 mm long. Seeds not seen. Phenology. — Flowering and fruiting: March. Distribution and habitat.— Endemic to Mexico where it is known only from the arid por¬ tion of the basin of the Rio Balsas of north-central Guerrero in a region of tropical deciduous for¬ est at 650 m elevation. Conservation Assessment. — Tetramerium vargasiae is known from a single collection from a population that was not observed during this study. The collector of the type indicated that only a few plants were observed at that site. Given the lack of additional information, this species is assessed as Data Deficient (DD) according to IUCN guidelines (IUCN 2014). Although Tetramerium vargasiae is incompletely known, it is highly distinctive. It pertains to Daniel’s (1986) sect. Tetramerium and resembles T. tetramerioides (Lindau) T.F. Daniel from southeastern Oaxaca (previously known only from the type, but recently rediscovered in the vicin¬ ity of the type collection) in several characteristics, including: shape of the bracts and bracteoles, marginal pubescence of the bracts, number of calyx lobes, color and shape of the corolla, and size and surface vesture of the capsule. These two species can be distinguished by the characters noted in the following key: la. Young stems ± evenly pubescent with antrorse to antrorsely appressed eglandular trichomes 0.05-0.2 mm long; leaves (immature) subsessile (petioles to 2 mm long), blades linear, length:width = 16.7-18.9; spikes 14-19 mm wide (measured flat) near midspike, rachis pubes¬ cent with eglandular trichomes; bracts 16-19 mm long, abaxial surface densely pubescent with antrorse to antrorsely appressed eglandular trichomes to 0.4 mm long; bracteoles 9-14 mm long; calyx 6.5-8.5 mm long; corolla 20 mm long; Guerrero. T. vargasiae lb. Young stems ± bifariously pubescent with an understory of glandular trichomes 0.05-0.2 mm long and an overstory (absent on some intemodes) of flexuose eglandular trichomes 0.2-0.7 mm long; leaves subsessile to petiolate (petioles to 15 mm long), blades lance-ovate, DANIEL & CRUZ DURAN: NEW SPECIES OF TETRAMERIUM FROM MEXICO 161 Figure 1 . Tetramerium vargasiae (from the holotype). A. Stem with exfoliating epidermis and young leaves. B. Branch with terminal inflorescence. C. Capsule. D. Young leaf. E. Branch with young leaves. F. Bracteole. G. Bract. H. Detail of bract showing trichomes on margin. I. Flower. J. Detail of calyx lobe showing trichomes. Drawn by Ramiro Cruz Duran. 162 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 4 length:width = 2.2-3.8; spikes 3-7 mm wide (measured flat) near midspike, rachis pubescent with glandular and eglandular trichomes; bracts 7-10 mm long, abaxial surface pubescent with an inconspicuous understory of glandular trichomes 0.05-0.2 mm long and a sparse overstory (especially along the midvein) of flexuose eglandular trichomes to 1.5 mm long; bracteoles 6-8 mm long; calyx 3.2^4 mm long; corolla 14-15 mm long; Oaxaca. T. tetramerioides This species is named in honor of the collector of the type, Andira Vargas Perez, who collect¬ ed plants for FCME between 1989 and 1990 during a study of the flora and vegetation in the east¬ ern part of the basin of the Rio Basas in Guerrero. Acknowledgments We thank Jose Antonio Hernandez Gomez (Facultad de Ciencias, UN AM) for digitizing the illustration and the curators of the following herbaria for making specimens available: CAS, FCME, and MEXU. We also thank two reviewers, Drs. Christine Anderson and Victor Steinmann, for their helpful comments. Literature Cited Daniel, T.F. 1986. Systematics of Tetramerium (Acanthaceae). Systematic Botany Monographs 12:1-134. Daniel, T.F. 2003. New and reconsidered Mexican Acanthaceae X. Flora del Bajio region. Novon 13:37-48. IUCN. 2014. Guidelines for Using the IUCN Red List Categories and Criteria, Version 11. Standards and Peti¬ tions Subcommittee of the IUCN Species Survival Commission, [accessed 16 November 2015]. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5, pp. 163-189, 4 figs. April 29, 2016 Avicennia (Acanthaceae: Avicennioideae) in North America and Mesoamerica Thomas F. Daniel Department of Botany, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118 U.S.A.; Email: tdaniel@calacademy.org A taxonomic revision of Avicennia, an acanthaceous genus of mangroves, in North America and Mesoamerica recognizes three species, two native (A. bicolor and A. germinans) and one locally naturalized in southern California (A. marina subsp. australasica ). Herbarium specimens from throughout the region were used to for¬ mulate descriptions of each species, obtain distributional and ecological data, and as a source for pollen in order to characterize and illustrate palynological features of each species. A distribution map, an illustration of the most common species (A. ger¬ minans), and photos showing diagnostic attributes of each species are provided. Una revision taxonomica de Avicennia, un genero de los manglares en la familia Acanthaceae, en Norteamerica y Mesoamerica reconoce tres especies, dos nativas (A. bicolor y A. germinans) y una naturalizada localmente en el sur de California (A. marina subsp. australasica). Los ejemplares de herbario de toda la region se uti- lizaron para formular descripciones de cada especie, obtener datos distributivos y ecologicos, y como fuente de polen con el fin de caracterizar e ilustrar caracteres palinologicos de cada especie. Ademas, se ofrecen un mapa de distribution, una ilus- tracion de la especie mas comune (A. germinans), y fotos que muestran los atributos diagnostics de cada especie. Avicennia has been treated previously in several families, most prominent among them Verbe- naceae and Avicenniaceae. Molecular phylogenetic analyses (Schwarzbach and McDade 2002; Borg et al. 2008; McDade et al. 2008) and studies of floral structure and development (Borg and Schonenberger 2011) reveal the genus to be monophyletic, nested among Acanthaceae s.l., and sis¬ ter to either subfamily Acanthoideae or more likely to subfamily Thunbergioideae. The genus is here treated as subfamily Avicennioideae. Although its taxonomic relationships have been contro¬ versial, Avicennia is well established as a genus of mangrove species. Indeed, they are usually treat¬ ed in an ecological sense among the so-called “true” mangroves (Tomlinson 1986), which taxa gen¬ erally occur exclusively in mangrove communities and exhibit both morphological and physiolog¬ ical specializations to the highly saline substrate. It is likely that morphological adaptations to a highly specialized environment have obscured its familial affinities. Other species of Acanthaceae in different tribes of subfamily Acanthoideae are considered to be true mangroves (e.g., Acanthus ebracteatus Vahl and A. ilicifolius L. of Acantheae fide Wang et al. 2011; but treated as mangrove associates by Tomlinson 1986) or mangrove associates {Bravaisia berlandieriana (Nees) T.F. Daniel and B. integerrima (Spreng.) Standi, of Ruellieae). The economic importance of mangrove plants in general, and species of Avicennia in particu¬ lar, was correctly pointed out by Moldenke (1960: 144) in citing and refuting another author’s statement that plants of Avicennia are of little economic importance; Moldenke stated, “...mem- 163 164 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 bers of this genus are of tremendous economic importance to man and his economy because of their constant battle with the sea and their great success in extending and eventually building up the sur¬ face of the land adjacent to the sea in subtropical and tropical climes.” Indeed, as discussed by numerous authors (e.g., Tomlinson 1986; Odum and Mclvor 1990; Costanza et al. 1997; Chivian and Bernstein 2008; Polidoro et al. 2010; and Cavanaugh et al. 2014) important ecological servic¬ es provided by mangroves include: serving as home and/or nurseries for numerous marine and ter¬ restrial organisms (e.g., fish, mammals, birds, crustaceans, worms, and insects), protecting coast¬ lines from storm surges (e.g., buffering wave action, preventing flooding and soil erosion, and decreasing saltwater invasion), waste treatment (e.g., nutrient recycling and pollution control), car¬ bon sequestration, and recreation (e.g., eco-tourism and sport fishing). The value of ecosystem services provided by mangrove communities worldwide has been estimated to be at least US $1.6 billion annually (Polidoro et al. 2010). The ecological and economic importance of mangroves, among which several species of Avicennia are noteworthy by their abundance and widespread dis¬ tribution, will only increase with both growth of human populations in coastal areas and sea-level changes associated with climate alterations due to global warming. Estimates for recent degradation and loss of mangrove vegetation worldwide is significant. Between 20 and 35 percent of mangrove habitats have been lost since about 1980, and the current rate of annual disappearance of this vegetation is estimated to be between one and eight percent (Polidoro et al. 2010). The loss of mangrove communities in Mexico has been estimated as from 15,000 sq. km. in the 1970s to ca. 5,000 sq. km. by the late 1990s (Spalding et al. 1997). Major anthropogenic threats to mangrove communities include coastal development (e.g., tourist/resident infrastructure, mariculture, agriculture, and conversion for harbor/industrial uses), pollution (e.g., sewage effluents, oil spills, and agricultural/industrial/urban runoff), fresh water diversions, and sea level changes due to global warming (e.g., Cintron M. and Schaeffer N. 1992). For the North American and Neotropical regions, distribution models under future climate scenarios (e.g., Ceron S. et al. 2015) predict a pole-ward shift for mangroves (including Avicennia germinans ), an over¬ all contraction of species distributions, and a decline in species richness. This regional taxonomic study used data from more than 650 specimens from North America and Mesoamerica (southern Mexico and Central America) in 22 herbaria to formulate descriptions of species, reproductive phenological periods, distributions, habitats, and other pertinent informa¬ tion. In the few instances where only an image of a specimen was studied, this is noted by “-image!” Pollen from each species was examined with scanning electron microscopy and charac¬ terized using terminology of Walker and Doyle (1975). Local names noted herein were obtained from herbarium specimens studied. Many additional local names and uses for these species can be found among references in the literature cited. Avicennia L., Sp. PI. 1: 110. 1753; Gen. PI. ed. 5, 49. 1754. Type .— Avicennia officinalis L. Bontia L., Sp. PI. 2: 638 (“938”). 1753. Type. — Bontia daphnoides L. Upata Adanson, Fam. 2: 201. 1763. = Avicennia L. Sceura Forssk., FI. Aegypt.-Arab. 37. 1775. Type.— Sceura marina Forssk. Halodendrum Thouars, Gen. Nov. Madagasc. 8. 1806. Type.— Halodendrum thouarsii Roem. & Schult. Hilairanthus Tiegh., J. Bot. (Morot) 12: 358. 1898. Type. — Not designated (two species cited). Shrubs or trees lacking cystoliths and with erect, aerial, and sometimes branched roots (pneu- matophores) up to 4 dm tall under and beyond canopy (and infrequently also with aerial stilt roots), cystoliths absent. Leaves opposite, leathery, entire, margin flat to revolute. Inflorescence of axil¬ lary and/or terminal dense (sometimes subcapitate) pedunculate dichasiate spikes or panicles of DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 165 dichasiate spikes; dichasia opposite, 1-flowered, sessile. Bracts ± leathery, concavoconvex. Bracte- oles concavoconvex. Flowers sessile, protandrous. Calyx 5-lobed, lobes ± free, concavoconvex, imbricate, equal to subequal in size. Corollas white to yellow to orange, tube expanded distally, shorter than limb, limb 2-labiate or actinomorphic, 4- or 5-lobed, lobes erect to reflexed, oblong to obovate, contorted (left contort aestivation) in bud. Stamens 4, at least 2 exserted from mouth of corolla (in ours), oriented in pairs below upper lip of corolla or oriented symmetrically around corolla with each equally distant from the others; anthers 2-thecous, thecae of a pair ± equally inserted, parallel, ± equal in size, lacking basal appendages, each dehiscing toward lower lip (i.e., flower nototribic) or toward each other (i.e., flower pleurotribic) by a longitudinal slit; pollen sub- oblate to euprolate, 3-colporate, exine reticulate; staminodes 0. Style not evident or elongating with age of flower; stigma 2-lobed, lobes slightly unequal. Capsule podlike, leathery, ± ellipsoid to ovoid (often asymmetric), ± compressed, unilocular, retinacula absent, dehiscence not explosive, usually occurring at time of detachment or soon after being shed. Seed 1 (-2) per capsule, filling fruit and consisting mostly of ripe embryo with 2 conspicuous folded cotyledons and a pubescent radicle (cryptoviviparous). Moldenke (1960, 1973) and Duke (1991) listed additional generic synonyms. The generic name is derived from Ibn Sina (=Avicenna), 980-1037, a Persian physician, philosopher, and naturalist. At least eight (e.g., Duke 1991; Duke et al. 1998) species of Avicennia are recognized in mar¬ itime regions of the tropics and subtropics worldwide. The genus is monophyletic, and includes some of the most prominent and widely distributed “true” mangroves (cf. Tomlinson 1986). Plants occur along shorelines and in tidally influenced waterways in two mostly tropical regions: Atlantic/Caribbean/eastern Pacific (3 spp.) and Indo-western Pacific (5 spp.). Although elevations up to 150 meters have been noted on herbarium labels of American plants, occurrences at eleva¬ tions much above sea level would appear to be exceptional (if accurately reported) or more likely erroneous. Leaves of Avicennia accumulate an external crystalline layer of excreted salt between rains. Three types of major roots are evident on plants: cable roots, anchoring roots, and pneu- matophores. Cable roots extend up to several meters horizontally underground from the base of the trunk and give rise to downward growing anchoring roots and upward growing pneumatophores. At least the cable and anchoring roots also give rise to finer roots within the substrate. Pneu¬ matophores (aerial, pencil-like roots that are evident at least at low tide and that facilitate the uptake of atmospheric oxygen) and cryptoviviparous seeds (in which the embryo swells and breaks through the seed coat, but not through the fruit wall, prior to the fruit falling from the plant) are characteristic of the genus. Macromorphological variation among several species of Avicennia is extensive, and often has been used in taxonomic circumscriptions. It has been shown that habitat conditions (e.g., variations in salinity and topographic position) and geography can correlate with morphology of the habit, leaves, and/or flowers (Sherrod and McMillan 1985; Duke 1991; Turner et al. 1995). Based on limited sampling (Fig. 1), pollen shape varies from suboblate to euprolate (i.e., with polar diameter: equatorial diameter [P:E] from 0.86 to 1.90). Much of this variation is evi¬ dent within a single sample (i.e., Salywon 1188 for A. germinans ) and probably represents harmo- megathic variation. Key to the Native and Naturalized Species of Avicennia in North America and Mesoamerica la. Flower actinomorphic; corolla yellowish to orangish, upper lip entire or bifid with the division to 0.2 mm long, the four corolla lobes ovate-triangular to ovate-elliptic; stamens inserted in distal half of corolla tube near base of lobes, oriented equally distant from each other and 166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 167 dehiscing toward the central gynoecium, 1.5-2 mm long, filaments 0.5-0.8 mm long; natural¬ ized in southern California. A. marina subsp. australasica lb. Flower ± 2-labiate; corolla white to cream (often with yellow in throat), upper lip bifid to 2-parted with the division to 3 mm long, the 4-5 lobes oblong to obovate; stamens inserted at midpoint or in proximal half of corolla tube, oriented in pairs adjacent to upper lip of corolla and dehiscing toward lower lip, 2-5 mm long, filaments 1.5—4 mm long; native in southeast¬ ern U.S.A., Mexico, and Central America.2 2a. Internodes of young stems glabrous; 1-flowered dichasia ± evenly distributed along rachis, internodes near midspike 3.5-7 (-9) mm long, rachis clearly visible; internal surface of corol¬ la lobes glabrous; 2 stamens exserted from mouth of corolla and 2 stamens included in corol¬ la tube (or only partly exserted from it); style not evident after corolla dehisces; capsule black¬ ish when dry. A. bicolor 2b. Internodes of young stems usually scurfy with shiny whitish trichome- or scale-like projections to 0.05 mm long; 1-flowered dichasia congested at or toward apex of rachis (± headlike), internodes near midspike mostly 0.8—4 mm long, rachis not or barely or only partially visible; internal surface of corolla lobes densely pubescent (at least in distal half); all 4 stamens exsert¬ ed from mouth of corolla; style usually conspicuous after (and often before) corolla dehisces; capsule usually grayish when dry. A. germinans 1. Avicennia bicolor Standi., J. Wash. Acad. Sci. 13: 354. 1923. Type. — PANAMA: Code: Aguadulce, outskirts of tidal belt, 5 XII 1911, H. Pittier 4968 (holotype: US!; isotypes: F-image!, K-image!, BM! NY! P! US!). Figures 1A-C, 2, 3D. Shrubs to 3 m tall or trees to 10 (-23) m tall. Young stems of reproductive shoots glabrous (tri- chomes sometimes persisting around leaf scars, but intemodes glabrous). Leaves petiolate, blades ovate to elliptic to broadly elliptic (to obovate), 62-175 mm long, 31-86 mm wide, length:width = 1.2-2.6, (emarginate to) rounded to subacute at apex, rounded to acute to subattenuate at base, sur¬ faces often conspicuously discolorous (abaxial surface lighter), punctate-pitted (sometimes incon¬ spicuously so abaxially), adaxial surface lacking trichomes, abaxial surface covered with a dense scurfy layer that sometimes includes longer ± appressed eglandular trichomes. Inflorescence of axillary (from distalmost leaves) and terminal pedunculate panicles of elongate spikes, panicles ± open, to 115 mm long (including peduncle and excluding corollas) and to 120 mm wide, peduncles 25—40 mm long, glabrous or distally becoming pubescent like branches and rachises, panicle branches subtended by triangular-concave inflorescence bracts (sometimes caducous) 1-3 mm long, pubescent like rachises, fertile portion of spike 12-35 mm long, rachises clearly visible, intemodes near midspike 3.5-7 (-9) mm long, densely pubescent with minute (< 0.05 mm long) Figure 1 (left). Pollen of Avicennia spp. A-C. Avicennia bicolor ( Sediles 461). A. Interapertural view. B. Apertural view. C. Polar view. D-G. Avicennia germinans (D, Palmer 484\ E-G, Salywon 1188). D. Interapertural view. E. Apertural view. F. Polar view. G. Interapertural view. H-K. Avicennia marina subsp. australasica ( Moran 28024). H. Interapertural view. I. Apertural view. J. Polar view. K. Apertural view. Scales = 5 pm. Pollen measurements (n = number of grains measured, P = polar diameter, and E = equatorial diameter). Many addi¬ tional grains from each specimen were examined, but data summarized below were made on a subset of those oriented to pemiit accurate measurements and photographic documentation. Avicennia bicolor — Sediles 461 (CAS, n = 5): P = 19-22 pm, E = 19-24 pm, P:E = 0.86-0.92. Avicennia germinans —Daniel & Araque 9478 (CAS, n = 2): P = 23 pm, E = 24-25 pm, P:E = 0.92; de Nevers et al. 6554 (CAS, n = 2): P = 25 pm, E = 26-28 pm, P:E = 0.89; Palmer 484 (CAS, n = 1): P = 46 pm, E = 24 pm, P:E = 1.90; Salywon 1188 (CAS, n =4): P = 27-42 pm, E = 24-29 pm, P:E = 0.93-1.75. Avicennia marina subsp. australasica — Moran 28024 (CAS, n = 4): P = 28-41 pm, E = 24-27 pm, P:E = 1.04-1.70. 168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 glandular trichomes and with antrorse (to flexuose) eglandular trichomes to 0.1 mm long. Bracts triangular-ovate to subcircular, 1-2.3 mm long, abaxially pubescent like rachis. Bracteoles similar to bracts. Flowers mostly 8-10 per spike (but up to 22 per spike). Calyx 2-4 mm long, lobes broad¬ ly ovate to subcircular, abaxially pubescent like rachis. Corollas 4.5-7 mm long, internally white to cream, externally glabrous (proximal 2/3 of tube) and densely pubescent with appressed eglan¬ dular trichomes to 0.2 mm long (distal 1/3 of tube and limb), tube 2-2.5 mm long, limb ± 2-labi¬ ate with 4-5 lobes, upper lip apically 2-parted (with division up to 2 mm long) and/or wider than lobes of lower lip, all lobes oblong to obovate, glabrous internally, 3—4.5 mm long. Stamens 4, inserted at midpoint or in proximal half of corolla tube, 2 exserted from mouth of corolla tube and 2 included in or only partially exserted from corolla tube, oriented in pairs near upper lip of corol¬ la with thecae opening toward lower lip, 2-2.5 mm long, filaments 1.5-2 mm long, anthers pre¬ sented at 2 heights, thecae 0.3-0.6 mm long; pollen suboblate to oblate spheroidal, polar diameter (P) 19-22 pm, equatorial diameter (E) 19-24 pm, P:E = 0.86-0.92. Style not evident, stigma lobes 0.2-0.4 mm long. Fruit greenish yellow, black when dry, ovoid to ellipsoid, 15-29 mm long, 7-17 mm wide, ± sparsely pubescent with antrorsely appressed eglandular trichomes to 1 mm long, these sometimes more or only evident distally on mature fruits. Phenology. — Flowering: September-May; fruiting: February-August. Distribution and habitats. — Pacific coast of southern Mexico (Chiapas, ca. lat. 16°02'26"N), Central America, to southern Panama (Los Santos and Darien, ca. lat. 7.98°N; Fig. 2); plants occur along and near shorelines in coastal mangrove swamps (mangals) and salt marsh¬ es at elevations at or near sea level. In addition to the provinces and departments of Central Amer¬ ican nations from which specimens have been examined (noted below), this species potentially also occurs or occurred in the following Pacific coastal political units: Guatemala (Escuintla, Jutiapa, Retalhuleu, San Marcos, Santa Rosa, Suchitepequez), El Salvador (La Libertad, La Paz, La Union, San Miguel, San Vicente, Sonsonate, Usulutan), Nicaragua (Carazo, Managua), and Panama (Chiriqui, Veraguas). Gibson (1970) included Avicennia bicolor Standi, in her account of the Guatemalan taxa because of its expected occurrence in the country. Although it undoubtedly occurred or still occurs there, the species has yet to be collected in Guatemala. The southernmost occurrence of Avicennia bicolor has been attributed to the Pacific coast of Colombia (e.g., Sanders 1997; Duke 2010; Aymard 2015). The sole collection cited by Aymard (2015; Forero & Gentry 794 at COL) and another Colombian collection identified as this species (Gentry & Juncosa 41115 at COL), both appear to pertain to A. germinans based on images of these collections supplied by COL. Unless other collections that conform to A. bicolor have been made in Colombia, the southern extent of this species appears to be in Panama at ca. 7.98°N, on both sides of the Gulf of Panama (i.e., in the provinces of Los Santos [Dwyer 5079A] and Darien [Duke 5488]). Local names. — “Madre sal” ( DeRiemer s.n.); “madresal prieto” ( Santamaria D. & Romero B. Id); “palo de sal” (Sediles 457); “palo de sal hoja ancha” (Sediles 455, 460). Conservation status. — The extent of occurrence (EOO) of Avicennia bicolor is 262,479 km 2 , although a major portion of that area consists of open ocean and inhospitable upland habitats. The species has been assessed as VU (vulnerable) by the IUCN (Duke 2010) based on a docu¬ mented continuing decline in population under criterion A. Discussion.— This species is readily recognized by the combination of its glabrous young stems, relatively remote dichasia in the inflorescence (Fig. 3C), bilaterally symmetric flowers with internally glabrous corolla lobes, and black fruits (when dry; Fig. 3D). The Mexican occurrence of this species has sometimes been overlooked (e.g., Breedlove 1986; Spalding et al. 1997) or minimized (e.g., treated as a synonym of A. germinans for practical pur- DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 169 Figure 2. Map showing distribution of Avicennia spp. in North America and Mesoamerica. The generalized distribution of A. germinans is shown. The distribution of A. germinans in Cuba and other West Indian islands is not shown. poses; Lopez R and Ezcurra 2002). Throughout its distributional range it often grows with A. ger¬ minans (e.g., Pittier 4968, the type, grows with Pitier 4969, a collection of A. germinans from the same locale). Rabinowitz (1978) noted that where these species are sympatric there is apparently no zonation between them. Like those of A. germinans, flowers of A. bicolor have been noted to be fragrant (e.g., Knapp 1244). Corollas are usually described by collectors as white to cream; they are sometimes noted to have a yellow throat (like A. germinans', e.g., Borg and Schoenenberger 2011). Williams 78 from Panama shows exceptionally long and floriferous inflorescences with the fertile portion of spikes to 65 mm long and with up to 22 flowers (vs. usually 4-8) per spike. Tomlinson (1986) treated A. tonduzii as a synonym of A. bicolor, and others have followed his influential taxonomic account (e.g., Aymard 2015). In his key to species of Avicennia and descrip¬ tion of A. bicolor, Tomlinson indicated that corollas of this species were conspicuously pubescent within (as they are on the type of A. tonduzii, but not on the type of A. bicolor). Distinctions, if any, between A. tonduzii and A. germinans are not readily apparent, and others have treated these names as synonymous. Additional information about A. tonduzii is provided below under A. germinans. Additional specimens examined.— Costa Rica. Guanacaste: Nandayura, Peninsula de Nicoya, Playa Bejuco, 09°49'56"N, 085°20'34"W, A. Fernandez 1762 (MO); Abangares, Cuenca del Abangares, San Buenaventura, 10°10'29.8253"N, 085°09'31.4524"W, L. Gonzalez & A. Garita 3909 (MO); Tamarindo, Playa Tamarindo, 10°18'N, 085°51'W, W. Haber & W. Zuchowski 8961 (F, MO); Port Parker, J. Howell 10242 (CAS); Refugio Silv. Tamarindo, Estero Tamarindo, Santa 170 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 Figure 3. Avicennia spp. A. Flower of A. gemiinans in Florida (photo by Bob Peterson, cropped, creative commons license, ). B Flower of A. marina subsp. australa- sica in Australia (photo by M. Fagg, cropped, source: Australian National Botanic Gardens at ). C. Inflorescence of A. biflora (Sediles 461, CAS). D. Fruits of A. bicolor (top; Barrera 8, CAS), A. germi- nans showing dehiscence (middle; Ferris 5396, DS), and A. marina subsp. australasica (bottom; Nickerson 6445a from New Zealand, CAS). DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 171 Cruz, 10°19'40"N, 85°49'20"W, Q. Jimenez 859 (K, MO); P.N. Santa Rosa, Canton de La Cruz, Peninsula de Santa Elena, Estero Grande, 10°54'59"N, 085°47'03"W, J. Morales 4142 (F, MO); P.N. Santa Rosa, Playa Naranjo, 10°47'53"N, 085°40'44"W, J. Morales et al. 1262 (F); Puerto Jesus, R. Pohl & G. Davidse 10588A (F, MO, US); Canton La Cruz, P.N. Santa Rosa, Peninsula de Santa Elena, Murcielago, 10°55'20"N, 85°44'15"W, F. Quesada 84 (K, MO); P.N. Santa Rosa, Playa Naranjo, N. Zamora et al. 1140 (F, MO). Puntarenas: Golfo de Nicoya Extemo, Isla San Lucas, Punta Canon, J. Morales & D. Santamaria 12387 (MO); Garabito, Cuenca del Jesus Maria, Garabito, alrededores de Playa Punta Loros, 09°51'26.3900"N, 084°41'29.8110"W, A Rodriguez & V. Ramirez 6688 (MO). El Salvador. Ahuachapan: without locale, Padilla 333 (US); Las Salinas, K. DeRiemer 1625 (US), 1626 (US); Las Chacaras, en La Barra de Santiago, K. DeRiemer s.n. (US). Honduras. Choluteca: Playas de Cedeno, F. Padilla 101 (BM). Valle: Puerto Soto, 12 kin from El Tular, A. Molina R. 21457 (BM, NY, UC, US); Puerto Soto, 15 km WSW de San Lorenzo, C. Nelson 1323 (MO). Mexico. Chiapas: Paderon, Tonala, E. Matuda 16353 (US); Mpio. Pijijiapan, Estero San Jose, 15°4339.50'TST, 093°29'50.20"W, S. Santamaria-Damian & E. Romero-Berny la (MEXU), lb (MEXU). Nicaragua. Chinandega: Mpio. El Viejo, Reserva Natural de Cosiguina, San Remigio entre El Congo y Bella Vista, 13°0nS[, 097°35'W, I. Coronado G. & R. Rueda 3626 (MEXU, MO); Mpio. Puerto Morazan, de Morazan hasta 5 km en direccion Ttonala, 12°49'N, 087°09'W, R. Rueda et al. 17322 (MEXU, MO). Leon: Isla del Venado, 3.3 km de Las Penitas, Barrera 8 (CAS); Las Penitas, Isla Juan Venado, 12°13'N, 086°53'W, I. Coronado G. & R. Rueda 4943 (MO); Las Penitas, Isla Juan Venado, 12°17'N, 086°53'W, I. Coronado G. et al. 6979 (MO); Isla del Venado, 8.2 km de Las Penitas, 60 m del Estero Las Penitas, Sediles 445 (NY), 456 (NY); 3.3 km de Las Penitas, Isla del Venado, Sediles 454 (MO), 459 (MO), 461 (CAS); 3 km de Las Penitas, en la Isla del Venado, Sediles 455 (F), 460 (K); 8.1 km de Las Penitas, en la Isla del Venado, 60 m del Estero Las Penitas, Sediles 457 (P); 8.3 lan del Las Penitas, en la Isla del Venado, 80 m del Estero Las Penitas, Sediles 458 (US). Rivas: “Marsella,” 11°16-17'N, 85°52-53'W, M. Araquistain 3828 (MO, P); San Juan del Sur, entre Las Playas de Marsella y Rivas, ll°17TSf, 085°54'W, R. Rueda et al. 1435 (MO). Panama. Code: ca. 2 km del Puerto, camino entre el puerto de Aguadulce hasta el pueblo, M. Correa A. 4314 (MO); Isla del Pozo, salinas of Rio Estero Salado, 08°1LN, 080°30'W, S. Knapp et al. 3401 (MEXU, MO); below Aguadulce, E. Tyson 7262 (FSU, MO). Darien: ca. 10 mi S of El Real on Rio Pirre, J. Duke 5488 (MO). Herrera: Cienega El Mangle, NE of Paris, S. McDaniel 8020 (FSU, MO). Los Santos: Monagre Beach, J. Dwyer 5079A (MO). Panama: Isla Casaya, J. Duke 10372 (MO); San Jose Island, ca. 55 mi SSE of Balboa, Playa Grande, I. Johnston 1259 (DUKE, MO, P, US); Porto Posada, R. Williams 78 (NY). Panama Oeste: Punta Chame, W. D’Ar- cy 10241 (MO); Punta Chame, 10-15 mi from Pan American Hwy., 08°40'N, 079°45'W, S. Knapp 1244 (MEXU, MO). 2. Avicennia germinans (L.) L., Sp. PI., ed. 3, 2: 891. 1764. Bontia germinans L., Syst. Nat., ed. 10, 2: 1122. 1759. Type.— “Habitat in Indiis” [JAMAICA], P. Browne s.n., LINN Herb. No. 813.2 (lectotype, designated by Stearn [1958: 35]: LINN-image!). Figures 1D-G, 2, 3A,D. Avicennia nitida Jacq., Enum. Syst. PI. 25. 1760. Avicennia officinalis var. nitida (Jacq.) Kuntze, Revis. Gen. PI. 2: 502. 1891. Hilairanthus nitidus (Jacq.) Tiegh, J. Bot. (Morot) 12: 358. 1898. Type. — See discus¬ sion. 172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 Avicennia tomentosa Jacq., Enum. Syst. PI. 25. 1760. Hilairanthus tomentosus (Jacq.) Tiegh, J. Bot. (Morot) 12: 358. 1898. Type. — See discussion. Avicennia tomentosa Jacq. var. campechensis Kunth, Nov. Gen. Sp. 2: 229 (ed. folio); 284 (ed. quarto). 1818 (“1817”). Type. — MEXICO. Campeche: “Crescit prope Campeche Mexicanorum,” F. Humboldt & A. Bonplands.n. (holotype: P-P00670135-image!). Avicennia floridana Raf., Atlantic J. 1: 148. 1832. Type. — Not designated (no specimens cited). Avicennia floridana Gand., Bull. Soc. Bot. France 65: 64. 1918, nomen illegit. Type. — U.S.A. Florida: Lee County, Fort Meyers, A. Hitchcock 270 (see discussion). Avicennia oblongifolia Chapm., FI. South. U.S. 310. 1860. Type. — U.S.A. Florida: Monroe County, Oct, Key West (fide protologue); pertinent specimens, if extant, would likely be in the Chapman herbarium at NY, but none have been located. Avicennia tonduzii Moldenke, Phytologia 1: 273. 1938. Type. — COSTARICA. Puntarenas: Punta Mala [ca. 09°03'54.48"N, 083°39'04.84"W], zone littorale du Pacifique, Mar 1892, A. Tonduz 6776 (holotype: BR- image!, fragment of holotype at NY!; isotypes: BM-image!, M-image!, MICH-image!, US!). Shrubs to 5 m tall or trees to 20 (-25) m tall. Young stems of reproductive shoots covered with dense ± antrorsely appressed or matted whitish and shiny trichomelike or scalelike projections to 0.05 mm long (scurfy) and sometimes also with longer appressed eglandular trichomes, the intern¬ odes infrequently glabrate. Leaves petiolate, blades narrowly to broadly elliptic (to obovate), 25- 160 (-185) mm long, 11-50 (-63) mm wide, (1.6-) 2.7-6.9 x longer than wide, acute to rounded to emarginate at apex, acute to subattenuate at base, surfaces often conspicuously discol- orous (abaxial surface lighter), punctate-pitted (sometimes inconspicuously so abaxially), adaxial surface lacking trichomes, abaxial surface covered with a dense scurfy layer that includes ± appressed longer eglandular trichomes or sparsely to densely scurfy without longer trichomes. Inflorescences of axillary and terminal pedunculate densely bracteate (± headlike) spikes or pani¬ cles of these, inflorescence branches (if present) subtended by triangular- to oblate- to subovate- concave inflorescence bracts to 5 mm long (sometimes caducous), peduncles (5-) 10-50 mm long, scurfy, pairs of flowers congested at or toward apex of rachis, inflorescence intemodes mostly 0.8^4 mm long near midspike, rachis not or only partially or barely visible, densely pubescent with mostly antrorse eglandular trichomes to 0.2 mm long. Bracts triangular to oblate to broadly ovate to subcircular, 1-4 mm long, abaxial surface scurfy to pubescent like rachis. Bracteoles similar to bracts. Flowers 8-14 (-26) per spike. Calyx 2^1 mm long, lobes broadly ovate to subcircular, abax- ially pubescent with antrorsely appressed eglandular trichomes to 0.4 mm long. Corollas (3.5-) 4-8 mm long, internally white to cream with yellow in throat, externally glabrous (proximal por¬ tion of tube) and densely pubescent with appressed eglandular trichomes to 0.5 mm long (distal portion of tube and limb), tube (1.5-) 2-3.5 mm long, limb ± 2-labiate with 4 or 5 lobes, upper lip bifid to ± conspicuously 2-parted at apex (with division up to 3 mm long) and/or wider than lobes of lower lip, all lobes oblong to obovate, densely pubescent (at least on distal half if not through¬ out) internally. Stamens 4, inserted at midpoint or in proximal half of corolla tube, exserted from mouth of corolla tube, oriented in pairs near upper lip of corolla with thecae opening toward lower lip, 3-5 mm long, filaments 2.2-4 mm long, anthers presented at ± same height, thecae 0.6-1 mm long; pollen oblate spheroidal to euprolate, polar diameter (P) 23-46 pm, equatorial diameter (E) 24-29 pm, P:E = 0.92-1.90. Stigma lobes (0.2-) 0.4-0.7 mm long. Fruit ovoid to broadly ellip¬ tic to obovoid, usually grayish (rarefy black) when dry, 12-32 mm long, 9-25 mm across at widest expanse, pubescent with antrorsely appressed eglandular trichomes to 0.7 mm long (especially when less mature) and scurfy (especially when more mature). Phenology.— Flowering: throughout the year; fruiting: April-January (especially in August, and likely throughout the year). DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 173 Figure 4. Avicennia germinans. A. Habit (composite from photos of living plants). B. Pneumatophore ( Schwartz & Nickerson 9119). C. Fertile shoot ( Calzada 434, Pipoly 9034, Sousa 3112). D. Inflorescence with flower in profile ( Lakela 29824, Sousa 3112). E. Flower, front-view ( Lakela 29824, Sousa 3112). F. Stamen ( Sousa 3112). G. Gynoecium ( Lakela 29824). H. Fruit with detail of surface ( Lundell 7009). Drawn by Tom Davis from specimens at CAS. 174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 Distribution and habitats. — Southern North America (southeastern USA and Mexico), Bermuda, West Indies, Central America, South America (Brazil, Colombia, Ecuador [including Galapagos Islands], French Guiana, Guyana, Peru, Surinam, and Venezuela), and tropical western Africa (see Daniel and Figuerido [2009] for distributional information on African plants, which often have been referred to as A. africana P. Beauv.). Figure 2 shows the generalized distribution of this species in North America and Mesoamerica. Plants occur in/on tidal flats, beaches, coastal mangrove swamps (mangals), salt marshes, sand dunes, and coastal grasslands and savannas at elevations at or near sea level. Common associates include Allenrolfea occidentalis, Batis mariti- ma, Bravaisia berlandieriana, Coccoloba uvifera, Conocarpus erectus, Distichlis litoralis, D. spi- cata, Echinochloa polystachia, Frankenia palmeri, Laguncularia racemosa, Maytenus phyllan- thoides, Rhizophora mangle, Salicornia biguelovii, S. virginica, Scaevola plumieri, and Spartina alterniflora. Avicennia germinans is the most widely distributed species of Avicennia in the New World. Its northern distributional limit there is in Bermuda (ca. 32°20'N; Lacerda et al. 2002), where another true mangrove, Rhizophora mangle L. also attains the northern extent of its distribution; these occurrences are the northernmost of true mangroves. The northernmost occurrence of A. germinans on the Atlantic Coast of North America is in northern Florida (St. Johns County, 30°06.618'N, 081°22.303'W, Williams & Eastman s.n. at USF). On the coast of the Gulf of Mexico the northern extent of the species extends as far as the Bay St. Louis, Mississippi (Hancock or Harrison county, ca. 30°2'N, Sanger s.n. at NY). Of the three true mangroves that are common on the American mainland (A. germinans, Laguncularia racemosa, and Rizophora mangle) A. germinans is the northernmost in distribution. Its northern limit in North America appears to be determined by the frequency and duration of sub¬ freezing temperatures, which can kill the above-water parts of the plants or cause mortality (Sher¬ rod and McMillan 1985; Cavanaugh et al. 2014). Tomlinson (1986) noted a close correlation of the limits of the distributions of mangroves with the 24° isotherm of sea surface temperatures. A decade later Riitzler and Feller (1996) indicated that like corals, mangroves cannot survive where an average water temperature falls below about 23 °C. In recent decades, freeze-free winters have led to an expansion of A. germinans into salt marshes at the northern extremes of its range in the southern United States (Guo et al. 2013). A poleward expansion of the distributional range of A. germinans in southern North America, likely correlated with warmer winter temperatures and less extreme freezes, was recently noted by Cavanaugh et al. (2014). In the United States, plants have been documented from most of the coastal counties in Flori¬ da (except for most of those in the panhandle on the Gulf Coast in the northwestern portion of the state), rarely reported from at least two of the three coastal counties in Mississippi (e.g., Moldenke 1960; Scheffel et al. 2014; the locality cited for Mississippi by Lowe in 1921, Door Point, is in Louisiana), and documented from seven coastal parishes in Louisiana. In Texas the species has been collected along the Gulf Coast from Jefferson County near the Louisiana border intermittent¬ ly southward to Cameron County on the Mexican border. Although it likely occurs (or occurred) in all coastal counties of Texas, based on specimens noted below and in literature reports (Turner et al. 2003 and pers. comm, in 2010 from updated but unpublished maps; Rosen and Zamirpour 2014), it has been collected in Aransas, Brazoria, Calhoun, Cameron, Galveston, Jackson, Jeffer¬ son, Kleberg, Nueces, Refugio, San Patricio, and Willacy counties. Sherrod and McMillian (1981, 1985) discussed the distributional history of mangroves, including A. germinans, in Texas and in the northern Gulf of Mexico. The species has not been reported from the Gulf coast of Alabama (e.g., Krai et al. 2011). Avicennia germinans is known from all 16 coastal states of Mexico (occurring along the Gulf DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 175 of Mexico, Caribbean Sea, Pacific Ocean, and Gulf of California). The species reaches its north¬ ern extent in western Mexico along the Gulf of California in Sonora (Puerto Lobos, lat. 30°15'N; Felger et al. 2001) and its northern extent on the Pacific coast in Baja California Sur (ca. lat. 26°N; Turner et al. 1995). Distribution along Pacific coast of North America does not extend as far north¬ ward as that along the coasts of the Gulf of California, Gulf of Mexico, and the Atlantic Ocean like¬ ly because of upwelling of cold waters off the western coast of the peninsula of Baja California and California that limits their northward spread. Turner et al. (1995) discuss the distributions and prob¬ able environmental factors effecting them for this species in regions around the Gulf of California. In Central America to the south of Belize, the extent of mangrove communities and the abun¬ dance of A. germinans are better developed along the Pacific than the Caribbean coast. Indeed no specimens have been seen from the Caribbean coast of Costa Rica, although A germinans has been reported from the mouth of the Rio Mom, north of Puerto Limon (Zamora et al. 2004). In South America, as in North America, A. germinans has a broader distribution on the Atlantic than on the Pacific coast. Lacerda et al. (2002) indicated that the southernmost occurrences of A. germinans in South America are in northwestern Peru (Rio Piura, ca. 5°32'S) on the Pacific coast and in south¬ eastern Brazil (Atafona, ca. 21°37'S) on the Atlantic coast. Like the northward limit of this species on the Pacific coast of North America, the southern distributional limit of A. germinans along the western coast of South America in Peru appears to have a similar cause due to the cold waters of the Humboldt Current and upwelling there (Lacerda et al. 2002). Illustrations.— Figure 4; see also Hepper (1963: 449, fig. 309); Gibson (1970: 178, fig. 32, which is based on Standley 87592 at F); Moldenke (1973: 151, fig. 1); Villiers (1973: 65, pi. 16); Correll and Correll (1982: 1253, fig. 540); Nash and Nee 1984: 13, fig. 2); Tomlinson (1986: 189, fig. B.8); Sanders (1997: 83, fig. 1); Proctor (2012: 586, fig. 218). Nomenclature.— The synonymy provided above is based on types from North America and Central America or names sometimes applied to plants from these regions. Additional synonyms were listed by Moldenke (1960, 1973). Based on information provided by Stearn (1958), author¬ ship of the combination A. germinans often has been attributed to “(L.) Stearn” (e.g., Tomlinson 1986; Jarvis 2007; Daniel and Figueiredo 2009). In his letter to D. Ward (17 April 1962 attached to a specimen in FLAS), Steam indicated that the correct name for the black mangrove is A. ger¬ minans (L.) L., a case subsequently made by Compere (1963), and now widely accepted. The lectotype of Bontia germinans at LINN also has inflorescence intemodes similar to those of A. bicolor (i.e., ca. 4^4.5 mm long near midspike). From the image of this specimen, I have not been able to discern other diagnostic characters used here to distinguish these species. Thus, I am unable to confirm to which species it pertains. Moldenke (1960) noted that the types of A. nitida and A. tomentosa are both Herb. Jacquin s.n. specimens at BM, which he annotated as such in 1936. Images of both of these very sparse specimens have been seen, and based on what is visible and available, they likely conform to A. germinans. Although no specimen was designated as the type of A. floridana Gand., Gandoger’s herbari¬ um is at LY in France, and it is possible that the holotype is there; specimens of the type number at F, MO, Cornell, NY, and US were cited by Moldenke (1960), but he did not indicate where the holotype was located. Local names. — “Aili sip” (Kuna; de Nevers et al. 6554); “arbol salado” (Zizumbo & Colun- ga 500); “black mangrove” (e.g., Hammel 105); “cajiel” {Zizumbo & Colunga 500); “istaten” {J. Gonzalez & A. Perez 193); “kahil” (Huave; A. Gerardo B. 4); “madre de sal” {Rico-Gray & Espejel 305); “madresal” {Flores M. 1101; Garcia B. 486); “madre sal” {Espejo & Hernandez 2904); “mangle de sal” {Rico-Gray & Espejel 305); “mangle negro” {Chazaro B. 3299, Penning- 176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 ton & Sarukhan K. 9131); “mangle prieto” (e.g., DIAAPROY s.n .); “mangle pullequi” ( Moran 7151); “mangle rojo” (. DIAAPROY s.n.); “mangle salado” (e.g., Arteaga et al. 130; Allen 5631), “mangle senico” ( Carter 2721); “palo de sal” (Elorsa C. 285); “pipi” (Salas M. & Torres B. 825); “puyece” (Ferris 5396); “puyeque” (Gonzalez O. 1131; Mexia 1008); “saladillo” (Garda P. & Estrada L. 1974; Magana 1232); “sahkab luk’” (Ucan 431); and “u najil tikin xiw” (Tapia M. 1508). Uses. — Firewood (Mexia 1008; Garda B. 486); construction materials (Garda B. 486); and used by natives for diseases of the throat (Choussy 1593). Additional construction, medicinal, and food uses of the species were provided by Moldenke (1960), Burkill (1985), Cheatham et al. (2000), and Alvarez C. (2008). Conservation.— The black mangrove is a widespread and often an abundant shrub or tree of tropical and subtropical shorelines and near coastal habitats where it is an important (often domi¬ nant) constituent of mangrove communities. Its conservation status was assessed as Least Concern (LC) by Ellison et al. (2010). Discussion.— This species is readily distinguished by the combination of its scurfy young stems, densely spicate (headlike) inflorescences, bilaterally symmetric flowers with internally pubescent corolla lobes (Fig. 3 A), styles usually conspicuously elongating near the end of anthesis or shortly thereafter as the fruits begins to form, and the grayish fruits (when dry; Fig. 3D). Rare individuals from Mexico (e.g., Martinez s.n. from Baja California Sur at MEXU; Rico- Gray 95 from Yucatan at UC) have remote nodes (to 7 mm distant, as in A. bicolor) near the mid¬ point of the spikes, but show their affiliation to A. germinans by their internally pubescent corolla lobes, scurfy pubescent young stems, trichomes of the rachis to 0.5 mm long, and thecae 0.8 mm long. The fragrant flowers of Avicennia germinans are commonly visited by honeybees, and they are reputed to be an excellent source of honey. Corollas are often noted to be 10-20 mm long (e.g., Moldenke 1960; Tomlinson 1986). My observations concur with those of Gibson (1970), who indi¬ cated that she had seen none more than 8.5 mm in length. Morphological variation is considerable in this species, especially with respect to habit and leaf size and shape. For example, although it is commonly a sizable tree in portions of its range, A. germinans becomes a dwarf shrub at the northern extent of its range in the Gulf of Mexico (Sher¬ rod and McMillan 1985). Moldenke (1960) discussed variation in this species with respect to these and other attributes. In a letter to Paul Standley dated 19 February 1938 (attached to Brenes 12222 at F), Moldenke indicated that A. tonduzii could be “most easily” distinguished from A. germinans (as A. nitida) by the size of the flowers. Indeed some of the specimens cited and/or annotated by Moldenke as A. tonduzii have calyces (2-2.5 vs. 2^1) and corollas (3.5-5 vs. 4-8) that are shorter than most specimens treated as A. germinans. Subsequently, Moldenke (1960, 1973) distinguished A. tonduzii from A. germinans on the basis of its short (shorter than the stamens) style (vs. style exserted from the calyx when corolla is shed, surpassing the stamens) and leaf blades mostly elongate-oblong, 3- 5 times as long as wide (vs. lanceolate or lance-oblong to elliptic or obovate, usually < 3 times as long as wide). Elongate styles are evident on the holotype of A. tonduzii at BR (and on isotypes at BR and US). The other putative distinguishing characters of A. tonduzii overlap those of A. germi¬ nans when plants from throughout the ranges of both species are considered. Tomlinson (1986) considered A. tonduzii as a narrow-leaved variant of A. bicolor and treated the name as synony¬ mous with that species. Others have followed Tomlinson’s influential account, and A. tonduzii has been recognized as a distinct species in some relatively recent literature (e.g., Lacerda et al. 2002). In the southern Caribbean region and along the Atlantic coast of South America, the distribu- DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 177 tion and taxonomy of A. germinans become entangled with those of A. schaueriana Stapf & Leech- man ex Moldenlce. Stearn (1958) distinguished S. schauerana from A. germinans by the characters in the following couplet: la. Leaves mostly acute at tip; corolla lobes tomentose on both sides; style elongate. . A. germinans lb. Leaves mostly rounded at tip, sometimes acute; corolla lobes glabrous internally, tomentose externally; style very short, the stigma nearly sessile. A. schaueriana Leaf tips are variable in A. germinans. While length of the style might be somewhat depend¬ ent on floral stage and pollination, it appears likely that styles of A. schaueriana , like those of A. bicolor, do not elongate (or at least not to the extent) as in A. germinans. Like A. germinans, A. schaueriana has crowded inflorescences, but pubescence of the corolla lobes would appear to be a useful character for distinguishing these species. Representative materials examined.— Belize. Belize: St. Johns College, J. Dwyer & R. Pippin 10030 (MO); shortcut from southern part of Belize City to Western Hwy., A. Gentry 8560 (L); Belize, C. Hammel 105 (K); Belize, C. Lundell 4719 (K); W de Tropical Park, T. Ramamoor- thy et al. 3609 (MEXU); Gale’s Point, just S of Bar River mouth, ca. 17°14 r N, 88°18'W, J. Ratter et al. 6560 (K); Caye Caulker (southern island), South Point, west coast, 17°43'47"N, 088°02T5"W, J. Rietsema & D. Beveridge 19906 (NY). Corozal: ca. 4 km SE of Sarteneja, ca. 18°21'N, 088°07'W, G. Davidse & A. Brant 32658 (USE); ca. 2 km S of Sarteneja, 0.5 km NE of La Isla, 18 o 20'19"N, 088°07'25"W, Z. Goodwin & G. Lopez 1583 (MO). Stann Creek: Northeast Cay, F. Fosberg & D. Stoddart 53854 (US); Man-O’-War Cay, W of Tobacco Cay Range, F. Fos- berg & D. Spellman 54204 (MO, US); North Silk Cay, F. Fosberg & D. Spellman 54272 (US); Lit¬ tle Water Cay, F. Fosberg & D. Spellman 54343 (F); Buttonwood Cay, F. Fosberg & D. Spellman 54421 (US); Scipio Cay, 16°28.135'N-16°28.295'N, 088°17.793'W-88°17.829'W, D. Lentz et al. 2355 (NY); Long Coco Caye, off Placencia, ca. 16°29.5 r N, 88°12.7'W, D. Lentz et al. 2833 (NY); Dangriga, G. Proctor 36197 (MO); All Pines, [16°47'00"N, 88°19'00"W], W. Schipp 625 (UC); South Water Cay, Spellman & Stoddart 2157 (US); Seal Cay, Spellman & Stoddart 2481 (MO); Rendezvous Cay, Spellman & Stoddart 2498 (MO, US); Coco Plum Cay, Spellman & Stoddart 2567 (MO, US); 4.5 km S of Southern Highway turnoff, D. Stevenson 1132 (K). Toledo: W of beach road, Punto Gorda, J. Dwyer et al. 631 (MO). Costa Rica. Guanacaste: vicinity of Playa Naranjo in P.N. Santa Rosa, F. Almeda et al. 4205 (CAS); Playa de Coco, R. Blaisdell 300 (FSU); P.N. Palo Verde, Valle de Tempisque, Estacion Catalina, Laguna Nicaragua, 10°21T0"N, 085°13'00"W, U. Chavarria 1243 (F); Puerto Jesus, Nicoya, O. Cook & C. Doyle 743 (US); ca. 1 km E of Rio Tempisque ferry, G. Crow 6116 (F); between Playa Comchal and Playa Brasilito, L. Durkee 76-386 (F); Port Parker, Salinas Bay, F. Elmore El (DS); Sitio Ojochal, P.N. Santa Rosa, 10°47'35"N, 85°39'00"W, A. Fernandez 315 (K, NY); Playa de Samara, al S de Nicoya, A. Jimenez M. 526 (F). Puntarenas: Golfo Dulce, vicin¬ ity of delta of Rio Esquinas, P. Allen 5631 (DS, F, FSU, US); Quepos, F. Almeda et al. 3398 (CAS, F, NY); Canton Golfito, Peninsula de Osa, P.N. Corcovado, Estacion Sirena, Rio Corcovado, 08°28'50"N, 83°35'30"W, L. Angulo 136 (K); Puerto Jimenez de Osa, A. Brenes 12222 (F); Isla de Caballo, A. Brenes 15696 (NY); Punta Morales, J. Gomez-Laurito 9980 (F); near mouth of Rio Coto Colorado, ca. 8 km S of Golfito, 08°37'N, 083°09'W, W. Kress & L. Cablk 94-3803 (US); between Caldera and Matalimon, E of Puntarenas, K. Lems 5026 (NY, US); cerca de Caldera, J. Leon 505 (F); near Rincon de Osa, R. Liesner 2203 (NY); vicinity of Puntarenas, W. Maxon & A. Harvey 7841 (US); Santo Domingo, Pittier 7109 (K, US), 7110 (F, K, NY, US); estuaires de Santo Domingo de Golfo Dulce, A. Tonduz 10060 (DS, NY, US); N side of Estero, Puntarenas, I. Wiggins & D. Porter 132 (CAS). 178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 El Salvador. Ahuachapan: Las Chacaras, La Barra de Santiago, K. DeRiemer 1645 (US); San Francisco Menendez, Garita Palmera, zanjon El Aguacate, 13°43'N, 090°04'W, D. Rodriguez & E. Escobar 1875 (MO); Santuario de las Aves, 13°42'N, 90°00'W, R. Villacorta y E. Montalvo 81 7 (K, MEXU). La Libertad: El Amatal, San Diego, 13°25'N, 089°14'W, J. Gonzalez 316 (MO); Estero de San Diego, J. Gonzalez & A. Perez 193 (MEXU). La Paz: Estero de Jaltepeque, P. Allen 7278 (US); near mouth of Rio Jiboa, F. Choussy 1593 (US); El Zapote, Costa del Sol, 13°2EN, 89°W, J. Gonzalez & M. Hernandez 326 (MEXU, MO). La Union: rocky beach, La Union, A. Bee¬ tle 26263 (K, UC); ca. 1 km S of Barrancones, 13°26'08"N, 087°47'32"W, G. Davidse et al 37355 (MEXU, MO); coast near La Union, V Grant 716 (F); vicinity of La Union, P. Standley 20786 (US). Sonsonate: S of Acajutla, P. Allen 6837 (F, US); Estero San Juan, K. DeRiemer 1617 (US). Usulutan: Jiquilisco, El Tercio, 13°15'N, 088°31'W, R. Carballo & L. Cabrera 831 (LAGU- image!). Guatemala. Escuintla: Iztapa, Canjon Moron, M. Arrecis 106 (CAS, MEXU, MO); San Jose, J. Donnell Smith 2510 (K, NY, US); San Jose, W. Maxon & R. Hay 3659 (US); Had. Las Fian- zas, G. Salas 367 (US); Puerto de San Jose, J. Veliz & M. Veliz 94.4076 (CAS, MEXU). Izabal: near Puerto Barrios, P Standley 72167 (F, NY). Jutiapa: Las Lisas, Barra el Ahumado, M. Arrecis 74 (CAS, MEXU, MO), 121 (CAS), 122 (MEXU); Iztapa, Zanjon Moron, M. Arrecis 105 (CAS). Retalhuleu: Rio Ocosito en limite Tilepa, Ocos, San Marcos y Manchon, M. Arrecis 47 (MO); Champerico, P Standley 66563 (F), 87592 (F, NY). San Marcos: Mpio Ocos, almendrales, Tila- pa, M. Arrecis 50 (MO), Ocos, J. Steyermark 37803 (F). Santa Rosa: Las Lisas, M. Lara s.n. (MO); Las Lisas, Camaronera Mayasal, 13.80723°N, 090.21703°W, J. Lopez & R. Jimenez 120 (CAS); Parque de la Barra Hawaii, Aldea el Dormido, 13.84113°N, 090.34751°W, J. Lopez et al. 100 (CAS). Honduras. Atlantida: ca. 5 km NE of Tela near Telatinza, ca. 15°48TSf, 087°26'W, E Daniel & J. Araque 9478 (CAS, NY). Choluteca: Raton Island, A. Molina R. 22779 (US), 23287 (DS, F, NY, US); Cedenos Beach, A. Molina R. et al. 31978 (F); Punta Raton, 70 km NW de Cd. Choluteca, C. Nelson et al. 3259 (MO). Colon: Rio Guaimoreto, 4.5 km NE of Trujillo on road to Castilla, 15°57'30"N, 085°54'30"W, J. Saunders et al. 625 (F, MO, NY). Gracias a Dios: Puerto Lempira, Laguna de Caratasca, A. Diaz Z. 212 (MEXU); Puerto Lempira, Laguna de Caratasca, M. Espinal 142 (MO). Islas de la Bahia: Isla de Barbareta, C. Nelson & G. Cruz 8411 (US); Isla de Roatan, playa al E de Roatan, C. Nelson & E. Romero 4591 (MO). Valle: cerca de Isla Zacate Grande, D. Hazlett 916 (MO); San Lorenzo, A. Molina R. 8635 (F); Golfo de Fonseca, Puerto Soto, 12 km from El Tular, A. Molina R. 21454 (F, NY). Mexico. Baja California: Bahia San Francisquito [28.40814°N, 110.57079°W], R. Moran r 12625 (SD-not seen). Observed by I. Wiggins at Bahia de los Angeles (see Turner et al. 1995). Baja California Sur: Playa Santispac, carr. transpeninsular, B. Arteaga et al. 130 (MEXU); Mag¬ dalena Bay, E Brandegee s.n. (DS, UC); El Mogote, peninsula extending into La Paz Bay, 24°8- ll'N, 110°19-26'W, A. Carter 2721 (DS, K, UC); Isla Carmen, vicinity of Las Salinas, ca. 25°59'N, 111°07'W, A. Carter 5924 (UC); Isla Carmen, Puerto Balandra, ca. 26°00.5'N, 111°10.5'W, A. Carter & R. Ferris 3733 (CAS, UC); Bahia Concepcion, ca. 14 mi S of Mulege, M. Dillon et al. 1954 (F); Bahia de La Paz, Chametla, 24°09'N, 110°06W, R. Dominguez C. 443 (MEXU); Sierra de la Giganta, ca. 12-14 mi S of Mulege, near Bahia Santispac, E. Elias et al. 10812 (F, NY); 16 mi S of Mulege at Conception Bay, R. Ferris 8680 (DS); Mpio. La Paz, Punta Prieta cerca de Pichilingue, 14 km NE de La Paz, F. Gonzalez M. et al. 8163 (MEXU); La Paz, I. Johnston 3045 (CAS); Coronados Island, 1. Johnston 3758 (CAS, K, UC); Carmen Island, Puer¬ to Balandra, I. Johnston 3821 (CAS, K, UC); San Evaristo Bay, I. Johnston 4089 (CAS, K, UC); Puerto Escondido, I. Johnston 4293 (CAS, K, UC); Magdalena Bay, H. Mason 1909 (CAS, DS, K); DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 179 Ballandra Bay, Carmen Island, 26°00.5'N, 111°10.5'W, R. Moran 3926 (DS, UC), 9168 (MEXU); El Mogote, peninsula in La Paz Bay, ca. 24°10'N, 110°20'W, R. Moran 7151 (CAS, DS, K); Con¬ cepcion Bay, F. Shreve 7099 (F); Isla San Jose, costa SW, 24°54'N, 110°38'W, M. Sousa P. 218 (MEXU); Isla del Carmen, lado W, Puerto Balandra, 26°0EN, 111°11'W, M. Sousa P. 242 (MEXU); Magdalena Bay near Medano Amarillo, J. Thomas 7940 (CAS, UC); Bahia Concepcion, Playa Los Cocos, 26.87763°N, 111.97499°W, D. Valvov 2005088 (MEXU); Bahia de Concepcion between Mulege and head of bay, I. Wiggins 5455 (CAS, DS, F, UC); Estero Salinas, arm of Alme- jas Bay, S of Magdalena Bay, I. Wiggins 11487 (CAS, DS, UC); 1.8 mi E of La Paz toward Pichilinque Bay, I. Wiggins 14563 (CAS, DS); S shore of Bahia de La Paz, I. Wiggins 16174 (DS); Puerto Escondido, 15 mi S of Loreto, I. Wiggins 17526 (DS); N side of Bahia Astiones, W side of Isla San Jose, I. Wiggins 17672 (DS); NE part of Isla San Francisco [24°50'32.36"N, 110°33'59.94"W], I. Wiggins 17765 (DS); Bahia de la Concepcion, beach at Punta Guadalupe, I. Wiggins & D. Wiggins 18017 (DS); N side of Santispaquis Cove, Bahia de Concepcion, I. Wig¬ gins & D. Wiggins 18239 (CAS, DS). Campeche: between Sabancuy and Cd. del Carmen, R. Burnham & R. Spicer 146 (MEXU); 30 km W de Hecelchakan, camino a Isla Jaina, E. Cabrera & H. de Cabrera 13345 (MEXU); 6 km NE de Champoton, carretera Cd. del Carmen, E. Cabrera et al. 8497 (MEXU); Isla de Jaina, ca. 54 km W de Hecelchacan, E. Cabrera C. et al. 11975 (MEXU); Mpio. Calkini, Isla Punta, 20°30'N, 90°W, C. Chan V.&J. Flores 420 (XAL); “Panga” de Zacatal, J. Chavelas P & C. Zamora S. ES-4752 (MEXU); Mpio. Hecelchakan, 6 km antes de la costa, en la carretera de Pomuch, Isla de Jaina, 20°14'N, 090°24'W, E. Gongora 546 (UC, XAL); Mpio. Campeche, Palmas, Cd. de Campeche, 19°52'N, 090°30'W, C. Gutierrez B. 5876 (XAL); Mpio. Cd. del Carmen, 5 kiuNE de Sabankuy, 19°03W'N, 091°08'00"W, C. Gutierrez B. 7390 (MEXU); Puerto Real, F. Menendez L. 467 (MEXU); S de Campeche, F. Miranda 7944 (MEXU); Mpio. Calkini, Isla Arena, 27 km de Takuche, 20°37'25"N, 090°25'W, M. Narvaez 1365 (MEXU); Mpio. El Carmen, Punta Cochinitos, Laguna San Francisco, 18°26TST, 091°46'W, D. Ocana N. & A. Nov- elo R. 158 (MEXU); carretera Champoton-Isla del Carmen, ca. 15 km de Champoton, T. Pen¬ nington & J. Sarukhan K. 9405 (K); carretera Champoton-Campeche, T. Pennington & J. Sarukhan K. 9623 (K); Mpio. Champoton, camino a El Zapote, 4 km desde el entronque con la carr. Campeche-Champoton, 19°20'N, 090°45'W, R. Rico G. 141 (XAL); 2 km N of Cd. del Carmen, J. Sauer 2440 (F); Champoton, W. Steere 1751 (CAS, MICH); Mpio. Tenabo, granja camaronera de Tenabo, entre KM 27 y el mar, 20°01'30"N, 090°13'06"W, P Zamora C. et al. 5824 (XAL); Mpio. Cd. del Carmen, Isla del Centro de Cayo Areas, SE parte, S. Zamudio 103 (MEXU, XAL). Colima: no collections seen, but species recorded from the state (see: http://www.projects- abroad.org/_downloads/uk/conservation-management-plan/mexico-conservation-management- plan-2014.pdf). Chiapas: Mpio. Tonala, W side of Mar Muerto opposite Paredon, D. Breedlove 20771 (DS); Mpio. Tonala, E shore of Mar Muerto, N of Paredon, D. Breedlove & R. Thorne 20806 (DS, NY); Mpio. Arriaga, balneario La Gloria, A. Espejo & S. Hernandez 2904 (MEXU); Acapetahua, cerca al Embarcadero Las Garzas, 15°12'38.7"N, 92°48'39.1"W, H. Gomez D. 2293 (K); Las Garzas, Acapet, E. Matuda 2728 (K, NY); Paderon, Tonala, E. Matuda 16279 (US); Mpio. Acapetahua, 0.5 km antes de La Palma, 15°12T6, 092°48'37"W, S. Ochoa G. et al. 4535 (MEXU); Mpio. Tonala, Col. Miguel Hidalgo, afueras de Puerto Arista, 15°55 f N, 93°50'W, V. Rico-Gray & I. Espejel 298 (F, MEXU); Mpio. Pijijiapan, Salina atras del Chocohuital, 15°30'N, 093°15'W, V. Rico-Gray & I. Espejel 305 (MEXU, XAL); Mpio. Tapachula, Estero de Puerto Madero, 14°45'N, 092°35'W, V. Rico-Gray & I. Espejel 346 (F, MEXU); Mpio. Puerto Madero, Puerto Madero, E. Ventura & E. Lopez 91 (MEXU, XAL); Las Margaritas, Pijiiapan, G. Zavala P. & M. Illescas 19 (MEXU). Guerrero: Mpio. Copala, Laguna de Chautengo, J. Almazan 248 (FCME); Mpio. Zihuatanejo, Playa La Ropa, Bahia de Zihuatanejo, 17°40'N, 101°34'W, G. Castil- 180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 lo G. 1137 (UC), 6273 (MEXU, XAL); Mpio. Jose Azueta, Barra del Potosi, 17°40'N, 101°34'W, G. Castillo C. & P. Zamora C. 6511 (XAL); Mpio. Zihuatanejo, entre la Punta Ixtapa y el Cerro El Rialito, 17°40TSf, 101°39'W, G. Castillo C. et al. 6566 (MEXU); Mpio. Cuajinicuilapa, Punta Mal¬ donado, N. Diego 2213 (FCME); Mpio. Acapulco de Juarez, El Arenal, Laguna de Tres Palos, N. Diego 4161 (FCME, MEXU); Mpio. Petatlan, Cerro Huamilule, (Morro de la Laguna Potosi), N. Diego & R. Oviedo 6636 (FCME); Mpio. Cruz Grande, W de Las Penas, R. Fonseca 1625 (MEXU); Mpio. Cruz Grande, Los Tamarindos, Laguna de Chautengo, R. Gutierrez 4 (FCME); Mpio. Tecopan de Galeana, Laguna Nuxco, extremo SE, F Lorea 5280 (XAL); Mpio. Atoyac de Alvarez, Arenal de Palos, Laguna de Mitla, L. Lozada P. 418 (FCME, XAL); Laguna del Potosi, F. Menendez L. 448 (CAS); Mpio. Petatlan, 4 km de Petatlan direccion Tecpan, A. Nunez 605 (MEXU, XAL); Mpio. Jose Azueta, Cerro Huamilule, en Barra el Potosi, 17°31'56"N, 101°27'08"W, S. Peralta 426 (MEXU, FCME); Mpio. Tecpan de Galeana, Nuxco, laguna, 17°15'07.6"N, 101°49T8"W, S. Peralta 444 (FCME). Jalisco: Puerto Vallarta, R. Acevedo R. 1487 (NY, XAL); Mpio. La Huerta, Laguna de Corte, 19°19'00"N, 104°56'20"W, G. Castillo C. et al 10745 (MEXU); Mpio. La Huerta, Playa Tenacatita, 19°17'00"N, 104°5E50"W, G. Castillo C. et al. 10848 (MEXU); Mpio. Tomatlan, Playa Chalacatepec, 19°38'50"N, 105°12'20"W, G. Castillo C. et al. 10967 (MEXU, XAL); Mpio. Tomatlan, Laguna Xola, 19°43T0"N, 105°15'20"W, G. Castillo C. et al. 10978 (XAL); Salina al N de Chamela, M. Gonzalez G. 152 (CAS); Mpio. La Huerta, La Manzanilla, L. de Puga 15542 (XAL); Barra de Navidad, J. Rzedowski 14605 (DS). Michoacan: Mpio. Aquila, Estero de Maquili, B. Guerrero C. 676 (XAL); Las Salinas [vie. of delta of Rio Balsas, fide McVaugh 1951], E. Langlasse 146 (K); Mpio. Coahuayana, Boca de Apiza, C. Soto N. et al. 7116 (MEXU). Nayarit: Mpio. Bahia de Banderas, Bahia de Banderas, 20°47'N, 105°15'W, G. Castillo C. 5818 (MEXU); Mpio. Bahia de Banderas, Laguna del Quelele, 20°44'N, 105°18'W, G. Castillo C. 6012 (MEXU); Mpio. Bahia de Banderas, Club de Golf Flamingos, cerca de Bucerias, M. Chazaro B. & R. Romero 8472 (XAL); SE of San Bias through Matanchen, SE toward Rio San Cristobal, C. Davidson 7608 (CAS); Pochote, Santiago Ixcuintla, 21°55'20.1"N, 105°30'30.9"W, DIAAPROYS.A. de C. V. 47383 (MEXU); Tres Marias Islands, Magdalena Island, F. Elmore 1132 (F); vicinity of San Bias, R. Ferris 5396 (DS); Mpio. Santiago Ixcuintla, Mez- caltitan, J. Gonzalez O. 5537 (DS); Tres Marias Islands, Isla Maria Magdalena, H. Mason 1793 (CAS, F, K, NY); Mexcaltitlan, Y. Mexia 1008 (CAS, UC); Mpio. Santiago Ixcuitla, Isla de Mex- caltitan, 21°50'36"N, 105°24'42"W, A. Miranda & G. Villegas 2053 (MEXU); Mpio. San Bias, ca. 2 mi E of San Bias on Hwy. 54, D. Norris & D. Taranto 13329B (CAS); Mexcaltitan, J. Ortega 5537 (K); Isla Maria Magdalena, O. Soils 9 (MEXU). Oaxaca: Mpio. Santa Maria Huatulco, Estero Cacaluta, 15°43'20"N, 096°09'40"W, G. Castillo C. et al. 9782 (MEXU, XAL); Chacahua Bay, F. Elmore D21a (DS), D22 (UC); Distr. Juchitan, Mpio. Chahuites, camino Chahuites-Las Salinas, A. Flores M. 1101 (CAS); Mpio. Chahuites, Rancheria Trejo, M. Garcia B. 486 (XAL); Mpio. Huamelula, 4 km por la carr. Pochutla-Salina Cruz, 4 km despues Huamelula, J. Garcia P. & E. Estrada L. 1974 (MEXU); Distr. Tehuantepec, Mpio. Salina Cruz, beach at La Ventosa, ca. 16°10'N, 095°09'W, R. Gereau & G. Martin 1921 (CAS); Laguna Superior, S of Juchitan, near Xandanl, R. King 1549 (NY, UC); Mpio. Tututepec, Chacahua, J. Magana 1232 (XAL); Puerto Angel, C. Morton & E. Makrinius 2624 (K); La Ventosa Beach, ca. 6 mi E of Salina Cruz, A. Reznicek & D. Gregory 304 (NY); Distr. Tehuantepec, Mpio. Santiago Astata, Laguna Colora¬ do 4 km W de Zaachilac, 15°57'39"N, 095°34'40"W, S. Salas M. & E. Torres B. 825 (MEXU, XAL); Distr. Tehuantepec, Mpio. Salina Cruz, 500 m W de Salinas del Marquez, 16°10'6.9"N, 095°14'24.3"W, S. Salas M. et al. 5596 (XAL); Distr. Tehuantepec, Mpio. San Pedro Huamelula, Rancho Paraiso, 15°51'49"N, 095°50'22"W, N. Velazquez R. et al. 265 (MEXU); Mpio. San Mateo, Huazantlan, La Salina, D. Zizumbo & P. Colunga 500 (MEXU). Quintana Roo: S de Punta Allen, DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 181 Cayo Cedros, Bahia de la Ascension, E. Cabrera 3405 (CAS, MEXU); 10 km N de Puerto More¬ los, camino a Punta Caracol, E. Cabrera & H. de Cabrera 3134 (MEXU, NY); 4 km N de la zona hotelera de Isla de Cozumel, camino a Isla de la Pasion, E. Cabrera & H. de Cabrera 13582 (MEXU); Isla Mujeres, camino al Puerto de Abrigo, E. Cabrera et al. 17200 (MEXU); Mpio. Isla Mujeres, Isla Mujeres, lado W de la marina, 21°19'N, 086°46'W, C. Chan et al. 1590 (XAL); Reserva Biosfera de Sian Ka’an, 6 km E de Ramonal, R. Duran et al. 1120 (MEXU); Mpio. Felipe Carrillo Puerto, La Laguna Xunyanche, 20°00'N, 087°40'W, J. Flores & E. Ucan 8352 (F); Mpio. Isla Mujeres, Isla de Contoy, 21°30'N, 086°49'W, J. Flores & E. Ucan 8855 (MEXU); Mpio. Othon P. Blanco, Cayo Centro en el Banco Chinchorro, 18°35'N, 87°20'W, J. Flores et al. 8959 (XAL); Cozumel Island, G. Gaumer 146 (K); Holbox Island, G. Gaumer s.n. (K); Cozumel Island, E. Goldman 653 (F, US); brecha a Punta Brava al S de Pto. Morelos, P. Moreno 843 (MEXU); Mpio. Isla Mujeres, Isla Mujeres, lado SE, 21° 14X1, 086°46'W, A. Puch et al. 865 (XAL); Mpio. Cozumel, Isla Cozumel, 20°30'N, 086°58'W, A. Puch et al. 1073 (XAL); Mpio. Benito Juarez, camino a Punta Nizuc desde el entronque de la carretera Cancun-Chetumal, 21°N, 086°50'W, V. Rico-Gray 123 (MEXU); Isla Mujeres, N end, J. Sauer & D. Gade 3265 (MICH); Mpio. Lazaro Cardenas, Chiquila, 21°23'N, 087°23'W, E. Ucan 431 (XAL); Mpio. Isla Mujeres, atras del Puer¬ to de Abrigo, zona del Sak Bajo, 21°15'N, 086°45'W, E. Ucan E. & J. Flores 1038 (MEXU, UC); Mpio. Othon P. Blanco, camino Blanco de X-Calak, rumbo a Majahual, 18°26X1, 087°56'W, E. Ucan E. et al. 620 (MEXU); Mpio. Felipe Carrilo Puerto, Vigia Chico, Reserva de la Biosfera Sian Ka’an, R. Villanueva 812 (MEXU). Sinaloa: Mpio. Escuinapa, Palmito, 9 km E al estero “Mez- cal,” J. Beltran M. 1014 (FCME); Mpio. Rosario, E edge of Mazatlan Bay, 0.7 mi W of Mex. 15, 8.6 mi N of Rio del Presidio, D. Breedlove 1577 (DS); Estero Ballena, old channel of Rio Fuerte (W of Los Mochis), R. Felger 8437 (CAS, MEXU); Topolobampo, A. Gibson & L. Gibson 2095 (FSU); Escuinapa, Arroyo de la Codojuiz, J. Gonzalez O. 1131 (K); Mpio. Rosario, Coacoyolitos, J. Gonzalez O. 6458 (CAS); 4 km W de El Toldo, 24°57'N, 107°57'W, V. Lopez S. 8.1 (MEXU); vicinity of Topolobampo, J. Rose et al. 13309 (NY); vicinity of Mazatlan, J. Rose et al. 14046 (F); N side of Topolobampo, D. Seigler & P. Richardson 11686 (MEXU); Mpio. Los Mochis, just W of Topolobampo, 25°35'N, 109°05'W, T. Van Devender et al. 2000-28 (NY). Sonora: Bahia Kino, mouth of Rio de Sonora, F. Drouet & D. Richards 3542 (F); 18.5 mi N of Bahia Kino Nuevo, J. Hastings & R. Turner 64-35 (DS); Estero Tastiola, NW de Guaymas, O. Holguin s.n. (DS); Tepoca Bay, I. Johnston 3288 (CAS); beach S of Guaymas, G. Lindsay 1154 (DS); ca. 1 mi W of Puerto Lobos on narrow peninsula, C. Lowe & R. Turner 3319 (DS); Sargento, T. Mallery & W. Turnage s.n. (DS); Guaymas, W. Phillips 3483 (CAS) Mpio. Hermosillo, 2.7 km N of Punta Chueca, 20°02'20"N, 112°10'W, A. Reina G.&T. Van Devender 96-635 (MEXU); Mpio. Caborca, Puerto Lobos, 30°16'16"N, 112°51T4"W, A. Reina et al. 97-265 (MEXU); vicinity of Guaymas, J. Rose et al. 12578 (NY); Bahia San Carlos, W of Guaymas, Weedons M-1091 (MEXU); bayshore at Empalme, I. Wiggins 6341 (DS); Isla Tiburon, Estero San Miguel, 28.968611°N, 112.20194°W, B. Wilder et al. 06-276 (CAS); Isla Tiburon, Punta Tormenta estero, 29°00'51.17"N, 112°H'54.21"W, B. Wilder et al. 08-329 (CAS); Isla Tiburon, Cyazim It, spit in estero at Punta Perla, 29.22442°N, 112.29345°W, B. Wilder et al. 08-377 (CAS). Tabasco: Mpio. Frontera, Playa Boqueron, 20 km E de Frontera, A. Guadarrama 876 (MEXU, NY); Mpio. Centla, Ejido Nuevo Centla, antes Playa Boqueron, 18°33T0.3"N, 092°30'48"W, M. Guadarrama O. et al. 6688 (MEXU, XAL); Mpio. Nacajuca, Laguna Bayazu, llegando por el Rio Gonzalez, A. Hanan A. et al. 981 (MEXU); Mpio. Centla, Paso San Roman, F. Ventura A. 20406 (MEXU, XAL); Mpio. Paraiso, Puerto Ceiba, Isla Dos Bocas, F. Ventura A. 20432 (XAL); Mpio. Paraiso, 4 km N de Mecoacan, S. Zamudio R. 77 7 (MEXU). Tamaulipas: Mpio. Altamira, Barra de Chavarria S, entra- da por el Barranco, E. de Dunas 743 (XAL); 1 km W de La Pesca, cerca de la Laguna Blanca, 182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 L. Hernandez 1567 (MEXU); Mpio. Aldama, Rancho Nuevo, 23°08'12"N, 097°46'01"W, D. Infante et al. 445 (XAL); Mpio. Matamoros, delta del Rio Bravo, 25°56'56"N, 097°09'07"W, D. Infante & J. Vazquez 656 (XAL); Mpio. Soto la Marina, campamento totuguero La Pesca, 23°47'27"N, 97°44'12"W, E. Martinez 39316 (MEXU); Mpio. Altamira, playa cerca del Puerto Industrial, A. Mora O. & J. Mora L. 5450 (MEXU); Mpio. Altamira/Aldama, Barra de Chavarria, P. Moreno C. et al. 743 (MEXU); vicinity of Tampico, E. Palmer 484 (CAS F, K, NY). Veracruz: Puente de Alvarado, J. Calzada 434 (F); Mpio. Coatzacoalcos, Laguna Ostion, camino Pajapan- San Juan Volador, 18°11'N, 94°36'W, J. Calzada 12657 (MEXU, XAL); Mpio. Panuco, alrede- dores de Laguna de Tamos, 22°13'N, 098°02'W, J. Calzada et al. 6268 (XAL); Mpio. Actopan, El Morro de La Mancha, La Laguna, 19°36'N, 096°24'W, G. Castillo C. 182 (F, UC, XAL); Mpio. Cosoleacaque, Polvorin, carretera Polvorin-Cosoleacaque, 17°59'48'N, 094°38T2"W, G. Castillo C. et al. 14838 (MEXU, XAL); Mpio. Agua Dulce, Rio Tonala, cerca de Arroyo Blasillo, M. Chaz- aro B. 3299 (XAL); Mpio. Actopan, Estacion Biologica El Morro de la Mancha, 19°36'00"N, 096°22'40"W, C.M. V.A. 2 (MEXU); Mpio. Cazones, Rancho Nuevo, Estero Boquilla, M. Cortes 455 (XAL); Mpio. Actopan, Laguna de la Mancha, carretera Cardel-Nautla, J. Dor antes 57 (MEXU); S de Laguna Salada, J. Dorantes et al. 1053 (F); 16 km S de Palma Sola, Laguna del Far- allon [19°39T9.58'TSf, 96°24'40.28"W], J. Dorantes et al. 1171 (CAS); alrededores de Laguna Verde (SW de La Planta), Alto Lucero, J. Dorantes et al. 5132 (NY); Mpio. Alto Lucero, Lugana de San Agustin, KM 71 carretera Cardel-Nautla, 19°55'N, 096°31'W, C. Guttierez B. 1311 (MEXU); Mpio. Zempoala, Estacion de Biologia “El Morro de la Mancha,” INIREB, carretera Cardel-Nautla, G. Ibarra M. 3b (FCME); horde sur de la Laguna Salada, A. Lot et al. 2055 (F); Mpio. Boca del Rio, Mandinga, 19°03'05.52"N, 96°04'43.56"W, F. Medina H. et al. 27 (MEXU, XAL); Laguna de Tampamochoco, cerca de Tuxpan, A. Mendoza s.n. (DS); Mpio. Tuxpan, 8 km de Barra de Tuxpan, L. Monroy et al. 147 (CAS, XAL); Mpio. Actopan, 50 m SW de la boca de la Laguna de la Mancha, 19°35'N, 096°22'W, A. Novelo 408 (XAL); Mpio. Alvarado, Laguna de Alvarado, 18°46T3"N, 095°45'38"W, R. Palestina et al. 1460 (XAL); Laguna de Sontecomapan, T. Pennington & J. Sarukhan K. 9131 (NY, K); Mpio. Coatzacoalcos, terraceria La Barrillas- Laguna Ostion, 17°45'30"N, 094°42'07"W, A. Rincon G. et al. 1731 (XAL); Barra de Tuxpan (20°58'N), N of Rio de Tuxpan mouth, J. Sauer & D. Gade 2981 (F); Rio Coscoapan, M. Sousa 3112 (F); Mpio. Cazones, Barra Cazones, S. Vargas P. 90 (XAL); Mpio. Actopan, La Mancha, F. Ventura A. 5226 (CAS). Yucatan: Mpio. Tizimin, 55 km en el camino a Las Coloradas, cerca el Puente del Rio Lagartos, J. Aguilar Z. & S. Diez M. 225 (MEXU); Mpio. Telchac, Laguna Rosa, 1 km de Puerto de Telchac, 21°20'N, 89°16'W, J. Calzada et al. 6611 (F, XAL); Pena, Chocarro & Jim 567 (BIGU); Mpio. Tizimin, 16 km E de Las Coloradas, 21°30'40"N, 87°50T5"W, R. Duran et al. 2575 (MICH); Mpio. Hunucma, 7.5 km E de Sisal hacia Celestun, E. Estrada 283 (FCME); Mpio. Telchac Puerto, 2 km E de Telchac Puerto, A. Feliciano K. 325 (MEXU); Mpio. Progresso, Isla Larga de los Arrecifes Alacranes, 22°26'N, 089°3LW, J. Flores & E. Ucan 9253 (XAL); Alacran Atoll, S end of Perez Islet, F. Fosberg 41866 (NY, US); Alacran Atoll, Pajaros Islet, F. Fos- berg 41904 (US); Las Bocas de Silam, G. Gaumer et al. 23340 (F, NY, US); Progreso, C. Lundell & A. Lundell 8140 (MEXU, MICH, NY); Mpio. Dzilam de Bravo, entre Santa Clara and Dzilam de Bravo, 21°25'N, 088°50'W, J. Palma & R. Allkin 300 (MEXU); 6 km W de Dzilam de Bravo, brecha a Pto. Telchac, H. Quero R. & R. Grether 2469 (MEXU); Mpio. Hunucma, 2 km E de Sisal, 21°10T5"N, 090°00'45"W, E. Reyes de los Santos 607 (MEXU); Sisal, Schott 361 (F); Celestun, Schott 473 (F); Mpio. Celestun, 8 km antes Celestun, viniendo de Kinchil, 20°53'N, 090°20'W, V. Rico-Gray 60 (F, MEXU, UC, XAL); ca. 4 km de Sisal, viniendo de Hunucma, Hunucma, 21°13'N, 090°03'W, V. Rico-Gray 75 (MEXU); Mpio. Progreso, 1.5 km S de Chelem, camino a Progreso, 21°15'N, 090°20'W, V. Rico-Gray 87 (MEXU); 1.5 km E de Dzilam de Bravo, 21°25'N, DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 183 088°50'W, V. Rico-Gray 95 (F, UC, XAL); Mpio. Telchac Puerto, 0.5 km S de Telchac Puerto, 21°20'N, 089°15'W, V Rico-Gray 101 (F, MEXU, UC, XAL); afueras de la cuidad de Rio Lagartos, 21°35'N, 088°10'W, V Rico-Gray 106 (F, MEXU, XAL); between Progresso and Telchac Puerto, J. Sauer & D. Gade 3207 (F); Progresso, W. Steere 3092 (MICH); Mpio. Celestun, 1 km E de Celestun, 20°51'30"N, 090°24'00"W, J. Tapia M. 1508 (MEXU); Tizimln, alrededores de El Cuyo, ca. 21°30'45"N, 087°40'46"W, M. Ventura 158 (F); Alacran Reef, Isla Perez, S end of island, B. Welch s.n. (DUKE, MEXU). Nicaragua. Leon: Poneloya, ca. 12°23'N, 087°03'W, R. Haynes 8617 (NY); 7.1 km de Las Penitas, orillas del Estero Las Penitas, Sediles 408 (K); Estero Las Penitas, 3.1 km de Las Penitas, 12°20'N, 086°59'W, Sediles 413 (CAS), 443 (CAS); Estero Brasil, ca. 2 km S of Hwy. 32 on road to Velero, ca. 12°10'N, 086°45'W, D. Stevens et al. 17293 (CAS). Managua: Masachapa, J. Atwood & D. Neill AN32 (NY). Region Autonomista Atlantico Norte (northern Zelaya): Pozo Verde, 10 km NE de Puerto Cabezas, 14°06'N, 083°20'W, E. Little 25401 (F, US). Region Auton¬ omista Atlantico Sur (southern Zelaya): Bluefields harbor, El Bluff, S. Marshall & D. Neill 6507 (USF). Rivas: Estero San Juan del Sur, 11°14-16'N, 085°51-53'W, M. Araquistain 3809 (CAS); San Juan del Sur, F. Seymour 1269 (MEXU, NY, UC). Panama. Bocas del Toro: Changuinola Valley, C. Cooper & G. Slater 81 (US); Water Valley, H. von Wedel 987 (US). Chiriqui: playa cerca del KM 3, J. Him 358 (US). Code: Aguadulce, out¬ skirts of tidal belt, H. Pittier 4969 (US). Colon: vicinity of Colon, J. Cowell 97 (NY); vicinity of Viento Frio, H. Pittier 4116 (US). Guna Yala: trail from Cangandi to dock by Mandinga airport, [09°27'39.50"N, 079°05'0.70"W], G. de Nevers et al. 6554 (CAS, US). Herrera: PN. Sarigua, 08°00'41"N, 080°29'03"W, I. Alvarez B4339 (US). Los Santos: Salinas de Chitre, W D’Arcy & T. Croat 4200 (F). Panama: Perlas Archipelago, San Jose Island, Naval Cove, C. Erlanson 120 (NY); Perlas Archipelago, San Jose Island, I. Johnston 1129 (US); Isla San Jose, H. Kennedy 2281 (F, US); Taboga Island, Gulf of Panama, H. Pittier 3614 (NY, US); Miraflores Locks, W. Stern et al. 50 (US). U.S.A. Florida: Brevard Co.: Merritt Island, R. Krai 4972 (FSU, UC); S end of Merritt Island, Banana River, Coquina, R. Whetstone 9116 (MO). Broward Co.: along Dania Beach Blvd., ca. 2 mi W of beach, S. Leonard 6924 (FSU). Charlotte Co., 4.2 km S of De Soto Co. along Peace River, 2.7 km W of US 17, 26°59'50"N, 081°59'10"W, A. Franck & B. Upcavage 1866 (USF- image!). Citrus Co.: Shell Island, near mouth of Crystal River, R. Long 1309 (USF). Collier Co.: near Naples, R. Godfrey 58071 (FSU); Everglades City, O. Lakela 29824 (NY). Flagler Co.: inland waterway in vicinity of Marineland, R. Godfrey 61686 (FSU). Franklin Co.: between St. George Sound and small tidal marsh just NE of Culpepper home on Cannonball Acres, L. Anderson 5580 (FSU). Hernando Co.: N of Aripeka on Fla. 595, J. Carlton s.n. (USF). Hillsborough Co.: Long Key, F. Lewton s.n. (NY). Indian River Co.: along Indian River Lagoon, Oslo Riverfront East Con¬ servation Area, Oslo Road E of US 1, ca. 3 mi. S of Vero Beach, 27°35.193'N, 080°21.902'W, S. Myers 1281 (USF-image!). Lee Co.: Tarpon Bay, eastern Sanibel, W. Brumbach 7904 (NY); Lit¬ tle Pine Island, H. Moldenke 929 (MO). Levy Co.: causeway to Cedar Key, R. Godfrey & P. Red- fearn 52828 (UC); near jet. A Street and 3 rd Street, Cedar Key, K. Murray 80-43-10 (NY); Cedar Keys, I. Wiggins 19342 (DS); E side of Seahorse Key, SE of lighthouse, I. Wiggins & D. Wiggins 19430 (DS). Manatee Co.: Palmetto, G. Nash 2450 (MO); Palma Sola, vie. of Manatee, J. Simp¬ son 80 (UC). Martin Co.: Jonathan Dickinson State Park (Girl Scout Camp), R. Woodbury & R. Roberts s.n. (USF). Miami-Dade Co.: Coral Gables, S end ofVee Lake, 25'MO.5^0.7TST, 080°16.1- 16.5'W, J. Abbott 24063 (FLAS-image!); Everglades Natl. Park, Flamingo Area, 25°08.38'N, 080°55.88'W, W. Hess et al. 8579 (MO, NY); Kampong, 4013 Douglas Road, Coconut Grove, W. Judd 5602 (FLAS-image!). Monroe Co.: Long Key, D. Cornell & H. Cornell 40150 (MO); Rode 184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 Harbor, Key Largo, C. Janish & J. Janish 447 (DS, MO); Big Pine Key, W Muenscher & R. Thorne 18073 (UC); Lower Matecumbe Key, J. Pruski et al. 2826 (NY); Key West, just S of airport, 24°33.162'N, 081°46.024'W, A. Salywon 1188 (CAS); E end of Packet Key, I. Wiggins 20081 (DS). Palm Beach Co.: Jupiter Island, G. Cooley et al. 4864 (USF). Pasco Co.: just W of Port Richey along inlet canal from beach, J. Ray et al. 9980 (FSU). Pinellas Co., Boca Ciega, bayside just S of Treasure Island causeway, R. Thorne 48355 (UC). Sarasota Co.: Sarasota, Marie Selby Gardens, 27°19'33"N, 082°32'28"W, H. Bizet 51 (MO, NY); Historic Spanish Point, in Osprey W of US 41, 27°12T5"N, 082°29'46"W, M. Nolan 62 (USF). St. Johns Co.: between Matanzas and Marineland, R. Godfrey 70653 (CAS, FSU, GA, UC); Anastasia State Park, Conch Island, ca. 4.75 km NNW of jet. FL A1A and FL 312, E of FL A1A; immediately S of the St. Augustine Inlet, 29°53'56"N, 081°17'21"W, J. Kunzer et al. 2146 (USF-image!); Crescent Beach, D. Seigler & D. Young 10211 (MEXU); Anastasia State Recreation Area, NW Conch Island, S of Vilano Point and St. Augustine Inlet, UTM-471684, 3308353, C. Slaughter et al. 16617 (FSU); Tolomato River, 30°06.618'N, 081°22.303'W, A. Williams & S. Eastman s.n. (USF-image!). St. Lucie Co.: Hutchinson Island, Blind Creek access area, off Indian River, G. Silberhorn s.n. (USF). Taylor Co., Jug Island [29°50'31.75"N, 83°36'58.05"W], R. Godfrey 60403 (UC). Volusia Co.: 9 mi S Daytona Beach, near Ponce de Leon Inlet, R. Norris 541 (FSU). County undetermined: Indian River, A. Curtiss 1972 (CAS, GA, MO), s.n. (NY); Tampa Bay, P. Rolfs 248 (MO). Louisiana: Cameron Parish: Monkey Island near mouth of Calcasieu Ship Channel at Calcasieu Pass, adjacent to ferry landing, W Vermillion s.n. (LSU-image!). Jefferson Parish: Grande Isle, J. Carlton s.n. (USF); Fifi Island, 29.255541°N, 089.978158°W, D. Atha 12910 (NY). Lafourche Parish: 0.6 mi S of Fourchon Road bridge (R22E, T23S, S24), A. Lasseigne 6146 (MEXU); S of end of La. 3090, S of Fourchon City, S of Leeville, R. Thomas et al. 103237 (MO, NY). Orleans Parish: New Orleans, Nuttall s.n. (K). Plaquemines Parish: ca. 40 mi (air) SSE of New Orleans, 29.40782, -89.79907, M. Bell s.n. (LSU- image!). St. Bernard Parish: North Islands in North Chandeleur Sound, NE of Venice, R. Thomas et al. 89768 (MEXU). Terrebonne Parish: E end of Isle Dernier, F. Givens 3733 (MO); Brush Island, F. Lloyd & S. Tracy 249 (NY). Mississippi: Harrison Co., Bay St. Louis [possibly Hancock Co.?], C. Sanger s.n. (NY); Cat Island, 30.23037°N, 89.08532°W, Scheffel et al. 2014 (living plant-image!). Jackson Co.: Ranger Lagoon, Horn Island, 30.24171°N, 88.67886°W, Scheffel et al. 2014 (living plant-image!). Texas: Aransas Co.: Redfish Bay, causeway to Port Aransas, Rte. 361, S. Hill 18296 (MO, NY); Port Aransas, B. Tharp 253 (CAS, MO, NY, UC). Cameron Co.: South Padre Island, between Old Causeway and Queen Isabella Causeway, F. Banda 81 (USF); Padre Island, just E of Port Isabel, D. Cornell et al. 25539 (UC); 8 mi SW of Port Isabel, inlet crossed by FR 1792 (NY), J. Crutchfield 2985 (NY); Clark Island, near Boca Chica, C. Lundell & A. Lundell 8760 (CAS, NY, UC); Point Isabel, H. Parks 2939 (MO); bay at Boca Chica, Brazos Santiago Island, R. Runyon 2812 (NY); Point Isabel in Lower Rio Grande Valley, R. Runyon 5897 (UC); boca de Rio Bravo, Schott 139 (NY); Boca Chica, G. Webster & R. Wilbur 3035 (GA). Nueces County: Aransas Pass, causeway between Aransas Pass and Port Aransas, P. Fryxell 5162 (MEXU). Refugio Co.: near Tivoli, J. Williams 415 (NY). 3. Avicennia marina (Forsk.) Vierh., subsp. australasica (Walp.) J. Everett, Telopea 5(4): 628. 1994. Avicennia tomentosa Sieber var. australasica Walp., Repert. Bot. Syst. 4: 133. 1845. Avi¬ cennia resinifera G. Forst., PL Esc. 72. 1786. Avicennia marina var. resinifera (G. Forst.) Bakh., Bull. Jard. Bot. Buitenzorg, ser. 3, 3: 210. 1921, nom illegit. (superfl.). Type.— Sheet 1460 in Thunberg’s herbarium (lectotype, designated by Everett [1994: 628]: UPS). Figures 1H-K, 2, 3B,D Shrubs to 2.3 m tall. Young stems of reproductive shoots covered with dense shiny granules or DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 185 scalelike projections to 0.05 m long (scurfy), soon glabrate. Leaves petiolate, blades ovate to ellip¬ tic (to obovate), 44-100 mm long, 19^41 mm wide, 1.7-2.8 (-3.6) times longer than wide, acute (to rounded or emarginate) at apex, subattenuate to attenuate at base, surfaces discolorous (abaxi- al lighter), punctate-pitted (sometimes inconspicuously so abaxially), adaxial surface lacking tri- chomes, abaxial surface covered with a dense scurfy layer. Inflorescences of axillary and terminal (sessile to) pedunculate ± headlike spikes, peduncles (0-) 1-40 mm long, scurfy or distally pubes¬ cent like rachis, rachis not or but barely visible, intemodes near midspike 1—4 mm long, scurfy and pubescent with ± antrorse eglandular trichomes to 0.2 mm long. Bracts opposite, broadly ovate to triangular, concavoconvex, 3^4 mm long, abaxial surface scurfy and often pubescent like rachis. Bracteoles similar to bracts except smaller. Flowers mostly 4-16 per spike, sessile. Calyx 3.5^4 mm long, lobes elliptic to broadly elliptic, concavoconvex, imbricate, abaxially pubescent with antrorsely appressed eglandular trichomes to 0.8 mm long, margin ciliate with similar but spread¬ ing trichomes. Corollas 3.5-6.5 mm long, internally drying dark or blackish proximally and light brownish distally (those from Australasia are usually described as yellowish or orangish and the color is often darker in the corolla tube), externally glabrous (tube and base of lobes) and densely pubescent with appressed eglandular trichomes to 0.2 mm long (remainder of lobes), tube 1.5-2 mm long, limb actinomorphic, 4-lobed, lobes ovate-triangular to ovate-elliptic, 2-4.5 mm long, apically entire (or 1 lobe sometimes slightly bifid apically with division to 0.2 mm long), internal¬ ly lacking eglandular trichomes (at least distally) but sometimes punctate-pitted (proximally). Stamens 4, inserted in distal half of corolla tube near base of lobes, exserted from mouth of corol¬ la tube, oriented symmetrically (i.e., equally distant from one another) around corolla with thecae opening toward central gynoecium, 1.5-2 mm long, filaments 0.5-0.8 mm long, anthers presented at same height, thecae 1-1.2 mm long; pollen prolate spheroidal to euprolate, polar diameter (P) 28-41 pm, equatorial diameter (E) 24-27 pm, P:E = 1.04-1.75. Style not evident, stigma lobes 0.2 mm long. Fruit ovoid to subellipsoid, proximally blackish and distally light brownish when dry, 15-24 mm long, 10-19 mm across at widest expanse, pubescent with erect to flexuose to antrorse eglandular trichomes to 0.3 mm long (especially when less mature) and scurfy (especially evident when more mature). 2 n = 64, 96 (Dawson 1989). Phenology. — Flowering: February, August-September; fruiting: February, November. Distribution and habitat. — Avicennia marina has the most extensive distribution among species in the genus; it is native to eastern Africa, southern Asia, Indian Ocean and western Pacif¬ ic Ocean islands, and Australia. Subspecies australasica occurs primarily in subtropical and tem¬ perate Australasia (i.e., southeastern Australia and northern New Zealand). It is the southernmost- occurring taxon among species of Avicennia in the Old World (to 38°45’S; Duke 2006), and the southernmost-occurring mangrove in the world. In southern California, where this taxon has been introduced and become naturalized (Fig. 2), plants occur in salt marshes with Batis, Juamea, Salicornia, Spartina, and Suaeda at or near sea level. Illustrations. — Munir (1986: 1179, fig. 546); Duke (1991: 314, fig. 7); Clarke and Myer- scough (1991: 285, fig. 1). Nomenclature.— The name “A. marina var. australasica (Walp.) Moldenke” has been used for this taxon, but as discussed by Everett (1994), this combination was not validly published at this rank, and other infraspecific taxa of A. marina are currently treated as subspecies. Avicennia marina subsp. australasica and the synonyms noted above are all based on A. resinifera. See Mold¬ enke (1960) and Duke (1991) for a full list of synonyms of Avicennia marina. Local name.— Gray mangrove. Conservation. — Avicennia marina has been assessed as a taxon of Least Concern (LC) by Duke et al. (2010). This taxon is not native in the New World, but has become naturalized locally 186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 and is potentially invasive. On his collection 28024 made in 1979, Moran noted that plants in Cal¬ ifornia had been introduced from Aukland, New Zealand about 1966-69. He also noted the pres¬ ence of about 100 or more flowering-size plants plus many seedlings in the wildlife reserve where his observations were made. Initial efforts to eradicate the species were unsuccessful. Discussion. — This species is readily distinguished from those native to the western Hemi¬ sphere by its actinomorphic flowers with yellowish to orangish corollas bearing ovate-triangular to ovate-elliptic lobes and its equidistant stamens that are inserted near the base of the corolla lobes, dehisce toward the center of the flower, and vary from 1.5-2 mm in length (Fig. 3B). Duke et al. (1998) provided genetic evidence that supported the morphological recognition of A. marina as a distinct species and that supported recognition of the three infraspecific taxa (treat¬ ed by him as varieties, but here recognized as subspecies): subsp. marina , subsp. eucalyptifolia (Valeton) J. Everett, and subsp. australasica. The latter subspecies would appear to consist of or contain polyploids based on the reported chromosome numbers of 2 n = 64 and 96 (Dawson 1989, as A. resinifera ). These numbers suggest a possible base number of x = 8 or x = 16. A chromosome number of 2 n = 36 was reported for an unspecified subspecies of A. marina by Subramanian (1988; without citation of voucher). These appear to be the only recent chromosome counts for both this species and for the genus. If these numbers are accurate, both polyploidy and dysploidy would appear to have played a role in the evolution of taxa in A. marina. The only other known chromo¬ some counts for Avicennia are 2 n = ca. 66 and n = ca. 33 by Raghavan and Arora (1958; with a mei- otic figure showing n = 33, but without citation of a voucher) for A. alba Bl. Sanders (1997) indi¬ cated a base chromosome number for Avicenniaceae of x = 18. Although no rationale was stated for this number, his conclusion was probably based largely on Subramanian’s count of 2n = 36 for A. marina. Avicennia marina subsp. australasica was distinguished by Duke (1991) from the other two varieties of A. marina by the fully (or nearly so) pubescent calyces (vs. pubescent only near the base), and the gray, fissured (vs. green, chalky smooth, and often flaky in patches) bark of the mature trunk. Specimens Examined.— U.S.A. California: San Diego Co.: Northern Wildlife Preserve, North Mission Bay, tidal area 100 m W of Rose Creek, 32.7949°N, 117.2247°W, I. Kay 29 (UCR); E edge of Kendall/Frost Marsh, Mission Bay, San Diego, ca. 32°47.5'N, 117°13.8'W, sea level, 9 September 1979, R. Moran 28024 (CAS, GH, MEXU, NY, UC, US); same locality, 12 June 1990, R. Moran 31036 (CAS, JEPS). Acknowledgments I thank Tom Davis, 2015 intern in biological illustration at the California Academy of Sciences, for the fine illustration of the black mangrove; photographers (M. Fagg and B. Petersen) for making their images available; and Judy Gibson (SD), Layla Hains (SD), Carlos Parra (COL), Rebecca Peters (CAS), Jon Rebman (SD), and Dagoberto Rodriguez (LAGU) for supplying images of pertinent herbarium specimens. The following herbaria generously provided access to their collections, which served as the basis for this study: BM, CAS, DS, DUKE, F, FCME, FLAS, FSU, GA, GH, JEPS, K, MEXU, MICH, MO, NY, P, UC, UCR, US, USF, XAL. DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 187 Literature Cited Alvarez C., N.S. 2008. Avicennia germinans (L.) L. In D. Louppe, A.A. Oteng-Amoako, and M. Brink, eds., Prota 7(1): Timbers/Bois d’oeuvre 1. Wageningen, Netherlands. [accessed 31 January 2016]. Aymard, G. 2015. Avicennia. In R. Bernal, S.R. Gradstein and M. 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Proctor, G.R. 2012. Flora of the Cayman Islands, ed. 2. Kew Publishing, Royal Botanic Gardens, Kew, UK. Rabinowitz, D. 1978. Dispersal properties of mangrove propagules. Biotropica 10:47-57. Raghavan, R.S., and C.M. Arora. 1958. Chromosome numbers in Indian Medicinal Plants-II. Proceedings of the Indian Academy of Sciences, Section B 47:352-358. Rosen, D.J., and S. Zamirpour. 2014. Avicennia germinans (Acanthaceae) gaining new ground in southeast Texas? Phytoneuron 2014-74:1-4. Rutzler, K., and I.C. Feller. 1996. Caribbean mangrove swamps. Scientific American 274(3):94-99. DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 189 Sanders, R.W. 1997. The Avicenniaceae in the southeastern United States. Harvard Papers in Botany 10: 81-92. Scheffel, W.A., K.L. Heck, J. Cebrian, M. Johnson, and D. Byron. 2014. Range expansion of black man¬ groves ( Avicennia germinans ) to the Mississippi Barrier Islands. Gulf of Mexico Science 2013(1-2): 79-82. Schwarzbach, A.E., and L.A. McDade. 2002. Phylogenetic relationships of the mangrove family Avicenni¬ aceae based on chloroplast and nuclear ribosomal DNA sequences. Systematic Botany 27:84-98. Sherrod, C.L., and C. McMillan. 1981. Black mangrove, Avicennia germinans, in Texas: Past and present distribution. Contributions in Marine Science 24:115-131. Sherrod, C.L., and C. McMillan. 1985. The distributional history and ecology of mangrove vegetation along the northern Gulf of Mexico coastal region. Contributions in Marine Science 28:129-140. Spalding, M., F. Blasco, and C. Field, eds. 1997. World Mangrove Atlas. The International Society for Man¬ grove Ecosystems, Okinawa, Japan. Stearn, W. T. 1958. A key to West Indian Mangroves. Kew Bulletin 13:33-37. Subramanian, D. 1988. Cytological studies of some mangrove flora of Tamilnadu. Cytologia 53:87-92. Tomlinson, P.B. 1986. The Botany of Mangroves. Cambridge University Press, Cambridge, England, UK. Turner, B.L., H. Nichols, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Botan¬ ical Miscellany 24( 1): 1-648. Turner, R.M., J.E. Bowers, and T.L. Burgess. 1995. Sonoran Desert Plants, an Ecological Atlas. Univer¬ sity of Arizona Press, Tucson, USA. V illiers, J.-F. 1973. Avicenniaceae. Flore du Gabon 22:63-66. Wang, L., M. Mu, X. Li, P. Lin, and W. Wang. 2011. Differentiation between true mangroves and mangrove associates based on leaf traits and salt contents. Journal of Plant Ecology 4:292-301. Walker, J.W., and J.A. Doyle. 1975. The bases of angiosperm phylogeny: Palynology. Annals of the Mis¬ souri Botanical Garden 62:664-723. Zamora, N., B.E. Hammel, and M.H. Grayum. 2004. Pages 91-216 in B.E. Hammel, M.H. Grayum, C. Herrera, and N. Zamora, eds., Manual de Plantas de Costa Rica, Volume I, Introduccion. Missouri Botanical Garden Press, St. Louis, Missouri, USA. 190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63 Page intentionally left blank PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 6, pp. 191-200, 15 figs., 6 tables April 29, 2016 A New Species of Serranus from Sao Tome and Principe, Eastern Atlantic (Pisces Teleostei, Serranidae) Peter Wirtz 1 and Tomio Iwamoto 2 1 Centro de Ciencias do Mar, Universidade do Algarve, Campus de Gambelas, PT 8005-139 Faro, Portugal. Email: petei~wirtz2004@yahoo.com. 2 Department of Ichthyology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118 U.S.A. Email: tiwamoto@calacademy.org Serranuspulcher is described from Sao Tome and Principe islands. It differs from all other eastern Atlantic Serranus species except S. heterurus (Cadenat, 1937) in a com¬ bination of the following characters: dorsal fin X,12, anal fin III,7; 42-50 pored lat¬ eral-line scales; interorbital area without scales; upper lobe of caudal fin pointed, lower lobe of caudal fin rounded. Serranus pulcher differs from S. heterurus in life color. S. pulcher probably also occurs off the coast of West Africa. It is the smallest known Serranus species in the eastern Atlantic. The percoid genus Serranus currently contains 14 valid species in the western Atlantic Ocean, one species {Serranus sanctaehelenae) endemic to the central Atlantic Islands of Ascension and St Helena, and six valid species in the eastern Atlantic (Eschmeyer and Fricke 2015). No Serranus species is amphi-Atlantic. Four of the six eastern Atlantic species ( S. atricauda, S. cabrilla, S. hepatus, and S. scriba) reach into the Mediterranean Sea. A seventh eastern Atlantic species of Serranus is here described from the islands of Sao Tome and Principe in the Gulf of Guinea. Material and Methods The specimens were obtained by SCUBA diving, with a hand-held aquarium net, at depths of about 2 to 30 m. They were preserved in ethanol or formol. Point-to-point measurements were taken on the left side of each specimen using a digital caliper with an accuracy of 0.01 mm and rounded to the nearest 0.1 mm. Body depth was measured at the beginning of the first dorsal fin. Body width was measured behind the insertion of the pectoral fins. Head length was measured from the tip of the snout to the end of the operculum, snout length from anterior edge of eye to tip of upper lip, pectoral fin length from ventral insertion to longest ray. Lateral-line scales were count¬ ed on the left side of the animal, where possible. Pored lateral-line scales are taken to the caudal- fin base and do not include those on the caudal fin itself. The last ray of the dorsal and anal fins is usually split to the base and is counted as one ray. Gill-raker counts are from the first arch and include all rudiments; counts of the upper arm are separated by a plus sign (+) from those of the lower arm, the raker whose roots span both the upper and lower arms of the gill arch is included in the count of the lower arm. The description of color patterns of living specimens is based on numer¬ ous underwater photographs by the first author and others (see Acknowledgments). Common abbreviations used include SL - standard length; TL - total length; HL - head length; D - dorsal; fin; A - anal fin; P - pectoral fin; V - pelvic [ventral] fin; LI - pored lateral line scales. 191 192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 6 Specimens mentioned in the present paper are deposited in the California Academy of Sci¬ ences (CAS); the Colegao Ictiologica Universidade Federal do Espirito Santo (CIUFES) at Vitoria, Brazil; the South African Institute for Aquatic Biodiversity (SAIAB); Stuttgart Natural History Museum (SMNS); Zoologisches Museum Hamburg (ZMH); and the Zoologische Staatssammlung at Munich (ZSM). Tissue samples of three specimens from Principe Island (ZSM uncatalogued) were sent to Benjamin Victor (Ocean Science Foundation) for DNA analysis. The results of his analysis will be published elsewhere. Comparative material.— The following specimens were used for direct comparisons with the type specimens of the new species. Additional material used for general comments are listed in a synopsis of the eastern Atlantic members of the genus Serranus (Wirtz, Heemstra and Iwamoto, In prep.): Chelidoperca africana: SAIAB 26564, Cameroon. Serranus accraensis : ZSM 32516, Angola, ZSM 32596 Angola, ZSM 32610 Angola. Serranus hepatus : ZSM 25637, Croatia, ZSM 41914 France. Serranus heterurus : SAIAB 65552 Angola; SAIAB 65682, Angola; ZMH 11056 Senegal; ZMS 43051, Cape Verde Islands; ZMS 43730, Cape Verde Islands; ZSM uncatalogued, Cape Verde Islands; CAS 231614, Sao Tome Island; CAS 231627, Sao Tome Island; CAS 234709, Guinea; CAS 234711, Guinea. Serranus scriba : ZSM 23526, Greece. Species Description Serranus pulcher Wirtz and Iwamoto, sp. nov. Figures 1-9, Tables 1-5. Serranus sp.(“Sao Tome comber“), Debelius 1998: 148. Kuiter 2004:162, figs. A-D. Serranus n.sp., Wirtz et al. 2007:8-9, fig. 8 Material Examined.— Holotype: ZSM 43868 (59 mm SL), Sao Tome Island (00°25.0997S1, 006°41.718'E), near wreck “Mar Vassa“ on coral rubble in about 6 m depth, Jan. 2015, formol pre¬ served, coll. Nuno Vasco Rodrigues. A small piece of the left pectoral fin is missing and the dorsal fin is slightly torn between the spinous and the soft-rayed part (figure la and b). Paratypes meas¬ ured: ZSM 43869 (66 mm SL) same data as holotype. ZSM 43879 “Mar Vassa”, Sao Tome Island (00°25.099'N, 006°41.718'E), Nov. 2014, coll. Nuno Vasco Rodrigues. ZSM 43880 “Mar Vassa”, Sao Tome Island (00°25.099'N, 006°41.718'E), Mar. 2014, coll. Nuno Vasco Rodrigues. CAS227751 (70.5 mm SL), Sao Tome Island, Kia Reef (00°25'0.01"N, 006°48'E), 25^10 ft [7.6- 12.2 m], 11 Jan. 2009, coll. J.E. McCosker, D. Catania, and J.-L. Testori. CAS227753 (73 mm SL, specimen labelled DC1009), Principe Island (01°41'09.3"N, 007°28'07.6"E), 40 ft [12.2 m], 23 Jan. 2009; coll. J.E. McCosker and D. Catania. CAS 227754 (4 spec., 45-74 mm SL), Principe Island, nw side Bom Bom Is. (01°41'44.0"N, 007°24'00.3"E), 48 ft [14.6 m], 20 Jan. 2009, coll. J.E. McCosker and D. Catania. CAS227755 (1 spec.), Sao Tome Island, Kia Reef (00°21'37.1"N, 006°43'08.5"E), 45-72 ft [13.7-21.9 m], 11 Jan. 2009, coll. J.E. McCosker, D. Catania, and J.-L. Testori. CAS227756 (2 spec., 75-78 mm SL), Sao Tome Island, Batalleo (00°22'05.7"N, 006°45'41.6"E), 13 Jan. 2009, coll. J.E. McCosker, D. Catania, and E. Milson. CAS227757 (spec¬ imen labelled DC999), Principe Island, Pedro Adalia (01°42'04.0"N, 007°25'42.1"E), 21 Jan. 2009, coll. J.E. McCosker, D. Catania, and R. Van Syoc. Paratypes not measured: CAS227752 (7 juve¬ niles), Principe Island, Pedro Adalia (01°42'02.3"N, 007°25'43.8"E), 52 ft [15.8 m], 19 Jan. 2009, coll. J.E. McCosker and D. Catania. CAS227757 (8 juveniles), Principe Island, Pedro Adalia (01°42'04.0"N, 007°25'42.1"E), 21 Jan. 2009, coll. J.E. McCosker, D. Catania, and R. Van Syoc. CAS227758 (3 spec.), Principe Island, Isla Santana, cave (00° 14'33.1"N, 006°45'36.1"E), 62 ft [18.9 m], 28 Jan. 2009, coll. J.E. McCosker, and J.-L. Testori.CIUFES 150 (pectoral fin taken for WIRTZ & IWAMOTO: NEW EASTERN ATLANTIC SPECIES OF SERRANUS 193 DNA sample), Sao Tome Island; CIUFES 155 (4 spec.), Sao Tome Island; ZSM uncatalogued (3 juvenile spec., tissue samples taken for DNA analysis), Principe Island. Diagnosis. — Dorsal rays X,12; dorsal fin notched between spinous and soft part; anal rays III,7; pectoral rays usually 15 (rarely 14 or 16); pelvic rays 1,5; gill rakers 6-9+12-14 (19- 23 total); pored lateral-line scales 42^19; cir- cumpeduncular scales 20-25, usually 22-24; interorbital without scales. Caudal fin truncate, the upper lobe slightly produced, lower lobe rounded. Dorsal, anal, and pectoral fins scaly near base. Scales ctenoid, not deciduous; 5-6 rows of scales from the beginning of the dorsal fin to the lateral line; three opercular spines, the middle one largest, the upper one often obscure, the lower one sometimes invisible to the naked eye; rear margin of anterior nostril forming a flap usually fringed with 4-6 long cirri that reach well past the rear nostril; posterior nostril a simple opening lacking a raised rim. Lips red with dark bands; a short moustache-like red streak behind end of maxillary, running across hind margin of dentary and almost meeting opposite streak at midventral line and enclosing ivory-white of mandibular rami; another red diagonal streak running from upper edge of maxillary, across lower edge of preopercle, across interopercle and branchiostegal rays, to base of pelvic fin, and enclosing white of preopercle, interopercle and chest. Size to at least 9 cm total length. Description.— Measurements and counts are presented in tables 1-5. Body relatively slen¬ der and compressed, width over pectoral bases about half of HL, greatest body depth about 2.7-3.2 in SL and less than HL, which is about 2.4-2.6 in SL. Dorsal and ventral profiles gently curved from tip of snout to caudal fin. Snout shorter in length than orbit diameter; both substantially more than interorbital width. Lower jaw projecting slightly beyond upper jaw; maxilla extending to below posterior half of pupil. Preopercle margin serrated with flattened spines, those at angle largest. Mouth large with upper jaw reaching to the level of the rear edge of the eye or beyond. Pre¬ maxillary teeth in a narrow band, the outer series spaced and slightly enlarged; one or more large canines at anterior end. Dentary with a band of small teeth flanked by a series of slightly enlarged outer teeth that become larger posteriorly. Vomerine tooth band broadly V-shaped, followed on each arm by narrow band of small palatine teeth. Dorsal rays X,12; first 4-5 spines of dorsal fin graduated, the 3rd to 5th spines longest, the spines following subequal; the soft rays slightly higher than longest spines, the 3rd to 5th ray longest; a slight notch in fin profile. Anal rays III,7; anal fin relatively high, its posterior tip some¬ what pointed; spines shorter than soft rays, the first spine more than half length of second and third spines, the second spine longer and stouter than the others. Pectoral 14-16, usually 15 soft rays; pectoral fin broad-based, its origin about on same vertical as those of dorsal and pelvic fins; the tip Figure 1 . Holotype of Serranus pulcher sp. nov. ZSM 43868 (59 mm SL) left side: a) in alcohol; b) radiograph (photos Dirk Neumann). 194 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 6 of pectoral fm extends to, or almost to, anus. Pelvic rays 1,5; the distal tips of the soft rays sometimes reach the level of the anus but often they are shorter. Caudal fin with usually 15 (one specimen with 17) branched rays; caudal fm truncate; dorsal lobe slightly produced, ventral lobe rounded at tip (Fig 2). Lateral line slightly arched over pectoral fin and from there on par¬ allel to dorsal profile. Scales on the nape from the beginning of the dorsal fm forward to the level of the serrated edge of the preoperculum, but not further forward (i.e., interorbital without scales); scales present on operculum (7 oblique rows of scales) and preoperculum (6 rows), but absent in all areas in front of eyes; six bran- chiostegal rays. Rear margin of anterior nostril forming a flap usually fringed with five (rarely four or six) finger-like cirri that reach well past the rear nostril; rear nostril a round hole without raised rim. Counts and meas¬ urements of the holo- type and paratypes in the ZSM collection are given in Table 1 and those of some speci¬ mens in the CAS collec¬ tion in Table 2. Addi¬ tional counts and meas¬ urements on further specimens in the CAS collection are given in Table 3. Counts and measurements taken by Phil Heemstra (SAIB) on 6 specimens from Sao Tome and Principe, collected by P. Wirtz in 2004, are given in Table 4; these specimens apparently are lost now. Counts and measurements on 5 specimens from Sao Tome Island taken by Francisco Reiner are given in Table 5; these specimens are apparently now lost. The paratype ZSM 43879 was dissected and turned out to have the genus-typical (Erisman and Hastings 2011) hermaphroditic gonad: the ovotestis is dominated by ovarian tissue with testicular tissue restricted to the posterior and ventral part. Color in alcohol (Fig. la): Alternating light and dark areas on upper and lower lip; head and upper half of body light brown; some lighter blotches on rear half of lower body; belly light; no dark spots on snout and fm membrane between the first two dorsal spines not black. Color in vivo: Color extremely variable (Figs. 2-9). The following patterns appear to be most Table 1: Counts and measurements (in mm) of Serranus pulcher holotype and ZSM paratypes. ZSM 43868 ZSM 43869 ZSM 43879 ZSM 43880 Holotype SL 59.2 65.6 68.4 70.5 D X, 12 X, 12 X, 12 X, 12 A III, 7 III, 7 III, 7 III, 7 P 14/14 15/15 15/15 15/15 V 1,5 1,5 1,5 1,5 LI 50 50 49 47 Circumpeduncular scales 21 22 21 24 Length V 16.4 15 15.8 15.3 Length P 17.2 16.9 18.1 18.1 Body depth 18.8 20 22.1 23.9 Head length 22 23.9 25.7 26.9 Snout length 4.9 5.6 6.8 6.9 Body width 9.3 10.3 10.8 11.7 Orbit diameter 5.7 6.1 6.3 6.2 DC B80 Figure 2. Serranus pulcher specimen from Principe Island, directly after capture (photo Dave Catania). WIRTZ & IWAMOTO: NEW EASTERN ATLANTIC SPECIES OF SERRANUS 195 Table 2: Counts and measurements (in mm) of several Serranus pulcher paratypes in the CAS collection. CAS227753 CAS227755 CAS227756 CAS227756 CAS227757 DC 1009 DC924 DC915 DC999 SL 62 57 61 59 53 D X, 12 X, 12 X, 12 X, 12 X, 12 A III, 7 III, 7 III, 7 III, 7 III, 7 P 15/15 16/16 16/16 16/16 18/- LI 44 - 47 45 47 Body depth Head length 18 18 19.5 19 16 23 22 23.2 23.3 20 Snout length 5.4 5.8 5.3 5 4.7 Orbit diameter 5.7 6 6.2 6.4 5.2 P length 15.9 14 16.5 15 14 V length 13.7 13 16 15.4 13 Table 3: Counts and measurements (in mm) of additional Serranus pulcher paratypes in the CAS col¬ lection. CAS227751 CAS227754 CAS227754 CAS227754 CAS227754 DC880 DC986 DC987 DC989 DC988 SL 56.6 60.2 47.6 37.2 39.8 D 12 12 12 12 12 A 7 7 7 7 P 15 15 15 15 14 LI 48 46 45 45 43 Circumpeduncular scales 21 2 5 23 24 Head length 23.6 23.8 19 15.1 16 Snout length 5 9 5.5 4.1 3.2 3.6 Interorbital width 2 5 3 9 3.5 2.6 2.4 Orbit diameter 5.8 5.8 4.5 3.9 4.8 Suborbital width 2 2 2 4 1.7 1.6 1.5 Postorbital length 12.5 12.9 10.4 8.2 8.5 Orbit to preopercle 8.6 8.4 6.6 5.3 5.6 Upper jaw 11.3 11.1 8.6 6.9 7.5 Predorsal length 23.1 23 18.7 15.7 15.8 Preanal length 36.8 41 30.4 25.3 25.6 Body depth 21.1 18.6 17 12 12.7 Length P 15.4 14.5 12.5 9 12 Length V 13.9 13.9 12 9 10.7 196 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 6 HI H2 H3 H4 H5 H6 SL 59 60 63 63 63 60 D X,12 X,12 X,12 X,12 X,12 X,12 A III,7 III,7 III,7 III,7 III,7 III,7 P 16/16 16/16 - 15/15 15/15 15/15 depth at D1 18.6 19.1 20.5 - - 19 head length 22.7 24.3 24.8 25.2 24 24 snout length 5.4 5 5.4 - 6.6 5.2 orbit diameter 6.1 6.2 6.4 7 6.6 6.4 P length 16.5 16.7 18 17 17.3 14 V length 14.5 16.3 17.1 16 17.3 16 LI 46 47 47 42 47 44 common: lips red to orange with dark bands; Table 4: Counts and measurements (in mm) of a short moustache-like streak behind end of Serranus pulcher non-type specimens, provided by Phil maxillary, running across hind margin of den- Heemstra. tary and almost meeting opposite streak at midventral line and enclosing ivory-white of mandibular rami; another diagonal streak running from upper edge of maxillary, across lower edge of preopercle, across interopercle and branchiostegal rays, to base of pelvic fin, and enclosing white of preopercle, interoper¬ cle and chest (Fig. 8); first rays of pelvic fms white. Juveniles can have orange, white, and dark stripes (Fig. 9). Habitat and distribution.— On hard bottoms (rock, gravel, coral rubble, or maerl) from about 1 m (juveniles) to at least 30 m depth. The new species is currently known with certainty only from the islands of Sao Tome and Principe, where it is apparently common in suitable habitats (Luiz Rocha and John McCosker, CAS, personal communica¬ tion). If the DNA analyses of an aquarium specimen taken off Ghana (Figs. 10-11) show that specimen to be the same as S. pulcher , the new species is also present on the mainland coast of Africa in the Gulf of Guinea. Comparison with other eastern Atlantic species or Serranus. — The species most similar in morphology to Serranus pulcher is S. heterurus Cadenat, 1937. The two species share the name-giving feature of S. heterurus , i.e. the upper lobe of the caudal fin is point¬ ed and the lower lobe of the caudal fin is rounded (also the case in several western Atlantic Ser¬ ranus species). Serranus pulcher and S. heterurus are the two smallest of the eastern Atlantic Ser¬ ranus species. Serranus pulcher differs from all others except S. heterurus in a combination of the following characters (see also Table 6): dorsal fin X, 12; anal fin III, 7; 42-50 lateral-line scales; interorbital area without scales; upper lobe of caudal fin pointed, lower lobe of caudal fin round¬ ed. Table 6: Main characteristics of the eastern Atlantic Serranus species. Table 5: Counts and measurements (in mm) of Serranus pulcher non-type specimens, provided by Francisco Reiner. D A V P Lateral line scales R1 X,12 III,7 1,5/1,5 16/16 47/47 R2 X,12 III, 7 1 , 5/1,5 16/16 47/48 R3 X,12 III, 7 1 , 5/1,5 16/16 47/46 R4 X, 12 III,7 1 , 5/1,5 16/16 47/47 R5 X,12 III,7 1 , 5/1,5 16/16 48/48 Serranus Dorsal fin Anal fin Pectoral fin Lateral line scales Total gill rakers TL (cm) accraensis X, 12-13 HI, 7 (-8) 17-18 45-48 18-21 20 atricauda X, 15-16 HI, (7-) 8 15-17 77-90 20-24 35 cabrilla X, (13-) 14-15 II, 7 (-8) 15-17 69-78 18-23 40 hepatus X, (11-) 12 (-13) HI, (6-) 7 15 (40-) 45-50 19-23 15 heterurus X, 12 HI, (6-) 7 (15-) 16-17 (-18) 44-47 (17-) 21-26 14 pulcher X, 12 III, 7 (14-) 15 (-16) (42-) 45-50 17-20 9 scriba X, (14-) 15 (-16) III, 7 (-8) 13-16 62-75 14-19 36 WIRTZ & IWAMOTO: NEW EASTERN ATLANTIC SPECIES OF SERRANUS 197 Figure 3. The most common color pattern of Serranus pulcher, near Santana Islet, Sao Tome (photo Peter Wirtz). Figure 4. Color of paratype ZSM 43880 shortly before capture (photo Nuno Vasco Rodrigues). Figure 7. Rare color pattern of Serranus pulcher from Principe Island (photo Dave Catania). Figure 9. Juvenile (about 4 cm TL) near Bom Bom Islet, Principe (photo Peter Wirtz). Figure 6. Rare color pattern (frightened animal) of Ser¬ ranus pulcher from near Rolas Islet, Sao Tome (photo Peter Wirtz). Figure 8. Throat color of Serranus pulcher, near San¬ tana Islet, Sao Tome (photo Peter Wirtz). 198 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 6 Figure 11. Serranus sp. from Ghana (photo Joe Figure 10. Serranus sp. from Ghana (photo J.F. Flem- Russo). dal). Figure 12. Serranus heterurus from the Cape Verde Islands (photo Rogelio Herrera). Figure 13. Serranus heterurus from Senegal (photo Sebastien Blache). Figure 14. Throat of Serranus heterurus from the Cape Verde Islands (photo Patrick Louisy). Figure 15. Serranus heterurus in alcohol; specimen from the Cape Verde Islands (ZSM 430516) (photo Dirk Neumann). WIRTZ & IWAMOTO: NEW EASTERN ATLANTIC SPECIES OF SERRANUS 199 The values of all morphological variables measured in nine specimens of S. heterurus (i.e., those mentioned in the description of S. pulcher, above) overlapped with those of S. pulcher. The two species differ in live color: S. heterurus has seven narrow white bars along a wine-red body, the first one on the opercle, the last one directly before the tail fin, a crescent-shaped light blue or white mark directly behind the eye and small blue spots on head and vertical fins (Figs. 12-14). In alcohol-preserved specimens of S. heterurus (Fig. 15) the white bars are often still visible, the area between the tip of the snout and the eyes bears dark spots, and the upper margin of the fin mem¬ brane between the first dorsal spines is often blackish; the crescent-shaped mark directly behind the eyes is often still visible but brown. Etymology. — pulcher, Latin, meaning beautiful. Proposed English common name.— Sao Tome Comber. Remarks. — In the aquarium literature, Hemdal (2009) described and figured a Serranus species from the coast of Ghana called “Peppermint basslef’. It is similar to S. pulcher (compare Figs. 10 and 11) and almost certainly belongs to the same species. A tissue sample from a speci¬ men collected for the aquarium trade at Ningo, east of Tema harbour in Ghana, at about 12 m depth (05°41.177'N, 000°17.510'E) will be analysed for its DNA sequence. Many years ago, the first author sent specimens and photographs of the new species to Phil Heemstra (SAIB), who agreed to describe it. But after several years during which he apparently made no progress on the description, Heemstra agreed to send all the specimens back. In the ensu¬ ing years, as collecting efforts using SCUBA on the islands of Sao Tome and Principe became more common (see Afonso et al. 1999 and Wirtz et al. 2007), many more specimens of the new species became available. The species was, in fact, discovered to be quite common in coastal waters of the islands. It also appeared to be a species taken by aquarium collectors on the mainland coast of Africa off Ghana, although the identity of fish from that country has yet to be confirmed. Because we are finalizing a manuscript reviewing all eastern Atlantic members of the genus (Wirtz, Heem¬ stra, and Iwamoto, in prep.), and because the species is common and frequently observed on the islands, we felt it necessary for us to provide a name and description of the species. Acknowledgements Dirk Neumann at the Zoologische Staatssammlung Miinchen took the photographs of the pre¬ served specimens. For the loan of specimens, we are grateful to Roger Bills (SAIAB), Dave Cata¬ nia (CAS), Ronald Fricke (SMNS), and Ulrich Schliewen (ZSM). Phil Heemstra (SAIAB) and Francisco Reiner (Centro Portugues de Estudo dos Mamiferos Marinhos, Lisbon) kindly provided data on specimens of S. pulcher and S. heterurus in their collections (Tables 4 and 5). Nuno Vasco Rodrigues captured four fresh specimens of S. pulcher at Sao Tome Island, including the holotype, which were essential for the description of the species. Nuno Vasco Rodrigues, Rogelio Herrera, Sebastien Blache, and Patrick Louisy provided photos of living specimens of S. pulcher and S. heterurus. Jay Hemdal kindly sent a photo of the Ghana Serranus which is likely to be pulcher. We thank John McCosker, Dave Catania, and Luiz Rocha (CAS) for collecting specimens and tis¬ sue of the new species and for use of their photographs of fresh and living specimens; McCosker also kindly reviewed the manuscript and provided sage advice. The second author (TI) gratefully acknowledges Jens-Otto Krakstad and Oddgeir Alvheim, Institute of Marine Research, Bergen, Norway, for facilitating his participation in fishery surveys of the R/V Dr. Fridtjof Nansen, in par¬ ticular those surveys in 2010 off Sao Tome and Principe (STeP) and in 2012 in the Canary Current Large Marine Ecosystem during which specimens of S. heterurus and other Serranus species were collected. We received necessary permits and cooperation from Arlindo Carvalho, Director-Gener- 200 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 6 al of the Ministry of Environment, and Joao Pessao, Director of Fisheries, Sao Tome and Principe (STeP). Virginia Carvalho and Jose Dias Sousa Lopes of STeP Fisheries were particularly helpful during the 2010 Nansen survey of the islands. Many thanks to Ulrich Schliewen and Dirk Neumann for providing facilities at the ZSM for the first author to examine specimens and for many helpful comments. Thanks also to Rick Feeney of the Natural History Museum of Los Angeles County for sending a copy of the unpublished Ph.D. thesis of M.R. Meisler (1987). The Centro de Ciencias do Mar (CCMAR) of the University of the Algarve co-financed three trips of the first author to Sao Tome and Principe Islands. Literature Cited Afonso, R, F.M. Porteiro, R.S. Santos, J.P. Barreiros, J. Worms, and P. Wirtz. 1999. Coastal marine fish¬ es of Sao Tome Island (Gulf of Guinea). Arquipelago Life and Marine Sciences 17A:65-92. Cadenat, J. 1937. Recherches systematiques sur les poisons littoraux de la cote occidentale d'Afrique, recoltes par le navire President Theodore-Tissier, au cours de sa 5e croisiere (1936). Revue des Travaux Institut Peches Maritimes 10(fasc. 4, no. 40):425-562. Debelius, H. 1998. Mediterranean and Atlantic Fish Guide. IRAN, Frankfurt, Germany. 305 pp. Erisman, B.E., and P.A. Hastings. 2011. Evolutionary transitions in the sexual patterns of fishes: insights from a phylogenetic analysis of the seabasses (Teleostei Serranidae). Copeia 2011 (3):357—364. Eschmeyer, W.N., and R. Fricke, eds. Catalog of Fishes, electronic version (accessed August 2015). Hemdal, J.F. 2009. Aquarium Fishes of the Tropical Eastern Atlantic Ocean. Advanced Aquarist’s Online Magazine. 8(6): Kuiter, R. 2004. Serranidae and Plesiopidae. A Comprehensive Guide to Basslets, Hamlets, Longfins and Relatives. Aquatic Photographies, Seaford, Australia. Meisler, M.R. 1987. Limits and relationships of serranine seabasses, with revisions of Serranus and Men- tiperca (Pisces: Serranidae). Unpublished Ph.D. thesis, University of Southern California, Los Angeles, California. 250 pp. Wirtz, R, C.E.L. Ferreira, S.R. Floeter, R. Fricke, J.L. Gasparini, T. Iwamoto, L. Rocha, C.L.S. Sam- paio, and U.K Schliewen. 2007. Coastal fishes of Sao Tome and Principe islands, Gulf of Guinea (East¬ ern Atlantic Ocean) - an update. Zootaxa (1523): 1-48. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7, pp. 201-268, 37 figs. April 29, 2016 Twenty-six New Species of Predaceous Ground Beetles (Coleoptera: Adephaga: Carabidae) from Ranomafana National Park, Madagascar David H. Kavanaugh 13 and Johanna Rainio 2 1 Department of Entomology?, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118, USA; Email: dkavanaugh@calacademy.org; 2 Department of Ecology and Systematics, University of Helsinki, PO. Box 65, FIN-00014 Helsinki, Finland; Email: kj ohannarainio@gmail.com 3 Corresponding author: David H. Kavanaugh ( dkavanaugh@calacademy.org ) As part of an inventory of the carabid beetle fauna of Ranomafana National Park, Fianarantsoa Province, Madagascar, we introduce 26 new species, for each of which the Park is designated as type locality: Brachypelus ranomafanae sp.nov., Caelosto- mus latus sp.nov., C. rotundiformis sp.nov., Dactyleurys minimus sp.nov., D. ranomafanae sp.nov., Trichinillus ( Mallopelmus ) ranomafanae sp.nov., Chlaenius kathrynae sp.nov., C. robertae sp.nov., Omphreoides ranomafanae sp.nov., Perigona ( Ripogena ) deuvei sp.nov., P. ( R .) ranomafanae sp.nov., Archicolliuris ranomafanae sp.nov., Thysanotus bimaculatoides sp.nov., Madecassina bimaculata sp.nov., M. quadrimaculata sp.nov., Pristacrus ranomafanae sp.nov., Eurydera ocellata sp.nov., E. oracle sp.nov., E. simplica sp.nov., Pseudomasoreus ranomafanae sp.nov., Assadecma basilewskyi sp.nov., Lebia laterolucida sp.nov., L. ranomafanae sp.nov., L. apicoviolacea sp.nov., Eunostus minimus sp.nov., and Erephognathus ranomafanae sp.nov. We provide information on type material, derivation of species name, fea¬ tures for recognition and known geographical and habitat distributions, as well as digital images of habitus and other key features for each species, all but two of which are currently known only from the type area. Keywords: Coleoptera, Carabidae, new species, Ranomafana National Park, Madagascar Madagascar is one of the world’s most important biodiversity hotspots (Myers 1998) and con¬ servation priorities (Mittermeier et al. 2004, Brooks et al. 2006). Most of the species living in Madagascar are precinctive (Brooks et al. 2002, Goodman and Benstead 2004). For example, almost all of Madagascar’s reptile and amphibian species, and all of its lemurs can be found nowhere else on the planet. The uniqueness of the Malagasy biota can be explained at least in part by its long period of geographic isolation (for more than 65 million years (Krause 2004)), its top¬ ographical variation, its diversity of climatic zones and its great diversity of habitats, from the deserts of the hot and dry southern region to the lush rainforests of the north and east. About 1350 carabid beetle species have been documented as occurring in Madagascar, with most of them treated by Basilewsky (1973, 1976, 1985) and Jeannel (1946, 1948, 1949). Of the known species, about 95% are precinctive. The known distributional ranges of many of the species are limited to a single or only few localities, and the fauna of much of the island remains inade¬ quately sampled and largely unknown. It’s highly probable that the Malagasy fauna, when better known, will exceed 2,000 carabid species. 201 202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Vohiparar. villaav Talatakely 0 Vatoharanana 0 Valohoaka Tsinjorano Ranomafana National Park Upper Vohiparara Lower Vo hi para ra 0Torotosy Ranomafana village 0 Mangevo 0 Study sites — Road Park area Figure 1 . Map showing location of Ranomafana National Park in Madagscar (yellow dot in inset) and Park area, col¬ lecting sites and other places named in the text. Scale line = 5.0 km. The geographical focus of this presentation is the carabid fauna of Ranomafana National Park (RNP), located in the southeastern part of the island, centered at about 21.25°S, 47.42°E (Fig. 1). Inaugurated in 1991 as the fourth national park in Madagascar, it is recognized both nationally and internationally as one of the world’s most important conservation areas because of its exceptional¬ ly high species diversity and the immediate threat from human activity if not protected. It was included as part of the Rainforests of the Atsinanana World Heritage site in 2007. The park lies on the edge of Madagascar’s High Plateau, with elevations ranging from 500 meters to 1,500 meters above sea level. The steepness of the slopes had preserved the park from exploitation before 1986. The range of altitudes allows for many different forest types, from lowland rainforest to cloud for¬ est and high plateau forest. The park is divided into a core protected zone of 41,500 hectares sur¬ rounded by a peripheral (buffer) zone in which some exploitation of the forest is permitted. The peripheral zone contains more than 100 villages totalling more than 25,000 residents, most of them subsistence farmers. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 203 Rainio and Niemala (2006) provided the first inventory of the carabid beetles of Ranomafana National Park. Their interest was to determine whether or not carabid beetles could be used as bioindicators of environmental change in and around the Park (Raino and Niemala 2003). Based on fieldwork between 2000 ansd 2005, they documented 127 carabid species in the fauna of the Park, including 37 species previously undescribed. Working independently, field teams from the California Academy of Sciences sampled the carabid fauna of the Park during several expeditions between 1998 and 2002 and raised the number of species known from RNP to more than 250, including nearly 50 new species. The purpose of this presentation is to make the names of 26 of these new species available for use by systematists, ecologists and resource managers though the presentation of economic and efficient treatments for each (see Erwin & Johnson 2000) that satisfy the requirements of the Inter¬ national Commission on Zoological Nomenclature (ICZN 2000) for availability and validity. The remaining undescribed species, mostly platynines, will be treated in a separate paper to follow. A full inventory of the known carabid fauna of RNP, with habitat distributions for each species included, is in preparation. Materials and Methods All but four of the new species described here were collected from Ranomafana National Park (RNP) or in the surrounding peripheral zone by our field teams during the period 1998-2005. With¬ in the Park itself, collecting sites were located in both secondary (Talatakely and “Upper Vohiparara”) and primary montane forests (Vatoharanana and Valohoaka). Talatakely is low sec¬ ondary montane forest, which was selectively logged in 1986-1989. The range of elevations at which specimens were collected in this area is from about 925 to 1000 m. “Upper Vohiparara” is our name given to a secondary montane forest site about 2 km northwest of Vohiparara village and accessible from there by both road and trails. The range of elevations at which specimens were col¬ lected in this area is from about 1140 to 1170 m. The Vatoharanana and Valohoaka sites are in pri¬ mary montane rain forest, four and eight km south of Talatakely and at elevations of about 980 and 1080 m, respectively. At Vatoharanana, there has been some logging, but Valohoaka is undisturbed. The peripheral zone consists of a varied mosaic of cultivations, abandoned tavy-fields and forest edges. Within this peripheral zone, specimens were collected from the following villages and their environs: Mangevo, Ranomafana, Torotosy, Tsinjorano and Vohiparara (see Fig. 1) Most specimens were collected by hand using the following methods: beating vegetation and accumulations of dead leaves in vegetation up to a height up to 3 m above ground; shaking trees and scraping bark and mossy and epiphytic coatings on them onto beating sheets; breaking into decaying logs; turning logs and stones on the forest floor. Collections were made during both day (DHK and JR) and night (DHK) hours using all of these methods. Pitfall trapping and sifting of forest litter with subsequent extraction using mini-Winkler extractors (Fisher 1998) were used as additional collecting methods. Materials. — This study is based on a total of 8,025 carabid specimens collected during the course of our respective field programs between 1998 and 2005 plus an additional 300 specimens from RNP borrowed from other collections (see below). About 265 different carabid species were represented in the material assembled, including the 26 new species described here. Except as noted in the text, holotypes were examined for all species mentioned in comparisons. Codens used in this report for collections from which specimens were borrowed and/or in which specimens, including primary types, are deposited are as follows: 204 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 CAS California Academy of Sciences, San Francisco, U.S.A. EMEC Essig Museum of Entomology, University of California, Berkeley, California, U.S.A. MZF Finnish Museum of Natural History, Helsinki, Finland MNHN Museum National d’Histoire Naturelle, Paris, France MTEC Montana State University, Bozeman, Montana, U.S.A. NMNH United States National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A. Examination of specimens. — Specimens were examined using a Wild M5 and Leica MZ9.5 stereoscopic microscope with a Proline 80 LED Ring Illuminator. Measurements. — Measurements were made using a Leitz stereoscopic dissecting micro¬ scope with a calibrated ocular grid with a scale interval of 0.1 mm. The measurements used in this report include standardized body length (SBL), which equals the sum of head length (HL), meas¬ ured from the apex of the clypeus to a point on the midline at the level of the posterior margin of the ocular swelling), pronotum length (PL), measured from the apical margin to the basal margin along the midline, and elytral length (EL), measured along the midline from the apex of the scutel- lum to the apex of the longer elytron. Readers should be aware that SBL is typically about 15% shorter than body length measured as a single distance from apex of the mandibles or labrum to apex of the elytra, and this difference should be considered when comparing length data presented here with length data in the literature. Nonetheless, we prefer SBL as a comparative of body length because it eliminates errors due to different alignments of body regions among different specimens. In distinguishing some species, we also use certain ratios as diagnostic features. These include the ratios of: pronotal width (PW) to pronotal length (PW/PL, with PW measured cross the pronotum at its widest point); elytral length to pronotal length (EL/PL); elytral length to elytral width (EW) (EL/EW, with EW measured across both elytra at their widest point); and standardized body length to elytral width (SBL/EW). Dissections. — Male genitalia were extracted from specimens relaxed in water immediately after it had boiled and to which a few drops of liquid detergent had been added. Genitalic prepara¬ tions were then cleared in hot 10% potassium hydroxide solution for five to 10 minutes, each con¬ stantly monitored to achieve only a useful degree of clearing. They were then rinsed briefly in 10% acetic acid and then repeatedly in distilled water. After examination, preparations were stored in glycerin in polyethylene microvials and pinned beneath their specimens of origin. Illustrations. — Digital images of whole specimens and particular structures were taken using a Leica imaging system including an M165C dissecting microscope, DFC550 video camera, and two KL1500 LCD light sources. Stacked images were captured and combined into single mon¬ tage images using the Leica Application Suite V4.2.0. Plates of images were created using Adobe Photoshop CS5. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 205 Systematics The arrangement of species treatments and suprageneric classification used here follows Lorenz (2005), the most current world standard, rather than the arrangement and classification of Jeannel (1946, 1948, 1949). Several of Jeannel’s suprageneric names are no longer in common use and the arrangement of taxa in his three-part work is different from that now recognized by most current carabid systematists. Tribe Scaritini Bonelli 1810 Subtribe Clivinina Rafinesque 1815 Genus Brachypelus Putzeys 1866 Jeannel (1946) included only two species in his treatment of this precinctive Malagasy genus, and Basilewsky (1976) described three additional species. Bulirsch et al. (2005) described eight new species and provided a key to all species known at that time. Finally, Bulirsch and Moravec (2009) described one more new species, B. janaki , bringing the total number of known species to 14. Members of the new species described below are easily distinguish from those of all the other described species on the basis of their unique combination of features compared with excellent key characters provided by Bulirsch et al. (2005). Brachypelus ranomafanae Kavanaugh and Rainio, sp. nov. Figures 2-3 Brachypelus n. sp.l; Rainio 2009: 31 (informal designation). Type material.— Holotype (Figs. 2A-B), a female, in CAS, labeled: “MADAGASCAR Ranomafana NP. Talatakely, X, 130 m. 15.1.2001”/ “collected in pitfall trap on forest floor, F. Rata- lata & J. Rainio collectors”/ “32.” [handwritten label]/ “HOLOTYPE Brachypelus ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, ranomafanae , is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition.— Large for genus, SBL = 6.8 mm. Members of this species (Fig. 2A) are most similar to those of Brachypelus obesus Putzeys (1866), with which they share large size, only six setae on the labrum, and a single discal setiferous pore on elytral interval 3 (at apical one-fifth). They differ from B. obesus members in the following: transverse fronto-clypeal suture present and deeply impressed (Fig. 3A) (absent from B. obesus members); frons with longitudinal frontal fur¬ rows narrower and shallower than in B. obesus ; posterior lateral margins of the eyes broadly round¬ ed (more narrowly and angularly rounded in B. obesus members); elytral base (Fig. 3B) straight between intervals 3 and obliquely sloped laterally from interval 4 (elytral base straight between intervals 4 and gradually rounded laterally from interval 3 in B. obesus members); elytra with lat¬ eral margin distinctly toothed at humerus (smoothly rounded at humerus, without a tooth in B. obe¬ sus members); and elytral intervals slightly convex apically (nearly flat in B. obesus members). Others species with members of comparably large size include Brachypelus rolandi Bulirsch et al. (2005), Brachypelus betsileo Bulirsch et al. (2005), Brachypelas fischeri Bulirsch et al. (2005) and Brachypelus janaki Bulirsch and Moravec (2009). However, B. ranomafanae members differ from those of B. rolandi in having the labrum with only six setae (seven in B. rolandi members), elytra widest at middle (anterior to middle in B. rolandi members), and elytral intervals moderately con¬ vex throughout (flattened apical in B. rolandi members). Members of both B. betsileo and B. fis- 206 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 MADAGASCAR Ranomafena MR. Talatafcety,X, 130 m. 16.1.2001 collected in pitlail trap on forest tlijpr. F. k.itakitj and 1. Raiino collectors B Figure 2. Digital images of holotype female of Brachypelus ranomafanae Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels. Scale line = 1.0 mm. cheri have seven labral setae (whereas those of B. ranomafanae have only six) and lack discal setif- erous pores (those of B. ranomafanae have a single setiferous pore in the apical one-fifth on inter¬ val 3. Finally, B. ranomafanae members differ from those of B. janaki in having only six setae on the labrum (seven labral setae in B. janaki members), the elytral margin with a distinct humeral tooth (tooth absent from B. janaki members), and only a single discal setiferous pore on elytral interval 3, with intervals 5 and 7 without such punctures (five or more such punctures on intervals 3, 5 and 7 in B. janaki members). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The unique holotype was collected in a pitfall trap placed in sec¬ ondary montane rain forest in the Talatakely area, at an elevation of 900 m, at the 130 meter mark along Trail “X”. Vegetation in that area consisted of bamboo, small young trees and some larger trees with diameters at breast-height (DBH) of 20-30 cm. Tree trunks and branches were mostly covered with epiphytes and lianas. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 207 Figure 3. Digital images of holotype female of Brachypelus ranomafanae Kavanaugh & Rainio sp. nov. A. Head, dor¬ sal aspect; B. Elytral base, dorsal aspect. Scale lines = 0.5 mm. Cratocerinini Lacordaire 1854 Phylogenetic relationships among the “Pterostichini” in the broadest sense and the cratocer- ines in particular remain largely unresolved (see Grzymala and Will (2014) for an example). We follow the classification as presented in Lorenz (2005). Substribe Drimostomatina Chaudoir 1872 Genus Caelostomus MacLeay 1825 Caelostomus latus Kavanaugh and Rainio, sp. nov. Figure 4 Type material.— Holotype (Figs. 4A-B), a female, in CAS, labeled: “MADAGASCAR Ranomafana NP. Talatakely, Trail C, 350 m. 10.5.2005”/ “HOLOTYPE Caelostomus latus Kavanaugh & Rainio sp. n. 2015” [red label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. 208 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 MADAGASCAR Ranomafana NP. TaUtakety, trail C 350 m. 10.5.2005 Figure 4. Digital images of holotype female of Caelostomus latus Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels; C. Head, left lateral aspect (arrow points to tranverse depression between eyes); D. Right anterior tarsus, dorsal aspect. Scale lines A = 1.0 mm, C-D = 0.5 mm. Derivation of species name.— The species epithet, latus , is the Latin adjective meaning broad or wide, a reference to relatively broad pronotum of the holotype specimen and, presumably, other members of this species. Recognition.— Size moderate for genus, SBL = 6.3 mm. Based on Jeannel’s (1948) key, this species is a member of subgenus Caelostomus s. str. It’s members (Fig. 4A) share with those of other members of the subgenus the following features: head with frons and frontal furrows smooth, impunctate; elytral base distinctly margined from humerus medially to base of stria 3 or 2; elytral striae 6 and 7 deeply impressed throughout, not effaced apically; lateral elytral intervals narrow, moderately convex; apical part of umbilicate series of setiferous pores not inserted in a deep groove; and female apical abdominal ventrite with (two or) three pairs of setae subapically. With¬ in the subgenus, the following features are shared only with members of two species of JeanneTs convexiuscuius group, namely Caelostomus convexiuscuius Tschitscherine (1899) and Caelosto- KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 209 mus alluaudi Jeannel (1948): SBL greater than 6.0 mm, elytra short and broad, ratio EL/EW = I. 19, elytral disc convex; and elytral intervals distinctly punctate throughout, including in apical part. The holotype female of C. latus is slightly larger than members of these other two species, has a distinct and broad transverse depression between the eyes seen best in lateral view (Fig. 4C) (depression absent from members of the other two species). The pronotum in the holotype of C. latus is much wider than in C. convexiuscuius members, slightly wider than in C. alluaudi mem¬ bers, and less depressed anterolaterally than in members of either of the other species; and the pronotal basal foveae are straighter and more sharply defined laterally than in members of the other two species. The elytra are less abruptly rounded laterally and apically than in C. alluaudi mem¬ bers but similar to shape in to those in C. convexius cuius members. Finally, anterior tarsomeres 1 to 3 of the female holotype of C. latus are distinctly toothed medioapically (Fig. 4D), a feature not seen in females of the other two species. Geographical distribution. — At present, known only from the type locality. Habitat distribution. — The unique holotype female was collected by hand in secondary montane forest in the Talatakely area. Caelostomus rotundiformis Kavanaugh and Rainio, sp. nov. Figure 5 Caelostomus n. sp. 7; Rainio 2009: 31, Rainio 2012: 62 (informal designation). Type material. — Holotype (Figs. 5A-B), a female, in CAS, labeled: “MADAGASCAR Ranomafana NP., Vatoharanana, Trail K at 50 m mark, 07.06.2000”/ “collected by hand in daytime, F. Ratalata & J. Rainio collectors”/ “13.” [handwritten label]/ “7.6.00 K 50” [handwritten label]/ “HOLOTYPE Caelostomus rotundiformis Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (only 1): a female (in MNHN) labeled “MADAGASCAR Ranomafana NP. Talatakely, C1450 II. 3.2004 Johanna Rainio leg.”/ “112.” [handwritten label]/ “PARATYPE Caelostomus rotundi¬ formis Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, rotundiformis, is an adjective derived from the Latin words, rotundus, meaning round, and forma, meaning form or shape. The name refers to the very short, broad, rounded form of members of this species. Recognition. — Size moderately small for genus, SBL = 5.4-5.5 mm (females only), body form (Fig. 5A) short and broad, ratio SBL/EW = 1.97-2.01. This species is undoubtedly a member of genus Caelostomus as conceived by Jeannel (1948), although we cannot confirm this based on his key because he used male features only in two couplets to distinguish Caelostomus members from those of several other genera and we have only female specimens. However, we examined specimens of all species in all the other Malagasy caelostomine genera while at MNHN and con¬ firmed that this species represents none of them. Observable female features, including body fonn short and broad, frontal furrows simple, not doubled, parascutellar striae of elytra indistinct, basal setiferous pore situated at base of elytral stria 3, tarsomeres not densely setose ventrally, and ante¬ rior tarsomere 4 not bilobed apically (Fig. 5C), are consistent with Caelostomus females. Within this genus, members of C. rotundiformis share the following features with those of other species of the nominate subgenus: head with frons and frontal furrows smooth, impunctate; elytral base dis¬ tinctly margined from humerus medially to base of stria 3 or 2; elytral striae 6 and 7 deeply impressed throughout, not effaced apically; lateral elytral intervals narrow, moderately convex; apical part of umbilicate series of setiferous pores not inserted in a deep groove; and female apical abdominal ventrite with (two or) three pairs of setae subapically. However, members of this species 210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 MADAGASCAR, Ranomafana NP., VatoharanihiL Trail K at 50 m mark, 07.06.2000 B Figure 5. Digital images of holotype female of Caelostomus rotundiformis Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels; C. Right anterior tarsus, dorsal aspect. Scale line A = 1.0 mm, C = 0.5 mm. confound Jeannel’s (1948) key to species of Caelostomus s. str. Their short and broad elytra (ratio EL/EW = 1.13-1.15) are shared with members of the convexiuscuius species group, but their ely- tral striae are impuncate, whereas those of other species group members are markedly punctate throughout their length. Within the group, their small size (less than 6.0 mm) is shared with Caelostomus humilis (Tschitscherine 1903), Caelostomus minisculus Straneo (1940), and Caelostomus minutissimus Jeannel (1948); however, members of these three species are even smaller (SBL = 5.0 mm or less) and, again, have markedly punctate elytral striae. The striae in C. rotundiformis members are smooth, without evident punctures, and the shape of the pronotum KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 211 (Fig. 5A) is markedly different, with lateral margins evenly arcuate throughout their length and hind angles markedly obtuse. This species probably represents a distinct species group of its own within subgenus Caelostomus in Madagascar. Geographical distribution. — At present, known only from the type locality. Habitat distribution. — The holotype of this species was collected by hand in daytime in primary montane rain forest in the Vatoharanana area at an elevation of 1200 m, along Trail K at the 50 meter mark. The paratype specimen was collected in a pitfall trap in secondary montane rain forest in the Talatakely area at an elevation of 900 m, along Trail C at the 1450 meter mark. Genus Dactyleurys Tschitscherine 1899 Jeannel (1948) recognized a single species in Madagascar, Dactyleurys anomalus Tschitscherine (1899) and no additional species have been described subsequently. The two new species described below bring the total for the genus to three species, all precinctive to Masdagas- car. Dactyleurys minimus Kavanaugh and Rainio, sp. nov. Figures 6, 8A, 9A, 9D Type material. — Holotype (Figs. 6A-B), a male, in CAS, labeled: “MADAGASCAR, Ranomafana NP, Talatakely, X at 130 m mark 12.4.2005”/ “collected by hand in daytime, F. Rata- lata & Johanna Rainio collectors”/ “HOLOTYPE Dactyleurys minimus Kavanaugh & Rainio sp. n. 2015” [red label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation or species name. — The species epithet, minimus , is the Latin adjective meaning smallest, a reference to fact that the known adult male of this species is smaller than males of any of the other known species that we have examined. Recognition.— Smallest adult known for the genus, SBL = 6.9 mm. The following combi¬ nation of features identify members of this species as coelostomine pterostichines of the genus Dactyleurys : antennae with antennomere 3 attached symmetrically or nearly symmetrically to antennomere 2; mentum with deep emargination, apex of median mental tooth distinctly posterior to apices of epilobes; elytron with epipleuron interrupted by internal plica subapically; parascutel- lar stria not evident; basal setiferious puncture situated at base of stria 3; tarsomeres 1 to 4 of all legs with dense pads of setae ventrally; front and middle tarsomeres 1 to 3 smoothly convex dor- sally; male front tarsomeres 1 to 4 wide, dilated, not latero- or medioapically toothed; aedeagus of male with right face dorsal in repose, right paramere conchoid, left paramere reduced, short, api- cally digitifonn. Members of D. minimus (Fig. 6A) differ from those of D. anomalus in the fol¬ lowing features: smaller size (SBL = 6.9 mm compared with 1.1-9A mm in D. anomalus ); prono- tum (Fig. 8A) relatively narrower, widest at middle (relatively wider (Fig. 8C) and clearly widest anterior to middle in D. anomalus ); elytra faintly but evidently iridescence and elytral microsculp¬ ture markedly transverse with sculpticells great than 2.5 times as wide as long (elytra not irides¬ cent and elytral microsculpture moderately transverse with sculpticells less than 2.5 times as wide as long in D. anomalus ); and median lobe of male genitalia (Fig. 9A) with shaft abruptly bent ven¬ trally in apical fourth with apex nearly straight in lateral aspect (shaft nearly straight ventrally (Fig. 8C) in apical fourth with apex slightly recurved dorsally in D. anomalus ), apex narrowly rounded in dorsal aspect (Fig. 9D) (more broadly rounded (Fig. 9F) in D. anomalus ). Members of D. min¬ imus differ from those of the other new species, Dactyleurys ranomafanae , in the following fea- 212 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 MADAGASCAR Ranomafaiia NP. Ta1atakely,*Trai1 X at 130 m mark, , 12.4.2005 __ collected by hand in daytime. * F. Ratalala and J. Raimo collectors B Figure 6. Digital images of holotype male of Dactyleurys minimus Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels. Scale line = 1.0 mm. tures: smaller size (SBL = 7.7-8.3 mm in D. ranomafanae ); pronotum (Fig. 8A) relatively nar¬ rower and with evenly rounded lateral margin (slightly wider (Fig. 8B) and with lateral margin slightly straighter in basal one-third in D. ranomafanae ); and median lobe of male genitalia (Fig. 9A) with apex nearly straight in lateral aspect (apex slightly recurved dorsally (Fig. 9C) in D. ranomafanae ), apical part of shaft (Fig. 9D) slightly deflected right in dorsal aspect (apical part of shaft (Fig. 9E) straight in D. ranomafanae). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The unique holotype was collected by hand in daytime in second¬ ary montane rain forest in the Talatakely area, at an elevation of 900 m, at the 130 meter mark along Trail “X”. Vegetation in that area consisted of bamboo, small young trees and some larger trees with diameters at breast-height (DBH) of 20-30 cm. Tree trunks and branches were heavily laden with epiphytes and lianas. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 213 Dactyleurys ranomafanae Kavanaugh and Rainio, sp. nov. Figures 7, 8B, 9B, 9E Dactyleurys n. sp. 1; Rainio 2009: 31, Rainio 2012: 62, Rainio 2013: 95 (informal designation). Type material.— Holotype (Figs. 7A-B), a male, in CAS, labeled: “CASENT 1009744”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 26 April 1998”/ “21°15.3’S/ 47°25.9’E, Stop #98-83 D.H. Kavanaugh col¬ lector collected by beating Pandanus tree boluses with debris”/ “HOLOTYPE Dactyleurys ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (a total of 35): 1 male (in MZF) labeled “MADAGASCAR Ranomafana NP. Mangevo, plot 0 m., 21.4.2005”; 1 female (in MZF) labeled “MADAGASCAR Ranomafana NP. Mangevo, plot 100 m., 21.4.2005”; 2 females (in CAS) labeled same as holotype except “CASENT 1009742” and “CASENT 1009743”, respec¬ tively; 3 males and 2 females (in CAS) labeled “CASENT 1009737”, “CASENT 1009738”, “CASENT 1009739”, “CASENT 1009740” and “CASENT 1009741”, respectively/ “MADAGAS¬ CAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 25 April 1998”/ “21°15.3’S/47°25.9’E, Stop #98-78 D.H. Kavanaugh collector beaten from vegetation and suspended dead leaves”; 3 males and 2 females (in CAS and MNHN) labeled “CASENT 1009734”, “CASENT 1009735”, “CASENT 1009736”, “CASENT 1009732” and “CASENT 1009733”, respectively/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanaugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Ravelo- son collectors”/ “collected in daytime by beating live understory vegetation and/or dead leaves and other suspended debris”; 1 male and 1 female (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, 26.2.2001”; 1 female (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail B 1150 m., in Pandanus sp. 8.6.2000”; 1 male and 1 female (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail BF 380 m., in Pandanus sp. 16.6.2000”; 1 male and 1 female (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail BM 200 m., in Pandanus sp. 16.6.2000”; 1 male and 2 females (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail FBF 50 m., in Pandanus sp. 6.6.2000”; 1 female (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail FBF 380 m., in Pandanus sp. 6.6.2000”; 2 females (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail SP 50 m., 22.3.2001”; 1 female (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail SP 50 m., 9.2.2005”; 1 male (in MZF) labeled “MADAGASCAR Ranomafana NP. Talatakely, trail X 130 m., 12.4.2005”; 1 male (in MNHN) labeled “MADAGASCAR Ranomafana NP. Valohoaka, trail F 900 m., 28.2.2001”; 1 male (in MZF) labeled “MADAGASCAR Ranomafana NP. Valohoaka, trail F 850 m., 16.10.2002”; 1 male (in MNHN) labeled “MADAGASCAR Ranomafana NP. Valohoa¬ ka, trail F 850 m down., 18.12.2004”; 1 female (in MNHN) labeled “MADAGASCAR Ranomafana NP. Valohoaka, trail F 850 m., 25.1.2005”; 2 males and 1 female (in MNHN and MZF) labeled “MADAGASCAR Ranomafana NP. Valohoaka, trail F 850 m., 26.1.2005”. All paratypes also bear the following label: “PARATYPE Dactyleurys ranomafanae Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, ranomafanae, is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition.— Size average for genus, SBL of males = 7.7-8.2 mm, of females = 7.8-8.4 mm . The following combination of features identify members of this species as coelostomine 214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 CASJENT 1009744 i MADAGASCAR, fianarantsoa Province. Ranomafana Nali ona) Pa rk, *! alalakety area. 900 m, mixed tropical foresl, 26 Annl 199K, _ 21 I5.CS 47 25.9'!-:. Sion ft 98-83 D.l 1. Kavanadkh collector collected by beating Pandanus tree boluses with debris Figure 7. Digital images of holotype male of Dactyleurys ranomafanae Kavanaugh & Rainio sp. nov. A. Habitus, dor¬ sal aspect; B. Labels. Scale line =1.0 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 215 pterostichines of the genus Dactyleurys : antennae with anten- nomere 3 attached symmetrically or nearly symmetrically to antennomere 2; mentum with deep emargination, apex of median mental tooth distinct¬ ly posterior to apices of epilobes; elytron with epipleuron interrupted by internal plica subapically; paras- cutellar stria not evident; basal setif- erous puncture situated at base of stria 3; tarsomeres 1 to 4 of all legs with dense pads of setae ventrally; front and middle tarsomeres 1 to 3 smoothly convex dorsally; male front tarsomeres 1 to 4 wide, dilated, not latero- or medioapically toothed; aedeagus of male with right face dor¬ sal in repose, right paramere con¬ choid, left paramere reduced, short, apically digitiform. Members of D. ranomafanae (Fig. 7A) differ from those of D. anomalus in the following features: pronotum (Fig. 8B) widest at middle (distinctly widest anterior to middle (Fig. 8C) in D. anomalus ); elytra faintly but evidently irides¬ cence and elytral microsculpture markedly transverse with sculpticells great than 2.5 times as wide as long (elytra not iridescent and elytral microsculpture moderately transverse with sculpticells less than 2.5 times as wide as long in D. anomalus ); and median lobe of male genitalia (Fig. 9B) with shaft abruptly bent ventrally in apical fourth (shaft nearly straight ventrally (Fig. 8C) in apical fourth in D. anomalus ), apex narrowly rounded in dorsal aspect (Fig. 9E) (more broadly rounded (Fig. 9F) in D. anomalus ). Members of D. ranomafanae differ from those of the other new species, D. minimus, in the following features: larger size, SBL = 7.7-8.3 mm (size smaller, SBL = 6.9, in D. minimus)', pronotum Figure 8. Digital images of pronota of Dactyleurys spp., dorsal aspect. A. Holotype of D. minimus Kavanaugh & Rainio sp. nov.; B. Holotype of D. ranomafanae Kavanaugh & Rainio sp. nov.; C. D. anomalus Tschitscherine. Scale lines = 1.0 mm 216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Figure 9. Digital images of aedeagus of male genitalia of Dact\>leury>s spp. A-C. left lateral aspect; D-F. dorsal aspect. A, D. Flolotype of D. minimus Kavanaugh & Rainio sp. nov.; B, E. Holotype of D. ranomafanae Kavanaugh & Rainio sp. nov.; C, F. D. anomalus Tschitscherine. Scale lines = 0.5 mm. (Fig. 8B) relatively broader and with lateral margins slightly straighter in basal one-third (slightly narrower and with lateral margins more evenly rounded (Fig. 8A) in D. minimus ); and median lobe of male genitalia (Fig. 9B) with apex slightly recurved dorsally in lateral aspect (apex (Fig. 9A) nearly straight in D. minimus), apical part of shaft (Fig. 9E) straight (apical part of shaft slightly deflected right (Fig. 9D) in D. minimus ). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— Specimens of this species have been collected in both primary and secondary montane rainforest throughout RNP, at elevations ranging from of 900 to 1180 m. Most of the specimens were collected by beating vegetation, especially plants of Pandanus sp., and sus¬ pended accumulations of dead vegetative debris in shrubs and understory trees. One was collected in a malaise trap, another sifted from forest litter at the primary forest site, and another by hand at the secondary forest site. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 217 Genus Trichinillus Straneo 1938 Subgenus Mallopelmus Straneo 1942 Mallopelmus Alluaud 1936, unavailable Alluaud (1936) described this taxon as a new genus with several included species, but he failed to designate one of them as type species. This error rendered the name unavailable (ICZN 2000, Article 13.3) until Straneo (1942) designated Mallopelmus dactyleuryoides Alluaud (1936) (the first named species in Alluaud’s (1936) paper) as the type species. Both Straneo (1938) and Jean- nel (1948) treated Mallopelmus as a distinct genus. Straneo actually treated Trichinillus as a sub¬ genus of Mallopelmus in his 1942 paper, apparently not realizing at the time that his designation of a type species for Mallopelmus changed the data of availability for that name, as well as its authorship, with the result that Trichinillus became the senior synonym for the genus. Mallopelmus is now considered a subgenus of Trichinillus (Lorenz 2005) and is precinctive to Madagascar. Trichinillus ( Mallopelmus ) ranomafanae Kavanaugh and Rainio, sp. nov. Figure 10 Mallopelmus n. sp.l ; Rainio 2009: 32, Rainio 2012: 74 (informal designation). Type material. — Holotype (Figs. 10A-B), a male, in CAS, labeled: “MADAGASCAR Ranomafana NP. Buffer zone, degraded forest 14.10.2002”/ “collected by hand in daytime, F. Rata- lata & J. Rainio collectors”/ “HOLOTYPE Trichinillus ( Mallopelmus ) ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (only 1): a female (in MNHN) labelled “MADAGAS¬ CAR Ranomafana NP. Talatakely, trail B, 600 m. 05.03.2004”/ “collected by hand in daytime from dead leaves in bushes, F. Ratalata and J. Rainio collectors”/ “PARATYPE Trichinillus {Mallopel¬ mus) ranomafanae Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, ranomafanae, is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition.— Slightly smaller than average for subgenus, SBL male = 7.2 mm, female = 7.5 mm. The following combination of features identify members of this species as coelostomine pterostichines of the genus Trichinillus, subgenus Mallopelmus : antennae with antennomere 3 attached symmetrically or nearly symmetrically to antennomere 2; mentum with deep emargina- tion, apex of median mental tooth distinctly posterior to apices of epilobes; elytron with epipleu- ron interrupted by internal plica subapically; parascutellar stria short but evident; striae smooth, impunctate; basal setiferious puncture situated at base of stria 2 + parascutellar stria; tarsomeres 1 to 4 of all legs with dense pads of setae ventrally; front and middle tarsomeres 1 to 3 with more or less faintly-defined longitudinal groove medially and depressed areas paralaterally on dorsal sur¬ face; male front tarsomeres 1 to 4 wide, dilated, not latero- or medioapically toothed; aedeagus of male with right face dorsal in repose, right paramere conchoid, left paramere reduced, short, api- cally digitiform. Of the three previously described species represented in JeanneTs (1948) key to Mallopelmus species, members of T. ranomafanae (Fig. 10A) are most similar to Trichinillus abacetoides (Alluaud 1936) members based on the following shared features: head with temporal area long, nearly as long as the diameter of the eye, and joined to the neck region at a very obtuse (about 165°) angle; eyes moderately convex and projected; pronotum with basal area distinctly and abruptly depressed relative to disc; and elytral base without margination medial to base of stria 6. Members of the other two species, Trichinillus dactyleuryoides (Alluaud 1936) and Trichinillus perrieri (Jeannel 1948), differ in having the head with the temporal area shorter, about one-third as 218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 MADAGASCAR Ranomafana NP. Buber zone' degraded forest , 14 . 10.200 2 . _ collected by hand in daytime, • F. Ra labia and J. Ramio collectors B D Figure 10. Digital images of holotype male of Trichinillus ( Mallopelmus ) ranomafanae Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels; C-D. Aedeagus of genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines A = 1.0 mm, C-D = 0.5 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 219 long as the diameter of the eye, and joined to the neck region at a moderately obtuse (about 135°) angle, eyes more convex and projected, pronotum not or less depressed, especially medially, and margination of the elytra base more varied, extended medially to the base of stria 3 in most indi¬ viduals). The apex of the median lobe of the male genitalia is also differently shaped in each of these species (compare with Figs. 191c-d and 191 f-g, respectively, in Jeannel 1948). Members of T. ranomafanae differ from those of T. abacetoides in having: slightly larger size (SBL = 6.4 mm in T. abacetoides ); pronotum with shallow but distinct subbasal sinuation of the lateral margin (absent from T. abacetoides members); elytra relatively shorter and wider, especially across the base, and humeri broadly rounded and not at all sloped (humeri more slope in T. abacetoides , and median lobe of male genitalia with apex straighter, slightly bent ventrally toward apex in lateral aspect (Fig. IOC) and slightly deflect right in dorsal view (Fig. 10D) (not recurved dorsally and then ventrally in lateral aspect and straight in dorsal aspect as in T. abacetoides males; compare with Fig. 19le in Jeannel 1948) Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The male holotype of T. ranomafanae was collected in degraded secondary montane rainforest at an elevation of 800 m, near a village in the peripheral zone of Ranomafana National Park aboutlOOO meters from the main gate. The female paratype was col¬ lected in secondary montane rainforest in the Talatakely area, at an elevation of 900 m, at the 600 m mark along Trail B. Vegetation in this area was dominated by young trees (DBH <10 cm) and guava ( Psidium cattleianum). Tribe Chlaeniini Brulle 1834 Genus Chlaenius Bonelli 1810 Jeannel (1949) treated Chlaenites Motschulsky (1860) as a distinct genus, although it is now considered a subgenus of Chlaenius (Lorenz 2005) and includes only two species, both from the Palaearctic Region. Both of the new species described below key to Chlaenites in Jeannel’s (1949) key to chlaeniine genera, but neither is a member of that subgenus as presently conceived. Chlaenius kathrynae Kavanaugh and Rainio, sp. nov. Figure 11 Type material.— Holotype (Figs.llA-B), a male, in CAS, labeled: “CASENT 1002782”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, 1150 m, mixed tropical forest, 2-22 January 2001,”/ “21.24032°S/ 47.39399°E, Stop# DHK-01-002, D.H. & K.M. Kavanaugh, R.L. Brett, E. Elsom, F. Vargas, Ranaivosolo,”/ “E.F. Randrianifirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected in daytime by treading marsh vegetation and shore”/ “HOLOTYPE Chlaenius kathrynae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 8): 4 males (in CAS, MNHN and MZF) and 1 female (in CAS) labeled same as holotype except “CASENT 1002783”, “CASENT 1002784”, “CASENT 1002786”, “CASENT 1002787”, and “CASENT 1002785”, respectively; 1 male (CAS) labeled “CASENT 1049008”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, 1050 m, mixed tropical forest, 23 April 1998,”/ “21°13.6’S/ 47°23.0’E,”/ “Stop # 98-71, D.H. Kavanaugh collector collected in soil cracks, under dirt clods, and under root clods in abandoned rice paddy area”; 2 females (in MTEC) labeled “MADAGASCAR: Fianaran. Pr. Ranomafana N.P. HQ. area 21°15’24”S 47°25’15”E 25NOV1994, at night M.A. Ivie & D.A. Pol¬ lack”. All paratypes also bear the following label: “PARATYPE Chlaenius kathrynae Kavanaugh & Rainio sp. n. 2015” [yellow label]. 220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 CASENT B 1002782 MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, 1150 m, mixed tropical Forest, 2-22 January 20 01* 2L24032°S/47:39399°E, Stop# DHK-01-002 D.H, Sl K.M. Kfcvanaugh, R.L, Brett, E* Elsom, F. Vareas. R Ranaivosolo. E,F. Randrianirina, N. Rasoaman an an a, T.J. Ravelomanana* and H.C, Raveloson collectors collected in daytime by treading marsh vegetation and shore Figure 11. Digital images of holotype male of Chlaenius kathiynae Kavanaugh & Rainio sp. nov. A Habitus, dorsal aspect; B. Labels; C-D. Aedeagus of genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines =1.0 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 221 Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation ol species name.— The species epithet, kathrynae, is a noun in the genitive case, derived from the first given name of Kathryn May Kavanaugh, daughter of DHK and one of the collectors of the type series. Recognition.— Size slightly below average for genus, SBL of males = 10.2-10.8 mm, of females 9.8-11.1 mm. Members of this species (Fig. 11 A) share the following features, which place them in genus Chlaenites sense Jeannel (1948): antennomere 3 longer than antennomere 4; maxillary palpi setose, penultimate labial palpomeres 4 or 5 setose; pronotum with basolateral setae inserted distinctly anterior to the hind angles; elytra with pubescence restricted to near the striae on disk (i.e., not present on the centers of intervals), except more generally and densely present on lat¬ eral intervals and near apices; and tarsi asetose dorsally, tarsomeres 5 with two rows of stout setae ventrally. In JeanneTs (1949) key to subgenera of Chlaenites, members of this species key best, although not fully, to subgenus Chlaeniostenus Kuntzen (1919), which is now considered a sub¬ genus of Chlaenius (Lorenz 2005). Shared with other Malagasy members of this subgenus are the following features: pronotum relatively narrow, more or less cordiform, narrowed basally, with the anterior angles not projected anterior of the basal margin and the anterolateral areas of the prono¬ tum distinctly curved ventrally; and the setose punctures along the lateral margins of the elytral intervals not or only faintly evident, most clearly evident in the apical half only. Jeannel included three other species in this subgenus; namely, Chlaenius attenuatus Klug (1833), Chlaenius subo- vatus Chaudoir (1876), and Chlaenius sellatus Dejean (1831). Members of C. kathrynae differ from those of the other species in lacking any trace of the pale lateral and apical elytral border dis¬ tinct in members of the other three species. They are also smaller than members of the other species (SBL in the latter ranges from 11.9 to 15.3 mm), have slightly broader and less cordiform pronota and males have genitalia (Figs. 11C-D) that are very different in form from those of any of the other species (compare with JeanneTs (1949) Figs. 388c-i) Geographical distribution.— At present, known only from the type locality. Habitat distribution.— Most specimens of the type series were collected in open habitats at the edges of mixed tropical forest in the Vohiparara area at an elevation of 1050 m. Several were collected by treading down vegetation at the edge of a marsh, and one specimen was found under loose dirt and root clods in an abandoned rice paddy area. Chlaenius robertae Kavanaugh and Rainio, sp. nov. Figure 12 Type material.— Holotype (Figs.l2A-B), a male, in CAS, labeled: “CASENT 1002781”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, 1150 m, mixed tropical forest, 2-22 January 2001,”/ “21.24032°S/ 47.39399°E, Stop# DHK-01-002, D.H. & K.M. Kavanaugh, R.L. Brett, E. Elsom, F. Vargas, Ranaivosolo,”/ “E.F. Randrianifirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected in daytime by treading marsh vegetation and shore”/ “HOLOTYPE Chlaenius robertae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (only 1): a female (in CAS) labeled “CASENT 1049009”/ “MADA¬ GASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, 1050 m, mixed tropical forest, 27 April 1998,”/ “21°13.6’S/ 47°23.0’E,”/ “Stop # 98-91, D.H. Kavanaugh collec¬ tor collected on bare soil at night in abandoned rice paddy area”/ “PARATYPE Chlaenius robertae Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, robertae, is a noun in the genitive case, 222 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 B CASENT 1002781 MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, i 150 m, mixed tropical forest, 2-22 January 2001, 21.24032“S/ 47.39399°E, Stop# DHK-01-002 D,H. & K.M. F&vanaugh, RX. Brett, E, Elsom, F. Vargas. R, Ranaivosolo, E.F. Randrianirina, N. Rasoamananana, T.J. Rave Ionian an a, and H,C. plaveloson collector" collected in daytime by treadinghmrsh vegetation and shore Figure 12. Digital images of holotype male of Chlaenius robertae Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels; C-D. Aedeagus of genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines =1.0 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 223 derived from the given name of Roberta L. Brett, former graduate student and technical assistant of DHK and one of the collectors of the type series. Recognition.— Sized moderate for genus, SBL of male = 12.9 mm, of female =13.3 mm. Members of this species (Fig. 12A) exhibit the following features, which place them in genus Chlaenites sensu Jeannel (1949): antennomere 3 longer than antennomere 4; maxillary palpi setose; penultimate labial palpomeres 4 or 5 setose; pronotum with basolateral setae inserted dis¬ tinctly anterior to the hind angles; elytra with pubescence restricted to near the striae on disk (i.e., not present on the centers of intervals), except more generally and densely present on lateral inter¬ vals and near apices; and tarsi asetose dorsally, tarsomeres 5 with two rows of stout setae ventral- ly. In Jeannel’s (1949) key to subgenera of Chlaenites, members of this species key to his new sub¬ genus Chlaenitidius, which is now considered a junior synonym of subgenus Amblygenius Lafer- te-Senectere (1851) (Lorenz 2005), in having the pronotum relatively broad and the setose punc¬ tures along the lateral margins of the elytral intervals slightly foveate, especially in the apical half apically. Member s of C. robertae differ distinctly from those of the each of the four species Jean¬ nel included in this subgenus. They differ from members of Chlaenius cupreolus Faimaire (1901) in having striae deeply impressed and intervals moderately convex (striae superficial only and intervals flat in C. cupreolus members), head with metallic green reflection but pronotum black without a trace of metallic reflection (both head and pronotum with green metallic reflection in C. cupreolus members), and elytra without a pale lateral and apical margin (a thin pale margin pres¬ ent in C. cupreolus members). They differ from members of Chlaenius allacteus Alluaud (1919), which is current classified in subgenus Oochlaenius Alluaud (1933) (Lorenz 2005), in having all elytral interval similarly and moderately convex (intervals 1, 3, 5 and 7 more convex and intervals 2, 4, 6 and 8 flat in C. allacteus members) and, again, elytra without a pale lateral and apical mar¬ gin (a wide pale margin present in C. allacteus members). They differ from members of Chlaenius inaequalis Faimaire (1901) in having the pronotum black without metallic reflection (metallic green reflection present on pronotum in C. inaequalis members), a longer and narrower elytral form (shorter and broader elytra in C. inaequalis members) and elytral epipleurae black (pale in C. inaequalis members). Overall, members of C. robertae are most similar to those the fourth species, Chlaenius lyperus Jeannel (1949), with which they share similar elytral shape, depth of striae and convexity of intervals. However, they differ from C. lyperus members in having a prono¬ tum broader basally than the latter and legs that are pale in color (dark in C. lyperus members). Males of C. robertae also have genitalia (Figs. 12C-D) that are markedly different in form those of the other four species (compare with Jeannel’s (1949) Fig. 386a-d). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— Both specimens of the type series were collected in open habitats at the edges of mixed tropical forest in the Vohiparara area at elevations ranging from 1050 to 1150 m. One was collected in daytime by treading down vegetation at the edge of a marsh, the other was found at night running on bare soil in an abandoned rice paddy. Tribe Hexagoniini Horn 1881 Genus Omphreoides Fairmaire 1896 This genus, precinctive to Madagascar, includes five previously described species. Jeannel (1949) described a new genus, Stenomphreoides, for Omphreoides quodi Alluaud (1910), but this generic name is now considered just a junior synonym of Omphreoides (Lorenz 2005). 224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Omphreoides ranomafanae Kavanaugh and Rainio, sp. nov. Figure 13 Type material.— Holotype (Figs.l3A-B), a female, in NMNH, labeled: “MADAGASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 900 m 17-22 February 1990 W. E. Steiner”/ “at black light in montane rainforest near river and stream”/ “HOLOTYPE Omphreoides ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, ranomafanae, is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition.— Size larger than average for genus, SBL =13.4 mm. The unique holotype female of O. ranomafanae (Fig. 13A) is clearly unlike members of any described species of Omphreoides, with several features intermediate between the four typical species and O. quodi. The head of the holotype (Fig. 13C) is longer and more slender than that of females of any other species except O. quodi, which has an even longer and narrower head (see Jeannel 1949, Fig. 364). In O. ranomafanae, the tempora are straight, parallel in anterior their two-thirds, slightly more than twice as long as the diameter of eye, distinctly delimited posteriorly by narrowly round angles. The dorsal longitudinal grooves typical of all members of this genus, except those of Omphreoides bispinus Fairmaire (1896), are extended as sharply-defined grooves only about halfway from the point of insertion of the anterior supraorbital setae to that of posterior supraorbital setae and not continued on to base of the head as they are in Omphreoides bucculentus Alluaud (1899a) and Omphreoides distinctus Alluaud (1936). Posterior to the sharply-defined segments, broad and shal¬ low depressions extend posteriorly, arcuately convergent and nearly joined in the midline near the back of the head. The pronotum (Fig. 13C) is subquadrate, longer than wide (ratio PW/PL = 0.75), with a wavy diagonal row of sparse, coarse punctures (seen also in O. distinctus and O. quodi mem¬ bers but absent from those of O. bucculentus and O. bispinus ) extended from near the anterior transverse impression paramedially to the basal foveae posteriorly; lateral pronotal margins with short but distinct sinuations anterior to slightly obtuse hind angles; median longitudinal impression deep, wider than in all other Omphreoides except O. quodi (in which the impression is even wider) and sparsely but coarsely punctate; lateral explanation narrow (narrower at middle than that in O. bucculentus members and similar to that in O. distinctus member. The elytra are relatively wider than in all other species; each elytron with a straight apical spine in line with interval 3. Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The unique holotype was collected at ultraviolet light in montane rainforest near the junction of the Namorona River and a small tributary stream. Tribe Perigonini Horn 1881 Genus Perigona LaPorte 1835 Subgenus Ripogena Jeannel 1941a This subgenus includes ten described species, including five from the African mainland and five from Madagascar (Basilewsky 1989). Jeannel (1948) treated this taxon as a separate genus, although he had described it originally as a subgenus of Perigona (Jeannel 1941a), and included the four species known to him in a key. Deuve (1998) described a fifth Malagasy species. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 225 MADAGASCAR: Prov. Fianaranteoa, 7 k m W Ranomafana, 900 m 17-22 February 1990 W* E- Steiner At black light in montane rain forest near river and stream Figure 13. Digital images of holotype female of Omphreoides ranomafanae Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels; C. Forebody, dorsal aspect. Scale lines A = 1.0 mm, C-D = 0.25 mm 226 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Perigona ( Ripogena) deuvei Kavanaugh and Rainio, sp. nov. Figure 14 Ripogena n. sp. 1; Rainio 2009: 33 (informal designation). Perigona ( Ripogena ) n. sp. 1; Rainio 2012: 73, Rainio 2013: 96 (informal designation). Type material. — Holotype (Figs.l4A-B), a male, in CAS, labeled: “MADAGASCAR, Ranomafana NP., Vatoharanana, Trail K at 350 m mark, 24.07.2005”/ “collected by hand in day¬ time, F. Ratalata & J. Rainio collectors”/ “HOLOTYPE Perigona {Ripogena) deuvei Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 4): 1 female (in MNHN) labeled “MADAGAS¬ CAR, Ranomafana NP, Vatoharanana, Trail J at 50 m mark, 6.10.2005”/ “collected by hand in day¬ time, F. Ratalata & J. Rainio collectors”/ “239.” [handwritten label]; 1 female (in CAS) labeled “MADAGASCAR, Ranomafana NP, Vatoharanana, Trail S at 600 m mark, 9.9.2005”/ “collected by hand in daytime, F. Ratalata & J. Rainio collectors”/ “239.” [handwritten label]; 1 male (in MZF) labeled “MADAGASCAR, Ranomafana NP, Vatoharanana, Trail S at 600 m mark, 24.7.2005”/ “collected by hand in daytime, F. Ratalata & J. Rainio collectors”/ “253.” [handwrit¬ ten label]; 1 female (in MZF) labeled “MADAGASCAR, Ranomafana NP, Vatoharanana, Trail S at 600 m mark, 6.10.2005”/ “collected by hand in daytime, F. Ratalata & J. Rainio collectors”/ “245.” [handwritten label]. All paratypes also bear the following label: “PARATYPE Perigona {Ripogena) deuvei Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, deuvei, is a noun in apposition, derived from the surname of Dr. Thierry Deuve of the Laboratoire d’Entomologie at the Museum Nation¬ al d’Histoire Naturelle in Paris. We are pleased to name this new species in honor of Dr. Deuve in thanks for his friendship and kind hospitality during our several visits to MNHN in the course of this study. Recognition. — Size average for subgenus, SBL of males = 3.5-3.8 mm, of females 3.8-3.9 mm. Members of this species (Fig. 14A) share with those of Perigona heterodera Alluaud (1936) and Perigona prasinus Alluaud (1936) the following features: pronotum with lateral margins with¬ out sinuation anterior to hind angles, posterior margin slightly and smoothly convex, and hind angles either obtuse or broadly rounded. These features distinguish members of these three species from those of Perigona bembidioides Alluaud (1936) and Perigona viridimicans Jeannel (1948), which have distinct sinuation of the lateral margins anterior to rectangular hind angles and a straight basal margin. Members of P. deuvei differ from those of P. prasinus in having smaller body size (SBL = 4.7 mm in P. prasinus), narrower body form, and dorsum without metallic reflection (dorsum with metallic green reflection in P. prasinus) but elytra with moderate but distinct irides¬ cence generated by microsculpture comprised of transverse microlines and marjkedly transverse sculpticells. They are most similar to members of P. heterodera in body form and size but differ from them in having the pronotum proportionately longer and male genitalia (Figs. 14C-D) with shaft markedly narrowed basally, markedly inflated in apical two-thirds, and apex longer, slender and ventrally bent in lateral view (Fig. 14C), slightly deflected left and slightly longer in dorsal view (Fig. 14D) than in P. heterodera (see Figs. 351b-c in Jeannel 1948). They differ from mem¬ bers of Perigona descarpentriesi (Deuve 1998) in having moderately large eyes (eyes markedly reduced eyes in size in P. descarpentriesi), pronotum distinctly narrower and with less rounded lat¬ eral margins, and elytra with only stria 1 deeply impressed, stria 2 and 3 shallowly impressed but evident on disk, striae 4 to 7 effaced, and stria 8 impressed only in apical one-third (striae 1-7 shal¬ lowly impressed but evident at least on disc and stria 8 deeply impressed throughout in P. descar¬ pentriesi members). Also, the apex of the median lobe of the male genitalia is narrower in lateral KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 227 MADAGASCAR. Ranomafana NP„ Vatoharanam, Trail K at 350 m mark, 24,07.2005 collected by hand in daytime, • t : . Ratalata and I. Rainio collectors Figure 14. Digital images of holotype male of Perigona (Ripogena) deuvei Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B Labels; C-D. Aedeagus of genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines A= 1.0 mm, C-D = 0.25 mm. 228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 aspect in P. deuvei males (Fig. 143C) than in those of P. descarpentriesi (see Deuve 1998, Fig. 2.). Finally, P. deuvei members differ from those of the new species described below in having small¬ er body size, pronotum with lateral margination very narrow throughout, fewer elytral striae evi¬ dent, and median lobe of the male genitalia with a longer and ventrally bent apex. Geographical distribution. — At present, known only from the type locality. Habitat distribution. — All specimens of this species were collected by hand in daytime in primary montane rainforest in the Vatoharanana area at an elevation of 1200 m. Perigona ( Ripogena) ranomafanae Kavanaugh and Rainio, sp. nov. Figure 15 Ripogena n. sp. 2; Rainio 2009: 33 (informal designation). Perigona f Ripogena ) n. sp. 2; Rainio 2012: 73, Rainio 2013: 96 (informal designation). Type material.— Holotype (Figs.l5A-B), a male, in CAS, labeled: ““CASENT 1049017*7 “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vatoharanana area, 1050 m, Abotovory stream, 29 April 1998*7 “21°16.7’S 47° 26.l’E, Stop # 98-100, D.H. Kavanaugh col¬ lector collected in large rotting log just under loose bark with cavities”/ “HOLOTYPE Perigona {Ripogena) ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, ranomafanae, is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition. — Size larger than average for subgenus, SBL = 4.4 mm. The holotype male of this species (Fig. 15A) shares with members of Perigona heterodera Alluaud (1936), Perigona prasinus Alluaud (1936) and P. deuvei sp. nov. the following features: pronotum with lateral mar¬ gins without sinuation anterior to hind angles, posterior margin slightly and smoothly convex, and hind angles either obtuse or broadly rounded. These features distinguish members of these four species from those of Perigona bembidioides Alluaud (1936) and Perigona viridimicans Jeannel (1948), which have distinct sinuation of the lateral margins anterior to rectangular hind angles and a straight basal margin. Members of P. ranomafanae differ from those of P. prasinus in having slightly smaller body size and dorsum without metallic reflection (dorsum with metallic green reflection in P. prasinus). They differ from members of P. heterodera and P. deuvei in having larg¬ er body size (SBL = 3.4 mm in P. heterodera, 3.5 to 3.9 mm in P. deuvei), pronotal hind angles more sharply angulate, lateral explanation of pronotum markedly widened and flattened in region of hind angles (pronotal hind angles rounded or at least less sharply angulate and lateral explana¬ tion only slightly wider near hind angles than anteriorly in P. heterodera and P. deuvei). The unique male holotype of P. ranomafanae is slightly teneral and the genitalia are very soft and only lightly sclerotized. However, the apex is sclerotized sufficiently to allow comparisons with those of P. het¬ erodera and P. deuvei males. In lateral view (Fig. 15C), the apex is slightly longer than in P. het¬ erodera males (see Jeannel 1948, Fig. 351b) and distinctly shorter and straighter than in P. deuvei males. Finally, they differ from members of P. descarpentriesi in having moderately large eyes (eyes markedly reduced eyes in size in P. descarpentriesi), pronotum widest anterior to middle and lateral margins nearly straight in posterior half (pronotum widest at middle and lateral margins evenly arcuate throughout P. descarpentriesi). Also, the apex of the median lobe of the male geni¬ talia is narrower and more pointed in lateral aspect in P. ranomafanae males (Fig. 15C) than in those of P descarpentriesi (see Deuve 1998, Fig. 2.). Geographical distribution. — At present, known only from the type locality. Habitat distribution.— The unique holotype was collected by hand in primary montane KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 229 CASENT * 1049017 \IA DAG A SCAR, hanaranlsoa Provi ikc. Ra noma fana National Park, Vatoharanana # area, 1050 m Amhalovory stream 29 April 1998. 21 16.7'S 47 f, 26.1 'F. Stop tt 98-100, D.ll. Kavanaugh collector collected in targe rotting log iusl under loose bark with cavities Figure 15. Digital images of holotype male of Perigona ( Ripogena ) ranomafanae Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B Labels; C. Aedeagus of genitalia, left lateral aspect [dissection not fully cleaned due to tener- al nature of holotype] ; D. Dorsal aspect. Scale lines A = 1.0 mm, C = 0.25 mm. rainforest in the Vatoharanana area at an elevation of 1050 m from under the loose bark of a large log on the forest floor. Tribe Odacanthini LaPorte 1834 In his treatment of the Malagasy fauna, Jeannel (1948) assigned three species to genus “ Cas- nonia ” [a misspelling by Latreille and Dejean (1824) of Cosnania Dejean 1821], a genus that is now considered to include only Western Hemisphere species. Each of Jeannel’s three “Casnonia” 230 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 species is now included in a different genus (Lorenz 2005). The species with members most simi¬ lar to those of the new species described below is now included in Archicolliuris Liebke (1931). The present supraspecific classification of odacanthines is basically that of Liebke (1931 and 1938), and no comprehensive taxonomic or phylogenic treatment of this taxon worldwide has been produced since then. Clearly, a phylogenetic analysis of relationships among and inclusiveness of odacanthine genera is needed. Pending such a study, we tentatively include our new species in genus Archicolliuris. Genus Archicolliuris Liebke 1931 Archicolliuris ranomafanae Kavanaugh and Rainio, sp. nov. Figure 16 Type material.— Holotype (Figs.l6A-B), a female, inNMNH, labeled: “MADAGASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 1100m 22-31 October 1988 W. E. Steiner’7 “Flight inter¬ cept-yellow pan trap, island in stream, montane rainforest”/ “HOLOTYPE Archicolliuris ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, ranomafanae, is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition.— Size large for genus, SBL = 8.2 mm. The holotype female of this species (Fig. 16A) is easily distinguished from members of all other Malagasy odacanthine species based on numerous features. The following features used by Jeannel (1948) in his key to genera are shared with the three species that he included in “ Casnonia”'. integument glabrous (without pubes¬ cence), pronotum long, narrow and tubular anteriorly, impunctate, with one or more setae present along each lateral margin. The holotype of 4. ranomafanae differs from members of Casnonia fair- mairei Gestro (1895) [currently included in genus Erectocolliuris Liebke (1931)] in having a pronotum black in color and with only a single pair of midlateral setae (pronotum rufotestaceous in color and with 5 or 6 pairs of lateral setae present in E. fairmairei members). It differs from members Casnonia coerulans Kiinckel d’Herculais (1887) [currently included in genus Protocol- liuris Liebke (1931) in having the pronotum only about twice as long as wide (three times as long as wide in P. coerulans ), the elytra without metallic reflection (elytra with a dark metallic blue reflection), and legs dark black to piceous, except trochanthers rufous and basal parts of all femo¬ ra pale (legs pale in P coerulans, except apical parts of femora dark). It is similar to members of the last of JeanneTs “ Casnonia ” species, Colliuris olsoufieffi Alluaud (1935) [currently included in Archicolliuris ] in having a shiny black dorsum (with the only pale areas present as elytral pale spots), the pronotum with transverse grooves and ridges in the basal half and anterior angles pro¬ jected laterally, and the elytra with a deep transverse depression at the basal one-fourth and with three or four discal setiferous pores on elytral interval 3. However, it differs with members of A. olsoufieffi in several features: body size larger (SBL = 6.8 mm in A. olsoufieffi)', pronotum impunctate (coarsely punctate laterally and basally in A. olsoufieffi), slightly less than twice as long as wide (2.5 times as long as wide in A. olsoufieffi), with lateral borders vaguely present but only as very faintly impressed lines (lateral borders absent from A. olsoufieffi members); elytra with only one pair of small pale spots, located at apical one-third on interval 4 (both subapical and sub- basal pairs of spots present in A. olsoufieffi) and with three or four discal setiferous pores also on interval 5 (absent from A. olsoufieffi members); and legs dark, except pale at base of femora and on trochanters (legs pale thoughout in A. olsoufieffi). In addition, the holotype of A. ranomafanae is unique among members of all Malagasy KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 231 MADAGASCAR: Prov, Fianarantsoa, 7 km W Ranomafana,• 1100m 22-31 October 1988 W. E. Steiner Flight intercept- yellow pan trap in Malaise trap** island in stream* montane rainforest r L r Figure 16. Digital images of holotype female of Archicolliuris ranomafanae Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels. Scale line = 1.0 mm. odcanthine species in having the elytra distinctly convex throughout, except in the area of the trans¬ verse depression, elytral intervals flat, and elytral striae 3 to 7 effaced except in the transverse depression and apically, where they are evident. Striae 1 and 2 are very shallowly impressed but evident (stria 2 less so) in and posterior to the transverse depression. The form and location of the transverse depression of the elytra and the nearly effaced striae are similar to these features of members and genus Mimocolliuris Liebke (1933) of the eastern Palearctic and Oriental Regions. Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The unique holotype was collected in a flight intercept-yellow pan trap on an island in a stream in montane rainforest at an elevation of 1100 m. 232 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Tribe Lebiini Bonelli 1810 Subtribe Pericalina Hope 1838 Genus Thysanotus Chaudoir 1848 Jeannel (1949) recognized 14 species in this genus precinctive Malagasy genus. Deuve (2010) described six additional new species and returned two species treated by Jeannel as junior syn¬ onyms to full species status. He also provided an excellent key to all the known species. Thysanotus bimaculatoides Kavanaugh and Rainio, sp. nov. Figures 17-18 Type material.— Holotype (Figs.l6A-B), a male, in CAS, labeled: “MADAGASCAR Ranomafana NP. Mangevo plot 1600m 20.4.2005”/ “240” [handwritten label]/ “HOLOTYPE Figure 17. A-D. Digital images of holotype male of Thysanotus bimaculatoides Kavanaugh & Rainio sp. nov. A-D. A Habitus, Dorsal aspect; B. Labels. Scale line =1.0 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 233 Figure 18. Digital images of aedeagus of male genitalia. A, C. Left lateral aspect; B, D. Dorsal aspect. A-B. Holotype male of Thysanotus bimaculatoides Kavanaugh & Rainio sp. nov.; B, D. Thysanotus spinosus Alluaud (from Ranomafana National Park). Scale lines = 0.5 mm. Thysanotus bimaculatoides Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 2): 1 female (in MNHN) labeled “MADAGASCAR Ranomafana NP. Mangevo plot 1600m 22.4.20057 “240” [handwritten label]; 1 female (in CAS) labeled “CASENT 8069538”/ “MADA¬ GASCAR: Province Fianarantsoa, Foret d’Ambalagoavy Nord, Ikonga, Abatombe Nov-2000”/ “21°49’39”S, 47°20’20”E Calif. Acad, of Sciences colls: R. Harin’Hala & M.E. Irwin, malaise trap 625m MA-01-12-01”. Both paratypes also bear the following label: “PARATYPE Thysanotus bimaculatoides Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation ol species name.— The species epithet, bimaculatoides, is an adjective derived from the Latin words, bis, meaning two, and macula, meaning spot or spotted, and the Greek word eidos, meaning like. The name refers to the similarity between members of this species and those of Thysanotus bimaculatus Deuve (2010). Recognition.— Size slightly less than average for genus, SBL of male = 5.7 mm, of females = 5.3 mm. Members of this species (Fig. 17A) key to Thysanotus bimaculatus Deuve (2010) in Deuve’s key based on the following combination of features: elytra shiny, intervals smooth, elytral microsculpture indistinct (comprised of faintly impressed microlines and markedly transverse sculpticells, each elytron with a large, pale macula near the base on intervals 2 to 4, also extended partially onto interval 1 and to scutellum at basomedial part of macula, elytral apex bluntly round¬ ed, not toothed. Members of T. bimaculatoides differ from the holotype of T. bimaculatus in hav¬ ing slightly larger body size (SBL of holotype female of T. bimaculatus = 4.9 mm), pronotum slightly narrower, ratio of PW/PL =1.1 (ratio of PW/PL = 1.2 in T. bimaculatus ), narrower basal- ly with subbasal sinuation of the lateral margin slightly longer and deeper and hind angles sharp¬ er, rectangular to distinctly acute (compare with Deuve 2010, Fig. 17). Deuve did not have a male specimen of T. bimaculatus, so we cannot compare male genitalia of these two species. However, for future comparative purposes, we provide digital photographs of the genitalia (Figs. 18A,C) of the holotype of T. bimaculatoides as well as those of a male of Thysanotus spinosus (Alluaud 1936c) (Figs. 18B,D) [males of this species previously unknown], to complement the illustrations of other Thysanotus species provided by Jeannel (1949) and Deuve (2010). Geographical distribution.— At present, known only from two localities, 52 km apart, in central Fianarantsoa Province. Both localities are on the eastern slope off the central plateau. 234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Habitat distribution.— The holotype and one paratype were collected by hand in the Mangevo area, about 1600 m distance in from the edge of montane rainforest, at an elevation of 1600 m. The other paratype was collected in a malaise trap in montane rainforest at an elevation of 650 m. Thysanotus spinosus Alluaud 1936 This species was described on the basis of a single female specimen (holotype in MNHN). Neither Jeannel (1949) nor Deuve (2010) has a male specimen of this species available for study. We take this opportunity to provide digital photographs of the median lobe of the genitalia (Figs. 18B,D) from a male collected in RNP to complement the illustrations of other Thysanotus species provided by Jeannel (1949) and Deuve (2010). Genus Madecassina Jeannel 1949 Jeannel (1949) recognized at total of ten species and one additional subspecies in this precinc- tive Malagasy genus and provided a key for identification of adults. No additional species have been described since Jeannel’s study. Madecassina bimaculata Kavanaugh and Rainio, sp. nov. Figure 19 Type material.— Holotype (Figs.l9A-B), a male, in CAS, labeled: “CASENT 1049022”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 13 April 1998, 21°14.9’S 47 0 25.6’E,’7 “Stop # 98-29, D. H. Kavanaugh col¬ lector collected by beating suspended clusters of dead leaves and twigs”/ “HOLOTYPE Made¬ cassina bimaculata Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 16): 1 female (in CAS) labeled “CASENT 1004001”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanaugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Ravelo- son collectors”/ “collected in daytime by beating live understory vegetation and/or dead leaves and other suspended debris”; 1 female (in CAS) labeled “CASENT 1049023”/ “MADAGASCAR, Fia¬ narantsoa Province, Ranomafana National Park, Talatakely area, 900 m, Trail FF, 21°14.9’S 47° 25.6’E,”/ “10-16 November 1998, beaten from vines, V. F. Lee & K. J. Ribardo collector”; 1 male (in CAS) labeled “CASENT 1049024”/ “MADAGASCAR, Fianarantsoa Province, Parc National Ranomafana, Talatakely area, 900 m, mixed tropical forest, 24 April 1998,”/ “21°15.3’S 47°25.9’E,”/ “Stop # 98-76, D. H. Kavanaugh collector collected at night on standing tree trunk”; 1 male (in CAS) labeled “CASENT 8068817”/ “MADAGASCAR: Province Fianarantsoa, Ranomafana National Park, radio tower at forest edge, elev 1130 m 9-20 March 2003”/ “21°15.05’S, 47°24.43’E collector: R. Harin’Hala California Academy of Sciences malaise, mixed tropical forest MA-02-09B-55”; 1 male (in CAS) labeled “CASENT 1049026”/ “MADAGAS¬ CAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area, 1150 m, mixed tropical forest, 18 April 1998, 21°12.8’S 47°23.0’E,”/ “Stop# 98-52, D. H. Kavanaugh collector collected by beating live vegetation and suspended dead leaves and twigs”; 1 female (in CAS) labeled “CASENT 1049025”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vatoharanana area, 1050 m, Ambatovory stream, 29 April 1998, 21°16.7’S 47° 26.l’E,”/ “Stop# 98-101, D. H. Kavanaugh collector collected by beating suspended clusters of dead leaves and KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 235 CASENT B 1049022 , M A DAG A SC' A R. li an aranisoa Province. Ra noma tana National Park. Tala take I v area. 900 m, mixed tropical • forest. I* April 1998, 2m4.9'S4 7 25.6*E _ Stop tt 98-29 D.n, Kavanaugh collector collected by beating • suspended clusters of dead leaves and twigs _ Figure 19. Digital images of holotype male of Madecassina bimaculata Kavanaugh & Rainio sp. nov. A. Habitus, dor¬ sal aspect; B. Labels C-D. Median lobe of male genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines A = 1.0 mm, C-D = 0.5 mm. twigs”; 2 males and 1 female (in CAS and EM) labeled “CASENT 8072308” or “CASENT 8072309” or “CASENT 8072307”, respectively/ “MADAGASCAR: Fianarantsoa Parc National de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana elev 1100 m 27-31 March 2003”/ “21°15’24”S, 47°26’00”E California Academy of Sciences coll. Fisher, Griswold et al. beating low veg., montane rainforest code: BLF8401”; 1 male (in MNHN) labeled “MADAGAS¬ CAR, Ranomafana NP., Vatoharanana, Trail K at 150 m mark, 25.7.2005”/ “collected by hand in daytime from dead leaves of bushes and trees, F. Ratalata & J. Rainio collectors”; 2 females (in MNHN and MZF) labeled “MADAGASCAR, Ranomafana NP., Talatakely, Trail SP at 50 m mark, 236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 28.07.2005”/ “collected by hand in daytime from dead leaves of bushes and trees, F. Ratalata & J. Rainio collectors”; 2 females (in MNHN and MZF) labeled “MADAGASCAR, Ranomafana NR, Valohoaka, Trail B at 555 m mark, 28.01.2005”/ “collected by hand in daytime from dead leaves of bushes and trees, F. Ratalata & J. Rainio collectors”; and 2 females (in MNHN and MNHN) labeled “MADAGASCAR, Ranomafana NR,Valohoaka, Trail B at 555 m mark, 16.11.2005”/ “col¬ lected by hand in daytime from dead leaves of bushes and trees, F. Ratalata & J. Rainio collector”. All paratypes also bear the following label: “PARATYPE Madecassina bimaculata Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation ol species name. — The species epithet, bimaculata, is an adjective derived from the Latin words, bis, meaning two, and macula, meaning spot or spotted. The name refers to the pair of subapical elytral spots seen in members of this species. Recognition. — Size average for genus, SBL of males = 4.7-5.5 mm, of females = 5.3-5.6 mm. Members of this species (Fig. 19A) key most closely, to Madecassina picta Alluaud (1897) in JeanneTs (1949) key based on the following combination of features: pronotum impunctate, ely¬ tra with two subapical pale spots and striae deeply impressed and legs testaceous in color. The only feature used in the key with which T. bimaculata members differ is the presence of is metallic reflection on the forebody—members of T. picta have evident aeneous or greenish metallic reflec¬ tion on the head and pronotum, whereas T. bimaculata members are without metallic reflection. These two species are otherwise very similar externally; however the pronotum is proportionately slightly longer and more distinctly narrowed basally and the hind angles more sharply defined and rectangular in T. bimaculata members than in T. picta members (which have the hind angles slight¬ ly obtuse). Also, the apex of the median lobe of the male genitalia is shorter, thicker and straighter in lateral aspect (Fig. 19C) and more bluntly triangular in dorsal aspect (Fig. 19D) in T. bimacula¬ ta males than in those of T. picta males, which have the apex longer, narrower and distinctly hooked dorsally in lateral aspect and more narrowly rounded in dorsal aspect (see Jeannel 1949, Figs. 474g and 474i). Geographical distribution.— At present, known only from the type locality. Habitat distribution. — Specimens of this species have been collected in primary montane rainforest in the Vatoharanana and Valohoaka areas and in secondary montane rainforest in the Talatakely and Vohiparara areas, at elevations ranging from 900 to 1150 m. They have been col¬ lected mainly by beating live vegetation and suspended clusters of dead leaves and twigs. Madecassina quadrimaculata Kavanaugh and Rainio, sp. nov. Figure 20 Type material.— Holotype (Figs.20A-B), a male, in CAS, labeled: “MADAGASCAR, Ranomafana NP, montane rainforest near Tsinjorano village, 100 m in from forest edge,”/ “21.09637°S 47.52085°E, 1050 m, 19.05.2005, collected by hand in daytime, F.J. Ratelolahy col¬ lector”/ “HOLOTYPE Madecassina quadrimaculata Kavanaugh & Rainio sp. n. 2015” [red label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, quadrimaculata, is an adjective derived from the Latin words, quattuor, meaning four, and macula, meaning spot or spotted. The name refers to the two (subbasal and subapical) pairs of elytral spots seen in members of this species. Recognition. — Size average for genus, SBL = 5.2 mm. The holotype male of this species (Fig. 20A) keys to Madecassina maculata Alluaud (1899b) in Jeannel’s (1949) key based on the following combination of features: body black, forebody without metallic reflection, pronotum KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 237 MADAGASCAR, Ranomafam NP, montane rainforest - near Tsinjorano village, JOG m in from forest edge, 2L09637°S 47.52085°E, 1050 m, 19,05.2005, collected by hand in * daytime, * F.j Ratelolihy collector Figure 20. Digital images of holotype male of Madecassina quadrimaculata Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels C-D. Aedeagus of male genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines A = 1.0 mm, C-D = 0.5 mm. 238 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 punctate laterally, and elytra shiny with four pale spots. It differs from members of M. maculata in having a smaller body size (SBL = 5.5-6.0 mm in M. maculata), the pronotum punctate laterally and laterobasally only, with conspicuous transverse grooves and ridges on disc (in M. maculata the pronotum is punctate on the disc also, between the ridges, in contrast to what Jeannel said in his key), elytral intervals shinier due to more shallowly impressed isodiametric microsculpture (elytral intervals duller due to more deeply impressed isodiametric microsculpture in M. maculata ), the elytral spots much more distinct (much less so in M. maculata ) and median lobe of male genitalia with apex hooked dorsally and thicker in lateral aspect (Fig. 20C) and slightly swollen and bent right in dorsal aspect (Fig. 20D) (apex hooked but thinner in lateral aspect and more parallel-sided and straight in dorsal aspect inM maculata males) (see Jeannel 1949, Fig. 474f). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The unique holotype was collected in montane rainforest near Tsin- jorano village at a distance of 100 m inside the forest edge at an elevation of 1050 m. Genus Pristacrus Chaudoir 1869 Jeannel (1949) recognized four species in this precinctive Malagasy genus and provided a key for distinguishing their respective members. In this key, Jeannel used two features with which we found problems. First, his statements about pronotal proportions—width in relation to pronotal length (i.e. ratio PW/PL)—appear to have been based on visual impressions only, rather than actu¬ al comparative measurements, because our measurements of these proportions in specimens (including types) of the four species have not corresponded to those stated by Jeannel. Unfortu¬ nately, this problem renders two of his three couplets (couplets 1 and 3) difficult to interpret. Sec¬ ond, he used shape of the sutural angle of the elytral apex (i.e., whether it is broadly rounded or sharp and toothed) as the primary comparative feature in his couplet 2. In fact, based on the exam¬ ination of many more specimens than were available to Jeannel, we found that members of all species except one, Pristacrus laticollis (Gory and Laporte 1837) have elytral apices that are sharp and toothed, including members of Pristacrus bonotatus (Klug 1833), which Jeannel cited as broadly rounded in his key. We point out these problems with Jeannel’s key as caution to those try¬ ing to use it, as we were, for identifications. No additional species have been described since Jean- nel’s study. Pristacrus ranomafanae Kavanaugh and Rainio, sp. nov. Figures 21-22 Pristacrus n. sp. 7; Rainio 2009: 33, Rainio 2012: 73 (informal designation). Type material.— Holotype (Figs.21A-B), a male, in CAS, labeled: “CASENT 1003833’V “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,’7 “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected at night on sur¬ face of rotting logs on forest floor or suspended just above it”/ “HOLOTYPE Pristacrus ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 19): 1 male and 1 female (in CAS) labelled “CASENT 1003832” or “CASENT 1003831”, respectively, otherwise labeled same as holotype; 1 female (in CAS) labeled “CASENT 1004470”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 239 CASENT ■ 1003833 MADAGASCAR, Fianarantm Province, Ranomafana National Pa rkj Talatakely area, 900 m, mixed tropical forest 2*22 January 2001, 2U504i°b/47.41945“E. Stop# DHK-01-001 D. H, 8l K.M.* Karnaugh, R.L Brett, E, Elsom, F. Vargas, R, Ranalvosolo. E. F. Randrianirina, N. Rasoamananana, T.J. Ravdomanana, and H.C. Rav eloson collec tors collected at night on surface of rotting logs on forest Hoot dr suspended just above it Figure 21. Digital images of holotype male of Pristacrus ranomafanae Kavanaugh & Rainio sp. nov. A. Habitus, dor¬ sal aspect; B Labels. Scale line =1.0 mm. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Rav- elomanana, and H.C. Raveloson collectors”/ “collected in daytime from large accumulation of dead leaf, twig, and branch debris on and extended up to 2 m above forest floor”; 1 male (in CAS) labeled “CASENT 1002838”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Ravelo¬ son collectors”/ “collected in daytime under loss bark of rotting log logs on forest floor or sus- 240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 pended just above it”; 1 male and two females (in CAS and MZF) labeled “CASENT 1003752”, “CASENT 1003751” or “CASENT 1003753”, respectively/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected in day¬ time on underside of rotting log logs on forest floor or suspended just above it”; 1 male (in CAS) labeled CASENT 1004313 / MAD A- Figure 22. Digital images of aedeagus of genitalia of GASCAR, Fianarantsoa Province, Ranoma- male holotype of Pristacrus ranomafanae Kavanaugh & fana National Park, Talatakely area, 900 m, Rainio S P- nov - A - Left lateral as P ec # B - Dorsal as P ect Scale mixed tropical forest, 2-22 January 2001,”/ lme 0 5 mm - “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelo¬ manana, and H.C. Raveloson collectors”/ “collected in daytime from leaf litter on forest floor”; 1 female (in CAS) labeled “CASENT 1051144”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 27 April 1998”/ “21°15.3’S 47°25.9’E, Stop # 98-85A D.H. Kavanaugh collector collected in and under rotten logs on forest floor”; 1 female (in CAS) labeled “CASENT 8068264”/ “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, Belle Vue at Talatakely, elev 1020 m 28 April - 5 May 2002”/ “21°15.99’S 47°25.21’E collector: R. Harin’Hala California Acad of Sciences malaise sec¬ ondary tropical forest MA-02-09C-27”; 1 female (in CAS) labeled “CASENT 8068264”/ “MADA¬ GASCAR: Province Fianarantsoa, Parc National Ranomafana, Belle Vue at Talatakely, elev 1020 m 15-22 November 2001”/ “21°15.99’S 47°25.21’E collector: R. Harin’Hala California Acad of Sciences malaise secondary tropical forest MA-02-09C-03”; 1 male (in NMNH) labeled “MADA¬ GASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 900 m 1-9 February 1990 W. E. Steiner”/ “at black light in montane rainforest near river and stream”; 1 male (in CAS) labeled “CASENT 8005762”/ “21.25041°S/ 47.41945°E MADAGASCAR, Fianarantsoa Prov Ranomafana National Park, Talatakely area, 900m, 10:1:2001 Col. K. Will headlamp search logs”; 5 males and 1 female (in EMEC and MNHN) labeled “CASENT 8005762”/ “21.25041°S/47.41945°E MADAGASCAR, Fianarantsoa Prov Ranomafana National Park, Talatakely area, 900m, 10:1:2001 Col. K. Will head¬ lamp search logs”. All paratypes also bear the following label: “PARATYPE Pristacrus ranomafanae Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation ol species name. — The species epithet, ranomafanae , is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition. — Size small for genus, SBL of males = 8.3-8.5 mm, of female = 8.6 mm. Members of this species (Fig. 21 A) are distinguished from those of all other species in the genus except Pristacrus binotatus Klug (1833) by their small body size (SBL greater than 10.0 mm in members of the three other species). They differ from members of Pristacrus semipiceus (Fair- maire 1887), in having much smaller body size (SBL = 13.6 to 14.5 mm in P semipiceus mem- KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 241 bers), a shinier dorsal surface, shorter and broadly rounded mandibles and an impunctate (but lon¬ gitudinally rugulose) frons (dorsal surface duller, mandibles distinctly longer and more tapered, and frons punctate as well as rugulose in P. semipiceus members). They differ from members of P laticollis in having slightly smaller body size (SBL = 10.2-11.1 mm), slightly shinier dorsal sur¬ face, relatively longer and narrower pronotum and elytra that are broadly ovoid with lateral mar¬ gin arcuate and lateral explanation broad throughout, especially at middle, and elytral apices dis¬ tinctly angulate and toothed (duller dorsal surface, relatively broader and shorter pronotum and ely¬ tra that are nearly parallel-sided at middle with lateral explanation narrower throughout and elytral apices broadly rounded in P. laticollis members). They differ from the unique type of Pristacrus rotundatus (Fairmaire 1892) in having smaller body size (SBL =11.9 mm in P. rotundatus type), shinier dorsal surface, elytra less broadly rounded and humeri less anteriorly projected (dorsal sur¬ face dull and elytra markedly broad and rounded and humeri more projected anteriorly in P. rotun¬ datus type). Members of P. ranomafanae are very similar to those of P binotatus in size and over¬ all form. Specimens of the former have the dorsal surface slightly shinier, the pronotum relatively broader basally, with subbasal sinuation of the lateral margins absent or only very faint and hind angles less sharply defined, and elytra with striae slightly more deeply impressed and intervals more convex and more roughened at the raised center than in most members of P binotatus. How¬ ever, at least a few specimens of P binotatus share one or more of these features with P. ranomo- fanae members. They only features that most reliably distinguish members of P ranomafanae from those of P. binotatus, as well as from members of the other species, are the male genitalia. In P. ranomafanae males (Fig. 22A-B), the shaft of the median lobe is inflated subbasally but lacks the large, well-defined dorsal bulge seen in males of P. binotatus (see Jeannel 1949, Fig. 486) and, in lateral aspect (Fig. 21 A), the apex is distinctly hooked dorsally (apex not hooked in P binotatus males. Males of the each of the other species also have the shaft of the median lobe uniquely shaped. Geographical distribution.— At present, known only from the type locality. Habitat distribution.— Specimens of the type series were all collected in secondary mon¬ tane rainforest in the Talatakely area at elevations of 900 to 1020 m. The area is replete with inva¬ sive guava (Psidium cattleianum). Specimens were found in daytime in masses of dead leaves, twigs and branch debris accumulated on the forest floor, in leaf litter, and under loose bark or on the underside of rotting logs on the forest floor. Several were found at night running on the surfaces of rotting logs on the forest floor; two were collected in malaise traps and one was attracted to and collected at an ultraviolet light. Genus Eurydera Laporte 1831 Jeannel (1949) recognized 24 species plus three additional subspecies in this precinctive Mala¬ gasy genus and a provided a key for identification of their members. Mateu (1973) described seven additional species. Type specimens for all of Mateu’s species are deposited in MNHN, so we were able to compare our material with reliably determined specimens of all described Eurydera species, including the seven not included in Jeannel’s key. Eurydera ocellata Kavanaugh and Rainio, sp. nov. Figures 23-24 Type material.— Holotype (Figs. 23A-B), a male, in CAS, labeled: “CASENT 8068800”/ “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, Bellevue at Telatakely, elev 1020 m 12-19 February 2002”/ “21°15.99’S, 47°25.21’E collector: R. Harin’Hala California 242 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 ICASENT 18068800 MADAGASCAR: Province* Fianarantsoa, Pare Nations* Ranomafana. Belie Vu© at Talatakeiy, eiev 1020 rr 12-19 February 2002 21" 15.99'S, 47* 25.21'E collector R. Harin’Hala ‘California Acpo of Sen tout malaise, secondary tropica' forest MA-02-09C-"i6 Figure 23. Digital images of holotype male of Ewydera ocellata Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Label. Scale line =1.0 mm. Academy of Sciences malaise, secondary tropical forest MA-02-09C-16”/ “HOLOTYPE Eurydera ocellata Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (only 1); a female (in NMNH) labeled “MADAGASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 900 m 1-9 February 1990 W. E. Steiner”/ “at black light in montane rainforest near river and stream”/ “PARATYPE Eurydera ocellata Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, ocellata, is the Latin adjective meaning having little eyes or marked with spots. The name refers to the small, faint dark spot on interval 3 within the subapical pale area of each elytron at the insertion point of the subapical discal setifer- ous pore. Recognition.— Size small for genus, SBL of male and female = 9.2 mm. Members of this species (Fig. 23A) key to Eurydera mormolycoides Coquerel (1851) in Jeannel’s (1949) key on the KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 243 Figure 24. Digital images of holotype male of Eurydera ocellata Kavanaugh & Rainio sp. nov. A. Head, dorsal aspect; B-C. Aedeagus of male genitalia; B. Left lateral aspect; C. Dorsal aspect. Scale lines = 0.5 mm. basis of the following combination of features: size small; head (Fig. 24A) with frontal furrows deeply impressed, smooth, divergent posteriorly and without a transverse impression uniting them, posterior part of frons smooth, narrow and triangular, without lateral grooves prolonged posterior¬ ly beyond the posterior border of the eyes; mandibles short and broad, lateral border not broadly explanate, lateral margin only slightly convex; pronotum cordate, distinctly narrowed basally, lat¬ eral margins angulate at insertions of midlateral setae; elytra short, round and black, each with a reddish subapical spot tangential to the medial suture as well as subhumeral and subapicolateral reddish areas and reddish apical spines, elytral intervals moderately convex; median lobe of male genitalia without a large, angulate projection on the ventral margin of the shaft near the base. They differ from E. mormolycoides members in having: dorsal surface slightly shiny (dull in E. mor- molycoides members); pronotum with lateral explanations pale testaceous (darker in E. mormoly¬ coides members), anterior angles broadly rounded (more narrowly rounded in E. mormolycoides members) and lateral margins with shallow but distinct sinuation anterior to hind angles (sinuation absent from or less distinct in E. mormolycoides members); elytra with larger but less distinct (less contrasting with rest of elytra) pale subapical macula with black eye spot on interval three (macu¬ la extended further laterally and lateroanteriorly than in E. mormolycoides members) and apical spine about half as long as spine in E. mormolycoides members. Finally, the median lobe of the male genitalia (Figs. 24B-C) has a thicker shaft throughout and an apex shorter and thicker in lat¬ eral view (Fig. 24B) and broader and straighter in dorsal view (Fig. 24C) than in E. mormolycoides males (see Jeannel 1949, Fig. 492b). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The holotype specimen was collected in a malaise trap in second¬ ary montane rainforest in the Talatakely area at an elevation of 1030 m. The paratype specimen was collected at ultraviolet light in secondary montane rainforest near the junction of the Namorona River and a small tributary stream. 244 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Eurydera oracle Kavanaugh and Rainio, sp. nov. Figures 25-26 Type material.— Holotype (Figs.25A-B), a male, in CAS, labeled: “CASENT 8005762”/ “21.25041°S/ 47.41945°E MADAGASCAR, Fianarantsoa Prov Ranomafana National Park, Talatakely area, 900m, 10:1:2001 Col. K. Will headlamp search logs”/ “HOLOTYPE Eurydera oracle Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 7): 1 male (in NMNH) labeled “MADAGASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 1100 m 1-7 November 1988 W. E. Steiner”; 1 female (in NMNH) labeled “MADAGASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 1100 m 27-31 October 1988 W. E. Steiner”/ “Malaise trap on island in stream, montane rain forest”; 1 female (in EMEC) labeled “CASENT 1003844”/ “MADAGASCAR, Fia¬ narantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Rav- CASENT 8005773 21,2504l o S/47.4l945“E MADAGASCAR to ixirgntsoo Prov Ranomafana National Park, Talatalcely area, 900m, 10;f:20Qi CoL!CWill,headlamp search logs Figure 25. Digital images of holotype male of Ewydera oracle Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Label. Scale line = 1.0 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 245 Figure 26. Digital images of holotype male of Eury>dera oracle Kavanaugh & Rainio sp. nov. A. Head, dorsal aspect; B-C. Aedeagus of male genitalia; B. Left lateral aspect; C. Dorsal aspect. Scale lines = 0.5 mm. elomanana, and H.C. Raveloson collectors”/ “collected at night on underside of shelf fungus on rot¬ ting log on forest floor or suspended just above it”; 1 female (in CAS) labeled “CASENT 1003739”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected in daytime on underside of rotting logs on forest floor or suspended just above it”; 1 female (in MNHN) labeled “CASENT 8068234”/ “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, Bellevue at Telatakely, elev 1020 m 10-14 January 2002”/ “21°15.99’S, 47°25.21’E collector: R. Harin’Hala California Academy of Sciences malaise, secondary tropical forest MA-02-09C-11”; 1 female (in MTEC) labeled “MADAGASCAR: Fianaran. Pr. Ranomafana N.P. HQ. area 21°15’24”S 47°25’15E 25NOV1994, at night M.A. Ivie & D.A. Pollack”; and 1 female (in CAS) labeled “CASENT 8068052”/ “MADAGASCAR: Province d’Antsiranana, Montaigne Francais, elev 150 m 6-20 March 2001”/ “12°19.5’S, 49°20’E California Acad, of Sciences R. Harin’Hala coll. Malaise along forested limestone ridge MA-01-06-0. All paratypes also bear the following label: “PARATYPE Eurydera oracle Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, oracle, is a noun in apposition and refers to Oracle Corporation, headquartered in Redwood City, California. We are pleased to name this new species for Oracle Corporation in recognition of and thanks for its generous support of the Cal¬ ifornia Academy of Science 1998 Madagascar expedition, which produced many of the specimens on which this study has been based. Recognition.— Size above average for genus, SBL of males = 13.9-15.3 mm, of females = 12.4-14.1 mm. Members of this species (Fig. 25A) key to the armata group in Jeannel’s (1949) 246 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 key based on the following combination of features: size medium; head (Fig. 26A) with posterior part of frons formed as a broad, flat, smooth, elevated plateau, reddish in color, laterally delimited by deep furrows extended posteriorly beyond the level of the posterior margin of the eye then medi¬ ally at a 45° angle and joined posteriorly (but distant from the midline) with a deep transverse fur¬ row across the middle of the head; mandibles moderately short and broad; median lobe of male genitalia without a large, angulate projection on the ventral margin of the shaft near the base. How¬ ever, within the armata group, they cannot be identified to species using Jeannel’s key. They dif¬ fer from members of Eurydera crispatifrons (Fairmaire 1896) in having a deep, sharply defined transverse furrow across the back of the head behind the raised frontal plateau (this furrow absent or very shallow and poorly defined in E. crispatifrons members). They differ from members of Eurydera rufotincta (Fairmaire 1868) in having the head large, nearly as wide as the pronotum (narrower in E. rufotincta members), black or piceous legs (rufous in E. rufotincta members) and in lacking transverse pale bands on the elytra (present in E. rufotincta members) and the deep oblong fossae present on the back of the frons in E. rufotincta members. They differ from Eurydera armata Laporte (1831) and Eurydera latipennis (Klug 1835) mem¬ bers in having the lateral furrows on the back of the head angularly convergent toward the posteri¬ or transverse furrow whereas these furrows are virtually parallel posteriorly in E. armata members or very slightly convergent posteriorly in E. latipennis members and joined with the transverse fur¬ row more laterally in members of both species. Also, the posterior part of the frontal plateau is much more elevated in E. oracle males than in those of these other two species as well as males of Eurydera cuspidata (Klug 1835), Eurydera ornatipennis (Fairmaire 1897), Eurydera sulcicollis Mateu (1973) and Eurydera ambreana Mateu (1973). Most members of E. armata and E. ornatipennis have pale bands or spots on the elytra, although some specimens lack all pale mark¬ ings. None of the specimens of E. oracle that we have examined have similar pale markings, although several of our specimens have the apical one-fifth of elytral interval 1 narrowly pale, and two specimens also have interval 2 more dimly pale in the same area. Overall, members of E. ora¬ cle are most similar to those of E. sulcicollis, but they differ from the latter in having greater sex¬ ual dimorphism, with E. oracle males having relatively larger heads, more prominent eyes, frons more wr ink led and with the posterior portion of the medial area more elevated, the transverse groove deeper, and the pronotum shorter and distinctly wider than in E. sulcicollis male. Females of these two species are very similar. Finally, the median lobe of the genitalia of E. oracle males (Fig. 26B-C) is distinct from that of males all other species in the genus (compare with Jeannel 1949, Figs. 489, 491-498, 500, and Mateu 1973, Figs. 1-8) in the combined forms of the shaft and apex. Geographical distribution.— We have examined specimens from only two areas. Seven specimens were collected in Ranomafana National Park, the type locality, on the eastern slope off the central plateau in the northeastern part of the southern half of the island of Madagascar. The eighth specimen, a female, was collected at Montagne des Frangais, in Antsiranana Province, at the northern end of the island and just over 1000 km distant from Ranomafana. No specimens have been recorded from intervening areas Habitat distribution.— The specimen from Montagne des Frangais was collected in a malaise trap set on a forest limestone ridge at an elevation of only 150 m. All of the specimens from Ranomafana National Park were collected in secondary montane rainforest in the Talatakely area at elevations ranging from 900 to 1100 m. They have been collecting using malaise traps or by hand. In daytime, they have been found hiding on the underside of logs on the forest floor; and at night they have been found on the underside of shelf fungi on rotting logs and stumps. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 247 Eurydera simplica Kavanaugh and Rainio, sp. nov. Figures 27-28 Eurydera sp.; Rainio and Niemela 2006: 227 (informal designation). Eurydera n. sp. 3; Rainio 2009: 31, Rainio 2012: 72 (informal designation). Type material.— Holotype (Figs.27A-B), a male, in CAS, labeled: “MADAGASCAR, Ranomafana NP.,Talatakely, 900 m, Trail SP at 50 m mark, 08.03.2004”/ “collected by hand in day¬ time in secondary montane rainforest, F. Ratalata & J. Rainio collectors”/ “106” [handwritten label]/ “HOLOTYPE Eurydera simplica Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes Figure 27. Digital images of holotype male of Eurydera simplica Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels. Scale line = 1.0 mm. 248 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Figure 28. Digital images of holotype male of Eurydera simplica Kavanaugh & Rainio sp. nov. A. Head, dorsal aspect; B-C. Median lobe of aedeagus of genitalia; B. Left lateral aspect; C. Dorsal aspect. Scale lines = 0.5 mm. (only 1): a female (in MNHN) labeled “"MADAGASCAR, Ranomafana NR, Talatakely, 900 m, Trail B at 1000 m mark, 15.06.2000’V “collected by hand in daytime in secondary montane rain¬ forest, F. Ratalata & J. Rainio collectors”/ “7/13” [handwritten label]/ “PARATYPE Eurydera sim¬ plica Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, simplica , is an adjective derived from the Latin word, simplex , meaning simple. The name refers to the relative simplicity of form and surface macro sculpture of member of this species compared with those of other species of Eury¬ dera. Recognition.— Size average for genus, SBL of male = 13.1 mm, of female = 12.4. Members of this species (Figs. 27A) are unlike those of any other species in the genus. In JeanneTs (1949) key species of Eurydera, they key to his unicolor species group, based on the following features: size medium; head (Fig. 2 8A) with posterior part of frons formed as a broad, flat, smooth, elevat¬ ed plateau, laterally delimited by deep furrows extended posteriorly beyond the level of the poste¬ rior margin of the eye then slightly and arcuately convergent posteriorly and terminated abruptly at the level of the posterior margin of the ocular swelling and just less than halfway from the midline to the lateral margin of the head, area between the furrow smooth and only slightly elevated; mandibles moderately short and broad; median lobe of male genitalia without a large, angulate pro¬ jection on the ventral margin of the shaft near the base. Jeannel included only Eurydera unicolor (Klug 1833) in this species group, but Mateu (1973)added another species, Eurydera fossulata Mateu to the group, in part based on the shared lack of sexual dimorphism in the front tarsi in mem¬ bers of these two species. Unlike males of most Eurydera species, those of E. unicolor and E. fos¬ sulata have tarsomeres 1 to 3 no wider than in females and without ventral adhesive setae. Mem¬ bers of E. simplica are easily distinguished from those of these two species, as well as from all other described Eurydera species, by the uniquely shaped pronotum (Fig. 27A), which is relative¬ ly small and short for the genus, with the disc smooth, lateral explanation very narrow, lateral mar¬ gins with very short and shallow sinuation anterior to markedly obtuse hind angles, and basal mar- KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 249 gin obliquely angulate laterally. Unlike males of E. unicolor and E. fossulata, those of E. simplica have front tarsi with tarsomeres 1 to 3 wider than in females and with at least a few adhesive setae ventrally, which may call into question a close relationship with these two these species. Members of E. simplica also differ from those of E. unicolor in having the pronotum with anterior angles much less projected anteriorly. They differ from members of E. fossulata in having the posterior part of the head, between the lateral furrows, smooth and slightly convex, whereas E. fossulata members have a deep, transverse medial depression, the bottom of which is flat and covered with deep transverse creases, on that part of the head. They also have the elytral intervals without the roughened surfaces typical of members of several species in genus, including E. fossulata (at least subapically), and the sutural spines are very long and straight (similar to those in E. unicolor and E. fossulata members). Finally, the median lobe of male genitalia of the holotype of E. simplica (Fig. 28B-C) has the shaft much narrower basally and the apical orifice shorter than in E. unicol¬ or males (see Jeannel 1949, Fig. 495). No male of E. fossulata has been available for comparison. Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The two known specimens of this species both were collected by hand in daytime in secondary montane rainforest in the Talatakely area at elevations ranging from 900 to 1000 m. Subtribe Cymindidina Laporte 1834 Ball and Hilchie (1983) considered the following two genera as subgenera of a single genus, Hystrichopus Boheman (1848); however we follow Casale (1998) and Lorenz (2005) in treating them as separate genera, pending a more comprehensive phylogenetic analysis of the Cymindidi¬ na. Genus Pseudomasoreus Desbrochers des Loges 1904 Jeannel (1949) included only three species in his treatment of this genus for Madagascar. Mateu (1980) added six new species and provided a key to the nine Malagasy species known at that time. Casale (1998) added one additional Malagasy species, Pseudomasoreus deuvei Casale. This genus presently includes a total of 20 species (Lorenz 2005) with a cumulative disjunct dis¬ tribution ranging from the Pyrenees of France and Spain south to southern Africa and Madagascar (Ball and Hilchie 1983). Pseudomasoreus ranomafanae Kavanaugh and Rainio, sp. nov. Figures 29, 30A, 30C Deuveilla n. sp. 1; Rainio 2009: 31 (informal designation). Lebiinae (n. sp. 1); Rainio 2012: 72 (informal designation). Type material.— Holotype (Figs.29A-B), a male, in CAS, labeled: “CASENT 1004471”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,’V “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected in daytime from large accumulation of dead leaf, twig, and branch debris on and extended up to 2 m above forest floor”/ “HOLOTYPE Pseudomasoreus ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 8): 1 male (in CAS) labeled same as holotype except “CASENT 1004472”; 2 males (in CAS) labeled “CASENT 1004003” and “CASENT 1004004”, respectively/ 250 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 CASENT 1004471 L _ M AD AC i A S C A R, Fianarants oa Province. Ranomafana National Park, Talatakely area. 900 m. mixed tropical forest 2-22 January 2001. 21.25041 f, S/ 47.41945 a K Slop# DHK-0l-JJ0i D. H, & K.M. Kavanaugb. R.L Brett. E. Elsom. F. Vargas. R. Ranaivosolo. E. F, Randrianirina. K. Rasoamananatv'- T.J. Rave lorn anana. and H.C. Ravelos on collectors collected in daytime from large accumulation of dead leaf, twig, and branch debris on and extended up to 2 m above forest floor Figure 29. Digital images of holotype male of Pseudomasoreus ranomafanae Kavanaugh & Rainio sp. nov. A. Habi¬ tus, dorsal aspect; B. Labels. Scale line A = 1.0 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 251 “MADAGASCAR, Fianarantsoa Prov-ince, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 2-22 January 2001,”/ “21.25041°S/ 47.41945°E, Stop# DHK-01-001, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected in daytime by beating live understory vege¬ tation and/or dead leaves and other sus¬ pended debris”; 1 male (in CAS) labeled “CASENT 1002681”/ “MADAGASCAR, Fianarantsoa Prov-ince, Ranomafana National Park, Vohiparara area, 1170 m, mixed tropical forest, 2-22 January 2001,”/ “21.22644°/ 47.36979°S, Stop# DHK-01-004, D.H. & K.M Kavanugh, R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo,”/ “E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Raveloson collectors”/ “collected in daytime by beating debris suspended in leaf whirls of Pandanus sp.”; 1 male and 2 females (in CAS and MNHN) labeled “CASENT 1049027”, “CASENT 1049028” and “CASENT 1049029”, respectively/ “MADAGASCAR, Fia¬ narantsoa Province, Ranoma-fana Nation¬ al Park, Vohiparara area, 1150 m, mixed tropical forest, 24 April 1998, 21°12.8’S 47° 23.0’E,”/ “Stop# 98-74, D. H. Kavanaugh collector beaten from vegeta¬ tion at forest edge”; 1 male (in MZF) labeled “MADAGASCAR, Ranomafana NR,Talatakely, 900 m, Trail BFA 75 m, 16.03.2001”/ “collected by hand in day¬ time, F. Ratalata & J. Rainio collectors”. All paratypes also bear the following label: “PARATYPE Pseudomasoreus ranomafanae Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fia¬ narantsoa Province, Ranomafana National Park. Figure 30. Digital images of Pseudomasoreus spp. A-C. Holotype male of P. ranomafanae Kavanaugh & Rainio sp. nov.; D, E. P. catalai Jean- nel (from Ranomafana National Park). A. Right labi¬ al palpus, ventral aspect; B E. Aedeagus of male genitalia; B, D. Left lateral aspect; C, E. Ventral aspect. Scale lines = 0.5 mm. 252 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Derivation of species name. — The species epithet, ranomafanae, is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition.— Size large for genus, SBL of males = 9.7-10.6 mm, of females = 10.3 mm. In Mateu’s (1980) key to the Malagasy species of this genus, members of P. ranomafanae (Fig. 29A) key to Pseudomasoreus decorsei Jeannel (1941b) based on the following combination of fea¬ tures: size large, SBL = 8.5 mm or more; elytra concolorous brown or piceous, without pale areas. However, they differ from the unique holotype of P. decorsei in having even larger body size (SBL = 8.5 mm in P. decorsei type), pronotum narrower overall and especially basally and with less rounded, more evident hind angles (pronotum broad, rounded, with hind angles effaced by smooth¬ ly rounded transition from lateral to basal margin in P. decorsei type), and elytral lateral margins nearly parallel (more evenly rounded in P decorsei type). Members of P ranomafanae are actual¬ ly more similar to those of P deuvei, which was not included in Mateu’s key. They share large size, general body form with long, parallel-sided elytra, and antennomere 3 glabrous except for the api¬ cal whorl of fixed setae (additional pubescence seen in apical part of antennomere 3 in most other species of genus). However, their body size is slightly smaller on average (SBL = 10.2-11.5 in P deuvei members), the pronotum is narrower overall and more narrowed basally, with the hind angles more distinct than in P. deuvei members and the lateral margins either straight or with slight sinuation anterior to the hind angles (lateral margins not at all sinuate in the latter). Also, the penul¬ timate labial palpomeres in all specimens of the type series have three or four setae (Fig. 29C), whereas members of P. deuvei have only two such setae, a feature particularly noted by Casale in his original description. Finally, the median lobe of male genitalia (Figs. 30A,C) has the shaft slightly less arcuate and the apex slightly longer on more rounded apically than in P deuvei males (see Casale 1998, Figs.4-5). For future comparative purposes, we also provide digital photographs of the median lobe of the genitalia of a male Pseudomasoreus catalai Jeannel (1949) (Figs. 30B,D) [male previously unknown] to complement the illustrations of other Pseudomasoreus species pro¬ vided by Jeannel (1949), Mateu (1980) and Casale (1998). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— Specimens of this species were are collected in secondary montane rainforest in the Talatakely and Vohiparara areas at elevations ranging from 900-1170 m. Adults were collected by hand, by beating live vegetation and suspended dead leaves and twigs, particu¬ larly dead leaves suspended is leaf whirls of Pandanus sp. Two specimens were found in daytime in masses of dead leaves, twigs and branch debris accumulated on the forest floor. Genus Assadecma Basilewsky 1982 This genus is precinctive to Madagascar and previously represented only by the type species, Assadecma madagascariensis Basilewsky (1982). Assadecma basilewskyi Kavanaugh and Rainio, sp. nov. Figure 31A-B, 32 Type material.— Holotype (Figs. 31A-B), a male, in NMNH, labeled: “MADAGASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 940m 21°16’S, 47°25’E 18 April 1994’V “pitfall traps in montane rainforest, E. Rajeriarison & R. Malenky”/ “HOLOTYPE Assadecma basilewskyi Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 2): 1 male (in NMNH) labeled same as holotype; 1 male (in CAS) labeled “CASENT 1051147”/ “MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area, 900 m, mixed tropical forest, 26 April KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 253 MADAGASCAR: Prov. Fianarant^oa, 7 km W Ranomafana, 940m 21*16*5, 47*25'E 18 April 1994 _ MADAGASCAR: Prov. Fianarant^oa, 7 km W Ranomafana, 940m Figure 31. Digital images of Assadecema spp. A-B. Holotype male of Assadecma basilewskyi Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels; C. A. madagascariensis Basilewsky female, dorsal habitus. Scale lines =1.0 mm. 254 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 1998.”/ “21°15.3’S 47°25.9’E, Stop # 98- 84 D.H. Kavanaugh collector collected at night on ground beside trail in forest”. Both paratypes also bear the following label: “PARATYPE Assadecma basi- lewskyi Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fia- narantsoa Province, Ranomafana National Park. Derivation ol species name.— The species epithet, basilewskyi, is a noun in apposition, based from the surname of Dr. Pierre Basilewsky, now deceased, who was for many years a lead entomologist at the Koninklijk Museum voor Midden Africa (Musee Royal de TAfrique Cen- trale) in Tervuren. We are pleased to name this new species in honor of Dr. Basilewsky’s many contributions to knowledge of the carabid beetle faunas of the Afrotropical Region in general and Madagascar in particular. Recognition.— Small for genus, SBL of males = 9.5-10.8 mm, female unknown. Basilewsky (1982) included a single species, A. madagascariensis in this genus. We have examined the holotype male of that species and found several fea¬ tures that differ with those found in specimens of A. basilewskyi (Fig. 31 A). The former is much larger (SBL = 12.9 mm), (about the same size as the A. madagascariensis female illustrated in Fig. 31C) than males of A. basilewskyi. Its elytral microsculpture is comprised of markedly transverse sculpticells, producing distinct iridescence, whereas in A. basilewskyi the sculpticells are only slightly transverse, producing no evident iridescence. Also the relative body shapes and proportions of males of the two species are distinctly different. In A. madagascariensis, the elytra are long (ratio EL/PL = about 2.5) and the elytral lateral margins are nearly parallel for most of their length (same as in the female, Fig. 31C). In A. basilewskyi, the elytra are relatively shorter (ratio EL/PL = about 2.3) and the elytral lateral margins are slightly rounded throughout (Fig. 31 A). Front tar- somere 4 in males of A. basilewskyi (Fig. 32A) is not as short and broad as that in A. madagas¬ cariensis (see Basilewsky, 1982:23, Fig. 3a). Finally, the median lobes of the genitalia of A. basilewskyi males (Fig. 32B-C) and A. madagascariensis males (see Basilewsky 1982:22, Fig. 2d) are markedly different in form. Geographical distribution.— At present, known only from the type locality. Habitat distribution.— The holotype and one paratype of this species were collected in a pitfall trap in montane rainforest at an elevation of 940 m. The second paratype was found at night walking on the ground beside a trail in the same general area. Figure 32. Digital images of holotype male of Assadecma basilewskyi Kavanaugh & Rainio sp. nov. A. Left front tarsus, dor¬ sal aspect; B-C. Aedeagus of male genitalia; B. Left lateral aspect; C. Ventral aspect. Scale lines = 0.5 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 255 Subtribe Lebiina Bonelli 1810 Genus Lebia Latreille 1802 As currently conceived, this genus is a megadiverse taxon with hundreds of species arrayed in 16 subgenera (Lorenz 2005) and with a worldwide distribution in tropical to temperate regions. Jeannel (1949) considered Lebia and Nematopeza Chaudoir (1870) as distinct genera, but the lat¬ ter is now generally treated as a subgenus of the former (Lorenz 2005). Subgenus Metalebia Jeannel 1949 Jeannel (1949) considered this subgenus to include most of the Lebia species of tropical Africa, Madagascar, and India. He reported 18 species in his treatment of the Malagasy fauna. Lorenz (2005) lists a total of 36 species for this subgenus. Lebia ( Metalebia ) laterolucida Kavanaugh and Rainio, sp. nov. Figure 33 Type material.— Holotype (Figs. 33A-B), a female, in CAS, labeled: “CASENT 1006164”/ MADAGASCAR Fianarantsoa Prov. Parc Nacional Ranomafana Bell Vue Trail, tropical forest 21°15.5’S 47°25.6’E 1000 m M.E.Irwin and E.I. Schlinger MEI 99-MA-7 21-XII-19997 “HOLO¬ TYPE Lebia (. Metalebia ) laterolucida Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (only 1): a female (in CAS) labelled “CASENT 1049030”/ “MADAGASCAR, Fia¬ narantsoa Province, Ranomafana National Park, Talatakely area, 850 m, 13 April 1998,”/ “21°14’S 47°22’E Malaise trap in mixed tropical forest, M.E.Irwin and E.I. Schlinger collectors, Stop 98- MAD-1”/ “PARATYPE Lebia ( Metalebia ) laterolucida Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, laterolucida, is an adjective derived from the Latin words, lateris, meaning side, and lucidus, meaning bright or full of light. The name refers to the pale, translucent lateral pronotal margins, which contrast markedly with the dark pronotal disc in members of this species. Recognition.— Size moderate for genus, SBL of females = 4.5-5.0 mm. Members of this species (Fig. 33A) key to subgenus Metalebia Jeannel (1949) and the madagascariensis group in Jeannel’s (1949) key: the basal elytral border is absent medially between the base of stria 3 and the scutellum; the elytral striae are deep and the elytral intervals convex and impunctate; the forebody (head plus pronotum) has no metallic reflection and the elytra are uniform in color; and the frons is slightly convex and without distinct frontal furrows between the eyes. Within the madagas¬ cariensis group, L. laterolucida members are distinguished from those of L. alluaudana Jeannel (1949), L. apicoviolacea sp. nov., L. mirana Alluaud (1936a), L. ranomafanae sp. nov., and L. tanala Jeannel (1949) by the absence of metallic reflection on the elytra and punctation of the anterior dorsum of the head (both features present in members of the other five species). They dif¬ fer from members of L. sulcipennis (Fairmaire) (1889), L. nana Jeannel (1949), and L. perrieri Jeannel (1949) in having the pronotum broad, ratio PW/PL = at least 1.5, and with lateral margins distinctly rounded at least in the anterior half pronotum (narrower and less rounded anteriorly in members of the three other species). From adults of L. rufa Jeannel (1949), they differ in having the pronotal basal lobe distinctly margined (absent from the former). Lebia laterolucida adults are most similar to those of the two remaining species in Jeannel’s key, L. brunneipennis Jeannel 256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 CASENT 1006164 MADAGASCAR Fianarantsoa Prov. Parc Nadonal Ranomafana Bell Vue Trail, * tropical forest 21*15 6 3 47* 25.6’E 1000 m M.E. Irwin and E.l. Schlinger ME! 99-MA-7 21-XI1-1999 Figure 33. Digital images of holotype female of Lebia ( Metalebia ) laterolucida Kavanaugh & Rainio sp. nov. A. Habi¬ tus, dorsal aspect; B. Labels. Scale line =1.0 mm. (1949) and L. madagascariensis Chaudoir (1850), but differ from them in having the dorsum of the head, pronotum and elytra dark piceous, with markedly contrasting pale yellow lateral pronotal margins (dorsum uniformly reddish brown and the lateral pronotal margins not or less markedly contrasting in color in the together two species). In addition, they are distinctly smaller than L. madagascariensis adults and have distinct elytral microsculpture (absent or at least very faint in L. brunneipennis adults). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— Both specimens of the type series were collected in mixed second¬ ary tropical forest in the Talatakely area in malaise traps. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 257 Lebia ( Metalebia ) ranomafanae Kavanaugh and Rainio, sp. nov. Figure 34 Type material. — Holotype (Figs.34A-B), a male, in NMNH, labeled: “MADAGASCAR: Prov. Fianarantsoa, 7 km W Ranomafana, 1100m 22-31 October 1988 W. E. Steiner”/ “Malaise trap in small clearing, montane rain forest”/ “HOLOTYPE Lebia ( Metalebia ) ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (only 1): a female (in CAS) labeled “MADAGAS¬ CAR: Prov. Fianarantsoa, 7 km W Ranomafana, 900m 23-28 February 1990 W. E. Steiner”/ “Malaise trap in small clearing, montane rain forest”/ “PARATYPE Lebia {Metalebia) ranomafanae Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality. — Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name. — The species epithet, ranomafanae , is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition. — Size slightly small for genus, SBL of male = 4.3 mm, of female = 4.2 mm. Members of this species (Fig. 34A) key to subgenus Metalebia Jeannel (1949) and the madagas- cariensis group in Jeannel’s (1949) key: the basal elytral border is absent medially between the base of stria 3 and the scutellum; the elytral striae are deep and the elytral intervals convex and impunctate; the forebody (head plus pronotum) has no metallic reflection and the elytra are uni¬ form in color (but not necessarily in reflection); and the frons is slightly convex and without dis¬ tinct frontal furrows between the eyes. Within the madagascariensis group, L. ranomafanae mem¬ bers can be distinguished from those of all species except L. alluaudana, L. apicoviolacea sp. nov., L. mirana and L. tanala by the presence of elytral metallic reflection (metallic reflection absent from the elytra of members of the remaining seven species). They are distinguished from members of L. mirana by their reddish labrum (black in L. mirana members) and from those of L. alluau¬ dana by their small size and relatively short and broad elytra, with ratio of elytra length to width = 1.3 (size larger, SBL of males = 4.6-5.0 mm, of females 4.6-5.3 mm, and elytra relatively longer, with ratio EL/EW= 1.4, in L. alluaudana members). They are distinguished from members of L. tanala in having very dark and relatively faint elytral metallic reflection, pronotum with anteri¬ or angles indistinct, smoothly rounded from apical to lateral margin and not at all projected anteri¬ orly beyond apical margin, lateral margins nearly parallel in posterior two-thirds and not or only very slightly sinuate anterior to obtusely angulate hind angles (elytral metallic reflection more dis¬ tinct, pronotum with anterior angles broadly round but slightly projected anteriorly beyond apical margin, lateral margins distinctly sinuate anterior to sharp, rectangular hind angles slightly pro¬ jected laterally in members of L. tanala). Finally, they can be distinguished from members of L. apicoviolacea by slightly larger size and the faint, concolorous dark blue-green elytral metallic reflection and (size slightly smaller, SBL of male = 3.8 mm, of female = 4.0 mm, and elytra with distinct tricolored metallic reflection, green basally and on medial two-thirds, brassy on lateral one- third and violet in apical one-fifth, in L. apicoviolacea members. Shape of the median lobe of the male aedeagus (Fig. 34C-D) is also diagnostic, with the shaft broadest at basal one-third, slightly and evenly arcuate on ventral margin in lateral view (Fig. 34C) and apex parallel-side basally and broadly rounded apically in dorsal view (Fig. 34D). Geographical distribution. — At present, known only from the type locality. Habitat distribution. — Both specimens of the type series were collected in malaise traps in secondary montane rainforest in the Talatakely area at elevations ranging from 900 to 1100 m. 258 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 MADAGASCAR; Pcov. Fianarant soa, 7 km W Ra noma f ana, ifoom 22-31 October 1968 W. E, Steiner Malaise trap in small clearing* montane rain forest B Figure 34. Digital images of holotype male of Lebia ( Metalebia ) ranomafanae Kavanaugh & Rainio sp. nov. A. Habi¬ tus, dorsal aspect; B. Labels; C-D. Aedeagus of male genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines = 0.5 mm. KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 259 Lebia ( Metalebia ) apicoviolacea Kavanaugh and Rainio, sp. nov. Figure 35 Type material.— Holotype (Figs. 35A-B), a male, in CAS, labeled: “MADAGASCAR: Fia- naran. Pr. Ranomafana N.P. HQ. area 21°15’24”S 47°25’15E 25NOV1994, at night M.A. Ivie & D.A. Pollack”/ “HOLOTYPE Lebia (Metalebia) apicoviolacea Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (only 1): a female (in NMHN) labeled “MADAGASCAR: Prov. Fia- narantsoa, 7 km W Ranomafana, 1100 m 6-21 October 1988 W. E. Steiner”/ “At black light in mon¬ tane rain forest”/ “PARATYPE Lebia (Metalebia) apicoviolacea Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, apicoviolacea, is an adjective derived from the Latin words, apex , meaning tip, and violaceus, meaning violet-colored. The name refers B Figure 35. Digital images of holotype male of Lebia ( Metalebia ) apicoviolacea Kavanaugh & Rainio sp. nov. A. Habi¬ tus, dorsal aspect; B. Labels; C-D. Aedeagus of male genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines A = 1.0 mm, C-D = 0.5 mm. 260 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 to the violet or purple metallic reflection of elytral apices, which contrasts distinctly with the green or bronze metallic reflection of the remainder of the elytra in members of this species. Recognition.— Size moderately small for genus, SBL of male = 3.8 mm, of female = 4.0 mm. Members of this species (Fig. 35 A) key to subgenus Metalebia Jeannel (1949) and the mada- gascariensis group in Jeannel’s (1949) key: the basal elytral border is absent medially between the base of stria 3 and the scutellum; the elytral striae are deep and the elytral intervals convex and impunctate; the forebody has no metallic reflection and the elytra are uniform in color (but not nec¬ essarily in reflection); and the frons is slightly convex and without distinct frontal furrows between the eyes. Within the madagascariensis group, L. apicoviolacea members can be distinguished from those of all species except L. alluaudana, L. mirana Alluaud, L. ranomafanae and L. tanala by the presence of elytral metallic reflection (metallic reflection absent from the elytra of members of the remaining seven species). They are distinguished from members of L. mirana by their reddish labrum (black in L. mirana members) and from those of L. alluaudana by their small size and rel¬ atively short and broad elytra, with ratio of elytra length to width = 1.2-1.3 (size larger, SBL of males = 4.6-5.0 mm, of females 4.6-5.3 mm, and elytra relatively longer, with ratio of elytra length to width = 1.4, in L. alluaudana members). They are distinguished from members of L. tanala in having the pronotum with lateral margins slightly sinuate anterior to slightly obtuse hind angles (lateral margins distinctly sinuate anterior to sharp, rectangular hind angles slightly projected laterally in members of L. tanala) and from members of both L. ranomafanae and L. tanala in having elytra with distinct tricolored metallic reflection, green basally and on medial two-thirds, brassy on lateral one-third and violet in apical one-fifth (elytral metallic reflection uni¬ formly faint or distinct metallic green or blue in members of L. ranomafanae and L. tanala). Shape of the median lobe of the male aedeagus (Fig. 35C-D) is also diagnostic, with the shaft very slight¬ ly arcuate, dorsal and ventral margins more or less parallel in basal two-thirds in lateral view (Fig. 35C) and apex moderate in length and relatively broader in both lateral and dorsal (Fig. 35D) views than in males of other species of this species group (compare with Jeannel 1949, Fig. 440a-l). Geographical distribution.— At present, known only from the type locality. Habitat distribution.— Both specimens of the type series were collected in secondary mon¬ tane rainforest in the Talatakely area at elevations ranging from 900 to 1100 m. Both were collect¬ ed at night, one by hand from vegetation, and the other attracted to ultraviolet light. Tribe Galeritini LeConte 1853 Genus Eunostus LaPorte 1835 This genus occurs only in Africa and Madagascar (Jeannel 1949) and includes 14 previously described species (Lorenz 2005), six of which are endemic to Madagascar. Jeannel (1949) provid¬ ed a key to the six Malagasy species known to him. Eunostus minimus Kavanaugh and Rainio, sp. nov. Figures 36-37 Eunostus n. sp. 1; Rainio 2009: 31, Rainio 2012: 72 (informal designation). Type material.— Holotype (Figs. 36A-B), a male, in CAS, labeled: “CASENT 8069195”/ “MADAGASCAR: Fianarantsoa Parc National de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana elev 1100m 27-31 Mar 2003”/ “21°17’24”S 047°26’00”E California Acad, of Sciences coll. Fisher, Griswold et al. sifted litter, montane rainforest collection code: BLF8400”/ “HOLOTYPE Eunostus minimus Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 261 CASENT 8069195 MADAGASCAR: Fianarantsoa Parc National de Ranomafana, Vatoharanana River, • 4.1 km 231* SW Ranomafana eiev 1100 m 27-31 Mar 2003 21*17 24’ S 047* 26'00" E California Acad, of Sciences coll. Fisher, Griswold et al. * sifted litter, montane rainforest collection code: BLF8400 1 Figure 36. Digital images of holotype male of Eunostus minimus Kavanaugh & Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels. Scale line A =1.0 mm. 2): 1 male (in MNHN) labeled “CASENT 8069129”/ “MADAGASCAR: Fianarantsoa, Ranomafana JIRAMA water works, 21°14.91’S, 047°27.13’E’7 “21-24 December 2001 collector: R. Harin’Hala California Acad of Sciences malaise trap near river elev. 690 m, MA-02-09D-08”; 1 female (in CAS) labeled “CASENT 1051146”/ “MADAGASCAR Ranomafana NP, Vatoharana trail S 600 24.8.2004”/ “collected by hand in daytime, F. Ratalata & J. Rainio collectors”. Both paratypes also bear the following label: “PARATYPE Eunostus minimus Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation of species name.— The species epithet, minimus, is the Latin adjective meaning smallest, a reference to fact that the known specimens of this species are smaller than those of any of the other known species that we have examined. Recognition.— Small for genus, SBL of male = 5.5 mm, of female 5.0-5.1 mm. Members of this species (Fig. 36A) do not key out in Jeannel’s (1949) key for Eunostus species. They are much smaller and shinier than members of any other described species. Their heads are slightly narrower, with the tempora more evenly rounded to the back of the head, their antennal scapes 262 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 Figure 37. Digital images of Eunostus minimus Kavanaugh & Rainio sp. nov. A-B. Apical part of left hind femur, oblique medial aspect; A. Holotype male; B. Female; C-D. Aedeagus of holotype male genitalia; C. Left lateral aspect; D. Dorsal aspect. Scale lines = 0.5 mm. shorter and thicker than in other species, thickest slightly distal to mid-length. Lateral margins of the pronotum are smoothly rounded in anterior half with a moderately short and deep subbasal sin- uation anterior to slightly obtuse to rectangular hind angles. The elytral setae are generally sparser than in other species and arranged in 2 more or less distinct longitudinal rows on each interval. Male (Fig. 37A) and female (Fig. 37B) hind femora are sexual dimorphic, each unique among Malagasy species for each sex. Median lobe of male genitalia Figs. 37C-D markedly different in shape from those of the other species for which males are known (see Jeannel 1949, Fig. 513d-e). Geographical distribution.— At present, known only from the type locality. Habitat distribution. — Specimens of the type series were collected in both primary mon¬ tane rainforest in the Vatoharanana area (at elevations ranging from of 1100 to 1200 m) and in sec¬ ondary montane rainforest 1 km north of Ranomafana village at an elevation of 690 m. The spec¬ imens were collected in a malaise trap and sifted from forest litter, respectively, at the primary for¬ est site and by hand at the secondary forest site. Tribe Helluonini Hope 1838 Genus Erephognathus Alluaud 1932 This genus is endemic to Madagascar and includes only two previously described species (Lorenz 2005). Jeannel (1949) provided a key to those species. Erephognathus ranomafanae Kavanaugh and Rainio, sp. nov. Figure 38 Type material.— Holotype (Figs.38A-B), a female, in CAS, labeled: “CASENT 806853 l’V “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, Vohiparara, at broken KAVANAUGH & RAINIO: NEW SPECIES OF BEETLES FROM MADAGASCAR 263 CASENT 8068531 MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana^ Vohtparara at broken bridge, el 1110 m 8 >15 November 2001 21° 13.57'S. 47° 22.19’ E collector: R. Harin’Hels California Acgd of Sciences maiaise trap in high altitude rainforest, MA-02-09A-02 B Figure 38. Digital images of holotype female of Erephognathus ranomafanae Kavanaugh and Rainio sp. nov. A. Habitus, dorsal aspect; B. Labels. Scale line = 1.0 mm. bridge, el 1110 m 8-15 November 2001”/ “21°13.57’S, 047°22.19’E collector: R. Harin’Hala Cal¬ ifornia Acad of Sciences malaise trap in high altitude rainforest, MA-02-09A-02”/ “HOLOTYPE Erephognathus ranomafanae Kavanaugh & Rainio sp. n. 2015” [red label]. Paratypes (total of 2): 1 female (in CAS) labeled same as holotype, except “CASENT 8068532”; 1 female (in CAS) labeled “CASENT 8068672”/; “MADAGASCAR: Province d’Antananarivo 3 km 41° NE Andra- nomay, 11.5 km 147° SSE Anjozorobe elev 1300 m 5-13 December 2000”/ “18°28’24”S 264 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 7 47°57’38”E coll. Fisher, Griswold et al. California Acad, of Sciences montane rainforest, malaise trap, coll, code: BLF2375”. Both paratypes also bear the following label: “PARATYPE Erephog- nathus ranomafanae Kavanaugh & Rainio sp. n. 2015” [yellow label]. Type locality.— Madagascar, Fianarantsoa Province, Ranomafana National Park. Derivation ol species name. — The species epithet, ranomafanae , is a noun in apposition, derived from the name of the national park in which the type was collected. Recognition.— Size moderate for genus, SBL of females = 7.5 - 8.9 mm. In Jeannel’s (1949) key to species of Erephognathus, members of E. ranomafanae key to E. coerulescens (Fairmaire 1903) based on their short antennae (extended posteriorly only to the basal one-fourth of the ely¬ tra or less) and the absence of the tuft of setae from the apex of antennomere 11 (terminal anten- nomere) found in E. margarithrix Alluaud (1936a) members. The pronotum (Fig. 38A) is distinct¬ ly longer, narrower and less convex and with anterior angles more projected, narrowed and angu- late than in members of E. coerulescens. Geographical distribution.— We have examined specimens from only two areas. Two female specimens were collected in Ranomafana National Park, the type locality, on the eastern slope off the central plateau in the northeastern part of the southern half of the island of Madagas¬ car. A third specimen, also a female, was collected near Anjozorobe, in Antananarivo Province, slightly north of the center of the island and just over 300 km north of Ranomafana along the east¬ ern slope off the central plateau. No specimens have been recorded from intervening areas. Habitat distribution.— Specimens of the type series were all collected in malaise traps in montane rainforest at elevations ranging from 1110 to 1300 m. Acknowledgements Fieldwork conducted by DHK and other staff from the California Academy of Sciences and their Malagasy colleagues was generously supported by a grant from Oracle Corporation, Redwood City, California. The Academy of Finland provided similar support for JR and her team during their work in Ranomafana and the F inni sh Cultural Foundation provided a grant for the descriptions of new species. We also gratefully acknowledge the Institute for the Conservation of Tropical Envi¬ ronments (ICTE) at Stony Brook University and Madagascar Institute pour la Conservation des environments Tropicaux (MICET) in Madagascar for providing superb logistical support and facil¬ itating permits for our work in Ranomafana National Park. Finally, we must thank Roberta Brett, Emily Elsom Cunningham, Charles Griswold, Kathryn Kavanaugh, Norman Penny, M.J. Raheri- lalao, E. Rajeriarison, R. Ranaivosolo, J.S. Randrianarisoa, E.F. Randrianirina, N. Rasoamananana, Francois Ratalata, Felix Jean Ratelolahy, T.J. Ravelomanana, H.C. Raveloson, Jere Schweikert, Darrel Ubick, Flor Vargas, Heidi Viljanen, Helena Wirta and Kipling Will for their help with col¬ lecting specimens. References Alluaud, C. 1897. Descriptions de Carabiques nouveaux de Madagascar. Annales de la Societe Ento- mologique de France 66:165-181. Alluaud, C. 1899a. 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Descrive i nuovi Caelostomus colasi, minusculus, oblongus e substriatus del Madagas¬ car ed enuncia nuovi caraterri dei Caelostomini e del gen. All. Revue Franqaise dEntomologie 7:36-41. Straneo, S.L. 1942. Revisione del Caelostomini Africani (Colopt. Carabid.). Memoire della Societa Ento- mologica Italiana 21:21-164. Tschitscherine, T.S. 1899. Memoire sur qulques Platysmatini nouveaux ou peu connus d’Afrique et de Madagascar. Annales de la Societe Entomologique de France 68:50-82. Tschitscherine, T.S. 1903. Nouvelles espouvelles especes de la tribu des Platysmatini. Horae Societatis Entomologicae Rossicae 36:38-64. CALIFORNIA ACADEMY OF SCIENCES INSTRUCTIONS TO AUTHORS Authors planning to submit papers for consideration for publication in the Academy’s Proceedings, Occasional Papers, or Memoir series must follow the directions given below in preparing their submissions. 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COMMENTS Address editorial correspondence or requests for pricing information to the Editor, Scientific Publications Office, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118 U.S.A. or via email to the Editor, Scientific Publications, at aleviton@caIacademy.org or gwilliams@calacademy.org Table of Contents Gary C. Williams and Odalisca Breedy: A New species of Whip-Like Gorgonian Coral in the Genus Swiftia from the Gulf of the Farallones in Central California, with a Key to Eastern Pacific Species in California (Cnidaria, Octocorallia, Plexauridae).1 Luis M. P. Ceriaco, Sango dos Anjos Carlos de SA, Suzana Bandeira, HilAria Valerio, Edward L. Stanley, Arianna L. Kuhn, Mariana P. Marques, Jens V. Vin- dum, David C. Blackburn, and Aaron M. Bauer: Herpetological Survey of Iona Na¬ tional Park and Namibe Regional Natural Park, with a Synoptic List of the Amphibians and Reptiles of Namibe Province, Southwestern Angola.15 Robert V. Dowell, Raymond J. Gill, Daniel R. Jeeske, and Mark S. Hoddle: Exotic Terrestrial Macro-Invertertebrate Invaders in California from 1700 to 2015: An Analy¬ sis of Records.63 Thomas F. Daniel and Ramiro Cruz Duran: Tetramerium vargasiae (Acanthaceae), a new species from the Basin of the Rio Balsas in Guerrero, Mexico.159 Thomas F. Daniel: Avicennia (Acanthaceae: Avicenioideae) in North America and Mesoamerica.163 Peter Wirtz and Tomio Iwamoto: A New Species of Serranus from Sao Tome and Principe, Eastern Atlantic (Pisces, Teoeostei, Serranidae).191 David H. Kavanaugh and Johanna Rainio: Twenty-six New Species of Predaceous Ground Beetles (Coleoptera: Adephaga: Carabidae) from Ranomafana National Park, Madagascar.201