36s AW ALAS, eR WHER Poe Th) ay Hs te haths RAArba nh VisMA be Pah em men Toe ev Mi tele uh te ahs Lusty De ay, a Afi tae r ote PFE WPL Mises nye sy Pi An Bos ra > SS MN DOM T aera Pe Fans SH ee on ae cmas ate eee ety SON Dt, 2: IS ate haint NE NAME Ma Bas ee een MeSH eRe in act, MR KE ieitene, eae ne ee Fe tv tyne Orn NEI at mays Tien, ots 5 oe | Vol SRA . = a | | ay . < e | RAN PR ANE DEC Ay Ue eh 5 ea pare " ) a ‘ Ww \ igiey uc a ae uy My ¥ , A ie 1 ' ah vn y Mey , te i mt Ay i hy Ke i‘ i M | i i ' | j My ci ; ny PROCEEDINGS OF THE California Academy of Sciences FOURTH SERIES Vol. XXXIX SAN FRANCISCO PUBLISHED BY THE ACADEMY 1972-1974 COMMJTTEE ON PUBLICATION GrorcE E. Lrnpsay, Chairman Diana R. Youne, Editor No. No. No. No. No. No. alle lz. CONTENTS OF VOLUME XXXIX HERTLEIN, LEo G. Description of a new species of Chlamys (Mollusca: Pelecypoda) from the Galapagos Nietegrmd pee 908 So 0 5 th a a ie Pe Oh Bn SmitH, ALLYN G. Three new land snails from Isla Santa Cruz (Indefatigable Island), Galapagos -_______________ HERTLEIN, Leo G. Pliocene fossils from Baltra (South Seymour) Island, Galapagos Islands ——_______ ABE, TOKIHARU, and WILLIAM N. ESCHMEYER. A new species of the scorpionfish genus Helicolenus from the north apiCHOccahee tt AONE Pe ood ee Neat oe eee ESCHMEYER, WILLIAM N., and K. V. Rama Rao. Two new scorpionfishes (genus Scorpaenodes) from the Indo-west Pacific, with comments on Scorpaenodes muciparus (Phe encica) eee iets, Bee 2 AR BE ZULLO, Victor A., DEA B. BEACH, and JAMEs T. CARLTON. New barnacle records (Cirripedia, Thoracica) —...__ CAsTEEL, RicHarp W. A key; based on scales, to the families of native California freshwater fishes AsHLOCK, PETER D. The Lygaeidae of the Galapagos islands< (Hemiptera; «Eleteroptera). 2d Brown, WALTER C., and Marjorie V. C. FALANRUW. A new lizard of the genus Emoia (Scincidae) from the ianataiicse USGS 9 aoe Mes ee a McCoskeEr, JOHN E. Two new genera and two new species of western Pacific snake-eels (Apodes: Ophichthidae) HERALD, Ear S., and JoHN E. RANDALL. Five new Indo- Ge SIC EISINC Sie eres en eee CHEMSAK, JOHN A., and E. G. Linstey. The genus Mecas ieConte (Colecptera:Cerambycidae) 2. == Pages 47-53 55-64 65-74 75-86 87-103 105-110 111-120 121-140 141-184 No. 13. No. 15. No. 19. Now ZL. No. 23 SMITH-VANIZ, WILLIAM F., and Jon C. STAIGER. Compar- ative revision of Scomberoides, Oligoplites, Parona, and Hypacanthus with comments on the phylogenetic position of Campogramma (Pisces: Carangidae) ComPAGNoO, L. J. V. Ctenacis and Gollum, two new genera of sharks (Selachu? Carcharimidat) ">" eae MAnpRA, York T., A. L. BRIGGER, and H1GHooHT MANDRA. Chemical extraction techniques to free fossil silicoflagellates from marine’ sedimentary rocks ==" =" ee ESCHMEYER, WILLIAM N., YosuHitsuGu Hrrosakti, and TOKIHARU ABE. Two new species of the scorpionfish genus Rhinopias, with comments on related genera and species —___ GarTH, JOHN S. The brachyuran crabs of Easter Island _ ESCHMEYER, WILLIAM N., and Kaza V. Rama Rao. Two new stonefishes (Pisces: Scorpaenidae) from the Indo-west Pacific, with a synopsis of the subfamily Synanceiinae _____ Compacno, L. J. V. Gogolia filewoodi, a new genus and species of shark from New Guinea (Carcharhiniformes: Triakidae), with a redefinition of the family Triakidae and a key*to Triakid ‘genera’ 4° eee Howarp, L. D. Muscular anatomy of the forelimb of the sea otter (Enhydra lutris) i... Oe ee BURGHARDT, GLENN FE. A _ new Hawaiian chiton Rhyssoplax linsleyi (Mollusca: Amphineura: Chitonidae) — Iwamoto,Tomio. Nezumia (Kuronezumia) bubonis, a new subgenus and species of grenadier (Macrouridae: Pisces) from Hawaii and the western North Atlantic — SCHRADER, HANS-JOACHIM. Revised diatom stratigraphy of the Experimental Mohole Drilling, Guadalupe Site Index to: Volume: XOX XIX$ 2) es eee 185-256 257-272 273-284 285-310 311-336 337-382 383-410 411-500 501-506 507-516 517-562 563-580 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXIX, No. 1, pp. 1-6; 5 figs. January 21, 1972 DESCRIPTION OF A NEW SPECIES OF CHLAMYS (MOLLUSCA: PELECYPODA) FROM THE GALAPAGOS ISLANDS By Leo G. Hertlein INTRODUCTION The known marine molluscan fauna of the Galapagos Islands in general is sparser than that of the mainland. Eleven species of scallops (Pectinidae) have been reported living in the Galapagos Archipelago. These are: Pecten (Flabelli- pecten) sericeus Hinds, Pecten (Oppenheimopecten) galapagensis Grau, Chlamys (Chlamys) lowei Hertlein, Chlamys (Argopecten) circularis Sowerby, Chlamys (Nodipecten) magnifica Sowerby, Cyclopecten (Cyclopecten) exquisitus Grau, Cyclopecten (Cyclopecten) pernomus Hertlein, Cyclopecten (Delectopecten) polyleptus Dall, Cyclopecten (Delectopecten) zacae Hertlein, and from very deep water, Cyclopecten (Cyclopecten) liriope Dall and Cyclopecten (Hyalopec- ten) neoceanicus Dall. Seven of these are known to occur along the adjacent mainland. In 1969, Mr. Anthony D’Attilio, San Diego Society of Natural History, sent six paired valves (four with the animal) and four single valves representing a species of Chlamys to Allyn G. Smith, Department of Invertebrate Zoology, California Academy of Sciences, with a request for identification of the species. These subsequently were submitted to me for study. A search of the Academy’s collection as well as of the literature failed to reveal any similar species described from the eastern Pacific. Upon receipt of this information Mr. D’Attilio re- quested me to describe the species which appears in the present paper. [1] Si a eae me JAN 3 1 1972 RMdie oo ale lala eee a ET Ges rine Bicsiogical L vor tary Ce 2 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. ACKNOWLEDGMENTS The writer here expresses his thanks to the following persons: to Mr. Anthony D’Attilio, San Diego Society of Natural History, for permission to describe the new species and to retain paratypes; to Mr. Allyn G. Smith, Cali- fornia Academy of Sciences, for advice concerning the specimens; to Mr. Barry Roth of the same institution for critical reading of the manuscript and for arrangement of the illustrations; to Dr. Thomas R. Waller, Division of Inverte- brate Paleontology, United States National Museum, for information concerning comparative species in the collections of the National Museum; to Mr. Maurice Giles, Staff Photographer, California Academy of Sciences, who prepared the photographs from which the illustrations were made. DESCRIPTION OF NEW SPECIES Family PECTINIDAE Rafinesque Genus Chlamys Roding in Bolten Chlamys (Chlamys) incantata Hertlein, new species. (Figures 1-5.) Dracnosis. A species of Chlamys differing from other west American species in possessing very narrow, compressed, rather low, spinose ribs. DeEscriIPTION. Shell averaging about 45 mm. in height, ovate, valves gently and nearly equally inflated, hinge line rather short. Right valve sculptured with about 25 major ribs which are narrow, compressed, rather low, and occasionally unequally spaced; on top of each of these ribs is a row of spines which are con- cave ventrally; submargins with 5 or 6 very fine riblets; interspaces vary in width but are much wider than the ribs, nearly flat-bottomed, and sculptured with 1 to 3 fine radial threads, each bearing a row of spines, the ribs and inter- spaces crossed by fine concentric imbricating lines of growth; auricles unequal, the anterior one the larger, sculptured with about 5 spiny radial riblets, the hinge line above the auricle with scaly sculpture, below the auricle a well developed byssal notch about half the length of the auricle, below this along the margin there are 4 pectinidial teeth; posterior ear short, slightly concave, sloping rather steeply downward, sculptured with about 6 radial riblets. Left valve sculptured similar to the right but lacking a byssal notch. Hinge with one pair of slight cardinal crura. Interior of valves lightly grooved corresponding to the external ribbing. Color of the exterior of the valves whitish, the spines rosy, lending a roseate appearance to the valves, more pronounced on the left one. Dimensions. Length 40 mm., height 45 mm., length of hinge line 21 mm., convexity (both valves together) approximately 13 mm. The largest specimen, a left valve, is 48 mm. high. TYPE MATERIAL. Holotype no. 52263, also paratypes, San Diego Society of Natural History, from off Academy Bay, Santa Cruz (Indefatigable) Island, VoL, XXXIX] HERTLEIN: A NEW SPECIES OF CHLAMYS 3 Ficure 1. Chlamys (Chlamys) incantata Hertlein, new species. Holotype no. 52263 (San Diego Society of Natural History), from off Academy Bay, Santa Cruz (Indefatigable) Island, Galapagos Islands, dredged in 200 meters. View of the exterior of the left valve; height 45 mm. Ficure 2. Chlamys (Chlamys) incantata Hertlein, new species. View of the exterior of the right valve of the holotype. Ficure 3. Chlamys (Chlamys) incantata Hert- lein, new species. Paratype (SDSNH), from the same locality as the holotype. View of the exterior of a left valve; height 48 mm. Ficure 4. Chlamys (Chlamys) incantata Hertlein, new species. Paratype (SDSNH), from the same locality as the holotype. View of the exterior of a right valve; height 39.5 mm. Ficure 5. Chlamys (Chlamys) incantata Hertlein, new species. View of the interior of the right valve shown in figure 2. Galapagos Islands; dredged in 200 meters; Mrs. Jacqueline DeRoy collector, July 25, 1969. Paratypes are deposited in the California Academy of Sciences, the Amer- ican Museum of Natural History, and in the United States National Museum. CoMMENTS. The shape, spinose sculpture, and rosy coloration of this new 4 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. species are somewhat similar to those of Chlamys hastata Sowerby (see Arnold, 1906, pl. 42, figs. 1, la, 2, 2a; Grau, 1959, plates 27, 28) from California, but the ribs are more numerous, lower, more compressed, and not arranged in pairs on the right valve. The fewer ribs (25) on the new species serve to separate it from Chlamys amandi Hertlein (1935, p. 305; Pecten australis Philippi, 1845, p. 56 “Patriae: Insulae Chonos.”; not Pecten australis Sowerby, 1842) from Chile, which has 30 to 34 ribs which are only slightly scaly toward the submargins. The sculpture of C. incantata, new species, bears a general resemblance to that of the species described as Pecten (Chlamys) coccymelus by Dall (1898, p. 741, pl. 34, fig. 1) from strata of middle Miocene age at Plum Point Maryland. Dall called attention to the similarity of the fossil form to juvenile Pecten madisonius Say, but believed it to be a distinct species. Mansfield (1936, p. 177) stated that it “may be an immature specimen representing a varietal form of P. madisonius,’ and Rowland (1936, p. 1008) suggested that it ‘““may be a case of arrested development.” The sculpture of the new species described here bears a general similarity to that of illustrations of Pecten (Chlamys) nympha Bavay (1906, p. 246, pl. 7, figs. 3 and 4). The type specimen of that species was described as only about 15 mm. high. The type locality was given as “Habitat Caribaeum Mare?”. The type specimen in the Muséum National d’Histoire Naturelle de Paris, was originally in a carton which also contained a specimen of Pecten antillarum Récluz, a typical inhabitant of Caribbean waters. Dr. Thomas R. Waller (written communication, December 30, 1969), who has given considerable time to a study of the Pectinidae of the western Atlantic, stated that he feels certain that Pecten (Chlamys) nympha is a synonym of Chlamys benedicti Verrill and Bush (in Verrill, 1897, p. 74; not Pecten benedicti Lamarck, 1819) described from “off Martha’s Vineyard, 1356 fath., dead; West Indies, in 25 to 72 fath., living.” Adult specimens of C. benedicti (Weisbord, 1964, pl. 14, figs. 8-11) are quite distinct from the new species described here from the Galapagos Islands. The specific name of this new species, “cantata,” is derived from the vernacular appellation, “Las Islas Encantadas” (the Enchanted Islands), some- times applied to the Galapagos Islands. LITERATURE CITED ARNOLD, R. 1906. The Tertiary and Quaternary Pectens of California. United States Geological Survey, Professional Paper no. 47, pp. 1-264, pls. 1-53, 2 figs. in text. Bavay, A. 1906. Sur quelques especes ou variétés nouvelles du genre Pecten. Journal de Conchyli- ologie, vol. 53, no. 3, pp. 243-247, pl. 7, February 20. Dati, W. H. 1898. Contributions to the Tertiary Fauna of Florida with especial reference to the Silex beds of Tampa and the Pliocene beds of the Caloosahatchie River including Vor. XXXIX] HERTLEIN: A NEW SPECIES OF CHLAMYS 5 in many cases a complete revision of the generic groups treated and of their American Tertiary species. Transactions of the Wagner Free Institute of Sci- ence of Philadelphia, vol. 3, pt. IV, pp. I-VIII, 571-947, pls. 23-35, April. Grau, G. 1959. Pectinidae of the eastern Pacific. Allan Hancock Pacific Expeditions, vol. 23, pp. I-VII, 1-308, pl. 1-57, September 25. HERTLEIN, L. G. 1935. The Templeton Crocker Expedition of the California Academy of Sciences, 1932. No. 25. The Recent Pectinidae. Proceedings of the California Academy of Sciences, Fourth series, vol. 21, no. 25, pp. 301-328, pls. 18-19, September 26. MANSFIELD, W. C. 1936. Stratigraphic significance of Miocene, Pliocene, and Pleistocene Pectinidae in the southeastern United States. Journal of Paleontology, vol. 10, no. 3, pp. 168- 192, 1 text fig., pls. 22-23, April. Rowranp, H. I. 1936. The Atlantic and Gulf Coast Tertiary Pectinidae of the United States. II. Sys- tematic Descriptions, continued. American Midland Naturalist, vol. 17, no. 6, pp. 985-1017, pls. 5-10, November. VerRRILL, A. E. 1897. A study of the Family Pectinidae, with a revision of the Genera and Subgenera. Transactions of the Connecticut Academy of Sciences, vol. 10, pt. 1, pp. 41-95, pls. 16-21, June. WEIsBorD, N. E. 1964. Late Cenozoic Pelecypods from Northern Venezuela. Bulletins of American Paleontology, vol. 40, no. 204, pp. 1-564, pls. 1-59, February 18. Pig ne ae Shy WY aa ae a aaeay ait) ei b 2 > &? wah is : 7] nao wy 7 (ar? a Tif se. — > Far liieei® ralicrw - - 7 b fh aut ® a 2175.9) or : 3/= y a oi == ' 2 ? - q ' Rhee é a! wf ee ys ° PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Volume XXXIX, No. 2, pp. 7-24; 25 figs. January 21, 1972 THREE NEW LAND SNAILS FROM ISLA SANTA CRUZ (INDEFATIGABLE ISLAND), GALAPAGOS ey Allyn G. Smith Associate Curator, Department of Invertebrate Zoology, California Academy of Sciences From late January to early March, 1964, I had the opportunity to collect land snails on one of the larger of the Galapagos Islands as a participant in the Gala- pagos International Scientific Project (GISP). This expedition was sponsored by the University of California, the California Academy of Sciences through the Belvedere Scientific Fund, and the Charles Darwin Foundation. Assistance, both financial and material, was provided by the government of the Republic of Ecuador, the United States Navy, the National Science Foundation of the United States*, the Shell Oil Company, and the California Maritime Commission whose training vessel, the Golden Bear, provided transportation of personnel and equip- ment to and from the Galapagos. The major portion of the five weeks in the Galapagos Islands was spent in collecting on Isla Santa Cruz (Indefatigable). The decision to concentrate on this island was made for several reasons. First, it is large, high, and well forested, containing all of the life zones occurring in the Galapagos group; second, the Darwin Research Laboratory close to the village of Academy Bay made an ideal headquarters; and third, the well developed trail from Academy Bay into the highland area, which was pioneered by the members of the California Acad- emy’s 1932 Expedition sponsored by Templeton Crocker, cut through all life zones and made good collecting spots accessible (see map, fig. 1). These reasons fortunately tied in with the fact that the late winter and spring of 1964 proved to be a good time to find land snails on Isla Santa Cruz. There was considerable * Through NSF Grant no. GE-2370 [7] Marine Biological Lebaratory LIBRA?CYy JAN 3 1 1972 Weods Hole, Mass. 8 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. DAPHNE MINOR © (Be ISLA SEYMOUR SKETCH MAP BY: NORTH CHANNELN David Q. Cavagnaro ned DAPHNE MAJOR @ Sigvart Horneman March, 1965 Based on U.S. Hydrographic Chart Nos. 5921 and 5939. Punto del Norte Punta Corrion BAHIA CONWAY A & ISLA SANTA CRUZ (INDEFATIGABLE) ° ROCAS PS GORDON SS istas Seoles|e Covagnaro Crater Qc es CENTRAL HIGHLAND CHAIN ae + No \ } Deep Crater W237 i ixe™, “* al - #2 - eae Crocker * Scalesia x Chimney.” 34 ** 4p Table Mountain Sonta)Rosci@). grassland Minow gee bonds ote DCroter : miconia grassland fi Camote NAMELESS ee {Le Copa © ISLAND aE PS ores: \miconla .~“ mora imoras. oo “ i “FARM ZONE =< @ Kastdalen Farmhouse ponds H “ss... Castro °,@Horneman Farmhouse El Chato Farmhouse Sow Bello Vista “\.@ El Rancho Puerto Nunez Transition c Transition Bahia Tortuga | Old Trail z New Trall “2*<.5 Coastal °s. Barranco ne Sei smograph = Coastal Way oT 20g ote 4.3 mm. = 1 km. Punta re lem. =23.14 km. Tomayo APPROXIMATE SCALE ~ Punta Nunez Darwin Research Station = © Jensen Island unto Ploya Estrada de los Perros "~ BAHIA ACADEMY Bahia Tortuga 1 Ficure 1. Outline map of Isla Santa Cruz (Indefatigable Island), Galapagos Islands, with some of the principal features sketched in. rainfall, even in the Arid Zone along the shores of the island, which promoted snail activity. The collections made through personal efforts were considerably augmented by other GISP scientists during their field work on other islands as well as Santa Cruz. To these appreciation is due. In addition, special thanks go to André and Jacqueline De Roy, a Belgian couple living at Academy Bay, for much pertinent information on possible productive collecting areas. Although not an active conchologist like his wife, M. De Roy became interested in the species of land snails to be found on his island and has made a number of special trips into uncollected areas in the highlands of Isla Santa Cruz in the past several years with signal success. The following new species discovered as a result of these efforts make a significant addition to the land snail fauna of the Galapagos. The symbols “CASG” and “CASIZ” represent separate collections maintained by the Califor- VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 9 nia Academy of Sciences’ Geology and Invertebrate Zoology departments, respec- tively, the latter consisting mainly of specimens preserved in alcohol. Naesiotus deroyi A. G. Smith, new species. (Figures 2-9.) DescripTION. Shell fairly large for the genus, elongate-conic, yellowish white at the apex graduating to pure white on the body whorl. Whorls about 7, gently rounded, the sutures well impressed. Aperture subquadrate, with 2 promi- nent denticles, the first a strong, rounded boss on the columella, and the second a less strong, somewhat laterally compressed, subtriangular, parietal denticle posi- tioned a short distance behind the plane of the aperture. Peristome thickened slightly but not reflected except on the basal portion, which partly covers a small shallow umbilicus. The peritreme is completed by a fairly heavy wash of callus. The columellar axis is simple, solid, and only slightly twisted. (See fig. 4.) Nuclear whorls 2’, erect, sculptured by many fine, closely spaced, occa- sionally anastomosing, slightly sinuate and protractive transverse lirae. The first postnuclear whorl is relatively smooth and marked by hardly noticeable lines of growth. Beginning with the second postnuclear whorl, a rough irregular but generally transverse wrinkling appears, becoming very heavy, somewhat warty, and more or less patternless on the penultimate and especially on the body whorl. Underlying this strong wrinkled sculpture are fine, closely spaced, spiral lirae, also beginning on the second postnuclear whorl, but not overriding the raised areas forming the wrinkles. The animals, in alcohol, are yellowish white in color with no apparent special markings on the mantles. Unfortunately, I did not see specimens alive. Hototype. An adult shell (CASG Type Collection, no. 13730) with animal in alcohol (CASIZ Type Series, no. 466), the shell measuring: height, 19.8; maximum diameter, 12.0; height of aperture, 8.8; maximum width of aperture, 7.6 mm. Number of whorls, 6%. The apical angle of the spire is about 50°. (Seedies: 2,3.) PaRATYPES. Seventy-two specimens collected with the holotype. These have been deposited in various museums having recognized type collections, including the CASIZ Type Series, and in several private collections. Type Locality. Isla Santa Cruz (Indefatigable Island), Galapagos, on the northwest side at an elevation of about 264 m. (870 ft.), on a species of thorny bush (locally called “mora,” the botanical name not available) by André De Roy, 18 February 1964. OTHER MATERIAL EXAMINED (ALL FROM ISLA SANTA CRUZ). (1) Three adult specimens (CASG no. 40129) found in a locality “1-2 hours walk west of the Horneman Farm,’’—at an elevation of about 200 m. (650 ft.). 10 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 4 2 Ficure 2. Naesiotus deroyi A. G. Smith, new species. Holotype. Height, 19.8 mm. Aper- tural view. CASG Type Collection, no. 13730. Ficure 3. Same. Back view. FIGURE 4. Same. Shell with cut-away section to show configuration of columella. Paratype. Height, 18.1 mm. CASG Type Collection, no. 13731. Ficure 5. Naesiotus cf. N. deroyi. Adult VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 11 These were the first ones found by André and Jacqueline De Roy in 1963 in the same general area as the type lot collected in 1964. Others are in the De Roy Collection. (2) Six adult shells, typical (CASG no. 40076), collected on the trail 6 miles to the west of the village of Bella Vista, on bushes, by André De Roy, 10 June 1964. (3) One typical but rather small subadult (CASG no. 40302), collected in an area to the north of the central chain of craters well north and slightly west of the village of Santa Rosa, on the ground in a dense forest of Scalesia pedunculata, by André De Roy, 26 November 1966. (4) One juvenile (CASG no. 40122), collected 5 miles northwest of Bella Vista, on “mora,” at about 265 m. (870 ft.) elevation by André De Roy, 18 February 1964. (5) Three rather small adults (CASG no. 40225), collected on the north side of the hill nearest to Santa Rosa Spring, on grass in a grassy glade, 200 m. (650 ft.) elevation, by André De Roy, 29 June 1965. (6) Fourteen adults and subadults (CASG no. 40022), found 2 km. north- east of the village of Santa Rosa, on the ground, by André and Jacqueline De Roy, 28 June 1965. This series is not typical. (7) Seven specimens (mostly subadults) preserved in alcohol (CASIZ Coll.), taken about 2 km. northeast of Santa Rosa, in open Scalesia forest hanging on the leaves of various plants and bushes, by André De Roy, 27 November 1966. This and the preceding lot are similar. Remarks. This is one of the largest and most strikingly sculptured of the Galapagos species of Naesiotus. No species closely similar has been described. While an average specimen is about 20 mm. high, with 7 whorls or slightly less, an unusually large one may be as much as 23 mm. in height, with the whorls numbering 7% to 72. It appears to be a terminal species in the group of Galapagos snails that have developed a heavily wrinkled sculpture on nearly all of the postnuclear whorls rather than on the last portion of the body whorl, which < shell from lot no. 6 with weaker wrinkled sculpture. Hypotype. Height, 17.4 mm. CASG Type Collection, no. 13733. Apertural view. FIGURE 6. Same. Back view. FIGURE 7. Naesiotus deroyi. Adult shell from lot no. 5. Hypotype. Height, 17.5 mm. CASG Type Collection, no. 13734. Apertural view. Ficure 8. Same. Back view. FicurE 9. Galapagos Naesiotus related to N. deroyi. Left to right: N. rabzdensis (Dall, 1917) from Isla Rabida (Jervis Island), hypotype, height, 13.5 mm. (CASG Type Collection, no. 13735); N. sculpturatus (Pfeiffer, 1846) from Isla Santiago (James Island), hypotype, height, 13.6 mm. (CASG Type Collection, no. 13736); N. darwini (Pfeiffer, 1846) from the same island, hypotype, height, 15.9 mm. (CASG Type Collection, no. 13737) ; N. deroyi, holotype, fig. 2. 12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. is the type of sculpture on such species as Naesiotus lycodus (Dall, 1917), N. ochsneri (Dall, 1917), and several others. So far, NV. deroyi has been collected in the highland area of Isla Santa Cruz, only toward the northwest side of the island; it seems to prefer the moister forested area characterizing the Scalesia Zone. The nearest relative is Naesiotus darwini (Pfeiffer, 1846) from James Island, which is smaller, chunkier, and has a finer and much less rude type of wrinkled sculpture. Other species in the group include \. sculpturatus (Pfeiffer, 1846) from James Island, and NV. rabidensis (Dall, 1917) from Jervis. (See fig. 9.) Snails from lots 6 and 7 listed under “other material examined” come from the same general locality as the others. However, they are smaller in size than those from the type locality, averaging 17.1 mm. in height, with a range of 15.7— 17.6 mm. Color is a light yellow-brown and all shells have a narrow, darker brown band encircling the periphery of the whorls; many have the entire base tinged with the same brown color. The wrinkled sculpture is much less strong, some shells having smooth patches with no wrinkles at all. (See figs. 5, 6.) As in some other Galapagos snail species, this particular population evidently represents an evolutionary trend of recent origin. Whether a subspecies is in the process of development is difficult to say until the range limits of both forms can be determined and other related factors studied. At present, it seems suffi- cient to call attention to the occurrence of another race closely allied to V. deroyi. It is with considerable satisfaction that I take this opportunity to name a striking new species of Galapagos land snail for M. André De Roy, who collected the first specimens as well as the type and other lots. Naesiotus cavagnaroi A. G. Smith, new species. (Figures 10-18.) DescripTION. Shell fairly large for the genus, smooth, broadly elongate- conic, with a fairly heavy texture and a tumid body whorl. Normal color pattern is reddish brown to chocolate brown with a narrow yellowish band coloring the sutures of the postnuclear whorls and encircling the body whorl slightly below its periphery. Whorls about 6’, rounded, the sutures impressed. Aperture ovate, white inside, with 2 well developed denticles, the first an elongate, rounded swelling on the columella, the second one smaller, arcuate, and parietal, set well inside the retractive plane of the aperture and forming a U-shaped bay with the columellar denticle. Peritreme rather sharp, thinned down toward its edge, not reflected, the outside edge yellowish in color. The peritreme is completed by a heavy layer of callus, especially in older shells. There is a small shallow umbil- icus partly covered by the basal reflection of the peristome. The columella is simple and only slightly twisted. Nuclear whorls about two, dimpled, appearing smooth to the naked eye but VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 13 under magnification revealing a sculpture of extremely fine, closely spaced, transverse lirations that are slightly sinuate and beaded at their summits as a result of being cut by excessively fine spiral striae. Postnuclear whorls sculptured by lines of growth and very fine, closely spaced, spiral striations, the latter best seen under considerable magnification. The surface of most shells exhibits a rather dull finish overall, with a tendency in some toward a more shining exterior. Occasional shells are yellowish in color with no suggestion of a revolving band. Animals in alcohol are tuberculate dor- sally and light grey in color, there being no appreciable color difference between those occupying normally colored or xanthic shells. Hototype. An adult shell preserved without the animal has been deposited in the CASG Type Collection, no. 13738. It measures: height, 22.7; maximum diameter, 13.9; height of aperture, 10.4; maximum width of aperture, 9.1 mm. Number of whorls, 6%. Apical angle, about 70°. (See figs. 10, 11.) ParaTyPeEs. A total of 52 specimens collected with the holotype. Of this total 30 have adult shells, 17 are subadults, and 6 are juveniles. Of this same total, 10 have xanthic shells (6 adults, 3 subadults, and 1 juvenile). About half of the type lot were collected alive; animals with their shells of a few of both color forms have been preserved in alcohol. A distribution of these paratypes will be made in a manner similar to that indicated for the previously described species (Naesiotus deroyt). Type Locatity. Isla Santa Cruz (Indefatigable Island), Galapagos, about 7 km. northeast of the farming village of Santa Rosa in the vicinity of a series of small volcanic craters in a Scalesia forest; collected under lava rocks and dead wood by André De Roy, 27 November 1966. OTHER MATERIAL EXAMINED. (1) Three dead, bleached, adult shells (CASG no. 40158) collected along the trail near the summit of Mt. Crocker at an elevation of 650-870 m. (2130- 2850 ft.) by Robert L. Pyle, 25 February 1964. (2) Seven dead, bleached, adult shells (CASG no. 40237) collected near an isolated rocky crater (subsequently designated as Cavagnaro Crater), in a Scalesia forest on the north slope of the island at an elevation of about 500 m. (1600 ft.) by David Q. Cavagnaro, 10 April 1964. (3) One dead, bleached adult shell (CASG no. 40512) found about 0.5 mile below Santa Rosa on the trail from Bella Vista by André and Jacqueline De Roy, 1 March 1965. (4) One dead but fresh adult shell, 1 subadult and 1 juvenile (CASG no. 40229) collected in the Scalesia forest near Cavagnaro Crater, elevation about 625 m. (2050 ft.), by André and Jacqueline De Roy, 2 March 1965. (5) Sixty-two dead adult shells (CASG no. 40221) collected on the ground 14 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER. 12 15 18 Ficure 10. Naesiotus cavagnaroi A. G. Smith, new species. Holotype. Height, 22.7 mm. CASG Type Collection, no. 13738. Apertural view. Ficurre 11. Same. Back view. FIGURE VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 15 in the vicinity of Cavagnaro Crater, elevation about 690 m. (2200 ft.), by André and Jacqueline De Roy, 2 March 1965. (6) One dead, bleached, adult shell (CASG no. 40228) collected on the ground near Santa Rosa school by André De Roy, 2 November 1965. (7) One dead, bleached, adult shell (CASG no. 40234) collected 2 miles from the Santa Rosa school on the trail from Bella Vista, by André De Roy, 2 November 1965. (8) Seven dead, bleached, adult shells (CASG no. 40295) collected in the vicinity of Cavagnaro Crater, by André De Roy, 23 November 1966. (9) Three bleached “bones” (CASG no. 40300) collected in a small lava cave to the west of Chimney Mountain, by André De Roy, 25 November 1966. (10) Thirty-eight adult shells, all xanthic (CASG no. 27537), collected “at the very top of the island” (7.e., summit of Mt. Crocker) by Templeton Crocker, 10 or 11 May 1932. (11) Forty-seven adult shells, all xanthic but darker in color than the pre- ceding lot (CASG no. 27538), collected “‘on the trip to top of Mt.,” by Templeton Crocker, 10 or 11 May 1932. (12) Two adults and 1 juvenile with color pattern reversed taken alive half way between Chimney Mountain and Santa Rosa Spring under low bushes in an open area, elevation about 600 m. (1950 ft.), by André and Tui De Roy, 27 June 1965. (13) Thirty-three mostly adult shells taken alive (CASG no. 43333; 15 in alcohol are CASIZ Collection) 2 miles west of Mt. Crocker at the foot of a volcanic crater on the ground among small trees, by André and Jacqueline De Roy, 10 May 1970. Remarks. This smooth brown snail with a yellowish revolving band has no close relatives of comparable size either on Isla Santa Cruz (Indefatigable Island) or on any of the other Galapagos Islands. Its range on Indefatigable is limited to the north slope of the island extending down from the summit of the <_ 12. Same. Shell with cut-away section to show configuration of columella. Paratype. Height, 22.0 mm. CASG Type Collection, no. 13739. Ficure 13. Same. Xanthic color- phase, adult, from type lot. Paratype. Height, 21.2 mm. CASG Type Collection, no. 13740. Ficure 14, Same. Xanthic shell, adult, with over-all greenish-brown color tone, from lot no. 11. Hypotype. Height, 23.1 mm. CASG Type Collection, no. 13741. Apertural view. Ficure 15. Same. Back view. Ficure 16. Same. Adult shell with color pattern reversed, from lot no. 12. Hypotype. Height, 18.8 mm. CASG Type Collection, no. 13742. Ficure 17. Same, Adult shell with normal color pattern reversed, from lot no. 13. Hypotype. Height, 26.7 mm. CASG Type Collection, no. 13743. Ficure 18. Naestotus duncanus (Dall, 1893). Adult dead shell from Isla Pinzén (Duncan Island). Hypotype. Height, 18.7 mm, CASG Type Collection, no. 13744. 16 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. main volcanic crater, Mt. Crocker, generally in the Scalesia Zone, at elevations above 500 m. (1600 ft.). It is a ground snail, frequenting moist habitats under lava rocks, Scalesia dead-falls, or equivalent cover. Naesiotus cavagnaroi is remarkably consistent in size and general configura- tion. Measurements of 15 adult shells from the type locality average 21.9 mm. in length and 13.4 mm. in maximum diameter, with about 6’ whorls. The cor- responding range measurements for this series are 21.0—23.5 and 12.4—14.6, with the number of whorls ranging from 5*4 to 6%, the latter number being the most frequent. Young shells are prominently keeled at the base of the developing whorls. The occurrence of xanthic shells along with normally colored ones is of particular interest. (See figs. 13-15.) No similar situation has been observed in any other Galapagos species of Naesiotus. According to André De Roy, who collected the type series, the ratio of xanthic to normally colored shells is about one to five, which is confirmed by the count of specimens sent by him for identifi- cation and study. There is no difference in the color of the animals and all xanthic shells seen fall within the range of measurements given above. Lot nos. 10 and 11 in the preceding list are all the bandless xanthic color form. They were collected by Templeton Crocker during the Academy’s 1932 Galapagos Expedition at the time he and his party pioneered the first ascent of the island’s main crater, later called Mt. Crocker in his honor. The series of shells he col- lected at the crater rim are of the same yellowish color as xanthic shells from the type locality of the species. The second lot, collected at a lower elevation on the way to the top, are darker in color with a greenish brown cast. Both of these lots evidently represent pure xanthic populations of the species; the exact localities where they were found are presently unknown and a search for them should be made. . The three specimens in lot no. 12 in the list of specimens examined have a color pattern that is the reverse of the normal one, being a light beige-brown overall with a dark brown encircling band. The two adult shells have red-brown nuclear whorls but in size and sculpture they are normal for the species. (See fig. 16.) Until recently this reverse color pattern was thought to be of rare occurrence, but an apparently pure colony of such shells was collected by the De Roys in the highland area in May 1970 (lot no. 13, preceding). Of the 33 shells sent in for study, 21 have the same reverse color pattern just mentioned except that the encircling brown band is bordered below by a whitish band of somewhat variable width (see fig. 17); 7 have weak or indistinct banding and thus ap- proach the xanthic form; and 5 are similar to the xanthic shells occurring in the type lot. Compared with the type series, shells in this lot average slightly higher, several being a little longer spired with deeper sutures. The largest shell mea- sures 26.7 X 14.5 mm. in height and maximum diameter. Mme. De Roy’s com- VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 17 ments on this lot (personal communication, 1 August 1970) are of such interest that they are repeated, as follows: Surprisingly, we found them living in an area where André had collected before, only that he searched a couple of hundred yards on either side. In fact, the area where this colony of snails is living is restricted (?) to a brushy patch surrounded by grass- covered hills and may be no more than 3 or 4 acres. Snails there are quite abundant but this could change before long, as barbed-wire fence already divides the place in two lots and the land will be converted into pastures. Although the reverse color phase may possibly turn out to be a different species when the relation between it and the color phase considered to be the normal one is better known, present indications are that there is only a single species in- volved, in spite of the fact that there are four color forms. Two described Galapagos species have some superficial resemblance to Naesiotus cavagnaroi. One is N. duncanus (Dall, 1893) from Duncan Island (Isla Pinzon), shown in figure 18, which has the same general shape and number of whorls although it is considerably smaller in size, has a less well developed parietal denticle, and lacks the columellar thickening or flange present on J. cavagnaroi. Unfortunately, NV. duncanus has not been reported as having been collected alive and may, in fact, be extinct. The other similar species is the rare N. jervisensis (Dall, 1917), which again is much smaller than NV. cavagnaroi, has a less well developed parietal denticle and columellar flange, but has the same type of fine spiral sculpture: it differs, however, in having the last whorl irregularly corrugated and in being unicolored and unbanded. The species is named for David Q. Cavagnaro, California Academy ento- mologist and a member of the GISP, who collected some of the first specimens and also a number of other land snails of considerable scientific value on Isla Santa Cruz and on several other Galapagos islands he was able to visit. Naesiotus scalesiana A. G. Smith, new species. (Figures 19-25.) DESCRIPTION. Shell of medium size for the genus, elongate to globose-conic, the apex rose-colored with the remaining shell whitish or tinged with beige, yellowish white, or very light red-brown. Whorls 5% to 6%, almost flat-sided, the sutures only moderately impressed. Aperture subovate, usually without denticles; the peristome thin and unflared, hardly reflected at the base of the columella, leaving open a small but permeable umbilicus, not joined across the parietal wall of the body whorl with any appreciably thickened wash of callus. Nuclear whorls nearly 3, erect, sculptured with extremely fine, closely spaced, nearly straight transverse riblets or lirae, extending all the way across them. Postnuclear whorls somewhat shining, the first one or two with coarse, irregular lines of growth, later ones with additional, irregulary placed, rather deep indenta- 18 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 24 Ficure 19. Naesiotus scalesiana A. G. Smith, new species. Holotype. Height, 14.1 mm. CASG Type Collection, no. 13745. Apertural view. Ficure 20. Same. Back view. FIGURE 21. Same. Shell with cut-away section to show configuration of columella. Paratype. Height, 13.6 mm. CASG Type Collection, no. 13746. Ficure 22. Same. Enlarged view of portion VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 19 tions and elongate impressions, which are most prominent on the body whorl. Animals in alcohol generally yellowish white, occasionally tinged with light gray on the dorsal surface, the foot usually being of a slightly lighter color tone. Rarely, an animal is quite dark-colored. There appears to be no correlation between the color of the animal and the color of the shell. Hototype. An adult shell measuring: height, 14.1; maximum diameter, 7.5; height of aperture, 6.4; width of aperture, 3.7 mm. Number of whorls, 6%. Apical angle, about 45°. CASG Type Collection, no. 13745. PaRATYPES. Ninety-two additional specimens preserved dry, deposited in various museums maintaining type collections including the CASG Type Collec- tion, and 4 preserved in alcohol in the CASIZ Type Series, no. 470. Type LocaLity. Isla Santa Cruz (Indefatigable Island), Galapagos Islands, in the Scalesia Zone, Horneman Farm area, at an elevation of about 244 m. (800 ft.), on Scalesia trees and tall bushes, collected by A. G. Smith, 19 February 1964. RANGE AND ECOLOGY. A total of 36 separate lots, in addition to the type lot, have been collected and are available for study. All of these have contributed to the present consideration of the species. They represent over 500 specimens, mostly preserved dry, although a representative series of animals were drowned and preserved in alcohol in an expanded condition. Naestotus scalesiana seems to be the most prevalent species throughout the Scalesia Zone on Isla Cruz. Its occurrence most often on the trunks and branches of Scalesia trees makes the name given to it an appropriate one. This habitat is shared with NV. lycodus (Dall, 1917), also pink-tipped but with a much more wrinkled shell, which is almost as common at the type locality although it does not climb as high up as NV. scalesiana. The latter species has been collected also, though sparingly, at the upper edge of the Transition Zone and in the lower part of the Miconia Zone but it does not seem to thrive outside the areas occupied by the forests of Scalesia pedunculata wherever they occur under conditions of heavier rainfall and hence of moister conditions than exist in the lower or the higher, somewhat dryer life zones. Along the “old” or original trail from Academy Bay village to the highland < of holotype to show sculptural detail. Ficure 23. Same. Group of five adult shells from Scalesia Zone below Bella Vista village, Isla Santa Cruz. Allyn G. Smith and Ira L. Wiggins, collectors, 19 February 1964. Hypotypes. Height (left-hand shell), 13.3 mm. CASG Type Collection, nos. 13747, 13747a, 13747b, 13747c, 13747d. From CASIZ Color Slide, no. 955. Ficure 24. Same. Group of five adult shells from type lot. Holotype at left (height, 14.1 mm.) ; the rest paratypes, CASG Type Collection, nos. 13748, 13748a, 13748b, 13748c. From CASIZ Color Slide no. 1937. 20 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. area on Isla Santa Cruz, Dr. Ira L. Wiggins and I first encountered NV. scalesiana in 1964 in the Transition Zone at an elevation of about 120 m. (400 ft.) (CASG no. 40083, 2 adult specimens). It became quite common at the lower end of the Scalesia Zone on young Scalesia trees, along with N. lycodus, and continued prevalent in the vicinity of the village of Bella Vista, the areas of the Horneman and Kastdalen Farms, and for a short distance into the Miconia Zone (CASG no. 40070, 3 adult specimens). Elsewhere in the Scalesia Zone M. André De Roy and others collected it toward the eastern side of the island in the Table Mountain area (CASG no. 40232), on the northern side in the general vicinity of Cavagnaro Crater (CASG no. 40221), around the village of Santa Rosa, and to the south of Santa Rosa near the end of the trail into the Tortoise Reserve at El Chato. The Scalesia snail lives in more of an arboreal habitat than any other Gala- pagos land snail with which I am familiar. On one collecting trip, Dr. Wiggins and I decided to sacrifice one large Scalesia branch to see just how far up the snail might go. We found two living adults (CASG no. 40231) at the very tip of the branch, about 35 feet from the ground, among the small sunflower-like Scalesia blossoms in flower at the time. It does not confine itself entirely to Scalesia plants and trees, however, but has been collected living on Darwin- iothamnus, a bush of medium height, and below Santa Rosa village in thickets of thorny shrubs called “mora” by the native Ecuadoreans in company with another new species of Galapagos land snail, Naesiotus deroyi, described earlier in this report. Unfortunately, nothing is known at present of the breeding habits of the species; its egg clusters were not found in 1964 and have not been reported subsequently. INDIVIDUAL VARIATION. The type lot of VV. scalesiana was selected for general consistency in size and color of the shells, and also for the accessibility of the type locality at the Horneman Farm, making it relatively easy for topotypical material to be collected. Color in the type lot ranges from almost pure white to an overall suffusion of beige or light brown; only one adult shell in the series has darker postnuclear whorls, with a body whorl having a distinctly brown and white banded pattern. (See fig. 24.) The rose-colored tip is consistent within the type lot and for most of the other shells at hand. Variation in size within the type lot is shown in the following table of mea- surements of 25 adult paratypes, selected more or less at random: Item Height (mm.) Max. Diam. (mm.) Apical Angle (°) No. Whorls Tallest shell i522 7.6 49 614 Shortest shell 115 6.2 51 5% Avg. of 25 1333) Til 50 5%, VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 21 Ficure 25. Naesiotus scalesiana. Shell with living animal. Collected by Robert L. Usinger and Earl G. Linsley, Bella Vista village, 4 February 1964. Author’s field photo taken at Academy Bay. From CASIZ Color Slide, no. 440. Shells from other localities on the island have measurements that are consistent with the preceding figures. A few adult shells from the vicinity of Santa Rosa have slightly more tumid whorls and lack the usual rose-colored apex. Most shells have simple apertures with no parietal or columellar denticles or denticular swellings; this is true of all specimens in the type lot and of others collected in the Horneman Farm area. A few shells from the general vicinity of Santa Rosa do have small to subobsolete parietal denticles, and in some but not all of these there are perceptible swellings on the columella where a columellar tooth would normally occur, were one present. Such denticular processes are not a character of this species although they are normal in the apertures of other species of Naesiotus from Isla Santa Cruz and other Galapagos Islands. Whether the rare occurrence of apertural denticulation represents an evolutionary trend toward development of prominent teeth in the aperture, or away from this, is a question that cannot be answered from the present sparse knowledge of the biology of the species and its probable ancestral relationships. A well rounded body whorl is characteristic of NV. scalesiana, although occa- 22 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. sionally an adult shell shows a barely perceptible angulation at its base near the periphery. As might be expected, this feature is seen more often on juvenile and subadult shells. The indented sculpture on the body whorl is peculiar; it cannot be said to be malleate in terms of small, rounded hammer marks. Rather, it consists of pits in combination with elongated, sometimes anastomosing furrows that have the appearance of impressions made by a blunt point or an elongated, blunt edge. This type of sculpture varies somewhat in intensity between populations from which specimens have been collected. (See figs. 22, 25.) On the whole, however, the shells of NV. scalesiana exhibit characters that are quite consistent throughout its range. CoMMENTS. It seems unusual that such an abundantly represented species of Galapagos land snail has escaped notice for so long a time. W. H. Dall did not recognize it as having been collected during earlier Galapagos expeditions; apparently neither Baur nor Wolf saw it. Reibisch (1892) reported nothing like it, his only large snail from Indefatigable Island (Isla Santa Cruz) being Naesiotus wolfi, an entirely different species which he described as new. It was not found by Snodgrass and Heller during the Hopkins-Stanford Expedition of 1897—98; nor was it collected by W. H. Ochsner during the California Academy’s 1905-06 Expedition, which spent quite a little time at Academy Bay. The California Academy’s 1932 Expedition did not find it even though in the first ascent to the rim of the main crater the party led by Templeton Crocker traversed the Scalesia Zone on the way to the top as well as on the way back. The United States National Museum has no specimens like it in its large representative col- lection of Galapagos land snails; however, it does possess a single adult specimen from an unidentified island, collected or obtained by Hugh Cuming, under the name Bulimulus calvus (Sowerby, 1833) (USNM no. 104,864) that is quite close to Naesiotus scalesiana. The first authentic collecting record known to me is a single shell found by Dr. Robert I. Bowman, “5 miles North of Academy Bay,” March 2, 1953, at an elevation of 775 feet (CASG no. 34649). All shells collected subsequently were taken during the Galapagos International Scientific Project of 1964 and since. Naesiotus scalesiana has no demonstrable relationship with any of the other 15 species of Naestotus from Isla Santa Cruz, including the two previously described as new in this report. I have not seen shells like it from any of the other islands in the Galapagos Archipelago. Closest relatives would appear to be certain species of Naesiotus from the mainland of South America (Weyrauch, 1956, 1967) but lack of appropriate mainland material at the present writing places any detailed comparison beyond the scope of this discussion. VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 23 ACKNOWLEDGMENTS Appreciation and thanks to André and Jacqueline De Roy of Academy Bay, Isla Santa Cruz, and to several scientists of the Galapagos International Scientific Project of 1964 have already been expressed. In addition, David Q. Cavagnaro, California Academy entomologist with the GISP, also should have special men- tion for his land-snail collecting efforts on top of a full-time entomological pro- gram. I am greatly indebted to the California Academy’s scientific photographer, Mr. Maurice Giles, for his excellent work in providing acceptable snail illustra- tions from actual specimens, and for his black-and-white reproductions of three of the author’s 35 mm. color transparencies. LITERATURE CITED DALL, WILLIAM HEALEY 1893. Preliminary notice of new species of land-snails from the Galapagos Islands, collected by Dr. G. Baur. Nautilus, vol. 7, no. 1, pp. 52-56. May. 1896. Insular landshell faunas, especially as illustrated by the data obtained by Dr. G. Baur in the Galapagos Islands. Proceedings of the Academy of Natural Sciences of Philadelphia, 1896, pp. 395-459, pls. 15-17. 1900. Additions to the insular land-shell faunas of the Pacific Coast, especially of the Galapagos and Cocos islands. Proceedings of the Academy of Natural Sciences of Philadelphia, 1900, pp. 88-105, pl. 8. 1917. Preliminary descriptions of new species of Pulmonata of the Galapagos Islands. Proceedings of the California Academy of Sciences, ser. 4, vol. 2, pt. 1, no. 11, pp. 375-382. San Francisco. December 31. 1920. On the relations of the sectional groups of Bulimulus of the subgenus Naesiotus Albers. Journal of the Washington Academy of Sciences, vol. 10, no. 5, pp. 117-122. Washington, D. C. March 4. DALL, WILLIAM HEALEY, AND WASHINGTON HENRY OCHSNER 1928. Landshells of the Galapagos Islands. Proceedings of the California Academy of Sciences, ser. 4, vol. 17, no. 5, pp. 141-185, pls. 8-9. San Francisco. June 22. Pirspry, Henry AUGUSTUS 1897-98. American Bulimulidae: Bulimulus, Neopetraeus, Oxychona, and South Amer- ican Drymaeus. Manual of Conchology, ser. 2, Pulmonata, vol. 11, pp. 1-339, pls. 1-51. Philadelphia. REIBISCH, PAUL 1892. Die concholiologische Fauna der Galapagos Inseln. Sitzungsberichte und Abhandlungen Naturwissenschaftlichen Gesellschaft Isis in Dresden, 1892 (January—June), pp. 13-32, 1 text fig. (map), pls. 1-2. Dresden. STEARNS, ROBERT EDWARDS CARTER 1893. Report on the mollusk fauna of the Galapagos Islands with descriptions of new species. Scientific Results of Explorations by the U. S. Fish Commission Steamer Albatross, no. 30. Proceedings of the United States National Museum, vol. 15, no. 942, pp. 353-450, pls. 50-52. Washington, D. C. August. WEYRAUCH, WOLFGANG 1956. The genus Naesiotus, with descriptions of new species and notes on other Peruvian CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER. 24 Bulimulidae. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 108, pp. 1-17, pl. 1. June 22. 1967a. Treinta y ocho nuevos gastropodos terrestres de Peru. Acta Zoologica Lilloana, tomo 21, pp. 343-455, pls. 1-9. Tucuman, Argentina. 1967b. Descripciones y notas sobre gastropodos terrestres de Venezuela, Columbia, Ecuador, Brasil y Peru. Acta Zoologica Lilloana, tomo 21, pp. 457-499, pls. 1-4. Tucuman, Argentina. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Volume XXXIX, No. 3, pp. 25-46; 39 figs. April 6, 1972 PLIOCENE FOSSILS FROM BALTRA (SOUTH SEYMOUR) ISLAND, GALAPAGOS ISLANDS By Leo G. Hertlein California Academy of Sciences, San Francisco, California 94118 INTRODUCTION The present paper deals chiefly with invertebrate fossils which I collected while a member of the first Expedition of the Velero I// to the Galapagos Islands in 1931-1932. A general account of the itinerary of this expedition appeared in a report by Fraser (1943, pp. 50, 260, 262, 272-273). A report dealing with fossils of Pleistocene age collected on this expedition was published by Hertlein and Strong (1939) and in a later paper the same authors (1955) reported on the Recent shells. Fossils believed to be of Pliocene age, chiefly mollusks, were collected during January, 1932. The pressure of other duties delayed completion of the present paper. In addition to the collection which I assembled, Joseph R. Slevin collected a few fossils of Pliocene age in 1927 when he visited the island on Captain G. Allan Hancock’s Oaxaca. A few other specimens received through the courtesy of Dr. A. Myra Keen of Stanford University, are included in the present paper. The author is grateful for having had the opportunity to accompany the expedition offered by the late G. Allan Hancock, owner of the Velero J/I. The author acknowledges aid received from two other individuals, now deceased: A. M. Strong, who identified the micro-gastropods and E. H. Quayle who con- tributed information concerning the corals. Mr. Barry Roth, Department of Geology, California Academy of Sciences, furnished information concerning the identification of the species in the family Marginellidae, contributed helpful comments concerning certain other species, and also aided in 1 the preparation Marine Biological Labor..tory LIBRAFRY [25] APR 14 1972 Woods Hole, Viass. : 26 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. the plates. The identifications of the corals listed in this paper were furnished by Dr. J. W. Durham, Department of Paleontology, University of California, Berkeley. The author is grateful to Carmen Angermeyer and to Jacqueline De Roy, residents of Academy Bay, Santa Cruz Island, Galapagos Islands, who presented many specimens of Recent shells from those islands to the California Academy of Sciences. These specimens have been very useful for comparison with fossil forms from that archipelago. Photographs used to illustrate the fossils were prepared by Mr. Maurice Giles, Photographer, California Academy of Sciences. Margaret M. Hanna kindly retouched some of the photographs. The manuscript was typed by Enid Cook. One new subspecies, Diplodonta subquadrata baltrana, is described in the present paper. GENERAL REMARKS Darwin visited the Galapagos Islands during the voyage of the Beagle in 1835. He mentioned (1844, p. 115; see ed. 3, 1896, p. 130) sea shells belonging to modern genera, apparently on Chatham Island, ‘embedded several hundred feet above the sea, in the tuff of two craters, distant from each other.” Later Wolf (1895, pp. 250, 254, 265) reported the occurrence of fossil shells in the islands in palagonite tuff at a height of 100 meters above sea level. W. H. Ochsner, during an expedition of the California Academy of Sciences to the Galapagos Islands on the schooner Academy, spent most of one year in the islands and discovered fossils on Isabela (Albemarle), Santa Cruz (Inde- fatigable), and Baltra (South Seymour) islands. On this expedition Ochsner collected the fossils of Pliocene age on Baltra (South Seymour) Island mentioned by Dall (1924) and later reported upon by Dall and Ochsner (1927, see espe- cially pp. 94-99). They listed 28 species (and four others identified only as to genus) from beds believed to be of Pliocene age, 24 species from the “upper zone” and 4 in the “lower zone”. Chesterman (1963, p. 344) reported on rocks of the Galapagos Islands and mentioned fossiliferous limestone and “fossiliferous tuffaceous sandstone” on Baltra (South Seymour), collected by Ochsner. He also described a specimen of andesite from the south end of the island. Baltra Island is composed chiefly of lava and other pyroclastic material with very minor amounts of intercalated limestone and fossiliferous tuffaceous beds. This island experienced faulting in comparatively recent time, the lines of fracture trending approximately east-northeast. This faulting resulted in alter- nating raised and depressed blocks. A depressed block separates Baltra (South Seymour) from Santa Cruz (Indefatigable) Island and a similar depressed block separates Baltra from Seymour (North Seymour) Island. Lewis (1956, p 290) remarked on the relatively recent faulting as did Williams (1966, pp. 55-70) Vot. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 27 who discussed the general geology of the Galapagos Islands. More recently, McBirney and Williams (1969, pp. 17-20, fig. 7) discussed the geology and petrology of Baltra, as well as that of other islands in the archipelago. The fossil-bearing bed on the south end of the bay on the west side of the island is about 3 meters (10 feet) thick and dips about 5° or 6° south. Large angular blocks of lava in the lower portion of the bed offer evidence of deposition very near the shore. The majority of the species in the present list were taken from the tuffaceous ashy bed. A number of small specimens were obtained by passing some of the white ashy material through a sieve. COLLECTING STATIONS Locality 1305 (CAS), cliff on southwest side of Baltra (South Seymour) Island, Galapagos Islands. Joseph R. Slevin collector, December 16, 1927. Pliocene. Locality 27249 (CAS), white and yellowish tuffaceous strata interbedded with lava on the south side of the bay on the west side of Baltra (South Seymour) Island, Galapagos Islands. Leo G. Hertlein collector, January 16-18, 1932. Pliocene. Locality 27251 (CAS), strata about 1.5 meters (5 feet) thick on the northern side of the bay, about the middle of the west side of the island, near the top of the plateau, Baltra (South Seymour) Island, Galapagos Islands. Leo G. Hertlein collector, January 14-15, 1932. Pliocene. Locality 31838 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Fossils received by Dr. A. Myra Keen who presented them to the California Academy of Sciences, March 22, 1943. LIST OF FOSSILS FROM BALTRA (SOUTH SEYMOUR) ISLAND* An asterisk * indicates that the species is extinct. COELENTERATA Pavona gigantea Verrill Psammocora (Stephanaria), species indeterminate ECHINOIDEA Encope micropora galapagana H. L. Clark, locs. 13051; 27249 (CAS) Eucidaris thouarsii Valenciennes, loc. 1305 (CAS) PELECYPODA Anatina (Raéta) undulata Gould, loc. 31838 (CAS) * Anodontia spherica Dall and Ochsner, locs. 1305; 27251 (CAS) Anomia peruviana d’Orbigny, loc. 27249 (CAS) 1 Reported from this locality as Encope micropora L. Agassiz by Grant and Hertlein (Publ. Univ. Calif. Los Angeles Math. Phys. Sci., vol. 2, p. 98, 1938). 28 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Arca pacifica Sowerby, loc. 1305 (CAS) Arca (Arcopsis) solida Broderip and Sowerby, loc. 27249 (CAS) Atrina cf. A. tuberculosa Sowerby, loc. 27249 (CAS) Barbatia reeveana d’Orbigny, loc. 27249 (CAS) Cardium elenense Sowerby, loc. 27249 (CAS) Chama species, loc. 1305 (CAS) Chione undatella Sowerby, loc. 27249; (cf.) 31838 (CAS) Chione species, loc. 1305 (CAS) Chlamys (Argopecten) circularis Sowerby, loc. 27249 (CAS) Chlamys (Nodipecten) magnifica Sowerby, loc. 27249 (CAS) Ctena galapagana Dall, locs. 1305; 27249 (CAS) Cuminga ctf. C. lamellosa Sowerby, loc. 27249 (CAS) Diplodonta subquadrata baltrana Hertlein, new subspecies, loc. 27249 (CAS) Florimetis cognata Pilsbry and Vanatta, loc. 27249 (CAS) Glycymeris maculata Broderip, loc. 27249 (CAS) Megapitaria cf. M. aurantiaca Sowerby, loc. 27249 (CAS) Megapitaria squalida Sowerby, loc. 31838 (CAS) Modiolus capax Conrad, loc. 27249 (CAS) Nioche cf. N. zorritensis Olsson, loc. 27249 (CAS) Ostrea megodon Hanley, loc. 27249 (CAS) Ostrea palmula Carpenter, loc. 1305 (CAS) Ostrea species, loc. 27249 (CAS) * Pecten (Pecten) slevini Dall and Ochsner, loc. 27249 (CAS) Protothaca (Tropithaca) grata Say, locs. 27249; 31838 (CAS) * Protothaca (Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner, loc. 27249 (CAS) Tagelus cf. T. dombeii Lamarck, loc. 27251 (CAS) (cast) Tagelus species, locs. 27249; 31838 (CAS) GASTROPODA Acanthina grandis Gray, loc. 1305 (CAS) Acteocina infrequens C. B. Adams, loc. 27249 (CAS) Alaba supralirata Carpenter, loc. 27249 (CAS) Alvania cf. A. galapagensis Bartsch, loc. 27249(CAS) Alvania ct. A. halia Bartsch, loc. 27249 (CAS) Alvania cf. A. hoodensis Bartsch, loc. 27249 (CAS) Alvania cf. A. nemo Bartsch, loc. 27249 (CAS) Amphithalmus trosti Strong and Hertlein, loc. 27249 (CAS) Anachis cf. A. tabogaensis Bartsch, loc. 27249 (CAS) Balcis cf. B. berryi Bartsch, loc. 27249 (CAS) Bulla punctulata A. Adams in Sowerby, locs. 27249; 31838 (CAS) Cancellaria cf. C. ovata Sowerby, loc. 27249 (CAS) Cantharus janellii Kiener, locs. 1305; 27249 (CAS) * Cantharus cf. C. scissus Olsson, loc. 27249 (CAS) Cerithiopsis curtata Bartsch, loc. 27249 (CAS) Cerithiopsis cf. C. galapagensis Bartsch, loc. 27249 (CAS) Cerithium adustum Kiener, locs. 1305; 27249 (CAS) Cheila equestris Linnaeus, loc. 27249 (CAS) Conus fergusoni Sowerby, loc. 27249 (CAS) Conus cf. C. lucidus Wood, loc. 27249 (CAS) VoL. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 29 Cylichna ct. C. defuncta Baker and Hanna, loc. 27249 (CAS) Cymatium lineatum Broderip, loc. 1305 (CAS) Cypraea nigropunctata Gray, locs. 1305; 27249 (CAS) Cytharella camarina Dall, loc. 27249 (CAS) Diodora alta C. B. Adams, loc. 27249 (CAS) Engina pyrostoma Sowerby, loc. 27249 (CAS) Erato (Hespererato) marginata galapagensis Schilder, loc. 27249 (CAS) Eulimostraca cf. E. galapagensis Bartsch, loc. 27249 (CAS) Fissurella cf. F. macrotrema Sowerby, loc. 27249 (CAS) Fissurella virescens Sowerby, loc. 27249 (CAS) Fusinus dupetitthouarsii Valenciennes, loc. 27249 (CAS) Gastrocopta munita Reibisch, loc. 27249 (CAS) [a land snail] Granula cf. G. minor C. B. Adams, loc. 27249 (CAS) Granula cf. G. polita Carpenter, loc. 27249 (CAS) Granula species, loc. 27249 (CAS) Granulina margaritula Carpenter, loc. 27249 (CAS) Hipponix pilosus Deshayes, locs. 1305; 27249 (CAS) Hipponix pilosus Deshayes, loc. 1305; 27249 (CAS) Tselica cf. I. kochi Strong and Hertlein, loc. 27249 (CAS) Malea ringens Swainson, loc. 1305 (CAS) * Mangelia cf. M. hancocki Hertlein and Strong, loc. 27249 (CAS) Mitra gausapata Reeve, loc. 1305 (CAS) Mitra lens Mawe, loc. 1305 (CAS) Modulus cerodes A. Adams, loc. 27249 (CAS) Nassarius nodicinctus A. Adams, loc. 27249 (CAS) * Nerita oligopleura Dall and Ochsner, loc. 27249 (CAS) Odostomia (Menestho) aequisculpta Carpenter, loc. 27249 (CAS) Odostomia (Miralda) galapagensis Dall and Bartsch, loc. 27249 (CAS) Odostomia (Chrysallida) rinella Dall and Bartsch, loc. 27249 (CAS) Oliva species, locs. 27249; 31838 (CAS) Pedipes angulatus C. B. Adams, loc. 27249 (CAS) Persicula imbricata Hinds, loc. 27249 (CAS) Persicula cf. P. phrygia Dall, loc. 27249 (CAS) Persicula species, loc. 27249 (CAS) Polinices dubius Récluz, loc. 27249 (CAS) Polinices uber Valenciennes, loc. 27249 (CAS) Pyramidella (Triptychus) ct. P. (T.) olssoni Bartsch, loc. 27249 (CAS) Pyramidella (Voluspa) species, loc. 27249 (CAS) Pyrene castanea Sowerby, locs. 1305; 31838 (CAS) Pyrene fuscata Sowerby, loc. 27249 (CAS) Pyrene haemastoma Sowerby, loc. 27249 (CAS) Rissoina cf. R. firmata C. B. Adams, loc. 27249 (CAS) Rissoina signae Bartsch, loc. 27249 (CAS) Strombina gibberula Sowerby, loc. 27249 (CAS) Tectarius galapagensis Stearns, loc. 27249 (CAS) * Tegula forbesi Dall and Ochsner, loc. 27249 (CAS) Tegula snodgrassi Pilsbry and Vanatta, locs. 1305; 27249 (CAS) Triphora cf. T. galapagensis Bartsch, loc. 27249 (CAS) Triphora cf. T. panamensis Bartsch, loc. 27249 (CAS) 30 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Trivia pacifica Gray, loc. 27249 (CAS) Trivia radians Lamarck, loc. 27249 (CAS) Turbo agonistes Dall and Ochsner, loc. 27249 (CAS) * Turbo vermiculosus Dall and Ochsner, locs. 27249; 31838 (CAS) Turbonilla (Chemnitzia) houseri Dall and Bartsch, loc. 27249 (CAS) Turritella broderipiana marmorata Kiener, loc. 27249 (CAS) Vermicularia eburnea Reeve, loc. 27249 (CAS) Volvarina taeniolata Morch, loc. 27249 (CAS) REMARKS ON THE AGE AND RELATIONSHIPS OF THE FAUNA This list contains 102 identified species, 1 coral, 2 echinoids, 26 pelecypods, and 73 gastropods. Of these, 7, and probably 8, are extinct. Twenty-eight species are only provisionally identified but are compared with known species. In addition to the 102 species, 8 forms are identified only as to genus. The percentage of extinct species in the present faunal assemblage is approxi- mately 7.8 percent. However, a greater number of extinct species evidently occur at the present locality as Dall and Ochsner described additional extinct species from apparently the same locality. They also described a number of extinct species from what they believed to be approximately correlative strata on Santa Cruz (Indefatigable) Island. Dall and Ochsner considered the age of the fauna which they reported from those two islands to be of probable Pliocene age. Five of the eight extinct species in the present faunal list have not been reported from fossil assemblages of Pleistocene age from either Isabela (Albemarle) or from San Salvador (James) Island. Furthermore, the occurrence of the fossils in ashy beds (which in some places are decidedly indurated), interbedded with volcanic flows, lends support to the viewpoint that these beds are of Pliocene rather than of Pleistocene age. McBirney and Williams (1969, p. 19) mentioned two lavas on Baltra Island reported to be 1.47 million years old. Another factor contributing to the belief that the present fauna is of Plio- cene age is the fact that Anodontia spherica Dall and Ochsner (reported from strata of Pliocene age in the Galapagos Islands) was reported by Pilsbry and Olsson (1941, p. 57) to occur abundantly and to be a characteristic species of their “Zone H” in strata referred to Pliocene age at Punta Blanca, Ecuador. Ten species in the present faunal list were included among those reported from that locality. Most of the Recent species in this assemblage are known living in Galapagos waters. Those not known from there are chiefly elements of the Panamic fauna, but further collecting may reveal their presence in the Galapagos Archipelago. A consideration of the habitat of the Recent species in this fauna leads one to infer that the fossil forms lived under conditions similar to those now prevailing in the Galapagos Islands, namely, warm, shallow water. Vot. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 31 An interesting occurrence among the fossil forms is that of a land snail, Gastrocopta munita Reibisch. This species now lives on Baltra (South Seymour) Island (Hertlein, 1932b, p. 69) as well as on most of the other larger islands in the Archipelago. SYSTEMATIC PALEONTOLOGY PELECYPODA Chlamys (Nodipecten) magnifica Sowerby. (Figures 5, 15, 25.) Pecten magnificus SOWERBY, variety a, Proc. Zool. Soc. London for 1835, p. 109, issued October 9, 1835. “Hab ad Insulas Gallapagos.” “A single specimen of var. a was found in a coral sand at a depth of six fathoms.” SowerBy, Thes. Conch., vol. 1, p. 65, pl. 15 (Pecten, pl. 5), fig. 114, 1842. “East Columbia.” [lapsus calami for West “Columbia.” Locality doubtful.] Rrrve, Conch. Icon., vol. 8, Pecten, species 9, pl. 2, fig. 9, 1852. “Tsle of Plata, West Columbia (in coral sand at a depth of from six to seventeen fathoms); Cuming.” [Locality doubtful.] Kopert, Syst. Conch—Cab. von Martini und Chemnitz, Spondylus und Pecten, Bd. 7, Abt. 2, p. 164, pl. 46, fig. 1, 1888. (Descrip- tion and illustration from Reeve.) Pecten (Lyropecten) magnificus Sowerby, GRANT and GALE, Mem. San Diego Soc. Nat. Hist., vol. 1, p. 182, pl. 9, fig. 1; pl. 10, fig. 6, 1931. Galapagos Islands. Nodipecten magnificus Sowerby, Grau, Allan Hancock Pac. Exped., vol. 23, p. 132, pl. 44, 1959. “Galapagos Islands. (Ecuador doubtful.)” Lyropecten (Nodipecten) magnificus Sowerby, Otsson, Mollusks of the Tropical Eastern Pacific (Paleo. Res. Inst.: Ithaca, New York), p. 161, pl. 22, fig. 1, 1961. (Illustration from Reeve, 1852). Two left valves, one nearly complete, the other only one half of a valve were collected at locality 27249 (CAS). The larger specimen is 126 mm. long and 122.3 mm. high. There are 13 or 14 radial ribs. The entire valve is covered with radial striae of which about 5 or 6 occur in each interspace. The ribs are decidedly nodose. The nodes are hollow and develop in the areas of “ledges” (Moore, 1934, p. 216) between concentric constrictions of the valve. The nodosity on the east American species, Chlamys (Nodipecten) nodosa Linnaeus, increases from north to south in its range into warmer waters. A small left valve of C. magnifica 49 mm. high, was reported by Hertlein and Strong (1939, p. 369) from a raised beach on San Salvador (James) Island. Two Recent right valves in the collections of the California Academy of Sciences, the larger one 127 mm. high, were collected by Ochsner on the beach at Baltra (South Seymour) Island. These are devoid of nodes. The color is bright red. One right valve 59.8 mm. long and 60 mm. high, in the Academy’s collection, received from Jacqueline De Roy, was dredged in 9 to 18 meters (5 to 10 fathoms) off Santa Fé (Barrington) Island. 32 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Anodontia spherica Dall and Ochsner. Hypotype, left valve, no. 13653 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 802 (CAS), about 1% miles northeast of the settlement of Vilamil, Isabela (Albemarle) Island, Galapagos Islands, elevated beach deposit, about 12 meters (40 feet) above sea level. Pleistocene. Length 65 mm. Ficure 2. Anodontia spherica Dall and Ochsner. Hypotype, right valve, no. 13654 (Calif Acad. Sci. Dept. Geol. Type Coll.), from locality 803 (CAS), from virtually the same locality as the Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 33 Anodontia spherica Dall and Ochsner. (Figures 1, 2, 6, 7.) Lucina spherica DALL AND OcusNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p. 121, pl. 3, fig. 8; pl. 4, figs. 2, 7, June 22, 1928; “from upper horizon (zone D) on east shore of Indefatigable Island, Galapages Greuv. Probably Pliocene.” Loripinus (Pegophysema) spherica Dall and Ochsner, Pitspry AND Otsson, Proc. Acad. Nat. Sci. Philadelphia, vol. 93, p. 57, 1941. Pliocene of Punta Blanca, Ecuador. Anodontia (Lissosphaira) spherica Dall and Ochsner, Orsson, Mollusks of the tropical eastern Pacific (Paleo. Research Inst.: Ithaca, New York), p. 222, pl. 30, fig. 2, 1961. (Figured specimen from Cabo Blanca, Ecuador, Pliocene. Reported as Recent from “Lower California to Ecuador. Columbia: Isla del Gallo.”) Also earlier records. Specimens assigned to this species in the present collection [locs. 1305 and 27251 (CAS) | are casts. Olsson reported this species living from Baja California to Ecuador but it has not been reported from the Panamic Province by other authors. I have not seen Recent specimens. The deeply depressed posterior dorsal area on Anodontia spherica easily serves to separate it from A. edentuloides from the Gulf of California. The presence of well differentiated, deeply impressed, dorsal areas was the basis for the proposal of the subgenus Lissophazra Olsson. Large specimens of A. spherica are 74 mm. long. Berry (1968, p. 71) men- tioned specimens of 4. edentuloides which are 75 mm. long. Marks (1951, pp. 67-70), discussed the shell characters of Anodontia and Pegophysema, and some of the species referred to those groups. Apparently, the species cited by Marks (p. 69) as “C. densata” Dall and Ochsner is referable to A. spherica. A large globose species, 82 mm. long, Anodota sphericula Basedow, was reported by Ludbrook (1959, p. 227, pl. 3, figs. 1, 2, 3; pl. 5, figs. 1, 4) from strata of Pliocene age in Australia. The posterior dorsal portion of the shell of that species lacks a depressed area: it is a typical Anodontia believed to be related to A. philippiana Reeve in the western Pacific. Diplodonta subquadrata baltrana Hertlein, new subspecies. (Figures 8 and 11.) DescripTIon. Shell subquadrate, somewhat anteriorly attenuated, thin, beaks very low, anterior dorsal margin only slightly sloping; sculptured only co specimen shown in figure 1. Length 58 mm. Ficure 3. Trivia radians Lamarck. Hypotype no. 13668 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Length 18 mm. FicureE 4. Trivia radians Lamarck. Ventral view of the specimen shown in figure 3. FicurE 5. Chlamys (Nodipecten) magnifica Sowerby. Hypotype, right valve, no. 13652 (Calif. Acad. Sci. Dept. Geol. Type Coll. ), from locality 23167 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Recent. W. H. Ochsner Collector. Height 85 mm. 34 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. 13 Ficure 6. Anodontia spherica Dall and Ochsner. Hypotype, right valve, no 13655 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 803 (CAS), about 1% miles northeast of the settlement of Vilamil, Isabela (Albemarle) Island, Galapagos Islands, elevated beach deposit, about 12 meters (40 feet) above sea level. Pleistocene. Length 54.5 mm. FIGURE 7. Anodontia spherica Dall and Ochsner. View of the interior of the specimen shown in figure Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 35 with fine concentric lines of growth. Dimensions: length 22.6 mm., height 20 mm., convexity (both valves) 14.6 mm. Holotype, no. 13656, and paratype, a left valve, no. 13657 (California Acad- emy of Sciences Department of Geology Type Collection), from locality 27249 (CAS), white and yellowish tuffaceous strata interbedded with lava, on the south side of the bay on the west side of Baltra (South Seymour) Island, Gala- pagos Islands. Late Pliocene. L. G. Hertlein, collector. CoMMENTS. Specimens of this new subspecies, in general features, resemble fossil forms of Diplodonta subquadrata Carpenter (see illustrations by Durham, 1950, pl. 19, figures 4, 4a, 1950) from Santa Inez Bay, Baja California, Mexico. The present fossils from Baltra Island differ from Carpenter’s species in the narrower, somewhat attenuated anterior end and in the more nearly horizontal anterior dorsal margin. The general shape of the type specimen of the new subspecies resembles that of the Recent specimen of D. subquadrata from the Galapagos Islands illustrated by Hertlein and Strong (1947, plate 1, figure 11), more than it does that of the fossils illustrated by Durham. The paratype, 21.2 mm. long, is less attenuated anteriorly than the holotype. An imperfectly preserved right valve also was collected at the type locality. Protothaca (Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner. (Figures 38, 39.) Twenty-one single valves of a venerid varying from 11 mm. to 39.4 mm. in length are present in the collection from locality 27249 (CAS), Baltra (South Seymour) Island. These vary greatly in shape and sculpture. < 6. Ficure 8. Diplodonta subquadrata baltrana Hertlein, new subspecies. Holotype no. 13656 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Length 22.5 mm. Ficure 9. Nioche cf. N. zorritensis Olsson. Hypotype, left valve, no. 13658 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Length 39.5 mm. Ficure 10. Nioche cf. N. zorritensis Olsson. View of the interior of the specimen shown in figure 9. Ficure 11. Diplodonta subquadrata baltrana Hertlein, new subspecies. Dorsal view of the specimen shown in figure 8. Ficure 12. Nerita oligopleura Dall and Ochsner. Hypotype no. 13666 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height 14.5 mm. View showing fine threadlets between major concentric cords. Ficure 13. Nerita oligopleura Dall and Ochsner. Hypotype no. 13667 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from the same locality as the specimen illustrated in figure 12. Apertural view. Height 17.5 mm. Ficure 14. Nerita oligopleura Dall and Ochsner. Dorsal view of the specimen shown in figure 13. Ficure 15. Chlamys (Nodipecten) magnifica Sowerby. Hypotype, left valve, no. 13676 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height 128 mm. 36 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 16. Turbo agonistes Dall and Ochsner. Hypotype no. 13671 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 38977 (CAS), Baranco, Punta Nunez, Santa Cruz (Indefatigable) Island, Galapagos Islands. Recent. Jacqueline DeRoy collector. Height 21 mm. Ficure 17. Turbo agonistes Dall and Ochsner. Dorsal view of the specimens shown in figure 16. Figure 18. Turbo agonistes Dall and Ochsner. Hypotype no. 13672 (Calif. Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 37 One small specimen in the present series is quite similar in outline and sculp- ture to the early stage of growth of the type specimen of Chione seymourensts Dall and Ochsner from the same area. The type specimen of Chione seymourensis Dall and Ochsner (1928, p. 123, pl. 3, figs. 1 and 5) was described “from upper horizon, Seymour Island, Gala- pagos Group. Probably Pliocene.” It is a right valve, ovately rectangular in outline (see figs. 36, 37), lacks an escutcheon, and the radial sculpture is some- what reduced in strength as a result of erosion. Dall and Ochsner mentioned a similarity between their species and Chione pertincta Dall, a Recent species living in the Galapagos Islands. Compared with C. pertincta, the type specimen of C. seymourensis is in general, more elongated, thinner, and it has finer radial sculpture. Compared with Protothaca grata Say, the type specimen of C. seymourensis is more elongate in outline, the radial sculpture is coarser, the lunule is narrower, and the median cardinal tooth in the right valve is larger. The variation in shape and sculpture in the present series of specimens is similar to that in a series of Recent specimens of Protothaca grata Say (the type species of the subgenus Tropithaca Olsson), and most of them are here referred to Say’s species. GASTROPODA Tegula forbesi Dall and Ochsner. (Figures 24, 29.) Tegula forbesi DALL AND OcHSNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p. 116, pl. 2, fig. 13, June 22, 1928; “from upper horizon on Seymour Island, Galapagos Group. Probably Pliocene.” < Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height 28.5 mm. Ficure 19. Turbo agonistes Dall and Ochsner. Dorsal view of the specimen shown in figure 18. Ficure 20. Turbo agonistes Dall and Ochsner. View of the exterior of the operculum of the specimen shown in figures 16 and 17. Ficure 21. Turbo agonistes Dall and Ochsner. View of the interior side of the operculum shown in figure 20. Ficure 22. Turbo agonistes Dall and Ochsner. Hypotype no. 13673 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from the same locality as the specimen shown in Figures 18 and 19. Height 33 mm. Dorsal view of an unusually large specimen. Ficure 23. Turbo agonistes Dall and Ochsner. Apertural view of the speci- men shown in figure 22. Ficure 24. Tegula forbesi Dall and Ochsner. Hypotype no. 13660 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Galapagos Islands. Late Pliocene. Height 26 mm. FuicurE 25. Chlamys (Nodipecten) magnifica Sowerby. Hypotype no. 13652 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 23167 (CAS), Baltra (South Seymour) Island, Galapagos Islands. W. H. Ochsner Collector. Recent. Height 85 mm. View of the interior of the specimen shown in figure 5. 38 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Ficure 26. Turritella broderipiana marmorata Kiener. Hypotype no. 13669 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height (incomplete) 53 mm. Ficure 27. Turritella broderipiana marmorata Kiener. Dorsal view of the specimen shown in figure 26. FIGURE 28. Turritella broderipiana marmorata Kiener. Hypotype no. 13670 (Calif. Acad. Sci. Dept. Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 39 A few specimens of this species were collected at locality 27249 (CAS). The largest specimen is about 31 mm. in height, the lest whorl is 48 mm. in maximum width. The concentric sculpture is much finer than that of Tegula aureotincta Forbes from California. This sculpture is more pronounced and regular on the upper surface of the whorls, and on the base much coarser, than that on specimens of T. rugosa A. Adams from the Gulf of California. Turbo agonistes Dall and Ochsner. (Figures 16, 17, 18, 19, 20, 21, 22, 23.) Turbo agonistes DALL AND OCHSNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p. 115, pl. 2, figs. 12, 16, June 22, 1928; “on east shore of Indefatigable Island, Galapagos Group. Probably Pliocene.” A number of fossil specimens of this species were collected at locality 27249 (CAS). The shell of this species is quite different from any other described form. Recently this species was found living in Galapagos waters. Four specimens from off Santa Cruz (Indefatigable) Island collected by Jacqueline De Roy; and one taken by Carmen Angermeyer 1’ miles west of Baltra (South Seymour) Island, were presented to the Academy. Recent shells are attractively colored, purplish on the base of the body whorl and orange or greenish above. < Geol. Type Coll.), from the same locality as the specimen shown in figures 26 and 27. Height (incomplete) 40 mm. Ficure 29. Tegula forbesi Dall and Ochsner. Hypotype no. 13660 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Width 24 mm. Basal view of the specimen shown in figure 24. Ficure 30. Polinices dubius Récluz. Hypotype no. 13662 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height 37.5 mm. Ficure 31. Polinices dubius Récluz. Apertural view of the specimen shown in figure 30. Ficure 32. Polinices dubius Récluz. Hypotype no. 13663 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 804 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height 40.5 mm. Ficure 33. Polinices dubius Récluz. Hypotype no. 13664 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from the same locality as the specimen shown in figure 32. Height 39.4 mm. Apertural view. Ficure 34. Polinices dubius Récluz. Apertural view of the specimen shown in figure 33, showing notch in callus of inner lip. Frcure 35. Turbo vermiculosus Dall and Ochsner. Hypotype no. 13661 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height 34.6 mm. Ficure 36. Chione seymourensis Dall and Ochsner. Holotype, right valve, no. 2970 (Calif. Acad. Sci. Dept. Geol. Type Coll.), “from upper horizon, Seymour Island, Gala- pagos Group Probably Pliocene.” Length 34 mm. Ficure 37. Chione seymourensis Dall and Ochsner. View of the hinge of the specimen shown in figure 36. Ficure 38. Protothaca (Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner. Hypotype, right valve, no. 13659 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Length 21 mm. Ficure 39. Protothaca (Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner. View of the hinge of the specimen shown in figure 38. 40 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Turbo vermiculosus Dall and Ochsner. (Figure 35.) Turbo vermiculosus DALL AND OcusNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p. 115, pl. 2, fig. 15, June 22, 1928; “frem upper horizon, Seymour Island, Galapagos Group. Probably Pliocene.” Three specimens of this species, none perfectly preserved, are present from locality 27249 (CAS), Baltra (South Seymour) Island. The largest specimen (figure 35) is 34.8 mm. high, the maximum diameter 23 mm. This species has an unusually high spire for the genus. The surface micro- sculpture “‘minutely vermiculately granulose and punctate,’ as mentioned by Dall and Ochsner, is quite different from any described species of Turbo from the western Americas. The shape and sculpture of the present species somewhat resemble that of Turbo caboblanquensis Weisbord (1962, p. 84, pl. 6, figs. 4, 5) described from strata of Pliocene age in Venezuela but the shell of that species is umbilicate. Polinices dubius Récluz. (Figures 30, 31, 32, 33, 34.) Natica dubia RéEctuz, Proc. Zool. Soc. London for 1843, p. 209, issued June, 1844 “Hab. Chile? H. Cuming.” Sowersy, Thes. Conch., vol. 5, p. 86, pl. 458 (Natica, pl. 5), fig. 56, 1883. “Chili.” Rrrvr, Conch. Icon., vol. 9, Natica, species 41, pl. 10, figs. 41a, 41b, 1855. “Hab. Chili?” Tryon, Man. Conch., vol. 8, p. 47, pl. 16, fig. 50, 1886. “Chili, Peru.” Polynices dubia Récluz, STEARNS, Proc. U. S. Nat. Mus., vol. 16, no. 942, pp. 401, 446, 1893. “Indefatigable Island.” One fairly well preserved specimen and four rather imperfect ones were collected at locality 27249 (CAS). These specimens agree exactly with the specimens identified by Dall and Ochsner under the name ‘“‘Neverita cf. reclusiana Deshayes” (see figs. 32, 33, 34), from locality 804 (CAS). That locality is approximately equivalent to locality 27249 (CAS). One of the specimens from locality 804 (CAS) retains most of the umbilical pad. The shape of the shell as well as the details of the callus agree closely with illustrations of Polinices dubius, a species reported by Stearns from the Recent fauna of the Galapagos Islands. Ranson (1959, p. 68) reported this species from an elevated beach at Guadalupito, Peru. The general shape of the Galapagos fossils is similar to that of the fossil species described as ‘‘Natica solida” by Sowerby (1846, p. 255; see ed. 3, 1896, p. 612, pl. 3, figs. 40, 41.) from “Navidad, Chile; Santa Cruz,ePataconiae? | Miocene: Herm, 1969, p. 87] but the callus on the Galapagos species is much more extensive and with an indentation on the umbilical margin. Sowerby’s species was renamed Natica darwinti by Hutton (1886a, p. 334; see also 1886b, p. 214) because of an earlier usage of the name WNatica solida by Blainville (1825, p. 251). Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 41 Nerita oligopleura Dall and Ochsner. (Figures 12, 13, 14.) Nerita oligopleura DALL AND OCHSNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p. 114, pl. 2, fig. 11; pl. 6, fig. 15, June 22; “upper horizon, Seymour Island, Galapagos Group. Probably Pliocene.” About 30 specimens of this species were collected at locality 27249 (CAS). The smallest one is 6 mm. high, the largest one 20.5 mm. high and 21 mm. wide. There are four coarse spiral ridges on the shell and, on well preserved speci- mens, two small spiral threads are visible in the interspaces between the ribs. The shell of this species is quite different from any described west American species, as pointed out by Dall and Ochsner. The Galapagos fossil species bears a general resemblance to Nerita asperata Dujardin from the Helvetian, Middle Miocene, of France, illustrated by Cossmann and Peyrot (1917, pl. 7, figures 83, 84). However, close relationship with that species is not postulated. Turritella broderipiana marmorata Kiener. (Figures 26, 27, 28.) Turritella marmorata KIENER, Spéc. Gén. et Icon. Coq. Viv., Famille Turbinacées, Twrritella, p. 23, pl. 8, fig. 1 (two figs.), 1843-1844. “Habite.” Over a dozen specimens of a Turritella, mostly incomplete, were collected at locality 27249 (CAS). These agree closely with Kiener’s illustrations of Turritella marmorata. Kiener’s figures depict a rather slender, elongated shell, the whorls nearly flat-sided, sculptured with fine, concentric threads, cream-colored with narrow wavy longitudinal brownish-violet lines or flecks. Reeve (1849, species 6, plate 2, figures 6a, 6b) placed T. marmorata in the synonymy of 7. broderipiana under which name two figures were shown. One (6a) represents a shell with fairly wide whorls, somewhat wider posteriorly below the suture, resulting in a slightly sinuous outline of the outer lip of the last whorl. The form shown in this illustration is similar to T. gonostoma Valenciennes. The other figure (6b) represents a slender, elongated shell with nearly flat-sided whorls comparable to Kiener’s illustrations of 7. marmorata. Turritella broderipiana d’Orbigny (1840, p. 388), was originally described but not illustrated from “les environs de Payta, ou elle a été péchée sur les fonds de sable.” The type specimen illustrated by Keen (1966, p. 3, pl. 1, fig. 21) reveals a slender, elongated shell with slightly concave whorls. Earlier, Keen (1958, p. 290) considered T. marmorata to be a variant of the extremely variable T. gonostoma Valenciennes (1832, p. 275; Kiener, 1843-1844, pp. 21-22, pl. 10, fig. 1). That species was originally described but not illustrated from “Habitat ad oras Americae australis in portum Acapulco Mexicanorum.” The posterior portion of the whorls on 7. gonostoma is often the widest; the color 42 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. of rather dense gray-black mottled markings in contrast to the much narrower flammules of 7. broderipiana and T. b. marmorata. A Recent specimen in the collections of the California Academy of Sciences, closely resembling Kiener’s illustrations of 7. marmorata, collected by D. L. Frizzell at Cabo Blanco, Peru, is 116 mm. long (incomplete), the body whorl 22.9 mm. in diameter. Other specimens collected at the same locality and at Paita Bay, and at the mouth of the Brazos River at San Ramon south of Sechura, Peru, have similar color markings but the shells are much less tapering with correspondingly wider apical angle. One such shell collected by J. G. Marks at San Pedro, Ecuador, is 133 mm. long (incomplete), the diameter of the last whorl 38 mm. Evidently the slender and the broad shells either represent ele- ments of a very variable series, or two distinct forms are living in the region of Ecuador and Peru. The fossils from the Galapagos Islands closely resemble the slender form with flat-sided whorls shown in Kiener’s illustrations of 7. marmorata. In this character they differ from the type specimen of 7. broderipiana (with slightly concave whorls) shown in Keen’s (1966) illustration. On that basis, in the present paper, the form described by Kiener is treated as a subspecies of T. broderipiana. Pilsbry and Olsson (1941, p. 43) mentioned that Turritella alturana Spieker, described from strata of Miocene age in Peru is ‘‘so close to the recent species | 7. broderipiana| that they cannot be consistently separated.” The subgenus Broderiptella Olsson was proposed to include the 7. broderipiana group represented in the northern South American region at least since middle Miocene time. This group of turritellas was discussed by Merriam (1941, pp. 50-51) and by Woodring (1957, pp. 110-112). LITERATURE CITED BANFIELD, A. F., D. St. Crarir, anp C. H. BEHre, Jr. 1956. Geology of Isabela (Albemarle) Island. Archipiélago de Colon (Galapagos). Bulletin of the Geological Society of America, vol. 67, no. 2, pp. 215-234, pls. 1-4, figs. 1-4 in text, February. BErRY, S. S. 1968. Some unusual mollusks, mainly Panamic. The American Malacological Union, Inc., Annual Reports for 1967, pp. 71-72, (1968). BLAINVILLE, H. D., DE 1825. Dictionnaire des Sciences Naturelles, vol. 34, p. 251, June. CauQuoiIn, MICHELE 1969. Mollusques récoltés par M. Hoffstetter sur les cotes de l’Equateur et aux Iles Galapagos. Mactridae. Bulletin du Muséum National d’Histoire Naturelle (Paris), 2° Sér., vol. 40, no. 5, pp. 1019-1030, fig. 1, 1968 (June 6, 1969). CHESTERMAN, C. W. 1963. Contributions to the petrography of the Galapagos, Cocos, Malpelo, Cedros, San Benito, Tres Marias, and White Friars Islands. Proceedings of the California Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 43 Academy of Sciences, Fourth Series, vol. 32, no. 11, pp. 339-362, 7 figs., September 10. [Baltra (South Seymour) Island, pp. 344-345.] Cuuss, L. G. 1933. Geology of Galapagos, Cocos, and Easter Islands. With petrology of Galapagos Islands by C. Richardson. Bernice P. Bishop Museum, Bulletin 110, pp. 1-67 [Chubb, pp. 1-44; Richardson, pp. 45-67], pls. 1-5, figs. 1-9 in text. CossMANn, M., AND A. PEyROT 1916. Conchologie Néogénique de L’Aquitaine. Scaphopodes et Gastropodes. Actes Societé Linnéenne de Bordeaux, tome 69, pp. 1-384, pls. 1-10, figs. 1-50 in text. [Also issued separately, tome 3, Livraison 1, December, 1917. |] DatL, W. H. 1924. Note on fossiliferous strata of the Galapagos Islands explored by W. H. Ochsner of the Expedition of the California Academy of Sciences in 1905-6. Geological Magazine, vol. 61, no. 723, pp. 428-429. Dat, W. H., anp W. H. OcHSNER 1928. Tertiary and Pleistocene mollusks from the Galapagos Islands. Proceedings of the California Academy of Sciences, Fourth Series, vol. 17, no. 4, pp. 89-139, pls. 2-7, 5 text-figs., June 22. Darwin, C. R. 1846. Geological observations on South America. Being the third part of the Geology of the voyage of the Beagle .... during 1832 to 1836. (London), pp. i-vii, 1-279, 5 pls. (1 geol. col.), 1 map, text illustr. [From the Catalogue of the British Museum, A-D, p. 422, 1903.] 1896. Geological observations. On the volcanic islands and parts of South America visited during the voyage of H. M. S. ‘Beagle’, edition 3, New York: D. Appleton and Co., pp. I-XIII, 1-648, figs. 1-40 in text, pls. 1-5. [Galapagos Archipelago, pp. 110-131]. DurHAM, J. W. 1950. 1940 E. W. Scripps cruise to the Gulf of California. Part II. Megascopic paleon- tology and marine stratigraphy. Geological Society of America, Memoir 43, pp. VII-VIII, 1-216, pls. 1-48, August 10. Eames, F. E. 1951. The type species of Anodontia Link 1807. Proceedings of the Malacological Society of London, vol. 28, pt. 6, pp. 232-233, June 8. FISCHER-PIETTE, E. 1969. Mollusques récoltés par M. Hofistetter sur les cotes de l’Equateur et des Iles Galapagos. Veneridae. Bulletin du Muséum National d’Histoire Naturelle (Paris), 2° Sér., vol. 40, no. 5, pp. 998-1018, pls. 1-3, 1968 (June 6, 1969). Fraser, C. McLEAn 1943. General account of the Scientific work of the Velero III in the eastern Pacific, 1931-41. Part I. Historical Introduction, Velero III, Personnel. Allan Hancock Pacific Expeditions I (1): 1-48, pls. 1-16, July 1943. Part II. Geographical and Biological associations. 1(2): 49-258, pls. 17-128, December, 1943. Part III. A ten year list of the Velero III Collecting Stations. 1 (3): 259-424, charts 1-115; an appendix of Collecting Stations of the Allan Hancock Foundation for the year 1942, pp. 425-431, December. Herm, D. 1969. Marines Pliozin und Pleistozan in Nord- und Mittel-Chile unter besonderer Beriicksichtigung der Entwicklung der Mollusken-Faunen. Zitteliana, Abhand- 44 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. lungen der Bayerischen Staatssammlung fiir Paldontologie und _historische Geologie, 2, pp. 1-159, pls. 1-18, figs. 1-46, September 1. HERTLEIN, L. G. 1932a. [Jn] A Pioneer Heritage, by Sam T. Clover, Los Angeles; Saturday Night Pub- lishing Company, pp. 206-207. 1932b. Gastrocopta munita on South Seymour Island, Galapagos Group. Nautilus, vol. 46, no. 2, pp. 69-70, October (issued October 22). HeErTLEIN, L. G., AND A. M. STRONG 1939. Marine Pleistocene Mollusks from the Galapagos Islands. Proceedings of the California Academy of Sciences, Fourth Series, vol. 23, no. 24, pp. 367-380, pl. 32, July 20. 1947. Eastern Pacific Expeditions of the New York Zoological Society. KXXIX. Mollusks from the west coast of Mexico and Central America. Part V. Zoologica, New York Zoological Society, volume 31, part 4, pp. 129-150, pl. 1, February 21. 1955. Marine mollusks collected at the Galapagos Islands during the voyage of the Velero III, 1931-1932. [Jn] Essays in the Natural Sciences in honor of Captain Allan Hancock on the occasion of his birthday July 26, 1955. (University of Southern California Press: Los Angeles), pp. I-XII, 1-345. (Hertlein and Strong, pp. 111-144, pl. A). HOFFsTETTER, R. 1956. Lexique Stratigraphique International, vol. 5. Amérique Latine. Fascicule 5a. Ecuador. (Appéndice Islas Galapagos o Archipiélago de Colon). Paris 8°, 191 pp. [Baltra (South Seymour) Island, pp. 155—157.] Hutton, F. W. 1886a. New species of Tertiary shells. Transactions and Proceedings of the New Zealand Institute, vol. 18, pp. 333-335, May. 1886b. The mollusca of the Pareora and Oamaru systems of New Zealand. Proceedings of the Linnaean Society of New South Wales, second series, vol. 1, pt. 1, pp. 205-237, May 25. Keren, A. Myra 1958. Sea Shells of Tropical West America. Marine mollusks from Lower California to Colombia. (Stanford University Press: Stanford, California), pp. I—XII, 1-624, 1709 figs. in text [unnumbered frontispiece (colored) p. XI], pls. I-X in color. 1966. West American types at the British Museum (Natural History). III. Alcide d’Orbigny’s South American collection. The Veliger, vol. 9, no. 1, pp. 1-7, pl. Is Vialby al. Lewis, G. E. 1956. Galapagos Islands (Archipiélago de Colon) Province. Geological Society of America, Memoir 65, pp. 289-291, June 15. [Baltra (South Seymour) Island, p. 290.] LupBrook, NELLIE H. 1959. A widespread Pliocene molluscan fauna with Anodontia in South Australia. Transactions of the Royal Society of South Australia, vol. 82, pp. 219-233, pls. 1-5, April. Marks, J. G. 1951. Miocene stratigraphy and Paleontology of southwestern Ecuador. Bulletins of Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 45 American Paleontology, vol. 33, no. 139, pp. 1-162, pls. 1 (43)-9 (51), figs. 1-12, December 20. McBrrney, A. R., AND WILLIAMs, H. 1969. Geology and Petrology of the Galapagos Islands. (With chemical analyses by Ken-ichiro Aoki.) Geological Society of America, Memoir 118, pp. I-XIII, 1-197, plates 1-21, figures 1-48 in text, tables 1-21. Menarp, H. W., T. E. CHASE, AnD S. M. SmitH 1964. Galapagos Rise in the southeastern Pacific. Deep-Sea Research, vol. 11, no. 2, pp. 233-242, figs. 1-4, April. Merriam, C. W. 1941. Fossil Turritellas from the Pacific Coast region of North America. University of California Publications, Bulletin of the Department of Geological Sciences, vol. 26, no. 1, pp. I-V, 1-214, pls. 1-41, figs. 1-19 in text, 1 map, March 8. Moore, H. B. 1934. On “lJedging” in shells at Port Erin. Proceedings of the Malacological Society of London, vol. 21, part 3, pp. 213-217, pl. 22, December. OcHSNER, W. H. 1906. Geology of the Galapagos Islands. Unpublished notes in the California Academy of Sciences. Otsson, A. A. 1964. Neogene Mollusks from northwestern Ecuador. Paleontological Research Insti- tution, Ithaca, New York, pp. 1—256, pls. 1-38, October 28. D’ORBIGNY, A. 1834-1847. Voyage dans l’Amérique Méridionale. Mollusques. Paris, vol. 5, pt. 3, pp. I-XLII, 1-758, atlas, pls. 1-85. Pirssry, H. A., anp A. A. OLsson 1941. A Pliocene fauna from western Ecuador. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 93, pp. 1-79, pls. 1-19, September 9. Ranson, G. 1959. Mollusques de la plaine cotiere soulevée de Guadalupito, récoltés par M. Bernardo Boit. Journal de Conchyliologie, vol. 99, no. 2, pp. 66-76, May 15. REEVE, L. 1849. Conchologia Iconica, vol. 5, Turritella, plates I-XI, and explanations, May to June. SLEVIN, J. R. 1959. The Galapagos Islands. A history of their exploration. Occasional Papers, Cali- fornia Academy of Sciences, no. XXV, pp. I-X, 1-150, figs. 1-31, December 22. SoweErBY, G. B., (Ist of the name) 1846. Descriptions of Tertiary fossil shells from South America. [Jn] Darwin, C. A., Geological observations on South America. Appendix to Part II. [See Darwin.] 1896. Edition 3, pp. 605-623, pls. 2-4. STEARNS, R. E. C. 1893. Scientific Results of Explorations by the U. S. Fish Commission Steamer Albatross. XXV.—Report on the mollusks of the Galapagos Islands, with descriptions of new species. Proceedings of the United States National Museum, vol. XVI, no. 942, pp. 353-450, pls. 51, 52 (map), September 29. VALENCIENNES, A. 1832. Coquilles univalves de l’Amérique Equinoxiale, recueillies pendant le voyage de MM. de Humboldt et Bonpland. [Jz] Humboldt, F. H. A., von, and A. J. A. 46 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Bonpland, Voyage aux régions equinoxiales du nouveau continent. Paris, pt. 2, Recueil d’observations de Zoologie et d’Anatomie comparée, vol. 2, pp. 262— SO), joll, S75 WEISBORD, N. E. 1962. Late Cenozoic Gastropods from northern Venezuela. Bulletins of American Paleontology, vol. 42, no. 193, pp. 1-672, pls. 1-48, March 5. Wi.iAMs, H. 1966. Wotr, T. 1895. Geology of the Galapagos Islands. [Jn] The Galapagos. Proceedings of the Symposium of the Galapagos International Scientific Project. R. I. Bowman, editor, University of California Press, pp. I-X VII, 1-318, illustrated. [Williams, Geology, pp. 65-70]. Die Galapagos-Inseln. Verhandlung der Gesellschaft fiir Erdkunde zu Berlin, Bd. 22, pp. 246-265, pl. 3 [map]. Wooprinc, W. P. 1957. Geology and Paleontology of Canal Zone and adjoining Parts of Panama. Geology and description of Tertiary mollusks (Gastropods: Trochidae to Turritellidae). United States Geological Survey, Professional Paper 306—-A, pp. I-IV, 1-145, pls. 1-23, figs. 14 in text. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XX XIX, No. 4, pp. 47-53; 2 figs.; 1 table April 6, 1972 A NEW SPECIES OF THE SCORPIONFISH GENUS HELICOLENUS FROM THE NORTH PACIFIC OCEAN By Tokiharu Abe Tokai Regional Fisheries Research Laboratory, Ministry of Agriculture and Forestry, Katchidoki 5-5-1, Kyobashi P. O., Tokyo 104, and Zoological Institute, Faculty of Science, University of Tokyo, Hongo, Tokyo 113, Japan and William N. Eschmeyer California Academy of Sciences, Golden Gate Park, San Francisco, California, U.S.A. 94118 Apstract: Helicolenus avius is described as a new species in the fish family Scor- paenidae, subfamily Sebastinae. The type locality is the northwestern Pacific Ocean between Japan and Midway Island at the southern end of the Emperor Seamount Chain, 32°40/N., 172°17’E. to 35°05’N., 171°46’E., at a depth between 450 and 600 meters. A description and 2 figures are provided. Remarks on the genus Helicolenus are given. INTRODUCTION In the summer of 1970, around 21 and 22 August, the Japanese vessel Daini- Oriento Maru of the Tokusui Company, Ltd., was conducting night-time trawling operations for Beryx spendens at the Emperor Seamount Chain between Japan and the Hawaiian Islands. At several stations a total of more than 250 kg. of specimens of a new scorpionfish of the genus Helicolenus was captured. The genus Helicolenus is virtually worldwide in distribution, containing temperate and tropi- 1 Contribution no. B552 from the Tokai Regional Fisheries Research Laboratory, Tokyo. Supported in part by U. S. National Science Foundation Grant GB-—15811. [47] 48 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. cal offshore species. According to Matsubara (1943), the genus Helicolenus, along with Sebastiscus, Hozukius, and Sebastes, constitutes the subfamily Sebas- tinae. Recently, Eschmeyer and Hureau (1971) suggested that the genus Sebastes may be a terminal North Pacific-evolving offshoot characterized by a reduced rather than an incipient suborbital stay. They suggested that the genus Sebastes may have evolved from a tropical ancestor, particularly one with a com- plete suborbital stay and one sharing the characters found in the worldwide tropi- cal and temperate genus Helicolenus and the Oriental genera Sebastiscus and Hozukius. The discovery of another species of the genus Helicolenus may help in understanding the evolution of the subfamily Sebastinae and the family Scor- paenidae. The new species is quite distinct from the other species referable to the genus Helicolenus, particularly in having reduced head spination, smaller scales, longer gill rakers, better developed toothed protuberances (dentigerous knobs) on the anterior ends of the premaxillaries, and a forked caudal fin. As a final introductory comment, we wish to mention that Helicolenus dac- tylopterus frequently is the main ingredient in French Bouillabaisse, or the Mediterranean fisherman’s stew. Perhaps this new species has similar qualities. Although some specimens were discarded, 245.8 kg. of the present new species was frozen quickly on board the collecting vessel and kept at —20°C. Mr. Tsujisaki, who collected the specimens, has informed us that the fish kept at this tempera- ture remain in good condition for sale for three months. Chemical analysis of flesh from one specimen measuring 21 cm. in standard length was made by Mr. Masa-aki Takeuchi (Tokai Regional Fisheries Research Laboratory) and showed the following: body weight 212.5 grams, edible part 27 percent, water of flesh 72.6 percent, crude protein 19.2 percent, crude fat 5.6 percent, crude ash 1.3 percent, and calories per 100 grams 126.2. Species of Helicolenus frequently are one of the dominant fishes in their habitat, and the new species may prove to be of commercial value. METHODS Methods of measuring follow Eschmeyer (1969) and are similar to methods used for other teleostean fishes with a few exceptions. Measurements originating from the anterior end of the upper jaw (head length, snout length, standard length, jaw length) are taken from the anterior end of the premaxillaries, includ- ing the dentigerous knobs; pectoral fin length is measured from the base of the first ray to the end of the longest ray, with the fin pointing back; caudal fin length is measured from the posterior end of the hypural plate to the most distal ray when the upper and lower caudal fin lobes are squeezed together. Termi- nology of head spines for the genus Helicolenus is the same as used by Matsubara (1943) and by Eschmeyer (1969). Abbreviations of depositories of specimens are as follows: ABE—personal VoL. XXXIX] ABE & ESCHMEYER: A NEW SCORPIONFISH 49 collection of the senior author; BMNH—British Museum of Natural History; CAS—California Academy of Sciences; USNM—United States National Mu- seum; ZIUT—Zoological Institute, Faculty of Science, University of Tokyo. ACKNOWLEDGMENTS The present writers take pleasure in expressing here their sincere thanks to Messrs. Hisateru Tsujisaki and Shoji Tamayama, Division of Offshore Fishing, Tokyo Branch, Tokusui Company, Ltd., Tokyo, for their cooperation with the sen- ior author during the past three years, and especially for providing the specimens used in the present study; and to Mr. Masa-ake Takeuchi for making the chemi- cal analysis of the flesh of one specimen of the new species. Comments on the manuscript were made by Dr. Lo-chai Chen and Mrs. Lillian Dempster. Mrs. Dempster also helped with literature work and the selection of the scientific name. Miss Pearl Sonoda aided in curatorial ways. Helicolenus avius Abe and Eschmeyer, new species. (Figures 1-2.) MATERIAL EXAMINED. All specimens were collected in the northwestern Pa- cific Ocean between Japan and Midway Island, at the southern end of Emperor Seamount Chain, between 32°40’N., 172°17’E., and 35°05S’N., 171°46’E., at a depth between 450 and 600 meters (probably most from 475 m.), with a bottom trawl, vessel Daini—Oriento Maru, around August 21-22, 1970. Holotype: ZIUT 52457 (211 mm.S.L.). Paratypes: ABE 15256 (1, about 200 mm. T.L. [used for description of color in life only|); BMNH 1971 12.14.1 (1, 173 mm. S.L.); CAS 13614 (2, 172-200 mm.S.L.) and CAS 13615 (1, 178 mm. S.L., cleared and stained) ; USNM 206327 (2, 176-181 mm. S.L.). DESCRIPTION. (Measurements and counts summarized in table 1; body shape and coloration in figures 1-2.) Dorsal fin with 12 spines and usually 13% (13’2- 14%) soft rays. Anal fin with 3 spines and 6’ soft rays. Pectoral fin with 18-20 rays, most frequently 19; 3rd through 11th or 12th rays branched in available specimens. Pelvic fin with 1 spine and 5 soft rays. Gill rakers total 31-34, 9 or 10 on the upper arch and 2224 on the the lower arch, increasing in size toward angle of gill arch; longest raker about '2 of orbit diameter. Vertebrae 25. Airbladder absent. Head spines mostly rudimentary or absent; preorbital (lachrymal) bone with two weak spinous points over maxillary, sometimes vir- tually absent; five preopercular spines present, second longest, all broad; supple- mental preopercular spine absent; nasal spines present; opercular bone with two spines; preocular spine well developed; supraocular spine weak or absent; post- ocular and tympanic spines small, mostly covered by skin or developed as ridges only; upper posttemporal spine poorly developed or absent; supracleithral spine broad; nuchal spine sometimes present; other spines, including the parietal, sphen- 50 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TABLE 1. Counts and measurements for the type specimens of Helicolenus avius. (Measurements are in millimeters; see standard lengths in Material Examined section for depositories and catalog numbers.) Standard length 172 173 176 178 181 200 211 Dorsal rays 12-+13% 124-13% 124-134% 12+-13%4 12-+413% 12 eeeeiae Anal rays 3+6% 346% 346% 346% 346% 346% 346% Pectoral rays 19, 19 19, 19 20, 20 20, 19 18, 18 19, 19 19,19 Pelvic rays 1+5 1+5 1+5 1+5 1+5 1+5 1+5 Vertebrae 25 25 Gill rakers 10423 9424 9:44:22 9423 9-223) “iosepamemtoses Head length 61.5 58.7 65.7 62.5 64.2 73.4 76.7 Body depth 48.8 49.1 53.4 50.3 50.7 56.9 57.4 Orbit diameter 18.0 17.3 17.9 18.5 17.8 2233 22.0 Snout length 15.6 13.6 16.0 14.7 16.4 16.8 17.9 Interorbital width 9.2 9.1 9.5 9.2 10.8 11.9 1222 Jaw length 30.0 29.6 28.3 30.1 30.8 34.7 3025 Predorsal fin length 61.3 57.8 61.5 59.3 62.8 68.7 (Bis: Length 3rd dorsal spine 17.5 18.9 20.2 — — 22.8 PAL Length 1st anal spine 8.0 6.9 6.3 7.8 8.0 8.9 8.7 Length 2nd anal spine 16.9 15.9 17.0 16.2 i7e5 tee 19.0 Length 3rd anal spine 17.2 16.2 18.1 15.6 18.5 18.2 18.6 Length pectoral fin 49.1 49.2 51.0 50.3 53.7 57.7 57.9 Length pelvic fin 31.4 29.0 S10 28.7 Sil 33.0 Basil Length caudal fin 39.8 42.3 43.2 42.0 44.2 46.9 51.5 otic, coronal, pterotic, lower posttemporal, and suborbital spine absent or present only as scarcely developed ridge. Scales on sides of body small, ctenoid; vertical scale rows from supracleithral spine to end of hypural about 100, difficult to count; pored lateral line scales about 50-55, plus 5 or more on the caudal fin; most of head with cycloid scales, including the maxillary, cheek, and interorbital area; snout unscaled; scales on belly, pectoral fin, and bases of vertical fins mostly cycloid. Premaxillary, dentary, vomer, and palatine toothed; longest teeth on the dentigerous knob of the premaxillary; all teeth short, conical, not arranged in definite rows. Color pattern of preserved specimens as in figures 1-2. Most conspicuous feature the dark spots above and below the lateral line anteriorly and on the back and dorsal fin. Buccal cavity black posteriorly, pallid anteriorly. Peri- toneum black. (Color in life, according to Mr. Tsujisaki, was unchanged when the frozen specimens were handed to the senior author. A color slide was made on 12 October, 1970, of one specimen (ABE cat. no. 15256) by the senior author soon after receipt of the specimens, and the following is taken from the slide. The coloration was also illustrated in Abe (1970) ). Body mostly red above and white ventrally. Dorsal surface of body and dorsal fin mottled with dark brown spots on a red background; spots arranged as in figure 1 of a preserved specimen. Pectoral and caudal fins red with yellowish tips. VoL. XXXIX] ABE & ESCHMEYER: A NEW SCORPIONFISH 51 Ficure 1. Lateral view of Helicolenus avius, CAS 13614, paratype, 172 mm. in standard length. Name. The specific name avius is the Latin word for “‘out of the way, remote or solitary.”’ This name is in reference to the type locality, an isolated seamount. A Japanese name, ‘“‘okikasago,” meaning Scbastes-like or Helicolenus hilgendor fi- like fish, was given to this species (Abe, 1970). DIsTRIBUTION. This species is known only from the type locality. We expect that it might occur at other seamounts in the northwestern Pacific and partic- ularly along the Hawaiian Island chain. COMPARISON AND REMARKS. The subfamily Sebastinae has been defined by Matsubara (1943). Two of the four genera, Sebastes (including Sebastodes) and Sebastiscus, have an incomplete suborbital stay which does not attach to the preopercle. The other two genera of the subfamily, Helicolenus and Hozu- kius, have a complete suborbital stay. Species of Helicolenus lack an airbladder and have 25 vertebrae; while the single species of Hozukius has an airbladder and 26 vertebrae; the species show cranial differences also. At most there are five or six closely related known valid species or sub- species belonging to the genus Helicolenus. Helicolenus dactylopterus (Dela- roche, 1809) has a wide distribution in the Atlantic Ocean extending barely into the Indian Ocean in South Africa; synonyms include H. maculatus (Cuvier, 1829), H. imperialis (Cuvier, 1829), H. maderensis Goode and Bean, 1896, H. thelmae Fowler, 1937, H. uruguayensis Fowler, 1943, and as a subspecies H. d. lahillei Norman, 1937 (see Eschmeyer, 1969). Helicolenus mouchezt (Sauvage, 1875) is a senior synonym of H. tristanensis Sivertsen, 1945, and is known from the south Atlantic at Tristan da Cunha and in the southern Indian Ocean at Saint Paul and Amsterdam islands (see Eschmeyer and Hureau, unr bo Ficure 2. Lateral and dorsal views of paratype, 172 mm. in standard length. the head of CALIFORNIA ACADEMY OF SCIENCES Helicolenus [Proc. 4TH SER. avius, CAS 13614, VoL. XXXIX] ABE & ESCHMEYVER: A NEW SCORPIONFISH 53 1971). The nominal Oriental species is H. hilgendorfi (Steindachner and Déoder- lein, 1884), although this species has been treated as H. dactylopterus by some authors. Helicolenus lengerichi Norman, 1937, is known from the southeastern Pacific. The Australian-New Zealand species is H. papillosus (Schneider, 1801) with the following as synonyms, H. percoides (Richardson, 1842), H. cottoides (Forster, 1849), H. barathri (Hector, 1875) and H. maccullochi (Phillipps, 1927) (see Whitley, 1968, p. 83). Helicolenus microphthalmus Norman, 1935, was shown to belong to the genus Sebastiscus (Wheeler and Eschmeyer, 1968), and H. rufescens Gilbert, 1905, from Hawaii belongs in the subfamily Scor- paeninae (Eschmeyer, 1969). Helicolenus avius agrees with these six species or subspecies in such features as counts of fin rays, general body shape, absence of an airbladder, condition of the suborbital stay, and 25 vertebrae. Helicolenus avius differs from the other species of Helicolenus in the following features. In H. avius the caudal fin is forked while the caudal fin in the others is “‘square- cut” or only slightly emarginate; the gill rakers are longer in H. avius; the tubed lateral-line scales number more than 50 in H. avius while they are fewer than 35 in the other species; the body scales are smaller in H. avius, about 100 vertical scale rows versus 80 or fewer; H. avius shows reduced spination, with some spines which are normally found in the other species either poorly devel- oped in H. avius, present as slight ridges, or absent; and H. avius, has the pro- tuberance or dentigerous knob at the end of each premaxillary bone much better developed. LITERATURE CITED ABE, TOKIHARU 1970. Shingao no sakana, 1970 nen ban [New faces of fishes, edition for the year 1970]. Publications of the Shigezo Ito Institute of Ichthyology, Tokyo. 1970, no. 2, 8 pp. (In Japanese). ESCHMEYER, WILLIAM N. 1969. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scorpaenidae). Occasional Papers of the California Academy of Sciences, no. 79, 130 pp., 13 figs. EscHMEYER, WILLIAM N., and JEAN CLAUDE HUREAU 1971. Sebastes mouchezi, a senior synonym of Helicolenus tristanensis, with comments on Sebastes capensis and zoogeographical considerations. Copeia, 1971, no. 3, pp. 576-579. MartsusBara, KiyoMATsuU 1943. Studies on the scorpaenoid fishes of Japan. The Transactions of the Sigen- kagaku Kenkyusyo, no. 1, 486 pp., 4 pls. WHEELER, ALWYNE, and WILLIAM N. ESCHMEYER 1968. The identity of the “British” scorpionfish Helicolenus microphthalmus. Jour- nal of the Linnean Society (Zoology), vol. 47 (no. 312), pp. 309-314, 1 fig. WHITLEY, GILBERT P. 1968. A check-list of the fishes recorded from the New Zealand region. The Australian Zoologist, vol. 15, part 1, pp. 1-102. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XX XIX, No. 5, pp. 55-64; 1 fig.; 1 table. April 6, 1972 TWO NEW SCORPIONFISHES (GENUS SCORPAENODES) FROM THE INDO-WEST PACIFIC, WITH COMMENTS ON SCORPAENODES MUCIPARUS (ALCOCK) By William N. Eschmeyer California Academy of Sciences, San Francisco, California 94118 and K. V. Rama Rao Zoological Survey of India, Calcutta-16, India ABSTRACT: Two new scorpionfishes of the genus Scorpaenodes are described from the Indian Ocean. Additional comments are given on Scorpaenodes muciparus (Al- cock). These three species are compared with each other and with S. tribulosus Eschmeyer, the fourth species of the genus Scorpaenodes known from offshore areas in the Indo-West Pacific. INTRODUCTION Eschmeyer (1969a) identified Indian Ocean specimens collected by the United States International Indian Ocean Expedition as Scorpaenodes muciparus (Alcock, 1889), offering a description and figures. Recently, we had the oppor- tunity to examine two specimens of S. muciparus in the collection of the Zoological Survey of India, and one of us (W.N.E.) was also able to examine the specimen collected by the Siboga which was identified as S. muciparus by Weber (1913) and also by de Beaufort (Weber and de Beaufort, 1962). It was then apparent that the specimens identified by Eschmeyer (1969a) as S. muciparus were not specimens of that species but represented a new species, a species which one of us (K.V.R.R.) was in the process of describing as part of current studies on the Marine Biological Laboratory LIBRA Pv APR141972 \VAlan nt | aT ee ee a { [55] 56 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. scorpionfishes of India. Examination of scorpionfishes in other museums has resulted in location of yet another closely related species, the description of which we include in this paper. These three species, along with S. tribulosus (Esch- meyer, 1969a), live in offshore level-bottom areas, while other members of the genus are shallow-water rock or reef inhabitants. ACKNOWLEDGMENTS The senior author acknowledges the support of National Science Foundation Grant GB-15811, which permitted him to examine scorpionfishes in most of the major fish collections. The following museums contained specimens used in this study, and we wish to thank the curators and staffs listed for courtesies extended during his visits to their museums: BMNH, British Museum of Natural History (N. B. Marshall, G. Palmer, A. Wheeler, P. Whitehead) ; CAS, California Academy of Sciences (L. Dempster, P. Sonoda); NMS, National Museum of Singapore (E. Alfred); USNM, United States National Museum (N. Gamblin, V. Springer); ZMA, Zoological Museum, Amsterdam (H. Nijssen); ZMC, Zoologi- cal Museum, Copenhagen (J. Nielsen) ; ZSI, Zoological Survey of India (A. G. K. Menon). Lillian Dempster, Pearl Sonoda, Kathy Smith, Terry Arambula, and Maurice Giles of the California Academy of Sciences, assisted in the study. We are grate- ful for comments on the manuscript made by Lillian Dempster and A. G. K. Menon. METHODS Counts, measurements, and terminology of head spines are as used by Esch- meyer (1969b, pp. 4-6). SPECIES ACCOUNTS The limits of the genus Scorpaenodes have been discussed recently (Esch- meyer, 1969a). Within the subfamily Scorpaeninae, the genus is characterized by the following combination of characters: normally thirteen dorsal spines, spinous procurrent caudal fin rays, and no palatine teeth. The genus contains species which are primarily tropical, shallow-water forms, and species of the genus are found in all warm oceans. The four species of Scorpaenodes treated in the present paper are the only ones of the genus occurring in offshore Indo-Pacific waters below a depth of about 20 fathoms. These four species may be separated as follows: 1. Vertical scale rows (above lateral line from first lateral line scale to end of hypural) OMOT oI) O1 CG ae ee 2 nee oe ee Se S. muciparus 1. “Vertical ‘scale rows about 50 or fewer 2.20 Eee 2 2. Soft dorsal rays 8%, scales strongly ctenoid with long ctenii, underside of head covered with ctenoid scales —___ ES GV ANE en Ye ee S. tribulosus Vor. XXXIX ESCHMEYVER & RAO: TWO NEW SCORPIONFISHES 57 2. Soft dorsal rays usually 914, scales ctenoid, underside of head naked or with cycloid SOPUBS en eed EE Se ee ee ae Se ee en ee a 3 Remarks. A new species, Scorpaenodes steinitzi, was described recently by Klausewitz and Frgiland (1970) from the Red Sea. Seventy-five additional speci- mens of this species are available to us. Two of the major differentiating char- acters used by these authors were that the dorsal and anal soft ray counts were 1 higher in S. steimitzi than in the other species, but this results from the fact that Klausewitz and Fr¢iland counted the last double ray as 2 rays, while recent authors count this as 1 or 1'2 rays. Despite this, Scorpaenodes steinitzi appears to be a valid species which is distinguishable from other species most easily by its coloration. We mention this because readers may wonder if S. steinitzi might not be one of the species treated here by us. Scorpaenodes steinitzi is a pale-colored species with some brown pigment on the sides and it usually has a dark spot near the posterior end of the spinous dorsal fin, but S. steinitzi does not have the lateralis canal head pores well developed and it is a shallow-living species. In S. steinitzi the spots on the caudal fin when present are arranged in rows while they are more scattered in S. smithi; and the dark pigment on the sides in S. steinitzi is not arranged in definite bars as in S. smithi, S. investigatoris, and S. muciparus. Scorpaenodes investigatoris Eschmeyer and Rama Rao, new species. Scorpaenodes muciparus, ESCHMEYER, 1969a, pp. 4-8, figs. la, b, table 1 (description; these specimens now referred to this new species, not S. muciparus (Alcock, 1889). Scorpaenodes varipinnis, SMiTH, 1957, p. 65 (in part; only the 50 mm. specimen taken from the stomach of a specimen of Pristipomoides microlepis Bleeker collected in 110 fathoms off Memba and tentatively referred to S. varipinnis). REMARKS. The specimens wrongly identified by Eschmeyer (1969a) as S. muciparus (Alcock) are designated as the types of this new species along with two specimens from the collection of the Zoological Survey of India. The holo- type (CAS 24264) and one paratype (CAS 24265) were figured by Eschmeyer (1969a, fig. la, b). TYPE MATERIAL. Holotype. CAS 24264 (1, 71.2 mm. S.L.), West Pakistan, 24°13’ N., 65°52’ E., in 93 fathoms, Anton Bruun Cruise 4B, station 267B, 9 December 1963. Paratypes. CAS 24266 (1, 87.4 mm. S.L.), West Pakistan, 25°02’ N., 56°52’ E., in 159 fathoms, Anton Bruun Cruise 4B, station 264A, 2 December 1963; CAS 24265 (2, 45.8-48.9 mm. S.L.), off western India, 22°32’ N., 68°07’ 58 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. E., in 31.5 fathoms, Anton Bruun Cruise 4B, station 221A, 18 November 1963; USNM 204030 (1, 72.4 mm. S.L.), off western India, 17°25’ N., 71°39’ E., in 52.5 fathoms, Anton Bruun Cruise 4B, station 202A, 13 November 1963; ZSI 916/1 (1, 86 mm. S.L.) and ZSI 917/1 (1, 67 mm. S.L.), Arabian Sea, off Rat- nagiri Coast, 17°27’ N., 71°41’ E., 56-58 fathoms, bottom of fine sand, Agassiz trawl haul, /nvestigator station 242, 11 October 1898. (Smith’s specimen men- tioned above in the Synonymy is regarded only as an additional specimen and not a type.) DeEscrIPTION. See Eschmeyer (1969a, pp. 4-8, figs. la and 1b, table 1) for a complete description of this species. The two ZSI specimens have the following counts: Dorsal rays XIII, 9% anal rays III, 5’2-6'2; pectoral rays 19. The vertical scale rows in the two specimens number 45-48, and the gill rakers num- ber 5—6 plus 12-13. Except for the specimen with 6’ soft anal rays, the counts fall within those given by Eschmeyer (1969a, table 1). DISTRIBUTION. Scorpaenodes investigatoris is known from the Gulf of Oman and in the Arabian Sea off western India and West Pakistan. The specimen re- ported by Smith (1957, p. 65) was taken off Memba, Mozambique. Depths of capture range from 31 to 159 fathoms. CoMPARISONS. This species very closely resembles Scorpaenodes smithi in scalation, counts, and coloration. Scorpaenodes investigatoris and S. smithi may be easily separated from the other two offshore species of Scorpaenodes: S. muciparus has over 70 vertical scale rows as opposed to about 50 in S. investigatoris and S. smithi, and S. tribulosus has strongly ctenoid scales on the underside of the head while the other species have the underside of the head mostly naked or with cycloid scales. The descriptions of S. investigatoris and S. smithi are almost identical. The two characters which may be used to separate them are pectoral rays and the coloration of the rear of the buccal cavity. In S. investigatoris the pectoral rays are usually 19 (18-20) while they usually number 17 or 18 (16-19) in S. smithi. In S. investigatoris the rear of the mouth, including the pharyngeal bones and the areas adjacent to them, is dusky in color- ation while this area is pallid in S. smithi. No other clear-cut differences were noted. Scorpaenodes investigatoris appears to live at deeper depths than S. smith. Etymo.Locy. The name is based on the Royal Marine Survey Steamer Investigator, which made important collections in the Indian Seas during the years 1884-1926; the name “‘investigatoris” was used by one of us (K.V.R.R.) for this species in his unpublished thesis on Indian Ocean scorpionfishes. Scorpaenodes muciparus (Alcock). Sebastes muciparus ALCOCK, 1889, pp. 298-299, fig. 3 on pl. 22 (original description; type locality Bay of Bengal, 26 miles N. by E. of Gopalpur, in 45 fathoms). Atcock, 1898, pl. 18, fig. 5 (good illustration). WerBER, 1913, p. 491 (brief description; one from Siboga station 306, Solor Straits, 8°27’S., 122°54’W., in 247 meters). Vor. XXXIX ESCHMEYER & RAO: TWO NEW SCORPIONFISHES 59 Scorpaenodes muciparus, de Beaufort in WEBER and DE BEAUFORT, 1962, pp. 34-35 (redescrip- tion of Siboga specimen described by Weber). MATERIAL EXAMINED. ZSI 1179/1 (1, 80 mm. S.L.) Gulf of Martaban, 14°46/N., 95°52’E., in 61 fathoms, on soft, green muddy and sandy bottom, Agassiz trawl haul, /nvestigator station 328, 7 March 1904; ZSI 12432 (1, 73 mm. S.L.) Bay of Bengal, Ganjam coast, 19°24’N., 85°E., in 23 fathoms, muddy bottom, Blake trawl haul, /nvestigator station 42, 7 March 1889; ZMA 110.246 (1, 146 mm. S.L.) Indonesia, Solor Straits, 8°27’S., 122°54’E., in 135 fathoms, Siboga station 306, 8 Feb. 1900. REMARKS. The reader is referred to the references listed above for a more complete description of this species. The holotype of this species, which was originally in the ZSI collection, has been lost. If a neotype is ever needed, we suggest that the specimen listed above from /nvestigator station 42 (ZSI 12432) would make the best neotype, as it is from the same general locality as the type and was identified as S. muciparus by Alcock. Scorpaenodes muciparus is charac- terized by a dorsal ray count of 13 spines and 9’ soft rays, pectoral rays of 18-19, and vertical scale rows of over 70. The other three species treated in this paper have a scale count of 50 or fewer. De Beaufort (in Weber and de Beaufort, 1962, p. 35) gives a scale count of 45—58 for the Siboga specimen. Re-examination shows that the specimen has a vertical scale row count of about 70-73 when the scale rows are counted above the lateral line from the supracleithral spine to the end of the hypural plate. Scorpaenodes tribulosus Eschmeyer. Scorpaenodes tribulosus ESCHMEYER, 1969a, pp. 8-10, fig. 1c, table 1 (original description; type locality East Africa, off Kenya, 02°42’S., 40°53’E., in 77 fathoms and off the Somali Republic, 11°24’N., 51°35’E., in 40-96 fathoms). REMARKS. The species is still known only from the western Indian Ocean, from the holotype (CAS 24267, off Kenya) and one paratype (USNM 204031, off the Somali Republic) as given above. The reader is referred to Eschmeyer (1969a) for a description of this species. This species is characterized by a dorsal ray count of 13 spines and 8% soft rays, pectoral rays 19, and vertical scale rows under 50. Scorpaenodes tribulosus is most easily distinguished from the other species by the presence of strongly ctenoid scales on the underside of the head; the other three species have this area covered with cycloid scales or mostly naked. Scorpaenodes smithi Eschmeyer and Rama Rao, new species. (Figure 1.) TYPE MATERIAL. Holotvpe. BMNH 1929.6.12.6 (52.2 mm. S.L.), Andaman Sea, off western Malaya, 5°45’N., 98°20’E., in 40 fathoms, Madras- Penang Cable Survey, Cable ship Patrol, 16 March 1929. [Proc. 4TH Serr. CALIFORNIA ACADEMY OF SCIENCES 60 ‘Bag uvUepUY ay} Woy “T'S “WU 7°75 “9°71°9'6761 HNN ‘edAjoj04 ‘satads Mau ‘ory ePWUIeY puR JaAIWIYISY 2y472WUWsS Sapouangs0I¢ ‘T qunorg Vor. XXXIX ESCHMEYVER & RAO: TWO NEW SCORPIONFISHES 61 Paratypes. BMNH1929.6.12.7-15 (12, 21.5-47.0 mm. S.L.), and CAS 13616 (4, 23.2-52.1 mm. S.L.) and ZSI F6254/2 (1, 46.0 mm. S.L.), formerly in BMNH 1929.6.12.6—-15, taken with the holotype; BMNH 1932.2.15.15-20 (6, 24.5-48.3 mm. S.L.), Andaman Sea, west of Malaya, 6°01’20’N., 99°03’05”E., in 46 fathoms, mud bottom, Madras-Penang Cable Survey, no date; BMNH 1972.1.17.1-9 (9, 22.0-51.3 mm. S.L.) and USNM 206501 (4, 22.4—45.6 mm. S.L.) west of Malaya, 80 miles south of Penang, in 24 fathoms, en- tangled in coelenterates found on cable, cable ship Patrol, May 1923; BMNH orot.4 (1, 35.6 mm. S.L.), Arafura Sea, 11°59’30’S., 126°38’E., in 60 fathoms, E. and A. Telegraph Company, collected in or before 1933; Mie P7982—-83 (2, 56.5 mm. S.L.), Gulf of Tonkin, 20°26’N., 108°09’E., in 28 fathoms, collected by Schgnau in or before 1895; ZMC P7978-81 (4, 46.2-60.8 mm. S.L.), China Seas, collected by FE. Svenson, in or before 1893; ZMC P79100-112 (13, 34.0-62.4 mm. S.L.), southwest of Hong Kong, 26°10’N., 121°00’E., in 44 fathoms, collected by H. Christiansen, 31 May 1912; ZMC P79113-132 (20, 28.7-46.8 mm. S.L.), off western Malaya, 5°09’08’”N., 99°48’10”E., in 32 fathoms, sandy mud bottom, collected by Store Nordiske, 24 Dec. 1935; ZMC P7991-92 (2, 32.1-58.8 mm. S.L.), Formosa Strait, ““Namoa” Island, collected by Capt. Svenson, in or before 1912; NMS uncataloged (10, 15.5-54.4 mm. S.L.), near Singapore 5°59’06”N., 99°08’33”E., in 40 fathoms, collected by Tweedie. A total of 89 specimens. DEscriPTION. Measurements and counts summarized in table 1; body shape and coloration in figure 1. Dorsal fin rays normally XIII, 9'%, rarely 8’ or 10’. Anal fin rays normally III, 5%. Pectoral rays usually 17 or 18 (16-19), rays 2 or 3 through 7 to 9 branched in larger specimens; smaller specimens with fewer branched rays; the branching of pectoral rays begins at about 25-30 mm. S.L. Spines on head well developed. Preorbital bone with second and third lobes (Matsubara’s, 1943, terminology) each as a broad spinous point extending over the maxillary, third lobe usually with additional spinous points; suborbital ridge with one row of spinous points, usually one and sometimes several spines on the lateral face of the preorbital bone in line with three or more spinous points on the suborbital bones, the posterior one frequently double. Tympanic spines present; coronal spines usually absent, sometimes present only on one side, rarely present on both sides. Interorbital ridges rarely ending in spines. Small spines (postfrontal spines of Smith, 1957) near midline between tympanic spines sometimes present. Supple- mental preopercular spine present. First preopercular spine long, second usually small, third present, fourth usually present, points ventrally, and fifth points down (sometimes virtually absent). Preocular, supraocular, and postocular spines well developed. Small spine below parietal spine usually absent. Other spines present include the nasal, parietal, nuchal, sphenotic, pterotic, upper (some- 62 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. TABLE 1. Counts’ and measurements of type specimens of Scorpaenodes smithi. (Measurements are in millimeters; numbers in parentheses are percentages of standard length.) BMNH 1929.6.12.6 CAS 13616 P7982-3 Standard length Sf S2el 48.7 44.2 23.2 56.5 56.5 Dorsal fin rays + XIII+8% XIII+9% XIII+9'4 XII+9% XIII+9% XIII+9% XIII+8% Anal fin rays ! TI+5% Wi+5% WI+5% IWI+5% IiI+5% I+6% Ii+5% Pectoral fin rays ! Vaats 1st; isseis | 1ssise es-ES 18-P1g) sels-oe Head length 22.6(43 ) 23.3(45) 21.2(44) 19.1(43) 10.7(46) 24.3(43) 22.0(39) Snout length 6.1(12) 6.5(12) 6.2(13) SAID) BAU) 6.5(12) 6.2(11) Orbit diameter 6.8(13) 6.8(13) 6.7(14) 5.9(13) 3.2(14) 6.2(11) 6.7(12) Interorbital width 3.3(06) 3.0(06) 2.8(06) 2.9(06) 1.7(07) 3.8(07) 3.2(06) Upper jaw length 11.9(23) 1320 C25) leo (24) O)7(22)) Sash(7S))) 12°42) 122222) Predorsal-fin length 20.7(40) 21.4(41) 20.0(41) 18.3(41) 10.3(44) 21.0(37) 20.4(36) Body depth 2222)\(43)) 21.5(41) 19.6(40) 17.0(38) 9.9(43) 22.7(40) 22.0(39) Pectoral fin length 17 (SS) 19.1(37) 16.5(34) 15.8(36) 7.4(32) 19.2(34) 21.5(38) Pelvic fin length 13.9(27) 135/(26)) 12:6 (26) 112526) 5.7(24) 14.2(25) 14.8(26) Caudal fin length 14.8(28) TAPAS) elo O28) aeliZesiC2.8)) 7.0(30) 14.8(26) 15.3(27) ZSI F6254/2 BMNH 1932.2.15.15-20 Standard length 46.0 48.3 46.6 48.1 47.8 32.2 24.5 Dorsal fin rays XIII+9'% XII+10% XIII+9'4 XIII+9% XII+9% XIII-+9% XIII-+9% Anal fin rays I+5% Wl-+5% I1-+5% I1+5% 145% II+5% IiI+5% Pectoral fin rays 17+18 17-+-17 18+18 17+17 17+17 18+18 18+18 Head length 19.8(43) 21.5(44) 20.3(44) 19.8(41) 19.7(41) 14.2(44) 11.4(46) Snout length 5.2(11) 5.7(12). 5.2011) 5,5(11) .S4(11). 4eCie ieee VoL. XXXIX ESCHMEYER & RAO: TWO NEW SCORPIONFISHES 63 (Table 1 Continued) ZSI F6254/2 BMNH 1932.2.15.15-20 Orbit diameter 5.7(12) 6.0(12) fl) 5.9(12) 6.0(12) 4.1(13) 3.6(15) Interorbital width 2.7(06) 3.2(07) 2.8( 06) 3.0(06) 3.2(07) 2.2(07) 1.8(07) Upper jaw length 10.0(22) PROCS) V1OSC2) 10422) e022) 7.5(23) 6.2(25) Predorsal fin length 18.8(41) 19.1(40) 18.6(40) 19.4(40) 19.0(40) 14.2(44) 11.2(46) Body depth 19.0(41) 18.8(39) 18.8(40) 19.8(41) 20.0(42) 13.2(41) 10.6(43) Pectoral fin length 14.6(32) 14.5(30) 14.4(31) 14.0(29) 14.4(30) 9.0( 28) 7.7(31) Pelvic fin length 12.2(26) IO AS) SAAT) UAE@aD) aC) 8.0(25) 7.0(28) Caudal fin length 13.0(28) 124 (26) 2.827) 12292) 126 (26) 8.6(27) 7.1(29) 1 Counts of 63 additional specimens give the following totals (12 small specimens not counted): Dorsal fin rays XIII+916(70 specimens), XIII+814(2), XIII+9(1), XII+1014(1), XIII+1014(2), and XIII4+-5%(1, abnormal); Anal fin rays III+514(75 specimens), III+614(1), I4614(1, abnormal); Pectoral fin rays 16+17(1 specimen), 17+17(22), 17+18(6), 18+17(7), 184+18(37), 184+19(1), 19+18(2), 19+19(1). times absent) and lower posttemporal, two opercular, cleithral, and supracleithral (double). Fourth suborbital bone isolated, bearing a few small spines (Gins- burg’s, 1953, postorbital spines). Tentacles and other fleshy appendages associ- ated with most head spines. Supraocular tentacle about % to ' of orbit diameter, sometimes reduced, with branches distally. A few tabs on upper part of eye, two usually larger than the others. Scales on flanks ctenoid; vertical scale rows about 40 to 50, usually in middle forties; lateral-line scales usually 23 plus 1 or 2 on caudal fin. Bases of fins with a few scales. Scales on chest and pectoral fin base mostly cycloid. Dorsal parts of head scaled; underside of head unscaled or with cycloid scales; snout unscaled. Gill rakers including rudiments 15-17; upper arch with 5—6 short spiny rakers; lower arch with 9-10 rakers on cerato- branchial and usually 2 rudiments on hypobranchial. Small slit present behind fourth gill arch. Head pores well developed. Color pattern as in figure 1. Most conspicuous feature is the black spot on the posterior part of the spinous dorsal fin. Body usually with four broad bands, first under anterior dorsal spines, second under posterior spinous dorsal fin, third between soft dorsal fin and anal fin, fourth at base of caudal fin. All fins with brown or black spots. Three brown bars radiating from eye; first extends forward across preorbital bone, second below eye, and third extends down and back from posterior ventral part of eye. DISTRIBUTION. Scorpaenodes smithi seems to be widespread in the area of 64 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr. the South China Sea and through the Straits of Malacca into the Andaman Sea. A wider distribution in the Indo-Australian Archipelago is expected. Depths of capture range from 24 to 60 fathoms, and the bottom type appears to be mud or sand. COMPARISONS. See the Comparisons section under the account of Scorpaenodes investigatoris. EtyMoLocy. We name this species in honor of the late Professor J. L. B. Smith for his noteworthy contributions to the knowledge of Indian Ocean scor- pionfishes. LITERATURE CITED Atcock, ALFRED 1889. Natural history notes from H. M.’s Indian Marine surveying steamer /nvesti- gator, Commander Alfred Carpenter, R.N., D.S.O., commanding. No. 12. Descriptions of some new and rare species of fishes from the Bay of Bengal, obtained during the season of 1888-89. Journal of the Asiatic Society of Bengal, vol. 58 (pt. 2, no. 17), pp. 296-305, pl. 22. 1898. Illustrations of the zoology of the Royal Indian marine surveying steamer Investigator, . . . Fishes—part 5. Calcutta, pls. 18-24. ESCHMEYER, WILLIAM N. 1969a. A new scorpionfish of the genus Scorpaenodes and S. muciparus (Alcock) from the Indian Ocean, with comments on the limits of the genus. Occasional Papers of the California Academy of Sciences, no. 76, 11 pp., 1 table, 1 fig. 1969b. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scor- paenidae). Occasional Papers of the California Academy of Sciences, no. 79, 130 pp., 13 figs., 15 tables. GInsBurG, ISAAC 1953. Western Atlantic scorpionfishes. Smithsonian Miscellaneous Collections, vol. 121, no. 8, 103 pp., 6 text figs. KLAUSEWITz, WOLFGANG, and @ysTEIN FROILAND 1970. Scorpaenodes steinitzi n. sp. von Eilat, Golf von Aqaba. Senkenbergiana Biologica, vol. 51, no. 5/6, pp. 317-321, 2 figs. MATSUBARA, KrvYOMATSU 1943. Studies on the scorpaenoid fishes of Japan. The Transactions of the Sigenkagaku Kenkyusyo, no. 1, 486 pp., 4 pls. SMiTH, J. L. B. 1957. The fishes of the family Scorpaenidae in the western Indian Ocean. Part 1. The subfamily Scorpaeninae. Ichthyological Bulletin, Rhodes University, no. 4, pp. 49-69, pls. 1-4. WEBER, MAx 1913. Die fische der Siboga-Expedition, Siboga Expedition, vol. 57, 710 pp., pls. 1-12, text figs. 1-123. WEBER, Max, and L. F. pE BEAUFORT. 1962. The fishes of the Indo-Australian Archipelago, XI. Scleroparei, Hypostomides, Pediculati, Plectognathi, Opisthomi, Discocephali, Xenopterygii. E. J. Brill, Leiden, 481 pp., 100 text figs. ee eee wee peer Marine Blological Laborate “se tote PROCEEDINGS | Wocds Hole, Mass. OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXIX, No. 6, pp. 65-74, 22 figs. September 5, 1972 NEW BARNACLE RECORDS (CIRRIPEDIA, THORACICA) By Victor A. Zullo, Dea B. Beach, and James T. Carlton California Academy of Sciences, San Francisco, California 94118 Balanus amphitrite amphitrite Darwin, 1854. (Figures 1-4.) In discussing the introduction of Balanus improvisus Darwin to the Pacific Coast of North America, Carlton and Zullo (1969) noted that although this species was relatively common in collections dating back to 1853, no specimens of B. amphitrite amphitrite were found in collections made prior to 1939, when it was first observed by F. L. Rogers in San Francisco Bay (Henry, 1942). The establishment of B. improvisus on the Pacific Coast appears to be tied to the commercial importation of the North American East Coast oyster, Crassostrea virginica (Gmelin). Both barnacles are frequent epizooites of this oyster, but as B. amphitrite amphitrite is a relatively recent addition to the fauna of the East Coast as well, Carlton and Zullo concluded that its absence in early Pacific Coast collections reflected a similar absence on the East Coast. How- ever, further inquiry into the history of the Pacific Coast oyster industry has revealed that all importations of Crassostrea virginica were from areas north of Cape Hatteras, North Carolina (the northern range limit of B. amphitrite amphitrite), thus virtually eliminating this oyster as a vehicle of introduction for B. amphitrite amphitrite. While processing miscellaneous collections of barnacles for the Academy of Natural Sciences of Philadelphia (ANSP), two lots of B. amphitrite amphitrite were found that had been obtained from the California coast prior to 1939. This material, collected by Charles R. Orcutt and originally identified by Henry A. Pilsbry, consists of four specimens with opercular valves from La Jolla, Cali- fornia, and of a single, large (20 millimeter basal diameter) specimen from San Diego Bay, California. Both lots bear the number “158/428”, but no date of [65] 66 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficures 1-4. Balanus amphitrite amphitrite Darwin. Figure 1, scutum, height 4 mm.; figure 2, shell, greatest diameter 20 mm.; figures 3-4, terga, height 4 mm. (Figures 1, 3, 4, La Jolla, California; figure 2, San Diego Bay, California). Ficures 5-6. Opercular valves of Balanus improvisus Darwin, CAS locality 41227, Delta Mendota Canal, California, height of scutum and tergum 3.8 mm. collection. Dr. Robert Robertson and Nancy Rulen of the Philadelphia Academy were kind enough to search their records, and were able to provide the following data: ANSP no. 2257. La Jolla, California, C. R. Orcutt collector, donated by the U. S. National Museum and cataloged December 6, 1921. ANSP no. 2426. San Diego, California, C. R. Orcutt collector, De- cember 9, 1927. The apparent establishment of B. amphitrite amphitrite in southern Cali- fornia by at least the early 1920’s and in the late 1930’s in northern California is likely attributable to transport by ships. Its occurrence may eventually be placed considerably earlier than this, for it had already arrived in Hawaii by the early 1900’s (Pilsbry, 1907, p. 190). Balanus amphitrite amphitrite is restricted to certain estuarine parts of the San Francisco Bay complex where mean annual temperatures are highest, but even in these areas Newman (1967) has noted that the temperature regime is clearly suboptimal for this subspecies. If the establishment of B. amphitrite amphitrite in San Francisco Bay was VoL. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS 67 indeed later than in southern California, the delay may be explained by the paucity of suitable environments for successful colonization. Balanus eburneus Gould, 1841. (Figures 8-15.) Balanus eburneus, whose natural range includes the Atlantic coasts of North America and northern South America, the Caribbean Sea, and the Gulf of Mexico, has been introduced by shipping to various parts of the world. Known introductions, emphasizing those in the Pacific Ocean, were summarized by Matsui e¢ al. (1964) and Utinomi (1966). Definite Pacific localities include Balboa, in the Bay of Panama, eastern Pacific, the Hawaiian Islands, and the Japan Sea coast of central Japan. Weltner (1897) reported B. eburneus on Fasciolaria with B. amphitrite from Manila in the Philippines, but this identi- fication has not been corroborated. A sample in the collection of the Department of Invertebrate Zoology of the California Academy of Sciences (CAS) taken by Mr. Vern Brock in July, 1967, from between Ansala and Vera islands in Eniwetok Atoll in the Marshall Islands was found to contain several individuals of B. eburneus in association with B. amphitrite amphitrite. These specimens represent the first record of B. eburneus from islands of the central West Pacific, and no doubt reflect the heavy traffic in American shipping to that area since the Second World War. The occurrence of this Atlantic American species at Eniwetok suggests that other highly frequented ports in the central Pacific islands might also harbor successfully introduced populations of B. eburneus. This species was also discovered recently at another, albeit more likely, locality in the western Atlantic. Mrs. Joleen Gordon, in the course of an ecologic study of Bermuda barnacles during the summer of 1967, found B. eburneus to be a common member of that fauna in association with B. amphitrite amphitrite and Chthamalus stellatus thompsoni Henry. The barnacle fauna of the Bermudas was investigated most recently by Henry (1958) on the basis of collections made by T. A. and Anne Stephenson in the early 1950’s. Balanus amphitrite hawaiiensis Broch (= B. amphitrite amphitrite) and Chthamalus stellatus thompsoni were reported there for the first time, but Balanus eburneus was not found. Judging from the abundance of B. eburneus and its association with the above-mentioned subspecies as seen in 1967, it is possible that this species has been introduced to the Bermuda fauna since the date of the Stephensons’ survey. Balanus improvisus Darwin, 1854. (Figures 5-7.) A living population of B. improvisus was discovered in December, 1962, on the concrete lining of a section of the Delta-Mendota fresh water irrigation 68 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ss & ° = . S & —_ (D ‘. Se , Antioch ~ Z e Oe*s o OES > Rae oO Z. ee e e w = . 3 . © # e iJ a—\ ea a 4 8 - Tracy =) oO 4 *e, e S % fe. os His tee 2 9 Oe, cos) ve Cy ‘. N 4 OF Ore e e 0 10 Miles *e == e Ficure 7. Relationship of Delta-Mendota Canal system to the San Francisco Bay- Delta area. Numbers indicate Tracy Pumping Plant (1) and collection site of Balanus improvisus (2). canal in central California. The barnacles, collected by G Dallas Hanna and Allyn G. Smith of the California Academy of Sciences (CAS locality 41227), were taken at mile post 20.62, or more than 20 miles downstream from the Tracy Pumping Plant, which is about 9 miles northwest of Tracy, San Joaquin County, California (figure 7). This section of the canal was discussed previously by Hanna (1966, p. 40, figs. 30-31) with respect to the phenomenal infestation of the Asiatic fresh water clam Corbicula manilensis (Philippi). The occurrence of barnacles in the canal was noted by Prokopovich (1968, p. 53, photo- graph 48). Irrigation water is pumped into the Delta-Mendota Canal by the Tracy Plant via a 2.3-mile-long canal from the Old River Channel of the San Joaquin River. This intake is well over 25 miles from the established salt water barrier of the Sacramento-San Joaquin estuary in the vicinity of Antioch, California. Vor. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS 69 Continuing water quality analyses made at the pumping plant indicate that total dissolved solids do not exceed 500 parts per million (0.5 parts per thousand) at the canal intake. Although a few species of barnacles are known to live in river estuaries where salinities are negligible for most of the year, there has been only one other report of an apparent occurrence in totally fresh water (Shatoury, 1958; B. amphitrite in a cement irrigation holding tank 64 kilometers south of the mouth of the Nile Delta). On the basis of observational and experimental data, it has been assumed that euryhaline barnacles, such as B. improvisus and B. eburneus, can exist in fresh water as adults provided that higher salinities prevail in the area during larval spawning and settlement. These outlying populations are apparently unable to reproduce, and must rely, therefore, on larval recruitment from breeding populations in higher salinities downstream. The larvae can be transported upstream in the encroaching salt wedge during the dry season, provided that this season (usually summer months) corresponds to spawning season. Laboratory studies of the salinity tolerances of stage I and II nauplii of B. balanoides (Linnaeus), B. balanus (Linnaeus), and B. crenatus Bruguiére were made by Barnes (1953), and Crisp and Costlow (1963) investigated the effects of varying salinities on the developing eggs and early naupliar stages of B. amphitrite amphitrite Darwin, B. eburneus, and Chelonibia patula Ranzani. The results from these two studies agree rather closely. Cytolysis of eggs was observed in all three species examined by Crisp and Costlow at salinities of 10 parts per thousand or less, regardless of temperature. Early stage nauplii of both studies died in salinities of 5 parts per thousand or less after only slight exposure, and could not withstand prolonged exposure to salinities below 10 parts per thousand. There was, however, some indication that larvae hatched from embryos developed at lowered salinities were better adapted to brackish conditions. The survival ability of B. improvisus larvae in lowered salinities may be greater than that of other estuarine barnacles. Mohammad (1962), in a field and laboratory study of the distribution of barnacle larvae in Newport River, North Carolina, indicated that B. improvisus larvae could tolerate salinity as low as O parts per thousand, whereas the lower limit for those of B. eburneus was about 8 parts per thousand. If larval salinity tolerance were the limiting factor in the distribution of barnacles in estuaries, then B. improvisus should be expected in fresh water. However, this is not the case under natural con- ditions, suggesting that other factors are also limiting. The only readily apparent physical difference between the two fresh water localities cited here and natural estuarine habitats is direction of current flow. Under natural conditions, larvae must counteract river current once outside the encroaching salt wedge, and Crisp and Costlow (1963) have noted that 70 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Ficures 8-10. Opercular valves of Balanus eburneus Gould, CAS locality 42617, St. Georges Island, Bermuda, height of scuta and tergum 7.5 mm. Ficures 11-15. Balanus eburneus Gould, CAS locality 42618, Eniwetok, Marshall Islands. Figures 11 and 13, scutum, height 4 mm.; figures 12 and 14, terga, height 4 mm.; figure 15, labrum, 100. larval swimming ability is greatly decreased in lowered salinity. In pumped irrigation systems, however, weak swimming larvae can be swept up from river channels and carried into normally unattainable environments. Whether Vor. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS 71 ot s ' ‘ae &: " : “a : 19 —_— Ficures 16-20. Creusia domingensis Des Moulins, CAS locality 42616, Bermuda. Figures 16-17, terga, height 2.5 mm., and figures 18-19, scutum, height 2.5 mm.; figure 20, shell in Porites astreoides, greatest diameter of orifice 0.9 mm. FicurEs 21-22. Shell of Hexacreusia durhami (Zullo), CAS locality 27229, Panama, greatest diameter 3.5 mm. Figure 21, uncoated shell showing radii; figure 22, shell coated with ammonium chloride to show external sculpture (apex of rostrum is damaged). or not flow reversal is an adequate explanation for these unusual occurrences, the possibility remains that barnacles may pose serious fouling problems in irrigation systems. Creusia domingensis Des Moulins, 1866. (Figures 16-20.) A coral-inhabiting barnacle of the genus Creusia Leach was also found by Mrs. Gordon in Bermuda in 1967. The specimens are in two small heads of the 72 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. coral Porites astreoides (Lamarck). The larger coral fragment, measuring about seven centimeters in greatest diameter, contains eleven barnacles, and the smaller, measuring about four centimeters, harbors three individuals. These barnacles were identified through the kindness of Mr. Arnold Ross of the Natural History Museum, San Diego, who is presently preparing a monographic study of coral-inhabiting balanids. He indicated that they probably represent Creusia domingensis; a species originally described in P. astreoides from Port-au-Prince, Haiti (Des Moulins, 1866), but which has gone unnoticed since that time. Further examination of specimens of P. astreoides in the collections of the California Academy of Sciences (CAS) and of the University of California Museum of Paleontology, Berkeley (UCMP) disclosed additional localities for C. domingensis from the vicinity of Biscayne Bay, on the east coast of Florida (CAS locality no. 42265) and from the Dry Tortugas, off southern Florida in the Gulf of Mexico (UCMP locality nos. A-8146, A-8149, B-3717). Previously, coral barnacles have been reported from the eastern Gulf of Mexico and the West Indies in the Tropical Atlantic region. The discovery of C. domingensis in the Bermudas and in southern Florida thus significantly increases the known range both of this little known species and of coralophilous balanids as a group. Hexacreusia durhami (Zullo, 1961). (Figures 21-22. This six plated coral barnacle was originally described as a species of Balanus Da Costa and placed in a new subgenus, Hexacreusia Zullo (Zullo, 1961). It was found in late Pliocene, Pleistocene, and Recent specimens of the coral Porites californica Verrill from various localities in the Gulf of California, and was later reported in extant specimens of the same coral from the Tres Marias Islands, Mexico, just south of the Gulf of California by Ross (1962). Although Hexacreusia is related to Balanus, and especially to the subgenus Armatobalanus Hoek, it is more closely allied to the genera Creusia Leach and Pyrgoma Gray, with which it shares a cup-formed basis, “creusioid’”’ rather than “balanoid” opercular valves, and an obligate coralophilous habitat. For these reasons, Hexacreusia is removed from the genus Balanus and raised to generic rank. It was noted earlier (Zullo, 1967) that certain specimens included by Darwin (1854) in his original description of the Australasian species Balanus allium were, in fact, Hexacreusia durhami. These barnacles were in a coral from Hugh Cuming’s collection purported to be from Australia, but which is most probably Porites californica from the Pacific Coast of tropical America. As Cuming had spent some time in Central America, but had never traveled Vor. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS ae north of the Gulf of Fonseca, Honduras, it was suggested that Darwin’s speci- mens came from an area south of its present-known range. Confirmation of the presence of Hexacreusia durhami in Central America is based on the recent acquisition of two lots from Panama. A beachworn piece of Porites californica collected by Dr. Leo G. Hertlein of the California Academy of Sciences on December 22, 1931, at Bahia Honda, Veragua, Panama (CAS-Geology locality no. 27229) contains three shells of Hexacreusia durhami without opercular valves. The second lot, made available by Professor J. Wyatt Durham of the University of California Museum of Paleontology (UCMP), consists of about two dozen specimens in a large fragment of a Porites question- ably identified as P. lobata (Dana). The coral was collected by Dr. Peter W. Glynn of the Smithsonian Tropical Research Institute (Canal Zone, Panama) from depths between 10 and 20 feet off the Las Secas Islands, Golfo de Chiriqui, Panama (UCMP locality no. D-4138). These barnacles were alive when taken and still retain opercular valves and bodies, although the latter are unsuitable for dissection because of bleaching and drying during processing of the coral. Hexacreusia durhami is probably present in Porites throughout the Panamic Province, but apparently is absent from the faunas of offshore Eastern Pacific islands. Extensive examination of hermatypic corals from the Galapagos Archi- pelago, Cocos Island (Costa Rica), and Clipperton Island has failed to yield coral barnacles, although a second species, to be described at a later date, has been discovered in ahermatypic corals from deeper water (90 meters) in the Galapagos. REFERENCES CITED Barnes, H. 1953. The effect of lowered salinities on some barnacle nauplii. Journal of Animal Ecology, vol. 22, no. 2, pp. 328-330. Cartton, J. T., anD V. A. ZULLO 1969. Early records of the barnacle Balanus improvisus Darwin from the Pacific coast of North America. Occasional Papers of the California Academy of Sciences, no. 75, pp. 1-6. Crisp, D. J., AND J. D. CostLow 1963. The tolerance of developing cirripede embryos to salinity and temperature. Oikos, vol. 14, no. 1, pp. 22-34. Darwin, C. 1854. A monograph on the sub-class Cirripedia, Balanidae, Verrucidae. Ray Society, London, viii + 684 pp., 30 pls., 11 text figs. Des Mouvutins, Cu. 1866. Liste des principaux fossiles recueilles par les membres de la Société a Cazeneuve dans le calcaire de Bazas. Actes de la Société Linnéene de Bordeaux, tome 26, ser. 3; tome 6, pp. 293-344. Hanna, G D. 1966. Introduced mollusks of Western North America. Occasional Papers of the California Academy of Sciences, no. 48, pp. 1-108, figs. 1-85, pls. 1-4. 74 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Henry, Dora P. 1942. Studies on the sessile Cirripedia of the Pacific coast of North America. Univer- sity of Washington Publications in Oceanography, vol. 4, no. 3, pp. 95-134, pls. 14, figs. 1-5. 1958. Intertidal barnacles of Bermuda. Journal of Marine Research, vol. 17, pp. 215-234, pls. 1-9. HERTLEIN, L. G. 1951. Invertebrate fossils and fossil localities in the San Francisco Bay Area. In Geologic guidebook of the San Francisco Bay Counties. State of California Division of Mines, Bulletin 154, pp. 187-192, figs. 1-2. Martsu1, T., G. SHANE, AND W. NEWMAN 1964. On Balanus eburneus Gould (Cirripedia, Thoracica) in Hawaii. Crustaceana, vol. 7, pt. 2, pp. 141-145, text fig. 1. MonammMap, Murap-B. M. 1962. Larval distribution of three species of Balanomorpha in relation to some chemico-physical factors. Proceedings of the First National Coastal and Shallow Water Research Conference, October, 1961, pp. 360-361. NEwMan, W. A. 1967. On physiology and behaviour of estuarine barnacles. Proceedings of Symposium on Crustacea, Marine Biological Association of India, pt. 3, pp. 1038-1066, figs. 1-9. Piussry, H. A. 1907. Hawaiian Cirripedia. Bulletin of the Bureau of Fisheries, vol. 26, pp. 179-190, pls. 4—S. ProxopovicyH, N. P. 1968. Organic life in the Delta-Mendota Canal, Central Valley Project-California. United States Department of the Interior, Bureau of Reclamation, Region 2, Sacramento, California, x + 126 pp., 5 appendices. Ross, A. 1962. Results of the Puritan-American Museum of Natural History Expedition to Western Mexico, 15. The littoral balanomorph Cirripedia. American Museum Novitates, no. 2084, pp. 1-44, figs. 1-24. SHATOURY, H. H. 1958. A freshwater mutant of Balanus amphitrite. Nature, vol. 181, no. 4611, pp. 790-791, fig. 1. UtTiInomiI, H. 1966. Recent immigration of two foreign barnacles into Japanese waters. Proceedings of the Japanese Society of Systematic Zoology, no. 2, pp. 36-39, text figs. 1-2. WELTNER, W. 1897. Verzeichnis der bisher beschriebenen recenten Cirripedienarten. Archiv fir Naturgeschichte, Jahrgang 1897, Band 1, pp. 227-280. ZWELEO IN ee Ne 1961. A new subgenus and species of coral-inhabiting barnacle from the Gulf of California. The Veliger, vol. 4, no. 2, pp. 71-75, pl. 17, text figs. 1-2. 1967. On the identity of some specimens assigned by Darwin, 1854, to Balanus allium Darwin (Cirripedia, Thoracica). Crustaceana, vol. 13, pt. 1, pp. 126-128. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XX XIX, No. 7, pp. 75-86, 4 figs; 3 tables. September 5, 1972 A KEY, BASED ON SCALES, TO THE FAMILIES OF NATIVE CALIFORNIA FRESHWATER FISHES By Richard W. Casteel Department of Anthropology, University of California, Davis 95616 Asstract: Much interdisciplinary interest has been shown with regard to fish scales in addition to their use in fisheries biology. To aid future workers, a key to the scales of the native California freshwater fish families is presented along with photo- micrographs of scales from each group. INTRODUCTION Fish scales have been used in fisheries biology and systematic ichthyology for many years. Within fisheries studies, emphasis has been placed upon the use of scales in age and growth studies (Cable, 1956; Cating, 1954; Chugunova, 1959; Cooper, 1951, 1952; Fry, 1943; Hile, 1936; Hogman, 1970; Jensen and Wise, 1961; Miller, 1955; Phillips, 1948; Rush, 1952; Schuck, 1949; Taylor, 1916; Whitney and Carlander, 1956; Meehean, 1935). Various keys, based upon the morphology of scales, have been published dealing with species identification within families and with the identification of families comprising regional fish faunas (Batts, 1964; Lagler, 1947; Koo, 1962). Fish scales have been used in palaeontological work (David, 1944, 1946a, 1946b), sediment analysis (Lagler and Vallentyne, 1956; Pennington and Frost, 1961; Soutar and Isaacs, 1969), and archaeology (Follett, 1967a, 1967b; Hubbs and Miller, 1948). Even within fisheries work, scales have been encountered during analysis of the stomach contents of various fishes (Greenfield, Ross, and Deckert, 1970; Kimsey, 1954). Based upon this evidently wide interdisciplinary interest in and use of fish scales, it is felt that a scale-based key to the families of native freshwater fishes of California would be a useful aid. [75] Marine Biological Laboratory LIBRARY SEP 15 1972 Woods Hole, Mass. 76 CALIFORNIA ACADEMY OF SCIENCES TABLE 1. Species examined. [Proc. 4TH SER. Species Source Thaleichthys pacificus Oncorhynchus tshawytscha O. kisutch Salmo gairdnerii S. g. gairdnerii S. g. stonei S. g. gilberti S. g. aquilarum S. clarkii clarkii S. c. henshawi S. c. seleniris S. aguabonita aguabonita S. a. whitei Salvelinus malma parkei Prosopium williamsoni Xyrauchen texanus Catostomus luxatus . platyrhynchus . Santaanae . rimiculus . latipinnis . occidentalis occidentalis . 0. humboldtianus . mniotiltus . tahoensis Rhinichthys osculus klamathensis Mylopharodon conocephalus Orthodon microlepidotus Pogonichthys macrolepidotus Lavinia exilicauda exilicauda L. e. harengus Ptychocheilus grandis PD WY Py OS) B} &) Hesperoleucas symmetricus symmetricus H. s. subditus H. s. venustus H. navarroensis — H.. parvipinnis ~ Gila bicolor bicolor ., -G~.b>, obesa psy GeO. pectinifera G. mohavensis >» G. crassicauda G. orcutti : CAS* VED CAS, UCD UCD UCD UCD CAS CAS CAS CAS CAS CAS CAS CAS CAS CAS CAS CAS CAS CAS CAS UCD CAS CAS CAS CAS CAS, UCD UCD UCD UCD CAS UCD UCD CAS CAS, UCD CAS CAS CAS CAS, UCD CAS CAS CAS CAS 1 CAS = specimens from California Academy of Sciences, San Francisco. 2UCD = specimens from the author’s personal collection, presently at University of California, Davis. VoL. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 77 TABLE 1. (continued) Species Source G. elegans CAS Richardsonius balteatus egregius CAS Cyprinodon macularius californiensis CAS C. nevadensis nevadensis CAS C. n. calidae CAS C. n. shoshone CAS C. salinus CAS Fundulus parvipinnis CAS Mugil cephalis CAS Archoplites interruptus UCD Hysterocarpus traskii CASTUCED For the purposes of this paper, native freshwater fishes will refer to those fishes which occur exclusively in freshwater or spend a significant portion of their life-cycle in freshwater and which occurred in California prior to the known introduction of exotic species during and after the nineteenth century (Kimsey and Fisk, 1960; Shapovalov, Dill, and Cordone, 1959; Walford, 1931). The only exception has been the inclusion of the Mugilidae because of their impor- tance in the Colorado River. METHODS AND MATERIALS The scales studied come from 54 species of native fishes (table 1) and represent specimens collected by the author or by members of the California Department of Fish and Game, and specimens from the California Academy of TABLE 2. Scale sampling locations. . Row anterior to dorsal fin. . Right side, below dorsal fin, above lateral line. . Left side, below dorsal fin, above lateral line. . Right side, below dorsal fin, below lateral line. Marine Biological Laboratory LIBRARY SEP 15 1972 Woods Hole, Mass. . Left side, below dorsal fin, below lateral line. . Right side, caudal penduncle, above lateral line. . Left side, caudal penduncle, above lateral line. ncluifoptel= Iles ipaleoiel i acial = . Right side, caudal penduncle, below lateral line. I. Left side, caudal penduncle, below lateral line. J. Row posterior to dorsal fin. 78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TaBLeE 3. Scale characteristics of California freshwater fish families. Family Ctenoid Scales Cycloid Scales Scutes Neither Petromyzonidae xX Acipenseridae x Osmeridae Salmonidae Catostomidae me mM Cyprinidae Gasterosteidae x Cyprinodontidae x x? Mugilidae Centrarchidae x Embiotocidae xe Cottidae x a 3 Scales of these families, while being cycloid, should be oriented as shown for ctenoid scales in figure 1 b. Sciences, San Francisco. Table III indicates the general scale characteristics of the fishes in this study. The fish were sampled for scales from ten different body locations (table 2) on the author’s specimens. For reasons of future use, the specimens from the California Acadamy of Sciences could only be sampled from six locations on the right-hand side of the fish. Wherever possible, samples were taken from several individuals of different sizes within a species in order to allow for ontogenetic variations. All scale samples were mounted in glycerine jelly on microscope slides (Weesner, 1960) and examined under a dissecting microscope at between ten and thirty magnifications. DEFINITIONS The terms used here to describe the surface features of scales are taken from Lagler (1947, pp. 150-151) and are illustrated in figure 1. Circuli — “Elevated markings on the outer surface; usually appearing as lines which more or less follow the outline shape of the scale.” Focus — “First part of scale to appear in growth; often central.” Radi — “Grooves, usually more or less radiating from focus to one or more margins.” Primary Radu — “Radii that extend from focus to margin.” Secondary Radi — “Radii that begin outward from, not at, focus.” Ctenu — “Tooth-like structures on posterior portion of scale.” Vor. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 79 CIRCULI LATERAL FIELD PRIMARY RADII SD ae Dy iy POSTERIOR ZB FIELD Zz ANTERIOR FIELD SECONDARY RADII LATERAL FIELD SECONDARY RADII Ay Bie S _/cTENII oe Die } ve a = ANTERIOR K nt Lae WEN 3 eS = FIELD am MEE gow Soe “Ss FOCUS aD ; “iy ae ae Ge aS De Dw p me Zee —- Soe bs SS POSTERIOR PRIMARY wy SIS ae RADII i WSS. z : ; - ma : —— Se LATERAL CIRCULI FIELD Figure 1. Top. Cycloid Scale. Cyprinidae. Mvylopharodon conocephalus. UCD 5040 C. Bottom. Ctenoid Scale. Centrarchidae. Archoplites interruptus. UCD 5021 E. 80 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. yi wy, FicureE 2. a. Cyprinodontidae. Cyprinodon macularius californiensis. UCD 5063 G. b. Centrarchidae. Archoplites interruptus. UCD 5021 E. c. Mugilidae. Mugil cephalus. UCD 5056 I. Fields — ‘Areas of the outer surface of the scale, either real as delimited by angulation of the ridges (circuli) at levels of the four principal corners or imaginary if the corners or configuration of the circuli are wanting. Adjectives of direction applied to fields are based on their positions when the scales are normally situated on the side of the fish.” Anterior Field — “Bounded by imaginary lines connecting the anterolateral corners, or their equivalent points on scales which are rounded (dorsal and ventral) with the focus.” Posterior Field — “Bounded by imaginary lines connecting the posterolateral corners (dorsal and ventral) with the focus.” Lateral Fields — “Dorsal and ventral fields remaining after delimitation of anterior and posterior ones.” Vou. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 81 Ficure 3. a. Salmonidae, Salmo gairdnerii gairdnerii. UCD 5002 D. b. Salmonidae. Prosopium williamsoni. UCD 5016 C. c. Catostomidae. Catostomus occidentalis occidentalis. UCD 5026 C. SCALE-BASED KEY TO THE FAMILIES OF CALIFORNIA FRESHWATER FISHES The following is a key to the family level with the exception of Prosopium williamsoni which is identified to species. The Cyprinodontidae are characterized by both ctenoid and cycloid scales in the same individual. Lagler (1947, pp. 156-157) classified the Cyprinodontidae of the Great Lakes as having cycloid scales and the same appears true of the genus Fundulus in California. However, the other members of this family may also possess ctenoid scales (Lagler, Bardach, and Miller, 1962, p. 114). For this reason the key identifies this family twice, once on the basis of ctenoid scales and again on the basis of cycloid scales. This same case appears true of the Centrar- chidae (Lagler, Bardach, and Miller, 1962, p. 114). Based upon data from my collections, however, I find cycloid scales to occur only once in 32 samples and 82 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 4. a. Cyprinidae. Mylopharodon conocephalus. UCD 5040 C. b. Cyprinodon- tidae. Cyprinodon macularius californiensis. UCD 5063 A. c. Embiotocidae. Hysterocarpus traskiz. UCD 5099 C. then only from restricted areas of the body in centrarchids (table 2, A). This is a rather low frequency and if one considers the total number of scales over a fish’s body, it appears that occurrence of cycloid scales in Archoplites interruptus will be even more rare. I agree, therefore, that ‘These fishes may still be considered as predominantly ctenoid in their squamation but the degree and extent of development of the ctenii varies from place to place on the body” (Lagler, Bardach, and Miller, 1962, p. 114). In the rare event that a cycloid scale from this family should present itself in isolation it will key out as representing the Embiotocidae instead of the Centrarchidae. Figures 2, 3, and 4 illustrate each of the families or species separated by the key. Each illustration is oriented with the anterior field to the observer’s left. iva) Ctentiy present omisposterion) eile cl ee ee) b)! (Ctenit “absent on ‘posterior field == ee eee 2. a) Ctenii numerous and evenly spaced VoL. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 83 b) Ctenii not numerous and irregularly spaced — eee Cyprinodontidae (figure 2, a) 3. a) Radii converge toward focus ae Se ee RR PowresNGAaD Centrarchidae (specifically Archoplites interruptus ; figure 2, b) I eacimerouchiveunarallel) <2. 2 ee ee Mugilidae (specifically Mugil cephalus; figure 2, c) Pe mSCHeMmCVClOllmwithouts Tag) 2 a Be 5 Peal CmGy Cloldenwiliwna Cie sso. 88 ee ee ee 6 5. a) Scale with anterior and posterior fields only or without fields coe neipene tee aE cee eee Osmeridae and Salmonidae (figure 3, a) b) Scale with four fields; focus centrally located Prosopium williamsoni (figure 3, b) 6. a) Primary radii on both posterior and anterior fields = Catostomidae (also includes the cyprinid genera Rhinichthys and Orthodon; figure 3, c) b) Primary radii absent on either anterior or posterior field = 7 7. a) Primary radii present on anterior field, but absent on posterior field — == 8 b) Primary radii absent on anterior field, but present on posterior field = = 3 nc pec ME cade oo RS DA PE 8 Cyprinidae (figure 4, a) Sa ewartewes circuly im lateral than in anterior field —22-- = Cyprinodontidae (figure 4, b) b) Number of circuli in lateral field approximately equal to number in anterior TT en eS a a Embiotocidae (specifically Hysterocarpus traskiz; figure 4, c) ACKNOWLEDGMENTS I wish to express my thanks to Mr. W. I. Follett and Mrs. L. Dempster of the California Academy of Sciences for their advice and cooperation and to Dr. W. G. Kinzey, City University of New York, for his support and encouragement during the initial phases of this work. My thanks also to Dr. R. W. Brocksen and Messrs. L. Courtois, W. Wurtsbaugh, and H. W. Li, Department of Animal Physiology, University of California, Davis; to Mr. Lyons, U. S. Department of Reclamation; and to Messrs. A. Calhoun, L. Fisk, E. Armstrong, M. Coots, E. P. Pister, R. Reavis, and J. Burns, California Department of Fish and Game, for their aid in obtaining specimens. I also appreciate the helpful criticisms of the manuscript given by Mr. W. I. Follett, California Academy of Sciences; Dr. D. L. True, Department of Anthropology, University of California, Davis; and Dr. J. D. Hopkirk, Department of Biology, Sonoma State College. Finally, my thanks go to my wife for her help during all phases of this project. LITERATURE CITED Batts, Bitty S. 1964. Lepidology of the adult pleuronectiform fishes of Puget Sound, Washington. Copeia, 1964, pp. 666-672. 84 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Caste, L. E. 1956. Validity of age determination from scales, and growth of marked Lake Michigan lake trout. U. S. Fish and Wildlife Service, Fish Bulletin, no. 107, pp. 1-59. CatInG; J: P- 1954. Determining age of Atlantic shad from their scales. U. S. Fish and Wildlife Service, Fish Bulletin, no. 85, pp. 187-199. Cuucunova, N. I. 1959. Age and growth studies in fish: a systematic guide for ichthyologists. Academy of Sciences of the U.S. S. R., 132 pp. Cooper, E. L. 1951. Validation of the use of scales of brook trout, Salvelinus fontinalis, for age determination. Copeia, 1951, pp. 141-148. 1952. Body-scale relationships of brook trout, Salvelinus fontinalis, in Michigan. Copeia, 1952, pp. 1+. Davin, Lore R. 1944. Use of fossil fish scales in micropalaeontology. Carnegie Institution of Washington Publication, no. 551, pp. 25-43. 1946a. Some typical Upper Eogene fish scales from California. Carnegie Institution of Washington Publication, no. 551, pp. 45-79. 1946b. Upper Cretaceous fish remains from the western border of the San Joaquin valley, California. Carnegie Institution of Washington Publication, no. 551, pp. 81-112. Fottett, W. I. 1967a. Fish remains from coprolites and midden deposits at Lovelock Cave, Churchill County, Nevada. Annual Report of the University of California Archaeological Survey, no. 70, pp. 94-115. 1967b. Fish remains from Salinas La Blanca, an archaeological site on the Pacific coast of Guatemala. Smithsonian Contributions to Anthropology, vol. 3, pp. 129-134. Fry, F. E. J. 1943. A method for the calculation of the growth of fishes from scale measurements. Ontario Fisheries Research Laboratory, Publication no. 61, pp. 5-18. GREENFIELD, D. W., S. T. Ross, and G. D. DECKERT 1970. Some aspects of the life history of the Santa Ana Sucker, Catostomus (Pantosteus) santaanae (Snyder). California Fish and Game, vol. 56, no. 3, pp. 166-179. Hirer, R. 1936. Age and growth of the cisco, Leucichthys artedi (Le Sueur), in the lakes of the northwestern highlands, Wisconsin. U. S. Fish and Wildlife Service, Fish Bulletin, no. 19, pp. 211-317. Hocman, W. J. 1970. Early scale development on the Great Lakes coregonids, Coregonus artedii and C. kiyiz. In C. C. Lindsey and C. S. Woods (editors), Biology of Coregonid Fishes. Winnipeg, pp. 429-435. Hupss, C. L., and R. R. Mier 1948. Correlation between fish distribution and hydrographic history in the desert basins of western United States. Jn The Great Basin, with emphasis on glacial and postglacial times. Bulletin of the University of Utah, vol. 38, no. 20, pp. 17-166. VoL. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 85 Jensen, A. C., and J. P. WIsE 1961. Determining age of young haddock from their scales. U. S. Fish and Wildlife Service, Fish Bulletin, vol. 61, pp. 439-450. Kinsey, J. B. 1954. The life history of the tui chub, Siphateles bicolor (Girard), from Eagle Lake, California. California Fish and Game, vol. 40, no. 4, pp. 395-410. Kinsey, J. B., and L. O. Fisk 1960. Keys to the freshwater and anadromous fishes of California. California Fish and Game, vol. 46, no. 4, pp. 453-479. Koo, T.S. Y. 1962. Differential scale characters among species of Pacific salmon. Jn T. S. Y. Koo (editor), Studies of Alaska Red Salmon. Seattle, pp. 123-135. LAGLeR, K. F. 1947. Scale characters of the families of Great Lakes fishes. American Microscopical Society Transactions, vol. 66, no. 2, pp. 149-171. LActer, K. F., and O. R. VALLENTYNE 1956. Fish scales in a sediment core from Linsley Pond, Connecticut. Science, vol. 124, no. 3217, p. 368. LAGLER, K. F., J. E. BArpAcH, and R. R. MILLER 1962. Ichthyology. New York, 545 pp. MEEHEAN, O. L. 1935. The life history of the bluegill sunfish [Helioperca incisor (Cuvier and Valenci- ennes) ] as determined from the scales. Proceedings of the Los Angeles Academy of Science, vol. 2, pp. 139-145. Muter, D. J. 1955. Studies relating to the validity of the scale method for age determination of the northern anchovy (Engraulis mordax). California Division of Fish and Game, Fish Bulletin, no. 101, pp. 6-34. PENNINGTON, W., and W. E. Frost 1961. Fish vertebrae and scales in a sediment core from Estwaite Water (English Lake District). Hydrobiologia, vol. 17, pp. 183-190. Purtrps, J. B. 1948. Comparison of calculated fish lengths based on scales from different body areas of the sardine, Sardinops caerulea. Copeia, 1948, pp. 99-106. Rus, W. A. 1952. Observations of age and growth in the menhaden (Brevoortia tyrannus) as determined by scale examination. Copeia, 1952, pp. 208-209. ScHuck, H. A. 1949. Problems in calculating size of fish at various ages from proportional measurements of fish and scale sizes. Journal of Wildlife Management, vol. 13, no. 3, pp. 298-303. SHAPOVALOV,L., W. A. Dirt, and A. J. CorDONE 1959. A revised checklist of the freshwater and anadromous fishes of California. California Fish and Game, vol. 45, no. 3, pp. 159-180. SouTar, A., and J. D. Isaacs 1969. History of fish populations inferred from fish scales in anaerobic sediments off California. California Cooperative Oceanic Fisheries Investigations, Reports, vol. 13, pp. 63-70. TAYLor, H. F. 1916. The structure and growth of the scales of the squeteague and the pigfish as 86 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. indicative of life history. U. S. Bureau of Fisheries, Bulletin, no. 34, pp. 285-330. WALForpD, L. A. Ny 1931. Handbook of common commercial and game fishes of California. California a \ Division of Fish and Game, Fish Bulletin, no. 28. 3°: WEESNER, F. M. 1960. General zoological microtechniques. Baltimore. Wuirtney, R. R., and K. D. CARLANDER 1956. Interpretation of body-scale regression for computing body length of fish. Journal of Wildlife Management, vol. 20, pp. 21-27. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXIX, No. 8, pp. 87-103. September 5, 1972 THE LYGAEIDAE OF THE GALAPAGOS ISLANDS (Hemiptera:Heteroptera) By Peter D. Ashlock University of Kansas, Lawrence With the publication of this paper, reports on the Heteroptera of the Gala- pagos International Scientific Project are nearly complete. Published papers cover the Anthocoridae (Herring, 1966), the Tingidae (Drake and Froeschner, 1967), the Saldidae and Veliidae (Polhemus, 1968a, b), the Cydnidae ( Froeschner, 1968), the Nabidae (Kerzhner, 1968), the Miridae (Carvalho and Gagné, 1968), and the emesine Reduviidae (Villiers, 1970). The families Pentatomidae, Corei- dae, Rhopalidae, Stenocephalidae, Berytidae, Pyrrhocoridae, Reduviidae, Gerri- dae, and Corixidae also are found in the Galapagos Islands, and members of the expedition collected all but Pyrrhocoridae and Stenocephalidae. Hopefully, the many new island records in these families will be summarized in another contri- bution. The collections of only the Rhopalidae may still prove to contain new species. Before the expedition, Dr. Usinger and I had hopes of greatly increasing the number of Lygaeidae recorded from the islands, especially in the subfamily Orsillinae, a group well known for its ability to colonize islands. Only four species of Lygaeidae, including two orsillines, had previously been recorded from the Galapagos (Linsley and Usinger, 1966). Herein I am able to report only nine species of lygaeids—five are orsillines—of which two, including one of the orsillines, are possibly introduced by man. While more lygaeids may be found in the archipelago, I seriously doubt that the known endemic fauna of the group will double again. Because the lygaeid fauna of Hawaii contains forty percent of the world orsillines, the sparse fauna of the Galapagos deserves some explana- tion. [87] Marine Biological Laboratory LIBRARY SEP 15 1972 Woods Hole, Mass. 88 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser. MacArthur and Wilson (1967) discuss the factors that determine the intro- duction and extinction rates of island faunas as they develop the concept of an island colonization rate. Further, they postulate that each island or archipelago has a fairly fixed capacity for numbers of species. Among islands populated from American sources, the Galapagos have a far better chance of receiving introductions than do most other islands of the eastern Pacific, primarily because they form an archipelago. The Hawaiian Islands, a larger archipelago, would have received fewer introductions because of its great isolation. Some examples will illustrate the point. The smallest orsilline faunas are those of Guadelupe Island, 180 miles off Baja California, and San Felix, 550 miles west of Chile, each having a single undescribed endemic species (belonging to Ortholomus and Nvysius, respectively). The Juan Fernandez Islands, two islands about 450 miles off the coast of Chile, have two endemic orsillines, V ysius baeckstroemi and Robinsonocoris tingitoides, the latter an endemic genus. The Galapagos Islands have five species, resulting from four introductions, including the endemic genus Darwinysius with two species. The Hawaiian fauna by contrast is rich. Of the 92 species of Orsillinae, the tribe Metrargini | probably of American origin (Ashlock, 1967) | contains 58 species (in six endemic genera) resulting from but two introductions. The remaining Hawaiian species (in Nysius and the endemic Nesomartis, a probable derivative of Nysius) are of unknown origin. Clearly, the Galapagos have a much higher introduction rate than Hawaii, but the larger, more hospitable Hawaiian Islands have a lower extinction rate than the dryer and smaller Galapagos. Thus, possible ecological situations on the Galapagos are nearly saturated by naturally introduced orsillines. Hawaii’s isolation has allowed few intro- ductions, but its low extinction rate, combined with the large number of ecological situations, permitted explosive radiation. Linsley and Usinger (1966) acknowledge those that made the G.I.S.P. expe- dition possible. I must add my own thanks to the late R. L. Usinger, who made my participation possible. Much of the work on this paper was done at the B. P. Bishop Museum, Honolulu, with financial help from two NSF grants (GB-3105 and GB—5860). Types of new species are deposited in the California Academy of Sciences, San Francisco, and specimens upon which the study was based will be found in that institution and in the Bishop Museum; the U. S. National Museum; the California Insect Survey, Berkeley; and the collections of J. A. Slater, G. G. E. Scudder, M. H. Sweet, and myself. All measurements are in millimeters, and the year of collection is 1964 unless otherwise noted. Kry To GALAPAGOS GENERA OF LYGAEIDAE 1. Suture between abdominal segments IV and V (second visible suture) curving anteriorly, not reaching lateral margin of abdomen —____ Rhyparochrominae: Myodochini — 2 Suture between segments IV and V not curving anteriorly, clearly reaching lateral margin Olgabdom ery es eee jee 3 VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 89 2. Head constricted behind eye; eye removed from base of head by a distance greater than NEI EOBITMeEO Teme UTM CV CMe eee eee ee ee Heraeus Head not constricted behind eye; eye removed from base of head by a distance less than [enc meO lame GE Cy Camere st Le ft. eee ee ee Pachybrachius 3. Scutellum bifid apically; clavus basally opaque, apically hyaline —— Cyminae: Af 111 neem me pt ee Cymoninus Scutellum not bifid apically; clavus not divided into distinct opaque and hyaline areas __ i @rsillinae yete 2 as a A 4. Costal margin of corium straight, exposing connexivum of abdomen laterally _ ai rr @rsillint 222 2 a ee ee Ori Volomus Costal margin of corium straight at most for a distance less than length of scutellum, then arcuately curved to apex; connexivum of abdomen completely covered — 5 5. Buccula not punctate; lateral margin of abdomen and corium without cross-striated Giei@haliitieg, 2 ee ee INST 22-2 eee ee ee Nysius Either buccula punctate or lateral margin of abdomen and corium with cross-striated SUINIGIUIT ic, Se Metrarr gin .2 222 i ate eee 6 6. Buccula punctate, not tapering, ending abruptly at base of head; antenniferous tubercle acutely produced; abdomen and corium without stridulatory structures —— oper ee rrr rr a ee aS ee Darwinysius Buccula impunctate, tapering to a low carina well before base of head; antenniferous tubercle not acutely produced; abdomen and corium laterally with cross-striated stridu- PROVES ENUGEU LC meee eeaeeee ae Se et ee eee Xyonysius Genus Nysius Dallas Nysius DALLAS, 1852, p. 551. Although two species from the Galapagos were originally described in the cosmopolitan genus \Vysius, both have since been placed in other orsilline genera in the tribe Metrargini. Nysius (Ortholomus) naso Van Duzee is placed in Xyonysius (Ashlock and Lattin, 1963) and Nysius (?) marginalis Dallas is placed in Darwinysius (Ashlock, 1967). The following is the first true Nysius to be reported from the Islands. Nysius usitatus Ashlock, new species. (Figures 1, 3.) Head nearly flat between eyes, densely punctate, sparsely clothed with appressed pale hairs, length 0.66, width 0.88, anteocular length 0.27, eye length 0.29, eye width 0.19, interocular space 0.48; buccula widest anteriorly, tapering posteriorly, and ending abruptly just before base of head; labium extending to between hind coxae, first segment not reaching base of head but just exceeding buccula, segment lengths from base 0.39, 0.39, 0.37, 0.29; antenna with first segment exceeding clypeus by nearly half its length, segment lengths from base 0.26, 0.60, 0.49, 0.49. Pronotum moderately clothed with fine curved subappressed hairs, disk densely punctate, distance between punctures from one-half to one diameter of a puncture, sides nearly straight; length 0.59, width 1.02. Scutellum arine Biological Laboratory LIBRARY SEP 15 1972 Woods Hole, Mass. 90 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. FicureE 1. Dorsal views of (left) Nysius usitatus and (right) Ortholomus usingeri. with vestiture and punctation similar to those of pronotum; Y-shaped carina appearing as a tumid pyramid; length 0.48, width 0.58. Hemelytron exceeding abdomen, clavus and corium with surface almost dull, moderately clothed with subappressed curved hairs, with additional hairs projecting laterally along base of costal margin, clavus obscurely punctate only at base of claval suture, costal margins subparallel and paralleling vein R+M to level of apical two-thirds of scutellum, then gently becoming divergent and arcuate to apex of corium, veins evident but not prominent; length of claval commissure 0.34; length of corium 1.50; membrane normal with veins evident but not distinct, basal length to level of corial apex 0.65, apical length from corial apex 0.77. Color. Light reddish brown, with broad stripe passing through each ocellus, margins of clypeus, three spots at base of clypeus, ventor except buccula, and spot below eye black. Buccula, labium, and antenna yellowish brown, first segment of antenna a little darker. Pronotum light yellowish brown, VoL. XXXIX | ASHLOCK: GALAPAGOS LYGAEIDAE 91 with partial indistinct median stripe, broad complete stripe behind each ocellus, and narrow lateral stripe dark brown, callosities black. Scutellum black or nearly so, apex white. Hemelytra light yellowish brown with indistinct brown spots on veins and apical margin. Membrane hyaline, embrowned medially except veins obscurely opaque white, ventor black except anterior and posterior margins of propleura, acetabula, scent gland area, metapleural plate, and spots laterally and apically on abdomen pale. Legs pale yellowish brown, with usual brown spots on femur, tibia with a brown knee band. Hototyre. 6, Santa Cruz Island, 1.5 mi. north of Academy Bay, 13 Febru- ary, under Portulaca (P. D. Ashlock). PaRATYPES. Santa Cruz Island, 9 ¢,10 2, same data as holotype; 1 ?, same data but 11 February; 32 ¢,17 2, same data but 15 February; 3 ¢,5 2, same data but 25 February; 3 2, Horneman Farm, 220 m., 2 April (D. Q. Cavagnaro). Most specimens of this species were collected at the foot of the barranca (cliff) on the new trail from Academy Bay, Santa Cruz Island, under Portulaca and in company with Darwinysius marginalis. Since Nysius usitatus was collected only near Academy Bay on Santa Cruz and not at all from the Portulaca habitat on all other islands visited, it may be a recent introduction to the Galapagos. Specimens fit Dallas’ (1852, p. 553) description of NV. nubilis from Colombia fairly well, but the V. nubilis type is not extant. Dr. Usinger searched in vain for it at the British Museum in 1964, and found (corre- spondence) that the British Museum copy of the Dallas description has a note: ) “type missing.” I have a short series of Nysius from Guayaquil, Ecuador, and another from Palmira, Colombia, that also fit Dallas’ description, but the spermathecae differ substantially from that of NM. usitatus (fig. 3). Since there is only slight evidence that VV. usitatus is introduced, and its identity with N. nubilis is open to question, I have described the Galapagos form as new. The problem of the identity of VV. nubilis can best be solved by the judicious choice of a neotype when adequate series of NV ysius are available from northwestern South America. Genus Ortholomus Stal Nysius (Ortholomus) STAL, 1872, p. 43. Ortholomus Stal, BAKER, 1906, p. 134. Ortholomus usingeri Ashlock, new species. (Figures 1, 3.) Head flattened between eyes, obscurely punctate, densely covered with flattened appressed hairs, vertex carina straight, eye prominent, slightly raised above vertex; length 0.85, width 1.11, anteocular length 0.44, eye length 0.22, eye width 0.29, interocular space 0.61; buccula widest anteriorly, 92 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 2. Dorsal views of (left) Darwinysius wenmanensis and (right) Xyonysius naso. gradually narrowing posteriorly to mid eye level without abrupt change in width; labium reaching between posterior coxae, first segment exceeding buccula, not reaching base of head, segment lengths from base 0.46, 0.43, 0.46, 0.39; antenna with fine erect pubescence, first segment slightly exceeding clypeus, segment lengths from base 0.29, 0.46, 0.44, 0.49. Pronotum clothed with appressed flattened somewhat silky hairs, with occasional short erect hairs, moderately punctate, distance between punctures from one-half to one diameter of a puncture, sides straight, slightly swollen, length 0.78, width 1.28. Scutellum with vestiture and punctation like those of pronotum, Y-shaped carina swollen on upper arms, stem not prominent, length 0.54, width 0.75. Hemelytron slightly exceeding abdomen, clavus and corium moderately clothed with short silky appressed hairs, with a few short erect hairs projecting VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 93 laterally along base of costal margin; veins evident but not prominent; length of claval commissure 0.49; length of corium 1.80; membrane irregularly wrinkled transversely; veins distinct, basal length to level of corial apex 0.77, apical length from corial apex 0.78. Abdomen with connexival spiracles raised, very prominent. Color. Head very dark brown, clypeus, median stripe on posterior half of vertex, buccula, labium, and antenna paler except antennal segment IV very dark brown. Pronotum and scutellum reddish brown, callosities and punctures a little darker. Hemelytron pale yellowish brown, darkened apically on clavus and corium and on obscure spots on veins; vestiture nearly white, silky. Connexival segments pale, darker around pale spiracles. Legs yellowish brown, femur with obscure, slightly darker spots. Ventor reddish brown, acetabula and scent gland paler. Hototype. ¢, Santa Cruz Island, grassland, 1,800 ft., north of Academy Bay, 20 February, on Hypericum pratense (P. D. Ashlock). PARATYPES. Santa Cruz Island, 6 ¢, 12 2,4 nymphs, same data as holotype; feo, ) 2, same data but 2,100 it.; 1 ¢, 1 9%, Bella Vista, 26 February (R. L. Usinger); 15 ¢, 15 2, Table Mountain, 440 m., 16 April (D. Q. Cavagnero). Floriana Island, 1 ¢, 1 2, 15 February, Verbena (R. L. Usinger); 57 6, 48 2, Whitmer’s Farm, 15 February, Verbena (R. L. Usinger); 1 ¢,2 2, same data but Cordia species; 1 ¢, 1 2, moist forest 200 m. above Black Beach, 15 February, Cordia tree with yellow flowers (R. L. Usinger); 7 ¢, 10 2, 18 February (R. L. Usinger). San Cristobal Island, 3 ¢,4 2, Progresso, 23 February, Verbena (R. L. Usinger). Ortholomus usingeri is less elongate than most of the described species in the genus, and the connexivum is more broadly exposed, being approached only by O. gibbus (Berg) in this respect. The spiracles are more prominent and enlarged than in any other species. All series of the species are variable in color, ranging from the pale form illustrated (fig. 1), to specimens that are highly maculated on the corium, clavus, and membrane. Some specimens have a very dark, contrasting claval apex. None of these variations are correlated with island of origin or with host plant. The new species of Ortholomus from Guadelupe Island (see Introduction) may be very closely related to O. usingeri. It is with devoted appreciation that I dedicate this Galapagos orsilline to R. L. Usinger, who was the leading authority in the Orsillinae, and whose leadership of the G. I. S. P. expedition he considered the high point of his life. Genus Darwinysius Ashlock Darwinysius ASHLOCK, 1967, p. 42. Darwinysius is the only endemic genus of Lygaeidae—and one of the very few in the Heteroptera—found in the Galapagos. It is closely related to 94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. D. wenmanensis N. usitatus O. usingeri X. naso C. notabilis Ficure 3. Spermathecae of the named species. Robinsonocoris in the Juan Fernandez Islands, differing principally in features correlated with the flightlessness of Robinsonocoris. At the time of the original description of Darwinysius for Nysius (2?) marginalis Dallas, it was mentioned that a second species had been found on the Galapagos. Key TO GALAPAGOS SPECIES OF DARWINYSIUS Larger species, male more than 3.2 long, female more than 3.8 long; clothed above with short, appressed hairs; vertex of head raised above level of top of eye; Wolf Island — Bren oN Ele Te eek D. wenmanensis Ashlock, new species VoL, XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 95 Smaller species, male less than 3.1 long, female less than 3.5 long; clothed above with mixed erect and appressed hairs; vertex of head not raised above level of top of eye; main islands __ _____... D. marginalis (Dallas) Darwinysius marginalis (Dallas). Nysius (2?) marginalis DALLAS, 1852, p. 556. Cymus galapagensis STAL, 1959, p. 252 (synonymy by Butler, 1877). Darwinysius marginalis (Dallas), ASHLOCK, 1967, p. 42. In the summer of 1964, Dr. R. L. Usinger checked the types of Cymus galapagensis Stal and Nysius (?) marginalis Dallas at my request. In Stockholm he confirmed Butler’s (1877) synonymy of Stal’s species with that of Dallas, but in London, he found Dallas’ series of eight specimens to contain two species: four of the eight belonged to Xyonysius naso (Van Duzee). Since Dallas did not select a holotype, and no lectotype had since been chosen, Usinger selected the best specimen of the remaining four as a lectotype. He wrote that the lectotype ‘Gs rather broad and well marked and has no antennae.’ The specimen is numbered 77—2 (as were the X. aso) and is from Charles Island (Floreana). The remaining specimens were from James Island (Santiago) and “were very poor and not suitable to be made a lectotype.” The Charles Island specimen (no. 77-2) is here formally selected as the lectotype of Nysius (?) marginalis Dallas. Published records of Darwinysius marginalis are from three islands: Floreana, Santiago, and Daphne Major. Members of the G. I. S. P. expedition collected well over 300 specimens and can add five islands to the list: Isabella, Fernan- dina, Rabida, Pinzon, and Santa Cruz. Almost all of my specimens were collected under Portulaca, where they were feeding on fallen seeds of the plant. Dr. Usinger collected 55 specimens on Euphorbia viminea at Black Beach, Floreana. Darwinysius marginalis is generally similar to D. wenmanensis; differen- tiating characters can be found in the key to species. A dorsal view of D. marginalis, a side view of the head, and drawings of the aedeagus and sperma- theca are given in Ashlock (1967). Darwinysius wenmanensis Ashlock, new species. (Figures 2, 3.) Head elevated above eye about one-third height of eye, sparsely clothed with flattened appressed hairs, length 0.71, width 0.80, anteocular length 0.46, eye length 0.20, eye width 0.15, interocular space 0.48, buccula widest anteriorly, narrow, but not tapering to base of head, slightly projecting past posterior point of attachment, not reaching base of head, densely punctate; labium just reaching posterior coxae, first segment not reaching posterior end of buccula, segment lengths from base 0.46, 0.43, 0.44, 0.32; antenna with flattened sub- appressed hairs on first segment, semierect fine hairs on fourth segment, second 96 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. and third segments intermediate, first segment exceeding clypeus by nearly half its length; segment lengths from base 0.39, 0.56, 0.48, 0.41. Pronotum with flattened appressed hairs, deeply and densely punctate, cica- trices depressed, area anterior to cicatrices somewhat inflated; length 0.78, width 1.19. Scutellum with vestiture and punctation similar to those of pronotum, with a strongly elevated Y-shaped carina, length 0.51, width 0.68. Hemelytron exceeding abdomen, clavus and corium moderately dotted with appressed hairs, without erect hairs at base of lateral margin, surface overall ridged, punctate on both sides of claval suture and outer edge of vein R+M for entire length, all veins prominent and elevated, length of claval commissure 0.54, length of corium 2.04, apex attaining abdominal segment VII; membrane with surface irregularly ridged transversely, veins prominent, basal length to level of corial apex 0.90, apical length from corial apex 0.54. Color. Head, labium, antenna very dark brown, apices of first to third antennal segments and lower edge of buccula pale. Pronotum, scutelum, and hemelytron mottled dark and yellowish brown; pronotum with a patch of white flattened hairs behind cicatrix, corium with lateral margin alternately light and dark, apical margin beyond intersection of vein Cu very dark brown, membrane including veins mottled brown and transparent. Ventor dark brown except acetabula, posterior lobe of propleura, mesothoracic scent gland, and lateral posterior angle of metapleuron pale, legs with femur very dark brown, apex pale, tibia and tarsi pale with dark band dorsally and subapically on tibia. HototyPe. 6, Wolf Island (Wenman). 31 January, under Portulaca (P. D. Ashlock). ParATyPeEs. Wolf Island, 15 6,15 2, same data as holotype; 1 4, 24 Septem- ber 1906 (F. X. Williams). Darwinysius wenmanensis is abundantly distinct from the type species of the genus as indicated by characters in the key. Both species are variable in color pattern, ranging from nearly immaculate on the clavus and corium to quite dark with few pale maculations, but in general D. wenmanensis tends to be darker. Such features of both species as the spotted costal margin, the acute antenniferous tubercles, the general body shape, and, in D. wenmanensis, the raised vertex of the head, make both look very much like miniature Nesoclimacias (Hawaiian Islands). This similarity was noted earlier (Ashlock, 1967, p. 35); however, I now think these similarities—of Robinsonocoris and Darwinysius with the Hawaiian Metrarga, Nesoclimacias, and Nesocryptias—represent parallelisms rather than a monophyletic origin. Specimens of D. wenmanensis from Wenman Island were the first members of the genus I collected, and it was on Wenman that the “under Portulaca” habitat of Darwinysius was discovered. Unfortunately, it was then too late to VoL. XXXIX |] ASHLOCK: GALAPAGOS LYGAEIDAE 97 search the Portulaca on Darwin Island (Culpepper), the most remote of the archipelago. Genus Xyonysius Ashlock and Lattin X yonysius ASWLOcCK and LATTIN, 1963, p. 702. Xyonysius naso (Van Duzee). (Figures 2, 3.) Nysius (Ortholomus) naso VAN DUZEE, 1933, p. 27. Ortholomus naso (Van Duzee), BARBER, 1934, p. 285. Nystus naso Van Duzee, USINGER, 1941, p. 131. Xyonystus naso (Van Duzee), ASHLOCK and LATTIN, 1963, p. 702. Xyonysius is a New World genus of metragine Orsillinae whose species are found from Canada to Argentina and Chile. The endemic Galapagos species X. naso is one of the more variable ones, ranging from forms with immaculate hemelytra to forms with three irregular dark brown longitudinal stripes on the corium, the middle one continuing onto the membrane, and dark brown claval apices. Figure 1 shows a specimen with intermediate coloration. This variation was found in all series of the species collected, and does not appear to be correlated with island or host plant. Unlike X. californicus (Stal), a wide-spread North American species found on many composites, X. naso seems to be confined to species of the endemic composite genus Scalesia. The species was described from a single specimen taken on Floreana (Charles Island). Dr. Usinger and I collected 59 specimens on the following islands and host plants: Santa Cruz (S. affinis, S. helleri), Fernandina (Scalesia species), Barrington (S. helleri), Isabella (S. affinis, S. gummifera), and Floreana (S. affinis). Nowhere was the species abundant. Genus Cymoninus Breddin Cymoninus BREDDIN, 1907, p. 38. Cymoninus notabilis (Distant). (Figures 3, 4.) Ninus notabilis D1isTaNT, 1882, p. 191. Cymoninus notabilis (Distant), VAN DUZEE, 1917, p. 163. This species is here reported from the Galapagos for the first time. Wide- spread, it ranges from the southern United States through Central America and the Antilles south to Argentina and Brazil. I collected 55 specimens on Santa Cruz Island on the trail north of Academy Bay and Bella Vista at about 1,300 feet on a sedge identified by Ira Wiggins as Cyperus confertus Swartz. D. Q. Cavagnaro also collected a few specimens on Santa Cruz at the Horneman Ranch, just north of Bella Vista. This species may be a recent introduction to the Galapagos. I collected two specimens in the harbor at Guayaquil, Ecuador, at light, on board ship. 98 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Genus Pachybrachius Hahn Pachybrachius HAHN, 1826, p. 18. Orthaea DALLAS, 1852, p. 580. Key To GALAPAGOS SPECIES OF PACHYBRACHIUS Smaller species, less than 4 long; usually brachypterous; corium with long erect hairs and an Oval whitejspot neaainner angle 2 ee P. nesovinctus Ashlock, new species Larger species, more than 4 long; never brachypterous, fully winged; corium with short appressed hairs and without an oval white spot near inner angle — Pachybrachius insularis (Barber). (Figure 5.) Orthaea insularis BARBER, 1925, p. 246. Pachybrachius insularis (Barber), SLATER, 1964, p. 1127. Barber’s Galapagos Pachybrachius has been recorded from Isabella, Santiago, Baltra, and Santa Cruz Islands. Members of the expedition collected the species from all of these but Isabella and Baltra, but also collected it on Fernandina and Floreana. Most of the material was collected from lights, though occasional specimens were collected from the forest floor. The two specimens from Floreana, both females (Kuschel and Usinger, collectors) were unusual, since both had the labium extending onto the abdomen. Barber (1925) described the labium as “reaching just past intermediate coxae.’” However, labial length in a 36-specimen sample varied as follows: past front but not to mid coxae, 5 4, O 2; between mid coxae, 14 6, 7 2; past mid but not to hind coxae, 3¢, 4 2: between hind coxae, 0 6, 1 2; behind hind coxae onto abdomen, 0 ¢, 2 2. This character is clearly variable, and females tend to have the longer labia. Although the two females with the longest labia came from the same island, there was no other correlation of labial length with island of origin. Pachybrachius nesovinctus Ashlock, new species. (Figures 4, 5.) Head slightly elevated between eyes, densely but shallowly punctate, sparsely dotted with inconspicuous subappressed and long erect hairs, length 0.71, width 0.75, anteocular length 0.37, eye length 0.17, interocular space 0.44; buccula high, short, bucculae joined as a single low carina at level of antenniferous tubercle; labium not quite reaching mid coxae, first segment not reaching base of head, segment lengths from base 0.44, 0.44, 0.27, 0.29; antenna with first segment exceeding clypeus by nearly half its length, segment lengths from base 0.34, 0.63, 0.49, 0.94 (fourth segment drawn too thick in figure 3). Pronotum with sparse short subappressed hairs and long erect fine hairs, with a single transverse row of widely spaced shallow small punctures on collar, VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 99 fi f ye Kj ih Cymoninus notabilis Pachybrachius nesovinctus Ficure 4. Dorsal views of the named species, and anterior view of fore femur of the holotype of P. nesovinctus. and similar punctures scattered over posterior lobe, length 0.83, collar length 0.09, width 0.56, anterior lobe length 0.29, width 0.88; divisions between collar and two lobes very distinct, anterior lobe globular. Scutellum with vestiture and punctation like those of hind lobe of pronotum; length 0.46, width 0.46. Brachypterous, hemelytron reaching midway onto abdominal segment VI, clavus and corium with surface subshining, vestiture as in pronotum and scutellum, clavus with three linear rows of punctures and a confused fourth row between medial and scutellar rows, corium with two linear rows of punctures 100 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. P. nesovinetus le, insularis Ficure 5. Spermathecae of the named species. VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 101 paralleling claval suture, randomly punctate laterally, with a few punctures posterior to level of apex of clavus; length of claval commissure 0.31, length of corium 1.36; membrane dull, veins not visible, basal length to level of corial apex 0.63, apical length from corial apex 0.07. Fore femur armed beneath with spines in two ranks as figured (fig. 4); fore tibia unarmed, slightly curved. Color. Head black, clypeus and juga dark reddish brown, antenna, buccula, and labium light yellowish brown. Pronotum black, collar anteriorly pale, posterior lobe reddish brown with dark median line. Hemelytron with clavus and corium light yellowish brown, punctures reddish brown, corium with white spot in inner angle and the black apical margin of P. vinctus group. Venter dark with anterior and posterior margins of propleuron, acetabula, scent gland auricle, and metapleural plate pale. Abdomen ventrally and dorsally dark reddish brown. Legs light yellowish brown, fore femur dark brown except apically. Hototyree. ¢, Santa Cruz Island, grassland, 2,100 ft., north of Academy Bay, 20 February (P. D. Ashlock). PaRATYPES. Santa Cruz Island, 10 ¢, 12 2, same data as holotype; 1 (macropterous), Table Mountain, 440 m., 16 April (D. Q. Cavagnaro). Fernandina Island, 1 2, southwest side, 1,000 ft., 4 February (P. D. Ashlock). This description adds another species to the confusing Pachybrachius vinctus complex, which badly needs revision. The new species is distinct in having long erect hairs on the dorsum. The New World P. vinctus (Say) and the closely related Pacific P. pacificus (Stal) both have small, inconspicuous appressed hairs on the dorsum. Pachybrachius nesovinctus is usually found in the brachypterous state (only one macropterous specimen has been collected), as are many populations of P. pacificus from the western and central Pacific. Brachypterous forms are less common in the Western Hemisphere species P. vinc- tus. The Galapagos species show more brown on the hemelytron than do speci- mens of P. vinctus from the North and South American mainlands. Genus Heraeus Stal Heraeus STAL, 1862, p. 315. Heraeus pacificus Barber. (Figure 5.) Heraeus pacificus BARBER, 1925, p. 21. Barber described this species from Santiago, and it has previously been recorded only from this island. Members of the expedition collected it in large numbers from Santa Cruz, and a single specimen was collected from Floreana. Many of the specimens were collected at light between Academy Bay and the Horneman Ranch on Santa Cruz, and I collected a large series in the so-called grassland area of the island at 1,800 feet under Jaegeria hirta. 102 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. LITERATURE CITED ASHLOCK, P. D. 1967. A generic classification of the Orsillinae of the world (Hemiptera—Heteroptera: Lygaeidae). University of California Publications in Entomology, vol. 48; pp. 1-82. AsHLock, P. D., and J. D. Lattin. 1963. Stridulatory mechanisms in the Lygaeidae, with a new American genus of Orsillinae (Hemiptera: Heteroptera). Annals of the Entomological Society of America, vol. 56, pp. 693-703. Baker, C. F. 1906. Notes on the Nysius and Ortholomus of America. Invertebrata Pacifica, vol. 1, pp. 133-140. Barser, H. G. 1925. Hemiptera-Heteroptera from the Williams Galapagos expedition. Zoologica, vol. 5, pp. 241-254. 1934. The Norwegian zoological expedition to the Galapagos Islands 1925, conducted by Alf Wollebaek. XI. Hemiptera—Heteroptera. Nyt Magazin for Naturvidens kaberne, vol. 74, pp. 281-289. BreEpDDIN, G. 1907. Berytiden and Myodochiden von Ceylon aus der Sammelausbeute von Dr. W. Horn (Rhynch. het). Deutsche Entomologische Zeitschrift 1907, pp. 34-47. Butter, A. G. 1877. Account of the zoological collection made during the visit of H.M.S. “Petrel’” to the Galapagos Islands. X. Lepidoptera, Orthoptera, Hemiptera. Proceedings of the Zoological Society, London 1877, pp. 86-91. CarRVALHO, J. C. M., and W. C. Gacne. 1968. Miridae of the Galapagos Islands (Heteroptera). Proceedings of the California Academy of Sciences, vol. 37, no. 7, pp. 147-219. DALLAs, W. S. 1852. List of the specimens of hemipterous insects in the collection of the British Museum, London, Pts. I and II. 592 pp. Distant, W. L. 1880-1893. Insecta. Rhynchota. Hemiptera-Heteroptera. Vol. I. Biologia Centrali- Americana. xx + 462 pp. 36 pls. DRAKE, C. J., and R. C. FROESCHNER. 1967. Lace bugs of the Galapagos Archipelago (Hemiptera: Tingidae). Proceedings of the Entomological Society of Washington, vol. 69, no. 1, pp. 82-91. FROESCHNER, R. C. 1968. Burrower bugs from the Galapagos Islands collected by the 1964 expedition of the Galapagos Scientific Project (Hemiptera: Cydnidae). Proceedings of the Entomological Society of Washington, vol. 70, no. 2, p. 192. Hann, C. W. 1826. Icones ad monographium Cimicum. Nurnberg, Lechner. HERRING, J. L. 1966. The Anthocoridae of the Galapagos and Cocos islands (Hemiptera). Proceedings of the Entomological Society of Washington, vol. 68, no. 2, pp. 127-130. KERZHNER, I. M. 1968. Insects of the Galapagos Islands (Heteroptera, Nabidae). Proceedings of the California Academy of Sciences, vol. 36, no. 4, pp. 85-91. VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 103 LINSLEY, E. G., and R. L. USINGER. 1966. Insects of the Galapagos Islands. Proceedings of the California Academy of Sciences, vol. 33, no. 7, pp. 113-196. MacArruur, R. H., and E. O. Wison. 1967. The theory of island biogeography. Princeton, New Jersey, Princeton University Press. xi + 203 pp. PoLHEMUS, J. T. 1968a. A report on the Saldidae collected by the Galapagos International Scientific Project 1964 (Hemiptera). Proceedings of the Entomological Society of Washington, vol. 70, no. 1, pp. 21-24. 1968b. A new Microvelia from the Galapagos (Hemiptera: Veliidae). Proceedings of the Entomological Society of Washington, vol. 70, no. 2, pp. 129-132. SLATER, J. A. 1964. A catalogue of the Lygaeidae of the world. Storrs, University of Connecticut. 1,668 pp. Span, (C- 1859. Hemiptera species novas descriptsit. Konglika Svenska Fregattens Eugenies resa omkring jorden. III (Zoologi, Insecter) 1859, pp. 219-298. 1862. Hemiptera Mexicana enumeravit speciesque novas_ descripsit. Stettiner Entomologische Zeitung, vol. 23, pp. 81-118; 273-281; 289-325; 437-462. 1872. Genera Lygaeidarum Europae disposuit. Ofversigt af K. Vetenskaps-akademiens for handlingar, Stockholm vol. 29, no. 7, pp. 37-62. Usincer, R. L. 1941. The present status and synonymy of some orsilline species (Hemiptera, Lygaeidae). Bulletin of the Brooklyn Entomological Society, vol. 36, pp. 129-132. Van DuzeE, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico. University of California Publications in Entomology, vol. 2, i-xiv, 1-902. 1933. The Templeton Crocker expedition of the California Academy of Sciences, 1934. No. 4. Characters of twenty-four new species of Hemiptera from the Galapagos Islands and the coast islands of Central America and Mexico. Proceedings of the California Academy of Sciences, vol. 21, no. 4, pp. 25-40. Viriters, A. 1970. Les Emesines des Iles Galapagos (Hemiptera, Reduviidae). Jn: Mission zoologique belge aus iles Galapagos et en Ecuador (N. et J. Leleup, 1964-1965), vol. 2, pp. 227-237. 8 TTP | ’ Leg) 7 A i he ais PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXIX, No. 9, pp. 105-110. September 5, 1972 A NEW LIZARD OF THE GENUS EMOIA (SCINCIDAE) FROM THE MARIANAS ISLANDS By Walter C. Brown Menlo College, Menlo Park, California, and Division of Systematic Biology, Stanford University. Research Associate, Department of Herpetology, California Academy of Sciences. and Marjorie V. C. Falanruw University of Guam, Territory of Guam. INTRODUCTION Brown (1956) recognized eight species of the Papuan-Oriental genus of lizards, Emoia, as occurring in Micronesia. Two were noted as apparently undescribed, of which one, a moderate-sized species, was represented by only four specimens from the Marianas Islands in the northwestern part of Micro- nesia. One of us (the junior author) has recently obtained a large series of the species from the same island group, and this larger sample permits a more adequate diagnosis of the species. It apparently has its closest affinities with Emoia boettgeri (Sternfeld) from central and eastern Micronesia and with Emoia arnoensis Brown and Marshall from the Marshall Islands in eastern Micronesia. Emoia atrocostata (Lesson), a widely distributed species-complex in the western Pacific basin; Emoia flavigularis Schmidt from the Solomon Islands to the south; and the very large Emoia nigra Jacquinot and Guichenot, the range of which extends from the Solomon to the Tonga Islands in the southern Island chain, are also representatives of the same evolutionary group. Measurements were made with a caliper to the nearest 0.1 mm. Middorsal scale rows were counted between the parietals and a point opposite the vent; [105] arine Biological Laboratory LIBRARY erp 15 1972 [Proc CALIFORNIA ACADEMY OF SCIENCES 106 r} ages oe S ve Ly) eee: ¢ A 4 bs Sy Ss =: ME Fx =: i ee nome aes ao! Vor. XXXIX] BROWN & FALANRUW: A NEW SPECIES OF EMOIA 107 midbody scale rows at a point approximately equidistant between the fore and hind limbs; fourth-toe lamellae from that proximal rounded lamella which is at least two-thirds the breadth of the toe to the most distal ventral scale inclusive. Emoia slevini Brown and Falanruw, new species. Hototyre. United States National Museum no. 192781, Cocos Island (a small island off the southern end of Guam), Marianas Islands, collected by M. V. C. Falanruw. PaRATYPES. United States National Museum nos. 122645-122646, Rota Island; 122470, Ritidain Point, Guam Island, collected by L. P. McElroy; 128028, Mt. Lasso, Tinian Islands, collected by H. K. Townes, Jr.; University of Guam nos. 547-552, 1305-1306, 1393-1395, 1448, 1450-1451; California Academy of Sciences nos. 129138—-129143; Museum of Comparative Zoology, Harvard University no. 128164; Field Museum of Natural History no. 171832; and British Museum of Natural History no. 1971.1027, from the same locality as the holotype. Dracnosis. An Emoia species of moderately large size, 58-85 mm. in snout-vent length for 18 mature specimens; length of hind limb less than 50 percent of the snout-vent length; simply rounded lamellae on under surface of digits, numbering 30-37 beneath the longest toe; 34-38 midbody scale rows and 61—74 dorsal scale rows between parietals and base of tail (for a sample of 27 specimens); prefrontals separated by the frontal; frontoparietals distinct; interparietal moderate in size; ground color (in life) of dorsum and upper lateral surfaces light to dark brown with variable scattered dark and light flecks. DeEscriPpTION. An Emoia species of moderate size; snout-vent length of 6 mature females 63-75 mm., of 12 males 69-84 mm.; snout round-pointed, of moderate length (36-44 percent of head length); supranasals much broader anteriorly than posteriorly, in contact with the anterior loreal which is shorter and broader than the posterior; prefrontals not in contact (separated by the frontal which is in contact with the rostral); length of the frontal about equal to that of the fused frontoparietals; interparietal of moderate size, length about equal to breadth at the base to nearly one and one-half times as great; 4 or 5 supralabials anterior to the enlarged one beneath orbit; 4 supraoculars plus a small one posteriorly; a single pair of large nuchals (fig. 1), dorsal scales smooth; midbody scale rows 34-38 for twenty-eight specimens; transverse rows along the middorsal line from parietals to base of tail 61-74; number of rows across the nape from ear to ear 12; 30-37 smooth, rounded lamellae < Ficure 1. Emoia slevini. a. Lateral view of head of paratype; b. Dorsal view of head of paratype, CAS. Biological Laboratory LIBRARY SEP 15 1972 Marine [Proc. 47TH Ser. CALIFORNIA ACADEMY OF SCIENCES 108 £7 07 8 9 ST N Orr'0 €cS 0 €cS'0 6870 O8r'0 W T8+0 LSS°0 Oss'O sTs0 cso qjSue] JU9A-yNous —£0+'0 -9LV'0 -L6+'0 -8S+'0 —cer'o u /quily pury jo yywueT suowtoeds 8I Os 9 £7 9 v LT N ~ = ainyeul JO (“Wwu) S8-8S VIT-SL TZ-8$ CL-S& s9-09 88-19 S8—L9 ud Y}sue] JUAA-JNOoUs Lé £C 6 CC 8 L 17 N qrey 299 Te9 Oss 6149 8°89 £99 STL W JO aseq 0} spejatied VLNO 99-09 c9-98 LL-S9 vL-S9 OL-¥9 vL-L9 u UWLOAT SMOT V[BIS LC eC 6 £7 6 L 1Z N sts ore i 4 97¢h TLv gee 6¢ W Le-0¢ Se-Te vy-8e Sb-6¢ CS-SY Se-Te Ty-Se u sel[9We] 90}-YJANOY Lé GC 8 £C 6 L 1Z N S9E TLE OLE C8e V8e 66¢ ese W 8t-ve Or-Ce OV-SE Or-Se Ov-9E Or-6¢ Cv9S ua SMOI a[eIS APOQpIyAL minays DABIU SLUDINSIADp SypqUats10 1498 190Q DIDJSOIOAJD — SISUBOUAD slaqeIeYy) a a a Oh GI Uh GL a a (suamigags {0 4aqunu = Ay ‘upam = py ‘adudA = Y) TWIAVS BIOWY 07 paznjad Sa19ags AOf SJuaMadANsDaUt JUuaUIJAag PUD SJUNOI a[DIS “| ATAVL, Vor. XXXIX] BROWN & FALANRUW: A NEW SPECIES OF EMOIA 109 beneath the fourth toe for 27 specimens; limbs rather well developed, but hind limb less than 50 percent of the snout-vent length (table 1) and usually slightly less than distance from axilla to groin. Cotor. The ground color dorsally (in life) is irridescent medium brown to dark brown with some darker flecks and occasionally light flecks, limbs, especially the hind limbs, often lighter with more numerous flecks; venter whitish to gray or cream anteriorly, more yellow posteriorly, sometimes orange about the vent; in recently preserved specimens the ground color is brown or somewhat reddish brown with scattered darker markings, in some specimens forming a vague pattern of narrow, broken, transverse bands. The yellow and orange of the venter fade rapidly in preservative. EtymoLocy. The species is named for Mr. Joseph R. Slevin, former Curator of Amphibians and Reptiles at the California Academy of Sciences. ComPARISONS. Emoia slevini is probably most closely related to Emoia boettgert which occurs in the Caroline Islands. The two species are very similar in color pattern, but F. slevini differs from the latter primarily in the much lower number of subdigital lamellae, 31-38 (mean = 33) for 27 specimens, instead of 45-52 (mean = 47) for 6 specimens of Emoia b. boettgeri from the Caroline Islands, and the somewhat shorter hind limbs relative to snout- vent length, less than 50 percent (usually greater than 50 percent for EF. boett- geri). Emoia arnoensis and E. flavigularis are also readily distinguished from E. slevini not only on the basis of very different color patterns, but also by the higher subdigital lamellar counts; for E. flavigularis the number of scale rows between the parietals and the base of the tail is usually less than 65, whereas it is usually greater than 65 for E. slevini; and the interparietal is usually fused with the parietals in E. flavigularis, or, if distinct, relatively small. Of the group of related species noted in the Introduction, only £. atrocostata is sympatric with E. slevini. Populations of both species are represented in collections from Cocos Island, with 23 examples of E. slevini and two of E. atrocostata. Unfortunately this widespread species, described by Lesson (1830) on the basis of a unique specimen from Oualan (= Kusaie) Island in the eastern Carolines, is still poorly represented in collections from most islands of Micronesia and probably does not occur in the Marshall Islands, at least in the eastern part. The data for E. atrocostata presented in this paper are based upon two examples from Cocos Island and several specimens from Ulithi Atoll and the Palau Islands. Using the limited data from these two small samples of E. atrocostata, aside from totally unlike color patterns, E. atrocostata would appear to be most readily distinguished from £. slevini on the basis of the slightly greater number of midbody scale rows (table 1); the longer, narrower interparietal (length one and one-half to two times its basal breadth) ; and the posteriorly more broadly truncate rostral. 110 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Hasitat. Based on observations of the population on Cocos Island, E£. slevini is a forest species. The forest on this island is predominantly tall Casuar- ina trees with some scattered coconut and other broadleaf trees. The understory growth is sparse and the canopy permits only diffuse sunlight to mottle the forest floor. The forested area is bordered toward the lagoon by Scaevola shrubs. Most of the lizards observed were active on the forest floor. Occasionally specimens were seen in low hollows of tree trunks with only their heads protrud- ing. Two were observed on the lower part of the trunk of Casuarina trees. In most instances escape was attempted by hiding under material on the forest floor. A few specimens were observed at the edges of the Scaevola shrubs. Emoia atrocostata on Cocos Island, as elsewhere (Brown and Alcala, 1967), occupies the various marginal beach areas or the open, often sparsely grassy or shrub-dotted areas adjacent to the beaches. KEY TO THE SPECIES OF EMOIA IN THE MARIANAS 1. Lamellae beneath the fourth toe greatly thinned, more than 50 E. cyanura Lamellae) rounded; not’ thinned, less than’ 50) =e 2 . Midscale rows between the parietals and the base of tail less than 60; maturity attained ae CH) ienbont, ie JESS tin, Selanne |esayetday Pt E. caeruleocauda Middorsal scale rows between the parietals and ihe. base of tail greater than 60; maturity not attained at less than 45 mm., rarely less than 50 mm., in snout-vent length 3 3. Midbody scale rows 38 to 40; rostral broadly truncate posteriorly; dorsum grayish, greenish, or tan with black or dark brown spots; upper lateral surface marked by iresular black vor Ganrke frown" bere ee E. atrocostata Midbody scale rows 34 to 38; rostral rounded or narowly truncate posteriorly; dorsal and upper lateral surfaces brownish with scattered darker markings occasionally forming bo a vague pattern of narrow, broken, transverse bands —_..____»_»___ E. slevini ACKNOWLEDGMENTS This study is part of the senior author’s program concerned with the herpeto- fauna of the Pacific Islands with emphasis on the Philippines. The latter program is currently supported by National Science Foundation Grant GB- 16972. Illustrations were prepared by Mr. Walter Zawojski, Stanford University. LITERATURE CITED Brown, WALTER C. 1956. The distribution of terrestrial reptiles in the islands of the Pacific basin. Proceedings of the Eighth Pacific Science Congress, vol. 3, pp. 1479-1491. Brown, WALTER C., and ANGEL C. ALCALA 1967. Population ecology of the tropical scincoid lizard, Emoza atrocostata, in the Philippines. Copeia, 1967, pp. 596-604. Lesson, R. P. 1830. Zoologie: In M. L. I. Duperry, Voyage autour du monde .. . sur la corvette de sa Majeste, La Coquille, pendant les années 1822-1825. Paris, vol. 4, 743 pp. and 157 pls. Marine B LIBRARY JAN® 1973 Woods Hole, Mass, PROCEEDINGS ‘ological Laboratory OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXIX, No. 10, pp. 111-120; 5 figs. December 27, 1972 TWO NEW GENERA AND TWO NEW SPECIES OF WESTERN PACIFIC SNAKE-EELS (APODES: OPHICHTHIDAE) ey) John E. McCosker! Scripps Institution of Oceanography La Jolla, California 92037 ABSTRACT. Two new species representing two new genera of ophichthid eels, sub- family Ophichthinae, are described from the western Pacific Ocean. Evips percinctus, from Palau, and Allips concolor, from Thailand, are described and figured. INTRODUCTION Examination of the extensive collections of Indo-Pacific fishes deposited at the California Academy of Sciences has disclosed the presence of two new species of ophichthid eels, subfamily Ophichthinae, each of which is distinctly different from any member of a known genus. Difficulty in obtaining specimens of sand-dwelling eels has been noted in recent works (McCosker and Rosen- blatt, 1972), yet the two individuals upon which this report is based represent but a small fraction of the numerous ophichthids, moringuids, and xenocongrids collected by the George Vanderbilt and Naga expeditions to the central and western Pacific Ocean. It is the author’s intent in this study to make these generic names available in preparation for his more thorough osteological comparison of genera within the family Ophichthidae. Measurements are straight-line, made either with a 300 mm. ruler with 1Contribution from the Scripps Institution of Oceanography, University of California, San Diego. Tilia] 112 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. a aU ee iy id JANG 1973 Wosds Hole, Mass. Vor. XXXIX] McCOSKER: NEW SNAKE EELS a | ee 0.5 mm. gradations (for total length, trunk length, and tail length) and recorded to the nearest 0.5 mm., or with dial calipers (all other measurements) and recorded to the nearest 0.1 mm. Head length is measured from the snout tip to the posterodorsal margin of the gill opening; trunk length is taken from the end of the head to mid-anus; body depth does not include the fin. Branchiostegal and vertebral counts (which include the hypural) were made using radiographs. Gill arches were prepared by means of the Taylor (1967) trypsin technique. Material used in this study is housed in the California Academy of Sciences (CAS) or the Marine Vertebrates Collection of the Scripps Institution of Oceanography. I thank Richard H. Rosenblatt for reading this manuscript critically, Carl L. Hubbs for suggestions, and William N. Eschmeyer of the California Academy of Sciences for permission to utilize specimens in his care. TAXONOMY Evips McCosker, new genus Driacnosis. Body nearly cylindrical, head and trunk longer than tail. Dorsal and anal low, ending before pointed tail tip; dorsal origin above and slightly behind gill openings. Pectoral rudimentary and less than eye dia- meter, its base in upper corner of gill opening. Gill openings lateral and shorter than isthmus breadth. Underside of snout not grooved. Anterior nostril tubular, posterior nostrils open into mouth. Teeth pointed; inter- maxillary teeth largest and depressible, vomerine teeth smaller and_ fixed. Eye large. Other characters those of the single species. Type species. Evips percinctus McCosker, new species. EtymMotocy. From the Greek «i (eu, latinized to ev for euphony before a vowel), good, and iy (ips, masculine), a worm, in reference to the general appearance of this charming eel. RELATIONSHIPS. Evips percinctus appears most closely related to the more generalized ophichthines such as species of Ophichthus, Microdonophis, and Pogonophis. Similarities among these genera include the retention of the pectoral and median fins, the lateral slightly restricted gill opening, the posterior nostril and head pore conditions, and the generally bold coloration. Osteologically, E. percinctus also appears similar to them in its pectoral girdle and hyoid arch conditions (viewed from radiographs and cleared and stained specimens) and in the condition of the gill arches, primarily the retention of the fifth ceratobranchial and the separation of UP; and UP, < Ficure 1. Evips percinctus McCosker, new species, CAS no. 13966, holotype, 125.5 mm. TL. Arrows indicate the origin and termination of the median fins. 114 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 2. Evips percinctus McCosker, new species, CAS no. 13966, holotype. Arrows indicate location of median interorbital and temporal pores. (Nelson, 1966; personal observation). The new genus is separable from these and other ophichthines on the basis of its rudimentary pectoral fin, robust body, and short tail. Evips percinctus McCosker, new species. (Figures 1-3.) Holotype and only known specimen, CAS no. 13966, a 125.5 mm. juvenile from Kayangel Island, Palau Islands, Southern Caroline Archipelago (8°02’18’N., 134°43’21”E.), collected in shallow water (0-4”) by H. A. Fehlmann and party during the George Vanderbilt Foundation 1956 Palau Islands Expedition, on 8 October 1956. DESCRIPTION. (Measurements are in millimeters.) Total length 125.5, head 14.0, trunk 60.0, tail 51.5, predorsal 17.2, body depth behind gill opening 5.2, at anus 4.5, snout length 2.8, upper jaw 4.8, eye diameter 1.4, interorbital width 1.5, gill opening height 0.9, isthmus width 3.1. Vertebrae 132; 69 to anal fin origin. Body stout (not exceedingly elongate as in many ophichthids) and nearly cylindrical. Depth behind gill openings 24 times in total length, and at anus 28; width behind gill openings 31 times in total length, and at anus 34. Snout blunt. Lower jaw included, its tip beneath a line drawn from anterior nostril base. Eye large, about 3.4 times in upper jaw, its center opposite midpoint of upper jaw. Anterior nostrils tubular, about 1.5 in eye diameter. Posterior nostrils open into mouth and lie ahead of anterior margin of eye. Tongue adnate. Branchial basket expanded and supported by numerous branchiostegals and jugostegalia which broadly overlap along the ventral midline. Numerous papillae on snout, beneath eye, and on anterolateral flanges of upper lip (fig. 2). Dorsal origin above and slightly behind gill VoL. XXXIX] McCOSKER: NEW SNAKE EELS 115 Ficure 3. Dentition of holotype of Evips percinctus McCosker, new species. opening. Pectoral fin rudimentary, a small semicircular flap, about equal in length to anterior nostril and attached above midpoint of gill opening. Dorsal and anal low, disappearing in advance of tail tip. Caudal rays lacking. Head pores conspicuous; preoperculomandibular, temporal, suborbital, postorbital, and supraorbital series present (fig. 2). A single median inter- orbital and temporal pore. Two postocular pores, lying within faintly pig- 116 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. mented spots. Left lateral line pores 125; 9 before gill opening and 73 before the anus, terminating 0.3 head length before tail tip. Teeth pointed and depressible (fig. 3). Four anterior peripheral inter- maxillary teeth covered by skin folds, flanking a median intermaxillary pair, followed by seven uniserial vomerine teeth. Maxillary teeth biserial, the outer row covered by a skin fold and larger anteriorly, the inner 4 prom- inent. Mandibular teeth uniserial, about 10 on each side and grading smaller posteriorly. Gill arches removed, and cleared and stained. First basibranchial os- sified, second cartilaginous, third rudimentary, fourth absent. Hypobranch- ials 1 and 2 ossified, 3 cartilaginous. Ceratobranchials 1—5 ossified, the fifth a slender filament. Second and third infrapharyngobranchials ossified. Lower pharyngeal tooth plate slender with about 25 conical biserial teeth. Upper pharyngeal tooth plate (UP; and UP, of Nelson, 1966) separate (unfused) and subrectangular, with about 15 conical teeth. Color in isopropanol yellow, overlain dorsally with 16 brown saddles. First saddle just behind occiput, does not reach sides of head; remaining 15 extend below lateral midline but do not meet ventrally. The type has retained traces of larval pigmentation, visible as 9 minute black pigment patches evenly spaced along the ventral midline of the trunk. Median fins unpigmented. EtymoLocy. From Latin, signifying banded throughout. Allips McCosker, new genus Dtacnosis. Body very elongate, nearly cylindrical for most of its length; head and trunk longer than tail. Dorsal and anal low and lying within a shallow groove, ending before bluntly pointed tail tip; dorsal origin well behind gill openings. Pectoral rudimentary, a tiny flap in upper rear corner of gill opening. Gill openings lateral and low on sides, separated by an isthmus wider than their length. Underside of snout grooved. Jaw teeth and vomerine teeth small and pointed. Eye minute. Other characters those of the single species. Type species. Allips concolor McCosker, new species. Etymo.ocy. From the Greek adAos (allos), another, and iy (ips, masculine), a worm. RELATIONSHIPS. Allips concolor appears most similar to extant species of Bascanichthys, Phaenomonas, and Gordiichthys. The last is a poorly known ophichthid excluded from the recent generic treatment of Rosenblatt and McCosker (1970). It will be redescribed on the basis of new material by James E. Bohlke (personal communication). Allips differs from Bascanichthys in having a more posterior dorsal fin origin, a more rounded snout, and a more cylindrical trunk and tail. The two genera are quite similar in having Vor. XXXIX] McCOSKER: NEW SNAKE EELS 117 Ficure 4. Head of holotype of Allips concolor McCosker, new species, CAS 13967, 375 mm. TL. small nearly fixed teeth, comparable gill arch configurations (although the fifth ceratobranchial, absent in Adlips, is very reduced but retained in certain species of Bascanichthys), low crescentic gill openings, rudimentary pectoral fins, a grooved snout, reduced eyes, highly rugose skin regions, a nearly uniform coloration (although certain species of Bascanichthys are often darkly pigmented dorsally), and similar head and body shapes and _ proportions. Phaenomonas differs in its extreme dorsal fin reduction, complete loss of the pectoral and anal fins, smaller eye, and the elongation of the trunk region. Allips concolor McCosker, new species. (Figures 4-5.) Holotype and only known specimen, CAS no. 13967, a 375 mm. specimen from Goh Phi, Ranong Province, Thailand (10°57’42’”N., 98°35'18”E.), north of Ban Parknam Ranong. Collected in shallow water (0-3) at the mouth of Pakehan river by H. A. Fehlmann and party during the 1959-1961 Naga Expedition, on 1 June 1960. DESCRIPTION. (Measurements are in millimeters.) Total length 375, head 21.0, trunk 205, tail 149, predorsal 48, body depth behind gill openings 5.6, at anus 5.0, snout length 3.0, upper jaw 5.2, eye diameter 0.6, interorb- ital width 2.6, gill opening height 1.7, isthmus width 2.6. Vertebrae 174; 96 to anal fin origin. Body elongate and nearly cylindrical, becoming laterally compressed only near tail tip. Depth behind gill openings 67 times in total length, and at anus 75; width behind gill openings 83 times in total length, and at anus 86. Snout subconical, rounded at tip. Lower jaw included, its tip behind anterior nostrils. Eye small, 8.5 in upper jaw, and faintly visible under skin; its midpoint closer to corner of mouth than snout tip. Anterior 118 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. \ busy a Ficure 5. Dentition of holotype of Allips concolor McCosker, new species. nostrils in a short tube; posterior nostrils open into mouth beneath eye. Tongue adnate. Branchial basket expanded and supported by 21 pairs of branchiostegals and jugostegalia which broadly overlap along ventral midline. Sides of head, throat, chin, and body flanks markedly rugose. Dorsal fin origin well behind gill openings. Pectoral fin rudimentary, a small flap attached to upper rear margin of gill opening. Dorsal and anal fins low, lying within a groove and disappearing 0.85 head length before finless caudal tip. The median and posterior caudal edges are covered with a band of numerous small papillae. Head pores reduced, but preoperculomandibular, temporal, suborbital, VoL. XXXIX] McCOSKER: NEW SNAKE EELS 119 postorbital, and supraorbital series present (fig. 4). A single median inter- orbital and temporal pore. Lateral line canal and pores present, but impossible to discern because of their reduced state and a waxy precipitate which has formed in preservation. Teeth small, pointed, and close set (fig. 5). Jaw teeth and posterior vomerine teeth uniserial. Gill arches removed, and cleared and stained. First basibranchial os- sified, second cartilaginous, third absent, fourth cartilaginous but rudimentary. Hypobranchials 1 and 2 ossified, 3 cartilaginous. Ceratobranchials 1—4 os- sified, 5 absent. Second and third infrapharyngobranchials ossified. Upper and lower pharyngeal tooth plates are subrectangular patches with conical teeth; UPs and UP, separate. Color in isopropanol nearly uniform brown although slightly darker on upper half due to small dark punctations. EtymoLtocy. From Latin, in reference to the nearly uniform coloration. LITERATURE CITED McCosker, JoHN E., and R1icHarp H. ROSENBLATT 1972. Eastern Pacific snake-eels of the genus Callechelys (Apodes: Ophichthidae). Transactions of the San Diego Society of Natural History, vol. 17, no. 2, pp. 15-24, 5 figs. NELSON, GARETH J. 1966. Gill arches of teleostean fishes of the order Anguilliformes. Pacific Science, vol. 20, no. 4, pp. 391-408, 58 figs. ROSENBLATT, RicHarD H., and Joun E. McCosker 1970. A key to the genera of the ophichthid eels, with descriptions of two new genera and three new species from the eastern Pacific. Pacific Science, vol. 24, no. 4, pp. 494-505, 8 figs. TayiLor, WILLIAM RALPH 1967. An enzyme method of clearing and staining small vertebrates. Proceedings of the United States National Museum, vol. 122, no. 3596, 17 pp. PROCEEDINGS 0005 Hole, Vase. OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXIX, No. 11, pp. 121-140; 6 figs.; 3 tables. December 27, 1972 FIVE NEW INDO-PACIFIC PIPEFISHES By Earl S. Herald Steinhart Aquarium, California Academy of Sciences San Francisco, California 94118 and John E. Randall Bernice P. Bishop Museum, P.O. Box 6037 Honolulu, Hawaii 96818 During the 1946 fish investigations at Bikini Atoll in the Marshall Islands, V. E. Brock and Herald carried out the first moderate-depth rotenone stations for the collection of fishes. Since this was before the development of scuba, the diving gear consisted of Brock’s modified Monson lung rebreather with other divers receiving air supplied through a standard dive line attached to a small compressor. It was soon evident from these collections that an important break occurred in the vertical distribution of the fishes. Many were confined to the intertidal zone or a few feet below; others were common in water of about 15 feet or greater and rarely, if ever, were seen in shallow water. The Syngnathidae provided a good example of this faunal break. Of the ten species of pipefishes (47 specimens from 26 stations) which were collected in the Marshall Islands, five were found only at depths below 15 feet, and three of those were undescribed. With the exception of Dentirostrum janssi, the other four species of pipe- fishes described as new herein appear to belong to the deeper water group, being taken at depths ranging from 20 to 160 feet. One species was collected in the Hawaiian Islands, one in the Marianas, the third in the [121] 122 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TABLE 1. Meristic comparisons for the Dunckerocampus complex. Num- Trunk ber of Rings Tail Rings Dorsal Fin Rays eee 15) 16117 16) 7 18 19: 20) 20) 22 2320022023678 550 Dunckerocampus multiannulatus* g* ipa i fF il 1? Seas D. pessuliferus 1 1 1 1 D. c. caulleryi 1 1 1 1 D. c. chapmani 3 I 2 LZ: ih D. dactyliophorus 59 1 58 10 37 10 2 21318 14 8 5 D. baldwini 20 20 Wo @ 1 ey 2 2 Dentirostrum janssi 34 34 1 426 3 1 3 16 14 ‘Includes literature holotype data for D. multiannulatus and D. ben-tuviae. It is probable that trunk ring counts for these two types should be recorded as 16 rather than 17. Ryukyus, and the fourth at Easter and Pitcairn islands. These are island groups in which considerable fish collecting has been carried out in shallow water in recent years. The following abbreviations are used for institutions in this paper: AMS (Australian Museum-Sydney); ANSP (Academy of Natural Sciences of Phil- adelphia); BM(NH) [British Museum (Natural History)]; BPBM (Bernice P. Bishop Museum); CAS (California Academy of Sciences); LACM (Los Angeles County Museum of Natural History); SMF (Senckenberg Museum) ; and USNM (United States National Museum). Technical assistance in making counts and measurements has been pro- vided by Mr. D. Anderson. THE DUNCKEROCAMPUS COMPLEX Seven long-nosed species of very spiny belly-pouch Indo-Pacific pipefishes are members of the Dunckerocampus complex. All of these including the related Doryrhamphus and Oostethus pipefishes have a first trunk ring, i.e., the one bearing the pectoral fins, that is twice as long as any of the remaining trunk rings. Although it is actually double, it is counted as a single ring. In meristic characters these species are very similar (table 1). Based on pigment patterns or the absence thereof, members of the Dunckerocampus complex fall into four categories: the first is the narrow- banded group whose two members usually have 4 or 5 very narrow pigment bands on the opercle, two or more on each trunk ring and sometimes on each tail ring. Dunckerocampus multiannulatus (Regan) 1903, was described from Mauritius and was characterized by a lack of banding on the snout. VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 123 Dunckerocampus ben-tuviae Fowler and Steinitz 1956, described from the Red Sea had the same meristic characters but did have banding on the snout. Dr. Eugenie Clark advises us that Red Sea pipefishes of the D. multiannulatus/D. ben-tuviae group have a wide range of snout banding variability so that D. ben-tuviae will need to be considered a synonym of D. multiannulatus. The remaining species in the narrow-banded group, Dunckerocampus pessuliferous, is distinct from other Dunckerocampus in having a high dorsal fin count of 30 as compared with a normal range of 20-25. It is known from the holotype taken by dredge at a depth of 144 feet at Sulade Island in the southern Philippines. The two species in the wide-band group usually have a single pigment ring for each trunk and tail ring. The most common species is the central and western Pacific D. dactyliophorus; it has a single ring around the opercle which differentiates it from the twin opercular rings of D. caulleryi Chabanaud known only from Amboina and New Caledonia. This latter species has two subspecies: D. c. caulleryi with 19 tail rings, and D. c. chapmani Herald with 16 or 17 tail rings. More material is needed to determine the status of these two. The third category is represented by the striped Hawaiian endemic described as new herein: Dunckerocampus baldwini. This species extends the generic range from the Austral Islands northward to the Hawaiian region, a distance of about 2800 miles. And, finally, the fourth category is that group of pipe- fishes that, unlike Dunckerocampus, developed the brood pouch folds; the new genus and species for this is Dentirostrum janssi described as follows: Dentirostrum Herald and Randall, new genus Diacnosis. Spiny, long-snouted Doryrhamphine pipefish with abdominal brood area protected by lateral membranous folds but without protecting plates. Lateral trunk ridge continuous with inferior tail ridge at anal ring; superior trunk and tail ridges discontinuous at end of dorsal fin; lateral tail ridge ending free at anal ring. Tail non-prehensile. Dorsal fin rays 21-24, anal 4, pectoral 19-21, caudal 10; trunk rings 16; tail rings 20- 23. Named Dentirostrum in reference to the magnificent spines present on the median snout ridge of males and to a lesser extent on females. Discussion. Dentirostrum is most closely related to Dunckerocampus from which it differs, (1) in having brood pouch flaps rather than having the eggs nakedly attached to the abdomen, (2) in its extremely spinose nose ridge, and (3) in the presence of a secondary spine behind the primary spine at the ring juncture of the superior and inferior trunk ridges. Den- tirostrum and Doryrhamphus are similar in that both have brood-pouch folds, but the latter differs in its small size and its short snout. 124 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. The type species of Dentirostrum, D. janssi, is new and is the only known member of the genus. The description of D. janssi based on 34 specimens (14 males 61-126 mm. SL and 20 females 52-110 mm. SL) from 19 localities in the Indo-Malayan region is as follows: Dentirostrum janssi Herald and Randall, new species. (Figure 1.) From the Patau IsLaAnps: 13 types, 6 localities. Hototype. CAS 14139, male 125 mm. SL, with 126 brood patch sockets— a few with eggs; Arappu Point of Koror to Ho Island, south side of reef; 0-50 feet; H. DeWitt; November 19, 1957 (GVF 1442). ParATYPES. CAS 14140, male 123 mm. SL; same data as holotype. CAS 14141, male 105 mm. SL; south entrance to small bay on west side of Ngalab Point, Koror Island; 0-50 feet; H. DeWitt; November 21, 1957 (GVF 1445). CAS 14142, female 89 mm. SL; Iwayama Bay, off Kaibakku Island; 0-45 feet; H. DeWitt; October 3, 1957 (GVF 1408). AMS 16144-001, male 103 mm. SL; female 105 mm. SL; LACM 32123-1, male 105 mm. SL; SMF 11425, male 111 mm. SL; channel between Sanryo Island and Ngatkumer Island, Iwayama Bay; 0-50 feet; H. DeWitt; October 31, 1957 (GVF 1433). CAS 14143, 3 females, 53 mm., 97 mm., 103 mm. SL; channel between Sanryo Island and Kamori Island, Iwayama Bay; 0-42 feet; H. DeWitt; November 18, 1957 (GVF 1439). BPBM 11937, male 102 mm. SL; female 83 mm. SL; western tip of Ngargol Island; 20-40 feet, coral; J. E. Randall, E. 5. Heliman, O. Custer; June 8, 1968. From THAILAND: 16 paratypes, 9 localities. BPBM 11938, female 91 mm. SL; NW. side of Koh Kroi Island about 5.5 miles ESE. of Ban Pae Fisheries Training Center, Rayong Province; 0-4 meters; H. A. Fehlmann; April 30, 1960 (GVF 2183). CAS 14146, male 77 mm. SL; NW. side of Goh Samet Island, Rayong Province; 0-2 meters; B. Bronson; April 29, 1960 (GVF 2180). CAS 14147, 3 females, 62 mm., 64 mm., 83 mm. SL; NW. side of Goh Raed about 1.5 miles ESE. of Prachuap Khiri Khan town, Prachuap Khiri Khan Province; 0-15 feet (photo 3-79 a-c); H. A. Fehlmann; June 19, 1961 (GVF 2651). USNM 206654, male 126 mm. SL; female 78.5 mm. SL; CAS 14148, 2 males 80 mm. and 61 mm. SL; female 59 mm. SL; W. side of Goh Luem ca. 3 miles SE. of Prachuap Khiri Khan Town, Prachuap Khiri Khan Province; 0-15 feet; H. A. Fehlmann; June 18, 1961 (GVF 2648). CAS 14149, 2 females 95 mm. and 102 mm. SL; W. side of Goh Luem, Prachuap Khiri Khan Province; 0-15 feet; H. A. Fehlmann; June 17, 1961 (GVF 2646). CAS 14150, female 111 mm. SL; on fringing reef at head of Gulf of Siam, Goh Sak Island, about 35 miles NE. of Prachuap Khiri Khan town, Prachuap VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 125 PAPA saay. aoaaeaaaners a a a ae mm Ficure 1. Dentirostrum janssi Herald and Randall, Holotype; male 125 mm. SL (CAS 14139); (a) lateral view; (b) expanded ventral view of brood patch area; (c) head detail male holotype compared with (d) head detail female 111 mm. SL paratype (CAS 14150). Drawing by H. Hamman. Khiri Khan Province; 0-20 feet; H. A. Fehlmann and R. Rofen; October 24, 1957 (GVF 1460). CAS 14151, female 58 mm. SL; SW. point on Goh Chorakhay, Chumphon Province; 0-16 feet; B. Bronson; May 25, 1960 (GVF 2199). CAS 14152, male 91 mm. SL; fringing reef on NW. side of Goh Maprao, Chumphon Province; 0-12 feet; H. A. Fehlmann; May 18, 1960 (GVF 2186). CAS 14153, male 105 mm. SL; Prond Bay at SW. 126 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. corner of Goh Samed, ca. 100 meters from shore, Chumphon Province; 0- 10 feet; H. A. Fehlmann; May 17, 1960 (GVF 2185). From OTHER LOCALITIES”: 5 paratypes. CAS 14145, 2 females 96 mm. and 107 mm. SL; Bay of Nhatrang SW. of Hon Long, Vietnam, South China Sea; 1-15 meters; R. Bolin; March 14, 1960 (GVF 2116). BPBM 11939, female 93 mm. SL; Bay of Nhatrang, Hon Long, Ving Damlon, Vietnam, South China Sea; 1-6 meters; R. Bolin; February 23, 1960 (GVF 2072). CAS 14144, male 93 mm. SL; Surabaja, Java, Indonesia; Hilmi Oesman; February 1961. ANSP 119933, female 99 mm. SL; Little Hope Island, NE. end, Queensland, Australia; 5-8 meters; J. Tyler, C. L. Smith, and G. Bettle; January 3, 1969. It will be noted that the distribution of Dentirostrum janssi does not include the Philippines although it is known from west, east, and south of that area. It is probable that it will be found there when rotenone collecting is conducted by diving methods using scuba. Diacnosis. Dorsal fin rays 21-24, usually 23-24; dorsal covering a total of 5—6% rings of which 1-2 are trunk rings and 4-5 are tail rings; usual count: 1 + 5. Trunk rings 16, tail rings 20-23, usually 22; pectoral 19- 21, usually 20; anal 4, caudal 10. Head-in-standard length 4.2—5.0; snout- in-head 1.5—1.9; dorsal-base-in-head 1.9-2.8; brood pouch folds extending from second trunk ring to anal ring, and occasionally as far as 4th tail ring. Eggs large, about .75 mm. diameter; largest number in belly brood patch: 195 (105 mm. male). Ridge system and brood patch folds typical of Dentirostrum. DescripTION. Nearly all ridges of head, trunk, and tail are spinose. The median snout ridge of the males has 15-21 large spines, and on each side there is a lateral ridge with smaller spines. The spines on females are less accentuated. The supranasal ridge is present. The anterior ocular ridge is lacking, and the posterior ridge is smooth. The orbit is ringed with small spines. The opercle has a smooth median ridge over its entire length, and beneath it may have 5-7 additional radiating ridges (compare figs. le and 1d). The pectoral cover plate has 1 or 2 spines on its upper anterior edge, and a single spine in the center. Superior and inferior trunk ridges have double spines at each ring juncture, but on the tail only a single spine at each ring. The lateral trunk ridge has double spines for the first 5-7 rings, and then a single spine for the next 9-11 rings. Between 62 and 80 mm. the future brood patch folds show their presence by a pair of darkly pigmented lines on the abdominal surface of the male. Among the 14 males only two had eggs attached to the brood patch: the 2A new male Dentirostrum janssi (106 mm. SL) has just arrived; it is a depth record (100 feet) captured at Kranket I., Madang, New Guinea; G. R. Allen, May 15, 1972. VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 127 125 mm. male holotype (126 brood patch sockets, a few with eggs) and a 103 mm. male (21 eggs and 135 empty sockets = 156). These were collected in October and November in the Palaus. Another male of 111 mm. collected at the same time and place showed 156 empty egg sockets. Two males, 91 and 105 mm. from Chumphon Province, Gulf of Thailand (May collection), showed 106 and 195 empty sockets. All 33 specimens had 16 trunk rings, and the male brood patch folds usually cover 15 of the 16. However, five of the 14 males had the folds extending upon the tail, the furthest distance being 3% tail rings. Although the range of brood pouch coverage was 15 to 19% rings there was no indication of vascularity on the tail portion of the pouches. It is probable that this small tail section is not used as a brood area. The overall color of Dentirostrum janssi is light brown with two pairs of dark brown lines extending from the head to the dorsal fin with one on each side of each superior trunk ridge. The first pair extends from above the eye dorsally on the inside edge of the superior trunk ridge, whereas the second pair extends from the center part of the eye over the opercle and along the upper lateral side of the superior trunk ridge. The tail fin has a clear center section surrounded by a dark reddish black area. The junior author’s field notes on two males collected in the Palaus at Ngargol Island indicate that the body is blackish anteriorly and posteriorly with orange in the middle. Also, the pigment around the tail fin shows a small amount of white on the outer edge. Discussion. From the upper west side of the Gulf of Siam (GVF 64: BanAangtong Bay; CAS 14155), we have three small specimens (33, 33.5, and 42 mm. SL) that have all of the characteristics of D. janssi except that they have a single spine on each ring of trunk and tail and are lacking the double spines of the trunk of D. janssi. Our smallest specimen of the latter species is 58 mm. SL (CAS 14151), and it has the physical characteristics of the adults. Among the syngnathids, spines are usually lost with growth, not added. Because of the lack of spines on these three specimens, we are faced with the possibility of another species of Dentirostrum existing in this area. However, we will not describe these as new at this time, but will await the collection of adult material from the same area. At this same GVF station 64, juvenile representatives of two additional species of syngnathids were collected: 8 specimens (average length 44 mm.) of Corythoichthys species (CAS 14318) and a 61 mm. specimen tentatively identified as Syngnathus maxweberi (CAS 14317). Named ‘janssi? in honor of Mr. Edwin Janss, Jr., whose keen interest in tropical marine biology has resulted in important field investigations in many regions. 128 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. FicurRE 2. Dunckerocampus baldwini Herald and Randall, Holotype; male 125 mm. SL (CAS 24734); cave SE. of Pokai Bay, Waianae coast, Oahu; 70 feet. Photograph by J. E. Randall. Dunckerocampus baldwini Herald and Randall, new species. (Figure 2.) Twenty type specimens from the Hawaiian Islands: Oahu (17), Hawaii (2), and Maui (1). Capture methods: by hand, quinaldine, and chemfish ichthyocide. Depth range 20-160 feet; size range 63-131 mm. SL. HototypPe. CAS 24734, male 120.5 mm. SL (125.5 mm. total length); cave SE. of Pokai Bay, Waianae coast, Oahu; 70 feet; J. E. Randall and S. Swerdloff; July 20, 1969. PARATYPES. CAS 24735, female 125 mm. SL; same data as holotype. LACM 32126-1, female 126 mm. SL; CAS 14138, 2 males 106.5 and 129 mm. SL; 3 females 129, 131, and 131 mm. SL; Waianae coast off Makaha Shores Condominium, Oahu; 45 feet; J. E. Randall and A. R. Emery; April 26, 1970. BPBM 11941, 2 females 63 and 128 mm. SL: Waianae coast off Lahilahi Point, Oahu; 40 feet; J. E. Randall and P. M. Allen; July 11, 1970. SMF 11426, female 128 mm. SL; Kawaihoa Point, Maunalua VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 129 Bay, Oahu; 35 feet, lava caves; Jane Culp; November 15, 1971. AMS 15607-001, female 123.5 mm. SL; CAS 14137, female 134 mm. SL; Kawaihoa Point, Maunalua Bay, Oahu; 35 feet, lava caves; Jane Culp and James Moore; December 18, 1971. USNM 204683, female 92 mm. SL; off north side at base of dropoff, Moku Manu, Oahu; 160 feet; J. E. Randall, T. Stark, W. Baldwin; October 9, 1969. BM(NH) 970.1.26:1, female (?) 72 mm. SL, Moku Manu, Oahu; 85 feet; J. E. Randall and W. Baldwin; October 6, 1969. BPBM 7783, male 124 mm. SL; female 125 mm. SL; Kanoehe Bay at channel entrance, Oahu; 95 feet; J. E. Randall, D. Chave, W. Hashimoto; October 10, 1969. BPBM 11940, male 102 mm. SL (tail broken at 11th ring; caudal regenerated); north end of Honaunau Bay, Kona coast, Hawaii; 150 feet; J. E. Randall, E. S. Hobson, J. R. Chess; August 16, 1969. BPBM 11942, male 130 mm. SL; first point north of Hanakahau Boat Harbor, Kona Coast, Hawaii; 100 feet; J. E. Randall; August 18, 1970. CAS 14979, female 117 mm. SL; 1% miles north of Lahina, Maui, Hawaii; 20 feet; Steinhart Divers; June 29, 1972. Diacnosis. Holotype belly brood area covering % 12 % trunk rings with 143 egg sockets—73 on right and 70 on left. Dorsal fin rays 21-23; pectoral 19-21; anal 4; caudal 10; dorsal fin covering %—1 trunk ring and 3% tail rings; usually “4 + 3; total rings covered by dorsal 3%—4%; trunk rings 16; tail rings 20-21; head-in-standard length 4.5—5.2; snout-in-head 1.5— 1.7; dorsal fin base-in-head 2.9-3.5. Ridge system typical of Dunckerocampus, i.e., lateral trunk ridge continuous with inferior tail ridge; median tail ridge extending forward to anal ring. Tail fin long, 5-7 mm. in adults of 100 mm. or larger. General appearance except color similar to other members of genus Dunckerocampus. Color in life reddish, fading to dark with preserva- tion. Red stripe along upper part of body from snout to tail. Edges of caudal fin white. DeEscriPTION. Median snout ridge with many serrations; orbital, opercular, nuchal, and pectoral cover plate crests faintly visible. Eye very large, the orbit diameter contained almost two times in minimum snout depth or % length of opercle. Intermedial scutella between rings small and equal in width to one-half distance between scutella. Body ridges well developed with sharp spine projecting at posterior edge of juncture of each ring. The following color description was made from 102 mm. Honaunau Bay male shortly after capture: “light pinkish gray with broad dorso-lateral bright orange-red stripe becoming blackish red and narrower on snout; also a broad orange-red stripe midventrally extending forward on snout as red and blackish red. Snout yellowish, white dorsally and yellow between lateral and ventral blackish red stripes; tail nearly entirely red (red stripes close together); caudal fin red with a narrow white edge at sides, and a black area distally on 2 upper central interradial membranes; dorsal colorless.”’ 130 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. In preservative the males apparently retain the color stripe along the head and trunk much better than do the females. The 125 mm. female from Kaneohe Bay (BPBM 7783) has a conspicuous blackish tip on the outer margin of the last five rays of the dorsal fin. Other specimens did not show this character. Discussion. Capture data and dive observation records indicate that specimens of Dunckerocampus baldwini are often found in pairs. In captivity longevity has been short, usually no more than 2 months. This compares with about three months longevity for the banded pipefish, D. dactyliophorus. In recent years this latter species has often been imported into the United States from the Philippines. Cleaning activity by Dunckerocampus baldwini has been observed by Dr. Edith Chave. While diving at Milolii, Hawaii (August 4, 1970), in a cave at a depth of 75 feet, she watched an adult redstripe pipefish clean a cardinal fish, Apogon evermanni, then a moray eel, Gymnothorax species, and finally the pipefish attempted to clean Dr. Chave’s wrist. Only two other species of pipefishes are known to act as cleaners. Randall (1962) reported D. P. Wilson’s observations at the Plymouth Aquarium of a John Dory being cleaned by the snake pipefish, Entelurus aequoreus. The junior author has also observed the flagtail pipefish, Doryrhamphus mela- nopleura, cleaning reef fishes, specifically a moray and a cardinal (Randall and Helfman, 1972). In recognition of his study of Pacific fishes, the redstripe pipefish, Dunckerocampus baldwini, is named in honor of Wayne J. Baldwin who with the junior author collected the first specimens of this new species. Dunckerocampus dactyliophorus (Bleeker) 1853. Banded Pipefish. Syngnathus dactyliophorus Bleeker, Nat. Tijdschr. Nederl. Indiee, vol. 4, p. 506, 1853 (type locality, Onrust Island, Djakarta [Batavia], Java). Herald (1953, p. 252) studied geographic variability of meristic characters in a small series of Dunckerocampus dactyliophorus. He postulated sexual dimorphism in the trunk color ring pattern with females having a greater number than males. Tables 2 and 3 present data on the 62 specimens from 22 localities now available. Unfortunately, there does not appear to be any recognizable pattern that can be correlated with sex or area. It will be noted that the trunk ring count is remarkably constant at 16 with only one of 62 having a different count, i.e., 15. Size range for the 62 specimens was 41 to 159 mm. SL. The smallest mature male showing egg sockets on the brood patch area was 90 mm. The banded pipefish has some interesting color ring variations. The typical VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 131 TABLE 2. Geographic variation of Dunckerocampus dactyliophorus. Trunk Rings Tail Rings Dorsal Fin Rays ye) ly iy IG Koy a aa 2s AON Bil BRP Bs ab DS Australs 2 2 2 Marshalls 6 5 gS Kapingamarangi 1 1 1 Guam 1 1 1 Ulithi 2 2 2 Palaus 1 29 9 21 o) 12 7 Philippines 6 tee v4 1 IT the Sh as Solomons 2 il 1 1 Celebes 6 to § 3 2 1 Java 1 1 New Guinea 5 5 Se, Totals (62) i “Oil 10 40 10 2} 2 SS 2a Oe pattern shows about 5 rings on the snout (range 4-7), 1 over, under, and rarely through the eye, another around the opercle, 1 at the pectoral fins, then (table 3) 6-13 on the trunk, and 7-14 on the tail. The pattern for any given specimen is recorded as 5-1-1-1-7-9 = 24. The width of the individual color band is usually 42 to “% of an individual ring. However, the two Ulithi specimens (BPBM 8746) have remarkably narrow color bands; in width they are only equal to about % to ™% of a trunk or tail ring. Another extreme is shown by a 91 mm. specimen from Urukthapel in the Palaus (BPBM 7352); the color bands were very wide, 1 to 1% rings in width. Dunckerocampus dactyliophorus ranges through the western and central Pacific. The type locality, Djakarta, is the westernmost point in the distribu- tional pattern, and until recently the easternmost was 4300 miles away at Rongelap Atoll in the Marshall Islands. However, in 1971 the junior author diving at a depth of 185 feet collected two specimens at Rurutu (lat. 24° S.) in the Austral Islands. This extended the range westward for another 2700 miles and southward for 900 miles from the latitude of the Solomons (lat. 9° S.). The northernmost locality is Guam (lat. 12° N.). Recently Dr. Eugenie Clark advised us that she has examined Dunckero- campus dactyliophorus from the Red Sea (Eilat, Gulf of Aqaba) which is some 8800 miles (by water) from Djakarta. This break in distribution is similar to that which occurs in Corythoichthys flavofasciatus with the sub- species of C. f. flavofasciatus in the Red Sea and C. f. conspicillatus many miles away in the Central Pacific (Herald, 1953, p. 275). 132 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TaBLe 3. Geographic variation of Dunckerocampus dactyliophorus. Trunk Color Bands Tail Color Bands 7 Be —@) lO. abl 22 ie 7 8 9) DOI Stas Australs i ail 1 1 Marshalls Se nlee2 1 292 1 Kapingamarangi 1 1 Guam 1 1 Ulithi i! 1 1 1 Palaus A S1S s6r 4: 5. 4)» 59 2 Philippines Wee 2e> 2k et See Solomons 2 2 Celebes a ae 1 31 ee 1 Java 1 1 New Guinea 1 4 2M aes Totals (62) aks ale alk alloy ao) 1 6 16° 18> PIS: The specific collection localities for Dunckerocampus dactyliophorus listed for the general areas of tables 2 and 3 are as follows: Marshall Islands: Rongelap, Bikini, and Eniwetok atolls; Palaus: Koror, Auluptagel, Babelthaup, Urukthapel, and Arakabesan islands; Solomons: New Georgia and Florida islands; Philippines: Cebu, Jolo, Pandanan Island, and Sibutu. Although the Philippine localities are all from the southern section, we have been advised by Mr. Earl Kennedy, who is the major Manila live fish jobber, that the species is fairly well distributed throughout the islands; most of his specimens come from Luzon (Batangas), Mindoro, and Palawan. Amboina is the only recorded locality from which we have not examined specimens. THE PIPEFISH OF EASTER AND PITCAIRN ISLANDS On the map of the world, Easter Island is a small dot in the eastern south Pacific some 2000 miles west of Caldera, Chile, and about 1100 miles east of Pitcairn, the nearest inhabited island. This isolated volcanic outcrop has a total area of 46 square miles, being about 14 miles long and 7 miles wide (27° 05’ S. Lat. and 109° 20’ W. Long.). Although the island is inter- nationally famed because of the giant stone statues, the published record of its ichthyological fauna and relationships is less well known (Randall, 1970). In recent years three field parties of biologists (1958, 1965, and 1969) have made collections at various sites around the island. All have used ichthyocides, and among them they have taken ten specimens from three localities of the first syngnathid to be collected in the area. In December 1971 the junior VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 133 author visited Pitcairn Island and was able to collect three additional specimens. The type series consists of 13 specimens: 5 males (69-85.5 mm. SL) and 8 females (67-95 mm. SL). Several years ago Dr. David K. Caldwell studied the Ramsey Parks 1958 series of 6 specimens at the Los Angeles County Museum of Natural History. He tentatively determined that they represented an undescribed species. This we are now able to verify, and we name this species in his honor. Syngnathus caldwelli Herald and Randall, new species. (Figure 3.) Hototyer. LACM 6560-3, 76 mm. standard length male with eggs in pouch. Anakena Cove, Easter Island; Ramsey Parks, Yacht Chiriqui; boulders and brown algae, rotenone to 4.6 m.; October 1, 1958. PARATYPES. (5 from same collection as holotype): LACM 6560-41, 69 mm. male and 82 mm. female; CAS 24202: 71 mm. male and 65.5 mm. female; and USNM 203409, 79 mm. female. Another collection made at type locality 6% years later—BC 65440, 84 mm. male (pouch empty); closed tidepools, depth % to 8 m., rotenone; Ian E. Efford and Jack A. Mathias, Jan. 15, 1964. Ten days later at nearby Vinapu on southwest side—BC 65449, 2 females 93 and 93.5 mm.; large boulders and rocks; depth 2-3 m., rotenone, also Efford and Mathias, Jan. 25, 1965. Finally, 10% years after first colection—BPBM 6596, 95 mm. female (fig. 2); southwest coast between Hanga-Roa and Hanga-piko, inshore; depth 61 cm.; boulder bottom with brown algae; chemfish; J. E. Randall and G. R. Allen; January 26, 1969. BPBM 10856, 82 mm. male and 80 mm. female. Pitcairn Island, off “the Rope” 20 ft., large boulders with brown algae, sand and small rocks in low places; J. E. Randall, Dean B. Cannoy, Steve Christian, and Noggie Young; December 23, 1970. CAS 13922, 75 mm. female, same data as BPBM 10856. Diacnosis. Dorsal fin: 28-31; 7 specimens with 28 rays; 3 with 29; 2, 30; and 1, 31; pectoral 14-16; usually 14; anal 3; caudal 10; dorsal fin covering 6%4—7% rings, i.e., 72-1 trunk ring and 6-6% tail rings usually 1 + 6; trunk rings 16-17; tail rings 34-36; head-in-standard length 8.5— 9.2; snout-in-head 2.2—2.6; head-in-dorsal fin base 1.06—-1.15; dorsal fin base-in-head .87—.95. Lateral body ridges typical for genus Syngnathus: i.e., lateral trunk ridge interrupted at anal ring, then subcontinuous with lateral tail ridge (65.5 mm. female has the two ridges continuous on one side only). Brood pouch covering first 13-15 tail rings; eggs large, 1 mm. diameter, arranged in 3—4 single rows across pouch (holotype) or 2 rows wide and 2 rows deep (69 mm. male). Holotype egg count 88; for 69 mm. male: 19 134 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 3. Syngnathus caldwelli Herald and Randall, Paratype; female 93 mm. SL (BPBM 6596); Easter Island between Hanga-Roa and MHanga-piko. Photograph by J. E. Randall. and 21 eggs for dorsal rows and 14-14 for ventral rows or about 68 for total pouch count. Brood pouch folds almost in contact but without overlap or indentation, merely slightly thickened at free end of individual flap (modified Open Brood Pouch Closure: O-BPC). Color in preservative, brownish some- times with many narrow dark lines running length of body. Eye may have dark band extending obliquely postero-ventrally over opercle. Dark spots sometimes present at juncture point between rings on inferior trunk ridge, and to lesser extent on lateral trunk ridge. Spots on Pitcairn pipefishes, larger and very pronounced; also with wide whitish areas on upper surface: about 4 on trunk and 7 on tail. DescrRIPTION. Median snout ridge smooth, extending forward from inter- orbital area for about half snout length. Superior ocular ridge extends VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 135 posteriorly for distance equal to eye width but is absent anteriorly. Eye very large, in diameter about equal to 2% of snout length. Nuchal plates bilobed, indistinct. Upper and lower pectoral ridge plates evident but not pronounced; opercular ridge extends over % to *% length of opercle. Body ridges evident, smooth, and not strongly pronounced. Lateral flanges of brood pouch slightly developed. Fretp Notes. The junior author and G. R. Allen collected the most recent Easter Island paratype and also made color and black and white photographs at the time of capture. From field notes the near-living color of Syngnathus caldwelli from Easter Island is: “light brown with row of red dots along anterior lateral ridge and full length of ventral flange; a small red spot at front and at rear of dorsal fin; ventral part of body light yellowish with a midventral row of red dots along trunk; large irregular light gray blotches on back and upper side; a dark brown band running from snout through lower half of eye where it broadens on lower opercle and continues as a broad zone on chest; caudal fin yellowish with brown rays; dorsal fin clear.” Any distinctive marks on the 1958 specimens have been largely lost following preservation. The two 1965 specimens from Vinapu are the only ones to show clearly the narrow lines on the body, about 12 on the top of the trunk and approximately 10 on the trunk sides. The 1965 Anakena specimen is very dark and its markings are suggestive of the 1969 specimen whose near-life colors are described above. Comparisons. In the Indo-Pacific from Africa to the Americas there are about 34 members of the genus Syngnathus, excluding Corythoichthys and Bombonia. None of these approach Syngnathus caldwelli in their numerical or other characteristics. The nearest relative is probably Syngnathus balli of Hawaii. However, the latter is a smaller species (58 mm.) with fewer tail rings (32 vs. 35-36) and fewer dorsal fin rays (21-23 vs. 28-31). Syngnathus banneri Herald and Randall, new species. (Figure 4.) HototyrEs. BPBM 8695, 39 mm. SL (40 mm. TL) undet. sex; Ryukyu Islands, Ishigaki; reef about ’2 mile off harbor of Ishigaki City; depth 20- 35 feet; chemfish ichthyocide; J. E. Randall and A. H. Banner, May 22, 1968. PaRATYPE. CAS 14375, 26.5 mm. SL (27.3 mm. TL) immature; Marshall Islands, Eniwetok Atoll, lagoon off Eniwetok Island; 25 feet, coral and rubble patch; quinaldine; J. E. Randall, March 31, 1972. Diacnosts. Dorsal fin rays 16-18 located on a total of 3'2—-4 rings (%4-— 1 trunk and 3-3% tail rings); pectoral 11; anal 2; caudal 10. Trunk rings 136 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 5 mm Ficure 4. Syngnathus banneri Herald and Randall, Holotype; imm. 39 mm. SL (BPBM 8695); reef near Ishigaki City, Ryukyus. Drawing by L. Sabre. 15; tail rings 27. Head-in-standard length 7.3—7.8; snout-in-head 2.5-3.2; dorsal fin base-in-head 1.7-1.8. Body ridge pattern typical of Syngnathus, i.e., lateral trunk ridge discontinuous with lateral tail ridge at end of anal ring; inferior trunk and tail ridges continuous and superior trunk and tail ridges discontinuous at end of dorsal fin. Brood pouch unknown. DEscrIPTION. Holotype median snout ridge with two triangular flanges just anterior to nostrils; orbital ridge markedly extended dorsally; prenuchal- nuchal ridges present but not accentuated. Opercular ridge extending over one-half (paratype three-quarters) of opercle. Superior and inferior pectoral cover plate ridges sharply pronounced. Trunk and tail ridges slightly raised and indented between individual rings so that ridgés have moderately scalloped profile. Paratype ridges for the most part less accentuated than holotype. Holotype color light tan with exception of dark brown spot pattern on anterior trunk; beginning behind pectoral, 6 spots on lower trunk ridge and 5 on lateral trunk ridge on left side; spots on right side less distinct; on dorsal surface 9 dark brown bars, one-half ring in width: 3 extending between superior trunk ridges and 6 between superior tail ridges. Paratype color whitish with dark area on lower sides of gill cover extending underneath pectoral fins. Discussion. Syngnathus banneri is a remarkable species for it has the lowest dorsal fin count of any member of the genus (16-18 as compared with a normal range of about 19-45), and with this is combined a very low tail ring count of 27. When egg-bearing males are available for examination, this species will probably prove to be a member of the subgenus Microsyngnathus Herald 1953. These are small Svngnathine pipefishes usually less than 100 mm. in length that have overlapping brood pouch closures (O-BPC). Named ‘banneri’? in honor of Dr. A. H. Banner whose welcome field efforts resulted in the capture of the holotype of this species. VoLt. XXXIX] HERALD & RANDALL: PIPEFISHES 137 Minyichthys Herald and Randall, new subgenus of Micrognathus Duncker 1915 Type Species. Micrognathus brachyrhinus Herald 1953. Diacnosis. Differentiated from other members of Micrognathus by in- creased number of trunk rings (19-21 rather than 13-17), very short snout, and small size (mature at less than 50 mm.). Named Minyichthys, small fish, from the Greek ‘“miny” meaning small. Discussion. Pipefishes of the genus Micrognathus have the lateral trunk ridge continuous with the inferior tail ridge and the lateral tail ridge is present. The tail brood pouch has everted type closure in which the outer lip of one flap is turned back upon itself, and the other flap overlaps it (Herald, 1959). There are three subgenera: (1) Anarchopterus Hubbs 1935, characterized by smooth body ridges and absence of the anal fin, has two species limited to the Gulf of Mexico and Caribbean; (2) Minyichthys, defined above, has two Pacific species; and (3) the type subgenus Micrognathus with its typical sharp body ridges, has two Atlantic American species and nine Pacific species. Micrognathus (Minyichthys) myersi Herald and Randall, new species. (Figures 5 and 6.) HorotyPe. CAS 13918, 42.5 mm. SL mature male (43.5 mm. TL); Guam, NW. Cocos Island, outside of reef, 70-100 feet depth; rotenone; June 30, 1969; J. E. Randall, e¢ al. PARATYPE. BPBM 8759, 41 mm. SL female (43 mm. TL); Guam; south of Uruno Point, about 10 mi. NE. of Agana; depth 60-90 feet; reef edge adjacent to sand; June 27, 1968; J. E. Randall and H. Kami. Diacnosis. Dorsal fin rays 29-31 covering 9-10% rings, i.e., 2%-3 trunk rings and 64-77% tail rings; trunk rings 19, tail rings 40-41; pectoral 11- 12; anal 2, caudal 8; head-in-standard length 7.2—8.92; snout-in-head 2.36- 2.7; dorsal fin base-in-head .75—.93; pectoral base-in-pectoral length 1.6. Lat- eral ridge pattern typical of Micrognathus, i.e., median trunk ridge continuous with inferior tail ridge; lateral tail ridge extended forward onto 2 trunk rings (holotype) or 2% (paratype); superior trunk and superior tail ridges interrupted at posterior edge of dorsal fin. Distinctive spike on dorso- median snout ridge just ahead of nostrils. Brood pouch covering first 13 tail rings with embryos to 10th ring; 7 large embryo sockets; pouch protecting plates slightly developed with pouch closure of everted type (E- BPC). Mouth extremely vertical. DEScRIPTION. With exception of internasal spike, all head and body ridges faint. Orbital ridge mildly accentuated with pronounced spine on anterior border (holotype) or moderate (paratype). Opercular ridge extending 138 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 5 mm Ficure 5. Micrognathus myersi Herald and Randall, Holotype; male 42.5 mm. SL (CAS 13918); NW. Cocos Island, Guam. Drawing by L. Sabre. over half of opercle. Pectoral cover plate with faint superior and inferior ridges. Trunk and tail ridges with indentation between rings, and ridges with finely toothed edge. Base of dorsal fin resting in slight ‘V.’ Color light tan: holotype with indication of 4 pigment streaks extending downward from opercle and dark area on side of snout just anterior to eye. Paratype with indication of color bands formed by occasional dark spots: 7 bands or bars on trunk and 17 on tail. Discussion. Although the two type specimens are almost the same size, the holotype snout length is much greater than that of the paratype, with snout-in-head values being 2.36 and 2.7. If the other characters were not so similar, one would be tempted to consider them as closely related but separate species. Short snouts are the mark of the subgenus Minyichthys as is shown even better by the two known specimens of the genotype M. (Minyichthys) brachyrhinus (snout-in-head 3.17). A comparison between the two species of Minyichthys follows: M. (M.) brachyrhinus M. (M.) myersi Dorsal fin rays 23-25 29-31 Rings covered 1%-1%+5%- 2%-3+6%4- by dorsal 6%4=74-8%4 74%2=9-10% Trunk rings 19-21 19 Tail rings 37-39 40-41 Snout-in-head 3.17-3.18 2.3-2.7 Max. known size 31 mm. 42.5 mm. Distribution Oahu; Ticao I. Guam Philippines This new species is named in honor of Stanford faculty member Dr. George Myers (retired) who has been the major professor for many of today’s ichthyologists. VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 139 5 mm Ficure 6. Micrognathus myersi Herald and Randall, Paratype; female 41 mm. SL (BPBM 7579); south of Uruno Point, Guam. Drawing by L. Sabre. COMMENTS ON OTHER MICROGNATHUS Since the senior author’s Micrognathus study of 1953, there have been two additional new species, one in the Atlantic and one in the Pacific. In 1964 Gilbert Whitley described Micrognathus boothae from Australia’s Lord Howe Island (off New South Wales). This species is closely related to M. brocki discovered at Bikini Atoll in the Marshall Islands, about 2570 miles north of Lord Howe. The two species differ mainly in the shorter snout of M. boothae along with its 42 rather than 37 tail rings. Three specimens are known for the two species: the female holotypes of each plus a second specimen of M. brocki captured by the junior author in 1968 at Ishigaki in the Ryukyu Islands. This 82 mm. female (BPBM 8755; 20-70 feet) represents a westward range extension of about 2800 miles. When more material is available of both species, it is possible that the two may prove identical. In April and May 1971 the junior author collected four heavily-banded specimens of Micrognathus edmonsoni in the Marquesas. Three were taken at Nuka Hiva at a depth of 70 feet (BPBM 10857, 71 and 70 mm. SL, and CAS 13977, 62 mm.) and one was captured at 115 feet in Vaitahu Bay at Tahuata (BPBM 11936, 72 mm. SL). This is startling because previously this species had been considered a rare Hawaiian endemic. Although it was described more than 42 years ago (1930) only 7 other specimens are known from 3 localities on Oahu and Maui. This new locality extends the range about 2500 miles southward from the Hawaiian archipelago. It is entirely possible that this species is one of the deeper water syngnathids, and thus has escaped collections made by non-diving ichthyologists. Another problem deals with a pipefish originally described as [chthyocampus annulatus Macleay 1878. The two type specimens are actually Micrognathus brevirostris as shown by reexamination in 1971 by Dr. John R. Paxton 140 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. at the Australian Museum. Unfortunately the artist used a specimen of Yozia species to make the illustration for the original description, and in some unknown way this fact escaped the notice of the author. LITERATURE CITED HERALD, EArt S. 1953. Family Syngnathidae: Pipefishes. In: Leonard Schultz et al. Fishes of the Marshall and Marianas Islands. United States National Museum, Bulletin no. 202, pp. 231-278, 9 figs. 1959. From pipefish to seahorse—a study of phylogenetic relationships. Proceedings of the California Academy of Sciences, ser. 4, vol. 29, no. 13, pp. 465-473, 3 figs. 1969. A new pipefish from the Virgin Islands, Micrognathus dawsoni. Occasional Papers of the California Academy of Sciences, no. 73, 4 pp., 1 fig. RANDALL, JOHN E. 1962. Fish service stations. Sea Frontiers, vol. 8, no. 1, pp. 47. 1970. Easter Island, an ichthyological expedition. Oceans, vol. 3, no. 3, pp. 48-59. RANDALL, JOHN E., AND E. HELFMAN In Press. Dipractacanthus xanthurus, a cleaner wrasse from the Palau Islands with notes on other cleaning fishes. Tropical Fish Hobbyist. SmMitH, J. L. B. 1963. Fishes of the family Syngnathidae from the Red Sea and western Indian Ocean. Rhodes University Ichthyological Bulletin no. 27, pp. 515-543. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXIX, No. 12, pp. 141-184; 11 figs. July 9, 1973 THE GENUS MECAS LECONTE (COLEOPTERA: CERAMBYCIDAE) By John A. Chemsak and E. G. Linsley University of California, Berkeley The genus Mecas is a group of lamiines generally distributed from Guatemala to southcentral Canada. A single species occurs in North America west of the Rocky Mountains, but none are known from northeastern North America. Based upon present knowledge, most of the species are associated with weedy com- posites. A few of them are also lampyrid and/or cantharid and lycid mimics. Although the genus has been recently revised (Breuning, 1955), available in- formation and material have necessitated a further reclassification of the group. CLASSIFICATION AND NOMENCLATURE The first named species of Mecas as currently recognized (M. pergrata, M. cana, M. cinerea) were assigned by their authors to the genus Saperda Fabricius (Say, 1824; Newman, 1840). LeConte (1852, 1859a) transferred S. pergrata to Stenostola Mulsant, a generic name he also used for his ‘saturnina’ (LeConte, 1859a), but left ‘cana’, which he did not know, in Saperda (he had no reason to refer to ‘cinerea’ which was from Mexico). In 1847, Haldemen described ‘femoralis’ in the genus Phytoecia Mulsant, which LeConte (1852) made the monobasic type of his genus Mecas. Twenty-one years later, LeConte (1873a) added a second species, M. marginella, and defined the genus (1873b) in such a manner that all of the above species could be included, although he did not mention them by name. Unfortunately, Lacordaire (1872) overlooked Mecas which would have fallen in his Tribe IV (Phytoecides), although he included the related Dylobolus Thomson (which Bates later regarded as a synonym of Mecas) in his Groupe IV (Arenicides) and Pannychis Thomson (which we regard as a [141] 142 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. subgenus of Mecas) in his Groupe III (Amphionychides). LeConte’s expanded concept of the genus was first applied to the North American species by Horn (1878) and to the Mexican species by Bates (1881). Casey (1913) provided a key to the species in his collection and Breuning (1955) published a revision of the genus. However, this revision is incomplete, since three of the species, M. inornata of authors, M. cineracea Casey, and M. bicallosa Martin were omitted. These were incorrectly transferred by Breuning to the genus Saperda, although their published characters (including the cleft or toothed claws in both sexes) and recorded host plants (weedy composites) are typical of the species currently and historically referred to Mecas (the species of Saperda are wood borers). Although the evolution of the generic concept of Mecas until the revision by Breuning was fairly straightforward, there has been considerable uncertainty regarding the identity of some of the species. The most serious confusion in- volves the name ‘Saperda inornata’ described from Missouri Territory by Say (1824), who used the broad concept of Saperda current in his time. LeConte (1852), not knowing what species Say had before him, included it in his treat- ment of Saperda, quoting from Say’s description as he did for other species which he had not identified. Obviously uncertain, he speculated that it might be the male of his S. concolor from Sante Fe, New Mexico, which he described in the same paper (LeConte, 1852). However, he apparently decided later that ‘imornata Say’ belonged to the genus Mecas, and the specimens standing under that name in his collection belong to this genus. Further, the name ‘in- ornata Say’ does not appear in his key to the species of Saperda (LeConte, 1873a). Horn (1878), in his key to the species of Mecas, included the name ‘inornata, applying it to the species previously described by LeConte as ‘satur- nina, which Horn regarded as a synonym of ‘inornata.’ However, Blanchard (1887) found two species differing in claw structure standing under the name ‘saturnina’ in his collection. These had not been differentiated in Horn’s Key. This discovery prompted Horn (1888) to apply the name ‘saturnina’ LeConte to the species with the claws moderately deeply cleft with the inner division lobe-like. On this basis of distinction Gahan (1888) placed Saperda cinerea Newman and Mecas senescens Bates as synonyms of Mecas inornata (Say) and Mecas saturnina (LeConte) as a synonym of Mecas cana (Newman). We agree that M. saturnina and M. cana are conspecific, although in the material before us the two appear to be allopatric. Mecas cinerea (Newman) and M. senescens Bates also appear to be conspecific, but are not closely related to any species from the United States and thus could not under any circumstances be called ‘inornata Say.’ Although various species of Mecas have been treated taxonomi- cally in the intervening years as ‘M. inornata Say’ (see synonymical bibiog- raphies), the species which most closely fits Say’s very brief description is M. confusa (described below) or, possibly, M. cineracea Casey. VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 143 In 1924, Martin named a species MW. bicallosa on the assumption that ‘inornata Say’ was Saperda in the modern sense and a senior synonym of S. concolor Le- Conte (see above), although the arguments advanced for this decision were not conclusive (Martin also assumed that he was formally naming the species that previously had been called M. inornata; however, the species before him was from the Great Basin and the ranges accorded to M. inornata by Horn and Blanchard are to the east of this area). Martin’s placement of S. concolor LeConte as a synonym of S. inornata Say was accepted by Breuning (1952) and Nord and Knight (1970). These latter appealed to the International Commission on Zoological Nomenclature to use its plenary powers to designate as neotype of Saperda inornata Say the type of Saperda concolor unicolor Felt and Joutel, and place ‘inornata’ on the Official List of Specific Names in Zoology. While such an action would eliminate some nomenclatural confusion in the genus Mecas, its impact on the literature of the North American Saperda will be unfortunate. Proposing to relegate the name ‘inornata Say’ to the list of nomina dubia might have been a preferable solution to the problem. DISTRIBUTION The known species of Mecas occur in the area from western Canada to south- eastern United States, Mexico, and Guatemala. Of the 15 species recognized by us, five are restricted to northern and central Mexico and another ranges into Guatemala; six occur in both the United States and Mexico and three are thus far known only from the United States. A single species, M. bicallosa, is found west of the Rocky Mountains, occupy- ing the Great Basin and extending into British Columbia and northern Baja California. Most of the species occurring in the United States are found in either the southeastern and/or southwestern portions of the country. Only three species, M. femoralis, M. marginella, and M. confusa, are not yet known from Mexico but probably only M. femoralis does not occur there. The most widely distributed species, M. rotundicollis, ranges from Costa Rica to Kansas ana Arizona. Of the remaining four species common to both countries, M. cana saturnina, M. pergrata, and M. cineracea apparently occur only in northeastern Mexico but are more widely distributed in the United States. The other species, M. menthae, ranges from Distrito Federal north along the western slope of the Sierra Madre to Arizona and New Mexico. Five of the six Mexican species, M. sericea, M. humeralis, M. cinerea, M. cirrosa, and M. ambigena are apparently largely restricted to central Mexico from Chihuahua and San Luis Potosi to Puebla. The sixth, WM. obereoides, extends into Guatemala. In many respects the distribution of Mecas is comparable to that of Elytro- leptus (Linsley, 1962; Chemsak and Linsley, 1965). Of the 17 species of Elytro- leptus, 9 are known only from Mexico, 7 from the United States and Mexico, 144 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. and one from the United States only. The majority of the species, as in Mecas, are found in central and northern Mexico and southwestern United States. Since nothing is known of the biology of the species of Elytroleptus, the significance of the distributional similarity, if any, is not evident. FORM AND COLORATION Most of the species of Mecas are concolorous with the integument black and densely clothed with pale recumbent pubescence. A few of the species, M. fem- oralis, M. pergrata, and M. cinerea frequently or always possess reddish femora and the latter two often have the elytra and/or pronotum partly reddish. Some of the members of the genus are distinctive by the narrow, densely pubescent white bands on the suture and epipleurae of the elytra. The most striking divergence from the typical appearance is to be found in M. sericea, M. rotundicollis, and M. obereoides. The first of these, M. sericea, is definitely lycid-like in aspect usually possessing two lateral dark bands at the base and apex of the elytra. The basal band may be reduced or absent and occasionally both bands are lacking. Since this species is mimetic, the variation in color is probably an expression of resemblance to different lycid models within the range of the species. Lycus sallei Gorham is a possible model with two dark elytral bands and we have another similarly marked Lycus from Sinaloa and Colima. Mecas rotundicollis is unquestionably a lampyrid mimic throughout most of its range, although in some areas a cantharid may be the model. As is the case in M. sericea, M. rotundicollis apparently mimics different species of models in different parts of its range. Variation is expressed primarily by the presence or absence of yellow pubescence on the apical abdominal sternites and yellow pubescent bands on the suture and epipleurae of the elytra. We have been unable to find a geographical trend in these characteristics but most of the available specimens from Arizona lack the yellow pubescence. The third species, M. obereoides, may be involved in a mimetic ring with a cantharid model. Very little variation in color is expressed in this species but it is one of the most distinctive in the genus. Field studies will be necessary to confirm involvement with mimicry. The remaining species of Mecas are all quite similar in form and coloration. Differences between species involve such characteristics as tarsal claw structure, number of glabrous calluses on the pronotum, and relative lengths of antennal segments. BIOLOGY Little has been recorded on the biology of species of Mecas. The most complete accounts known to us are those of Baerg (1921) and Stride and Warwick (1962). Baerg’s report describes injury to Jerusalem artichokes in Arkansas and probably VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 145 refers to M. cana saturnina (LeConte) or the species named below as Mecas confusa, new species. Baerg’s account, under the name “Mecas inornata Say” is as follows: This beetle, half an inch long, of a light gray color, is a girdler that attacks arti- choke (Helianthus tuberosus). The beetles begin ovipositing early in July. The females when laying eggs girdle the main stem about six inches from the top. Two girdles are made, about 1-144 inches apart. Immediately above the lower girdle is the egg punc- ture. This is exactly similar to the method followed by the Raspberry cane girdler (Oberea bimaculata). The girdles are not clean cuts such as we find in woody plants, but rather a series of holes encircling the stem. Apparently one female will deposit in a large number of plants. In spite of the fact that only a few beetles could be located, practically all the plants in the field were attacked in the course of a few days. As a result of the injury, the leader in the plant dies and the plant develops a bushy type of branching. The young larvae upon hatching begin to feed between the girdles and later proceed towards the base of the plant. They confine their injury largely to the pith. Apparently under certain weather conditions the artichoke is not well fitted as a host plant. In only one out of four or five plants showing egg punctures was there a full grown larva. In most of the other plants the larva had begun to feed and some time later died, presumably it had been injured by the growing stalk. The larvae attain full growth, that is about seven-eights of an inch in length, some time in November. At this time the larvae are found at the very base of the stalk, about two inches below the surface of the ground, in an enlargement of the tunnel which has been padded with fine bits of pulp. The pupal stage has not been observed but since the adults appear early in July, the larvae will presumably pupate some time in May or early in June. It seems reasonable to assume that this species will attack most of the species in the genus Helianthus. None of these were near the artichokes, and no data have been secured. The only host plant other than artichoke that could be located is the common ragweed (Ambrosia artemisiifolia) . The observations of Stride and Warwick were made on M. saturnina in Aus- tralia where that species had been introduced as a biological control agent for Xanthium (Wilson, 1960). The habits as they reported are almost identical with Baerg’s account and an additional observation was the plugging of the oviposition hole with a gummy substance after the single egg had been laid. Stride and Warwick advance a hypothesis whereby the double-girdling behavior of M. saturnina may be regarded as a device originally evolved to permit the use of succulent green shoots of otherwise woody plants as food for cerambycid larvae. Presumably, it has been retained in M. saturnina because it promotes advan- tageous changes in the herbaceous host plant attacked, possibly increased pithiness. Earlier reports also refer to stem- and root-boring habits of Mecas (Riley, 1880; Beutenmuller, 1896; Leng and Hamilton, 1896). Most aspects of adult be- havior are lacking from the older literature but Townsend (1884) reported that 146 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ‘Mecas inornata’ takes wing and flies away when it observes someone approach- ing, but drops to the ground and feigns death when unexpectedly disturbed. Adults of M. menthae, new species were found in numbers by their collectors on the upper leaves of the mint, Agastache, during the day. Host RELATIONSHIPS Precise host data for most species of Mecas are lacking. In part this results from uncertainty regarding the identity of the species of Mecas associated with published host records and in part from the fact that most records, both pub- lished and unpublished, are based upon collections of adults from plants and not upon reared material. Nevertheless, there is an interesting consistency among the records that are available. With very few exceptions the plants involved are weedy, herbaceous composites which have special chemical characteristics expressed in terms of aromatic, medicinal, irritant, or toxic properties. The list of known or suspected hosts based upon field collections, with notations of some of their properties as reported by Blake (1951) and Kingsbury (1967) is as follows: COMPOSITAE Ambrosia (Ragweed). A. artemisiifolia Linnaeus (Common Ragweed). One of the most widespread causes of hayfever. Artemisia (Sagebrush). A. tridentata Nuttall (Big Sage). Used medicinally by Indians and early white settlers in the West; a hay fever plant; toxic to livestock if eaten in excess. Aster (Aster). A. tanacetifolius Humboldt, Bonpland, & Kunth (Tansyleaf Aster). Used medic- inally by the Indians; some species of aster absorb Selenium and become toxic. Baileya (Baileya). B. multiradiata Harvey & Gray (Desert Marigold). Causes mortality in sheep, par- ticularly, but not exclusively, on over-grazed land. Gaillardia (Blanket-flower) . G. pulchella Fougeroux de Bondaroy. A related species, G. pinnatifida Torrey. was used by the Hopi Indians as a diuretic. Guardiola (Guardiola). G. tulocarpus Gray. Helenium (Sneezeweed). H. hoopesti Gray (Orange Sneezeweed). Contains a toxic glucoside (dugaldin) which causes spewing sickness in sheep. H. microcephalum WDeCandolle (Sneezeweed). Toxic to livestock. H. tenuifolium Nuttall (Bitter Sneezeweed). Toxic to livestock and suspected of poisoning humans. Helianthus (Suntlower). H. annuus Linnaeus (Common Sunflower). Toxic to cattle in large amounts; the seeds are edible. H. tuberosus Linnaeus (Jerusalem Artichoke). Roots edible. VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 147 Verbesina (Crown-beard). V. encelioides (Cavanilles) Bentham & Hooker (Golden Crownbeard). Used by Indians and White pioneers in the West for treatment of boils and skin disease; Hopis reported to bathe in water in which plant has been soaked to relieve pain of spider bites. Xanthium (Cocklebur). X. spinosum Linnaeus (Spiny Cocklebur). Seeds and seedlings contain a glucoside (Xanthostrumarin) poisonous to swine and poultry. LABIATAE Agastache (Giant-hyssop) A. species (Horsemint). Genus Mecas LeConte Mecas LEContTE, 1852, Jour. Acad. Sci. Philadelphia, vol. 2, no. 2, p. 155; 1873, Smith- sonian Misc. Coll., no. 265, p. 347; Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; Bates. 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 203; LeContTEe and Horn, 1883, Smithsonian Misc. Coll., vol. 507, p. 332; Lrenc and Hamiurton, 1896, Trans. American Ent. Soc., vol. 23, pp. 151, 152; Casry, 1913, Memoirs on the Coleoptera, vol. 4, p. 360; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 138. Form elongate, usually parallel. Head with front convex, interantennal area usually concave; mandibles rather short, apices curved, acute; palpi slender, maxillary pair longer than labial; eyes rather small, finely faceted, deeply emar- ginate, upper lobe small; antennae usually slender, sparsely or densely fringed with long hairs beneath, particularly on basal segments; third segment usually longer than first, fourth subequal to or shorter than third, outer segments short or long. Pronotum wider than long, sides usually rounded; disk variably pubescent, often with glabrous calluses; prosternum short, intercoxal process narrow, expanded at apex, coxal cavities closed behind; mesosternum with coxal cavities open; metasternum with episternum broad in front, narrowing behind. Legs short; intermediate tibiae with a dorsal sinus; tarsal claws bifid. Abdomen normally segmented; last sternite deeply impressed in the male, linearly im- pressed in the female. Type spectEs. Phytoecia femoralis Haldeman (monobasic). This genus is distinctive from others in its tribe by the proportions of the antennal segments, the frequent presence of dorsal calluses on the pronotum, the shape of the metepisternum, and by the nature of the impressions of the last abdominal sternite, which are concave in the male and linear in the female. Many of the species resemble Saperda but the bifid claws will readily separate them. There are at present 15 known species of Mecas assignable to three subgenera, all occurring in the New World. 148 2(1). 4(3). 5(4). 6(5). 7(6). 8(4). CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. KEY TO THE KNOWN SPECIES OF THE GENUS Mecas Pronotum with sides rounded or subparallel; elytra not expanded apically behind! middle; appearance’ not ly ciionn) 22 Ea 2 Pronotum with sides obtusely produced at middle; elytra expanding slightly toward apices; integument yellow and black; appearance lycid-like. Mecas (Pannychis). 14-20 mm. Chihuahua to Veracruz _..__ SS M. sericea Pronotum not densely fringed with short, erect, golden pubescence, erect hairs moderately long; elytral apices broadly rounded or rotundate-truncate; abdomen with pubescence of sternites unicolorous. Mecas, sensu stricto. 3 Pronotum densely fringed with short, erect, golden pubescence with scattered long setae intermixed, dorsal surface with a pair of longitudinal vittae on each side of middle composed of short, appressed, golden pubescence; integument usually concolorous golden yellow, less commonly mottled or vittate with black, rarely wholly black; elytral apices angulate, obliquely truncate or emarginate; abdomen often with last three sternites margined laterally with longitudinal bands of yellowish-white pubescence suggesting luminescent organs of a lampyrid. Mecas (Dylobolus). 8-14 mm. South- western WUmited states! to) Costar Rica ses ee eee M. rotundicollis Pronotal pubescence dense, appressed, obscuring surface, intact or broken by well defined black polished callosites and usually also a median impunctate area. On ISK... i225 22 oe eee 4 Pronotal pubescence very sparse, erect, not obscuring surface which has five tubercles, one median, two antemedian, and two lateral, the surface red to yellow, concolorous or margined laterally with black and/or with black spots on the median or median and lateral tubercles. 8-10 mm. Sinaloa and San) JouismPotosiatoy Guatemalae = ee ee M. obereoides Pronotum with pubescence intact, not interrupted by polished black callosities. 5 Pronotum with at least two polished black callosities and usually a median elongate impunctate “area on) isk) eee 8 Pronotum and elytra with concolorous pubescence which obscures the surface; Stemnum) juniformly, densely, pubescent) = ———————E—EEs 6 Pronotum and elytra with longitudinal bands of dense, appressed, often yel- lowish pubescence at middle and sides, remaining pubescence not completely obscuring surface; sternum margined with a row of dense, appressed, yellowish pubescence. Length, 6.5-8 mm. Southeastern United States to IN GW, VICxXI COM == 28 ee ee eee M. marginella Tarsal claws with inner tooth much smaller than outer one. Smaller species, 6=0iimm-ain length: (fig. Ub), 2-2 Ee ee if Tarsal claws with inner tooth almost as long as outer one. Length, 10-14 mm. INansasMto.wMexas) (lig. 1a)).p «tet Eee M. confusa Femora always reddish; pubescence finer, not completely obscuring surface. Length, 6-8 mm. Southeastern United States. SS M. femoralis Femora always black; pubescence thick, obscuring surface. Length, 6-11 mm. Southeastern United States to Arizona, Colorado, and northeastern Mexico, etait AS ata cote he le ee M. cineracea ANCA: <2 82... 5 eee 9 Pronotum with two rounded glabrous calluses in addition to elongate median imnipunctate area. ee EEE EE = UG) VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 149 a b Cc FicurE 1. Some variations in the form of the tarsal claws in the genus Mecas. 9(8). Form elongate, elytra about 3 times as long as broad; antennae at least as long as body; elytra uniformly pubescent, margins and suture not distinctly pubes- cent. Length, 8-13 mm. San Luis Potosi to Morelos and Nayarit. __. M. cinerea Form rather stout, short, elytra about 244 times as long as broad; antennae shorter than body; elytra with margins and suture densely pale pubescent. Length, 6-12 mm. Great Plains to southeastern United States, New Mexico; and northeastern: Wiexico. 2 == ene M. pergrata 10(8). Tarsal claws with inner tooth short, obtuse, lobe-like (fig. Ic), 0. 11 Marcoleclaws with inner tooth acute: spine=likes eee ee 12 11(10). Antennae shorter than body, densely clothed beneath with long curved hairs, segments robust, flattened; elytra uniformly gray pubescent. Length, 10.5- AS emiinen Gea Wat oven! @Uuereter@: se eee eee M. cirrosa Antennae longer than body, sparsely clothed beneath with long erect hairs, segments slender; elytra broadly clothed with brownish pubescence along epipleurae; humeri glabrous. Length, 10.5-13 mm. Jalisco. —___ M. humeralis 12(10). Antennae much shorter than body, outer segments short. — 13 Antennae at least as long as body, outer segments elongate. = ti«i2“SH 13(12). Appressed pubescence gray, uniform on elytra; pronotum ieee Sains punctate; elytra lacking long erect dark hairs over apical one half. Length, 10-15 mm. Washington to northern Baja California and Colorado. __ M. bicallosa Appressed pubescence brownish, denser on margins and suture of elytra; pronotum coarsely, confluently punctate; elytra densely clothed with dark erect hairs over apical one half. Length, 9-10 mm. Durango to Mexico. tet ee, | a a ee ee eee M. ambigena 14(12). Elytra sparsely, separately punctate, lacking long erect hairs; pronotal calluses Siiallbemediansampunctates areas Via Sle. ee 15 Elytra coarsely, contiguously punctate, densely clothed with long erect hairs; pronotal calluses large, median impunctate area distinct. Length, 8-13 mm. AmizonaeiomNayanitrandmDistrito) Hecdetals seems sess ss een M. menthae 15(14). Elytra with distinct longitudinal pubescent bands along margins and suture. encth-— 10-16) mma shloriday = ee M. cana cana Elytra uniformly grayish or yellowish pubescent, suture and margins without pubescent bands. Length, 10-16 mm. Eastern United States to South Dakotasangenortheastenmel lexi Cp eee M. cana saturnina Subgenus Pannychis Thomson Pannychis THomson, 1864, Systema cerambycidarum, p. 127;1868, Physis, vol. 2, p. 197; LAcorDaIRE, 1872, Genera des coléoptéres, vol. 9, pp. 882, 889; BaTEs, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205; GimMmour, 1962, Rev. Biol. Trop., vol. 10, pp. 125, 137. 150 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. XG : Ss Ficure 2, Mecas (Pannychis) sericea (Thomson), ¢. VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 151 Form moderate to large, lycid-like. Antennae short, not extending beyond third abdominal segment, segments not gray pubescent. Pronotum broader than long, sides obtusely produced at middle; disk convex, sparsely punctate, pubes- cence not obscuring surface. Elytra slightly expanded behind middle, disk costate, pubescence bicolored. Legs with tarsal claws with long inner tooth. Abdomen with last sternite impressed at apex in females. Type SPECIES. Pannychis sericeus Thomson (monobasic). The lycid-like form and coloration and the obtusely produced pronotal sides make this subgenus very distinctive. A single species is known from Mexico. Mecas (Pannychis) sericea (Thomson). (Figures 2, 3.) Pannychis sericeus THOMSON, 1864, Systema cerambycidarum, p. 127; 1868, Physis, vol. 2, p. 197; Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205; Gimrmour, 1962, Rev. Biol. Trop., vol. 10, p. 137. Pannychis ducalis BATEs, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205; Gitmour, 1962, Rev. Biol. Trop., vol. 10, p. 137. New synonymy. Mate. Form moderate sized to rather large, elytra slightly expanding toward apices; integument yellowish, antennae black, legs variable, tibiae usually black, femora often partially black, head usually with 3 black bands over vertex and behind eyes, pronotum usually with 3 longitudinal bands joining at base and broad black bands at sides beneath, elytra yellow or with basal and broad apical black bands, basal bands often reduced or lacking, beneath variably colored. Head with front convex, coarsely, shallowly punctate; interantennal area con- cave, median line deep; vertex shallowly punctate; pubescence dense, appressed on cheeks, sparse, short, and suberect on front; antennae extending to about third abdominal segment, segments all clothed with very short, dark, depressed pubescence, basal segments with a few long erect setae beneath, third segment subequal in length to first, fourth shorter than third, remaining segments grad- ually decreasing in length, eleventh appendiculate, segments from fifth with vague longitudinal poriferous areas. Pronotum broader than long, sides obtusely produced at middle; disk convex, almost impunctate; pubescence dense, golden, appressed at sides, a little sparser on disk with a few long, erect hairs inter- spersed; prosternum sparsely pubescent; meso- and metasternum moderately densely pubescent, distinctly punctate at sides. Elytra about 3 times as long as broad, slightly expanding behind middle; each elytron lightly bicostate, epipleurae vertical; punctures at base coarse, dense, becoming fine and sparser toward apex; pubescence short, dense, subdepressed, colored as integument, longer suberect hairs present basally; apices rounded. Legs finely pubescent; tarsal claws with inner tooth slightly shorter than outer. Abdomen finely moderately pubescent; last sternite deeply impressed for most of its length. Length, 14-19 mm. FEMALE. Form more robust, elytra slightly more expanded behind middle. ‘(uosmoy) varzdas (s1yIKuung) SpIayY JO BOUAIINIIO UMOUY “¢ AWAY ‘uolsstded Aq paonpoiday ‘osPITYD Jo AjIsIOATUL) 24} Aq L¢61 WysAdoo dew aseg Sir NOILD3°Ou¥d V3NY-1VNDI S/3NNOB YNOLNOD 1004 OOO! » S¥313NOTIM OOF ooc o0z 201 = [Proc. 4TH SER. = > s37/Mo0e 00; Se 90) 31vos CALIFORNIA ACADEMY OF SCIENCES 152 VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 153 Antennae extending to about first abdominal segment. Abdomen with last sternite linearly impressed, concave at apex, apex broadly V-shaped. Length, 14-20 mm. TYPE LocALITy. Of ‘sericeus,’ Mexico; ‘ducalis,’ Orizaba, Mexico. RANGE. Chihuahua, Mexico to Veracruz (fig. 3). FLIGHT PERIOD. July to October. Remarks. This striking species undoubtedly is one of the lycid mimics. It is variable in coloration and, as is found among members of Elytroleptus (Chemsak and Linsley, 1965), all yellow individuals occur as well as ones with only the apices of the elytra black or with both the apices and base black. It would not be surprising to encounter individuals with all black elytra. MATERIAL EXAMINED. MEXICO. Chihuahua: 1 2, 3 miles SE. of Temoris, VII-25-69 (R. C. Gardner, C. S. Glaser, T. A. Sears); Jalisco: 1 ¢, 10 miles SW. of Tecalitlan, X-10-64 (A. E. Michelbacher); 1 ¢, Cuidad Guzman, Jalisco, IX-15-63 (M. C. Colorado); Morelos: 1 4, Tequesquitengo, VII-15-61 (R. & K. Dreisbach); Michoacan: 1 2, near Morelia, IX-5-52 (G. H. Dieke); Guerrero: 1 2, Iguala, VII-21-62 (H. E. Milliron); 1 2, 10 miles N. of Mercula, VIII-26-58 (E. L. Mockford); 1 6, Thaxmalac, IX-22-42 (W. F. Fosberg) ; Puebla: 1 6, 19 miles NW. of Cacaloapan, VII-30-65 (W. A. Foster). Subgenus Dylobolus Thomson Dylobolus THomson, 1868, Physis, vol. 2, p. 195; LAcorpatrE, 1872, Genera des coléopteéres, vol. 9, pp. 897, 900. Form slender, elongate. Antennae slender, third segment slightly curved. Pronotum with sides rounded, disk densely fringed with short, erect, golden pubescence. Elytra with apices angulate, usually obliquely emarginate. Legs with tarsal claws with inner tooth almost as long as outer one. Abdomen frequently with yellowish appressed pubescence at sides of apical sternites. Type species. Dylobolus rotundicollis Thomson (monobasic). This subgenus differs from the others by the pubescence of the pronotum and emarginate or truncate elytral apices. The single known species is a lampyrid mimic. Mecas (Dylobolus) rotundicollis (Thomson). (Figures 4, 5, 6.) Dylobolus rotundicollis THomson, 1868, Physis, vol. 2, p. 196. Mecas rotundicollis, Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 148. Mecas ruficollis Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205; Lenc and Hamirton 1896, Trans. American Ent. Soc., vol. 23, pp. 152; CAsEy, 1913, Memoirs on the Coleoptera, vol. 4, Das02: Mecas ruficollis morpha mediomaculata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 149. New synonymy. 154 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 4. Mecas (Dylobolus) rotundicollis (Thomson), ¢. VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 155 Mecas rotundicollis morpha ruficollis, BREUNING, 1955, Mem. Soc. Roy. Belgique, vol. 27, p. 149. Mecas laticeps Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204; BrREUNING, 1955; Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 151. New synonymy. Mecas laticeps morpha sutureflava BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 151. New synonymy. Mecas laticeps morpha mediopunctata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 151. New synonymy. Mecas mexicana BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204. Mecas rotundicollis morpha mexicana, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 149. Mecas vitticollis CAsEY, 1913, Memoirs on the Coleoptera, vol. 4, p. 362. Mecas laticeps morpha vitticollis, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, Deez. Mate. Form moderate sized, elongate, sides parallel; color black, head and pronotum orange, usually with dark spots or bands, legs often orange, thoracic sterna often orange, abdomen usually with broad bands of yellow appressed pubescence at sides of last three sternites, elytra frequently with narrow bands of appressed yellowish pubescence down suture and epipleurae. Head rather small; front convex, median line extending from clypeus to neck; interantennal area concave; vertex moderately coarsely, densely punctate; pubescence dense, yellowish, appressed with short, dark, erect hairs numerously interspersed; antennae shorter than body, scape finely, very densely punctate, third segment longer than first, fourth shorter than third, fifth subequal to first, segments from sixth gradually decreasing in length, scape rather densely clothed with short subdepressed hairs, underside of segments densely clothed with short, pale, appressed pubescence, basal segments with a few long erect hairs beneath. Pronotum usually broader than long, sides rounded; disk convex, sparsely to rather densely punctate; pubescence usually dense, consisting of short, dense, subappressed, longitudinal bands, one on each side of middle and at lateral margins, longer erect hairs numerously interspersed; prosternum densely pubes- cent; meso- and metasternum finely, densely punctate at middle, coarsely at sides, pubescence dense. Elytra over three times longer than broad; suture and epipleurae usually with narrow bands of appressed yellow pubescence; punctures rather coarse, dense, becoming finer and sparser toward apex; surface clothed with fine appressed pubescence, short, recurved hairs numerously inter- spersed, apices obliquely truncate. Legs finely, densely pubescent; tarsal claws with inner tooth almost as long as outer. Abdomen elongate, densely pubes- cent; last three sternites usually clothed with broad yellow bands at sides; last sternite deeply impressed for most of its length. Length, 9-16 mm. FEMALE. Form similar, more robust. Antennae slightly shorter than in male. Abdomen with last sternite linearly impressed for its entire length; last tergite strongely, obtusely conical at apex. Length, 10-19 mm. 156 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. to 8) 4 ~ ise) u —- a >} = SI S ae} > — 3 u vo ~~ {a} 4 Orr uw) vu !- ~~ ee g & a > Eg om po 3:} Sl 4s me fe io) =o one Sa Sto Si ~~ 2 oS a o ee el rare OH Lr mE So no» See 6b wo Ee = 8 fe i na gy = wz Oo = 2's my fy oO a) ~ VoL. XX XIX] CHEMSAK AND LINSLEY: MECAS 157 Type LocaLity. Of ‘rotundicollis, Mexico; ‘ruficollis, Texas; ‘laticeps,’ Guanajuato, Mexico; ‘mexicana,’ Izucar, Mexico; ‘vitticollis, Durango City; ‘mediomaculata, Guerrerro; ‘sutureflava, Temax, Yucatan; ‘mediopunctata,’ Mexico. RANGE. Oklahoma to Arizona, Texas, and south to Costa Rica (fig. 6). FLIGHT PERIOD. May to December. Host pLrants. Adults have been collected on flowers of Guardiola tulocarpa (Compositae) and on Eysenhardtia polystachya (Leguminosae). Remarks. This species is a lampyrid mimic, resembling different models in different parts of its range, as is evident by the polychromatism expressed in the specimens at hand. These color differences are expressed especially in the presence or absence of yellowish longitudinal sutural and epipleural bands on the elytra and in the yellowish apical sternites of the abdomen. There are varying combinations of these characters but we have been unable to correlate them geographically, although this might be possible with larger series of specimens and model-mimic associations over the entire range. In addition to the above differences, individuals also vary considerably in size, color of head, pronotum, sternum, and legs. MATERIAL EXAMINED. UNITED STATES. Oklahoma: 1 ¢, 1 2, Edmond, VII-9-57 (D. Alexander); Arizona: 2 ¢4, Dry Canyon, Sands Ranch, SE. end Whetstone Mts., Cochise Co., VII-10-52 (H. B. Leech, J. W. Green) ; 1 2, Tombstone, VIII-13-40 (E. S. Ross); 1 2, Madera Canyon, Pima Co., IX-1-54 (Menke & Stange); 1 6, 5 mi. W. Portal, Chiricahua Mts., VIII- 12-58 (P. Opler). Texas: 1 6, New Braunfels; 1 ¢, 1 2, San Antonio, May, VI-11-36 (C. D. Orchard); 1 4, 13 mi. W. of San Marcos, Comal Co., VI-24- 25-61 (R. L. Westcott); 1 ¢, Brownsville; 1 ¢, Van Horn, V-23-32 (E. G. Linsley). MEXICO. Tamaulipas: 1 ¢, Ciudad Victoria, VI-8-51 (P. D. Hurd); 1 ¢, Rio Corona, 21.3 mi. N. of Cuidad Victoria, X-25-65 (G. E. Ball, D. R. Whitehead). Nuevo Leon: 1 6,2 22, Monterrey, XI-30-65, X-12-52. San Luis Potosi: 1 2°, El Huizache, VIII-22-61 (R. & K. Dreisbach). Veracruz: 3 66,1 2, Veracruz, XI-1-57, IX-24-61 (R. & K. Dreisbach). Chihuahua: 1 ¢, 3 mi. E. of Temoris, VIII-26-69 (Sears, Gardner, Glaser). Durango: 1 ?, Encino, VII-27-47, 6200 ft. (Schramel); 14, 11 mi. W. of Durango, VIII-2-64, 7000 it. (L. Kelton); 1 6, 7 mi. W. of Durango, VII-23-64 (W. R. M. Mason); 1 4, 25 mi. S. of Durango, VII-24-64 (L. Kelton); 1 6,2 22,8 mi.S. of Canutillo, VII- 9-51, on flowers of Guardiola tulocarpa (P.D. Hurd). Hidalgo: 1 4, Ajacuba, VI- 21-37 (M. A. Embury); 2 66,2 22, Pachuca, VI-15-35, VII-10-37 (Embury), VII-31-35 (Embury); 1°, Zimapan, VII-14-68 (M. Wasbauer, J. Slansky). Dis- trito Federal: 2 22, Temascaltepec, 1931 (G. B. Hinton). Nayarit: 6 6¢, 5 2 2, Tepic, IX-13-57 (R. & K. Dreisbach), IX-15-17-53 (B. Malkin), IX-24-47 (B. Malkin); 1 2, Campostella, IX-16-57 (R. & K. Dreisbach). Jalisco: 1%, 13 158 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. SCALE 200 600 800 Jo0o MILES = T om oS 400 600 800 1000 1200 1400 KILOMETERS LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION Base map copyright 1961 by the University of Chicago. Reproduced by permission. Ficure 6. Known occurrence of Mecas (Dylobolus) rotundicollis (Thomson). VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 159 mi. N. of Chapala, VIII-1-63 (P. J. Spangler); 1¢, San Juan Lagos, VII-27-51, on Eysenhardtia polystachya (P. D. Hurd); 1 2, Tlaquepaque, VI-28-45 (N.C. H. Krauss); 1 2, Ajijic, VIT-28-54 (Cazier, Gertsch, Bradts); 1 6, 17 mi. SSW. of Guadalajara, VIII-22-70 (M. & J. Wasbauer). Colima: 1 ¢, Pine zone, SE. Slope Mt. Colima, XII-48 (E. S. Ross). Michoacan: 1 ¢, 5 km. W. of Zacapu, VII-13-51 (H. E. Evans); 1 ¢, Morelia, [X-30-45; 1 ¢, 6 mi. NW. of Quiroga, VII-11-63 (Parker, Stange); 1 4, 12, Quiroga, VII-15-56 (R. & K. Dreis- bach); 2 6¢, 1 @%, Tuxpan, ITX-6-03 (McClendon), IX-18-57 (Scullen). Morelos: 2 ¢¢,1 2, Hujintlan, VIII-22-56 (R. & K. Dreisbach); 1 2, Lake Tequesquitengo, IX-13-57 (Schullen); 2 ¢4, Cuernavaca, X-02, X-29-57 (Dreisbach); 1 2, Morelos, Oct. Guerrero: 1 2, Rio Balsas (Wickham). Puebla: 1 6, 3 mi. N. of Petalcingo, VIII-21-63 (Parker, Stange); 1 6, 55 mi. S. of Acatlan, VII-30-63 (J. Doyen); 1 ¢, Tehuacan, VI-23-51 (Evans); 1 4, 19 mi. NW. of Cacaloapan, VII-30-63 (Foster). Oaxaca: 8 44,1 2, Oaxaca, VII-20-37 (Embury); 2 64,1 2, Oaxaca, IX-13-20-47 (Malkin); 1 4,1 2, Monte Alban, VIII-3-54 (P. & C. Vaurie) , X-12-63 (A. E. Michelbacher) ; 2 2 2, 18 mi. NW. of Totolapan, VII-28-63 (Doyen, Foster); 1 2, Temescal, VII-5 -65 (G. H. Nelson). Chiapas: 1 ?, Jct. Hwys. 190-195, VI-6-69 (H. F. How- den); 3 64,2 22, San Jeronimo, Volcan Tacana, X-1-10-70, XI-7-70 (E. C. Welling). GUATEMALA. 1 2, El Salto, Escuintla, 1934 (F. A. Bianchi). COSTA RICA. 1 ¢, La Pacifica, 4 km. NW. of Canas, Guanacaste, XI-17-71 Gera. Opler). Subgenus Mecas sensu stricto Mecas LEConreE, 1852, Jour. Acad. Nat. Sci. Philadelphia, (2) vol. 2, p. 155; 1873, Smith- sonian Misc. Coll., vol. 11, no. 265, p. 347; BratcHLEy, 1910, Coleoptera—in Indiana, p. 1090; Knutr, 1946, Ohio Biol. Survey, Bull. vol. 39, p. 274. Form moderate sized, parallel; body usually densely clothed with appressed pubescence. Pronotum with or without dorsal calluses, sides broadly to narrowly rounded. Elytra parallel, apices rounded, disk not costate. Legs with tarsal claws variable, inner tooth long or short. TypeE species. Phytoecia femoralis Haldemen (monobasic). The members of this subgenus are easily recognizable by the densely pubes- cent body, subcylindrical and usually densely pubescent pronotum which fre- quently has glabrous dorsal calluses. Thirteen species are presently known. Mecas obereoides Bates. (Figure 7.) Mecas obereoides BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204, pl. 15, fig. 16; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 150. Mecas laminata Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 150. [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 160 ‘sayegq Saploadago spdapy JO 9dUIAIINIIO UMOUY ‘/ ANSI ‘uoisstujad Aq psonpoiday ‘oseoIyD Jo AjIsIaAIUL_) 9Y} Aq LEG] JYsIAdoD dew ase gq NOILD3FO¥d W3Y¥—-WNDI S/3NNOB YNOLNOD 1004 000! Swaiano7iM oor s371M00e VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 161 Mecas laminata morpha rufobasalis BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 150. New synonymy. Mecas laminata morpha discopunctata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 150. New synonymy. Mecas laminata morpha discoimpunctata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 150. New synonymy. Mate. Form moderate sized, elongate, sides parallel; color black, vertex of head and pronotum orange, pronotum usually with three black spots, one median and two lateral. Head with front convex, densely clothed with gray appressed pubescence which covers basal half of mandibles, erect dark hairs numerous on front; interantennal area convex; vertex moderately coarsely punctate; antennae slightly longer than body, densely gray pubescent beneath with a few long erect hairs present on basal segments, scape slightly shorter than third segment, fourth equal to third, fifth shorter than first, segments six to nine subequal, remaining two segments shorter. Pronotum usually a little broader than long, sides narrowly rounded; disk convex, middle with a usually black glabrous callus, frequently a vague callus present on each side of middle anterior to median one, two black glabrous spots present laterally; punctation irregular, moderately coarse; pubescence sparse, fine with a few long erect hairs present at sides; prosternum densely clothed with gray recumbent pubescence; meso- and metasternum densely clothed with gray recumbent pubescence, sides moderately coarsely punctate. Elytra about three times as long as broad; each elytron costate down middle, area between costae and suture impressed; punctures coarse, dense, becoming finer toward apex; pubescence short, dense, appressed, with longer suberect hairs numerously interspersed; apices rounded. Legs densely pubescent; tarsal claws with inner tooth slightly smaller than outer. Abdomen densely clothed with gray appressed pubescence; last sternite shallowly impressed for most of its length. Length, 10-14 mm. FEMALE. Form similar, slightly more robust. Antennae about as long as body. Abdomen with last sternite linearly impressed for its entire length, shallowly concave at middle at apex. Length, 10-15 mm. Type LocaLity. Of ‘obereoides, Cuernavaca, Mexico; ‘laminata, not restricted; ‘rufobasalis, Mexico; ‘discopunctata, Mexico; ‘discoimpunctata,’ Mexico. RANGE. San Luis Potosi and Sinaloa, Mexico to Guatemala (fig. 7). FLIGHT PERIOD. June to December. Host pLants. Unknown. Remarks. The sparsely pubescent orange pronotum and the dark pronotal spots will readily separate this species from other known species of Mecas. The coloration is almost constant in the series at hand. However, specimens from the eastern parts of Mexico tend to have dark appressed elytral pubescence while in those from the western portions to Guatemala this tends to be grayish. MATERIAL EXAMINED. MEXICO. San Luis Potosi: 1 2, 30 mi. N. of 162 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Tamazunchale, X-6-57 (H. A. Scullen). Veracruz: 2 ¢6¢, 3 2%, Veracruz, VIII-1-6-61 (R. & K. Dreisbach); 1 2, Jalapa, IX-28 to X-3-61 (R. & K. Dreisbach); 1 ¢, 1 2, Puente Nacional, VIII-3-56 (R. & K. Dreisbach), VII- 23-24-65 (Flint & Ortiz); 2 22, San Andreas Tuxtla, X-25-57 (R. & K. Dreisbach); 1 ¢, 12, Hueyapan, X-30-57 (R. & K. Dreisbach); 1 4, Cerro Venado, Los Tuxtlas Range, XII-29-62 (Edmonds, Robinson). Sinaloa: 1 ¢, Venedillo, VII-31-18; 2 6 3, 20 mi. E. of Villa Union, VIII-19-64 (M. E. Irwin, E. I. Schlinger). Nayarit: 2 64,1 2,15 mi. N. of Tepic, VII-25-54 (Cazier, Gertsch, Bradts); 1 ¢, 1 2, 18 mi. N. of Tepic, VIII-16-60 (D. C. Rentz); 3 66, Tepic, [X-21-24-53 (B. Malkin), IX-13-57 (R. & K. Dreisbach); 1 ¢, 14 mi. E. of San Blas, VII-21-54 (Schlinger). Colima: 1 ¢, Colima (Conradt). Jalisco: 1 2, Puerto Los Mazos, 9 mi. NW. of Autlan, VIII-28-70 (M. & J. Wasbauer). Guerrero: 2 ¢¢4,1 2, Teloloapan, VIII-15-21-57 (D. Douglas) ; 1 6,1 2,3 mi. S. of Acahuizlotla, XI-17-46 (E. C. Van Dyke); 1 ¢, Taxco, VITI-16-18-56 (A. E. Lewis); 1 ¢, Highway 95, 23 mi. N. of Acapulco. VII- 30-65 (Cornell U. Mex. field party); 1 ¢, 5 mi. W. of Mex. 92, Cacahuamilpa Caves, VITI-16-18-56 ( A. E. Lewis). Morelos: 6 6¢,2 22%, Hujintlan, VII- 22-56 (R. & K. Dreisbach); 1 ?, Lake Tequesquitengo, IX-13-57 (H. A. Scullen); 1 ¢, Tequesquitengo, VII-15-61 (Dreisbach); 1 ¢, Xochicalco, VII-13-61 (Dreisbach); 5 64, 1 2, Cuernavaca, VII-15-52 (G. M. Boush), VITI-9-13-38 (L. Lipovsky), VIII-1-6-21 Sept. (Barrett), IX-8-90 (D. De- long); 1 6,1 2, 22 mi. S. of Cuernavaca, [X-10-57 (Scullen); 1 ¢, 45 mi. S. of Cuernavaca, IX-12-57 (Scullen). Puebla: 1 ¢, 5 mi. S. of Izucar de Mata- moros, VIII-1-63 (Parker, Stange). Oaxaca: 1 ¢, 10 mi. NE. of Huajuapan de Leon, VI-27-65 (Doyen). Chiapas: 1 ?, Santo Domingo, 15 mi. SE. of Simojovel, VII-8-15-58 (J. A. Chemsak); 6 64,9 2, San Jeronimo, Volcan Tacana, VIII-10 to X-12-70 (E. C. Welling). GUATEMALA. 1 2, El Salto, Escuintla, 1934 (F. Bianchi). Mecas marginella LeConte. Mecas marginella LECoNTE, 1873, Smithsonian Misc. Coll., vol. 11, no. 264, p. 239; Horn, 1878, Trans. American Ent. Soc., vol. 23, p. 152; Bratcutry, 1910, Coleoptera—in Indiana, p. 1090; CAsry, 1913, Mem. Coleoptera, vol. 4, p. 361; BreuNntNG, 1955, Mem. Soc. Roy. Belgique, vol. 27, p. 147. Mate. Form small, subparallel; color black, pronotum with three longi- tudinal bands of yellowish to whitish appressed pubescence, elytra with narrow bands of pale pubescence down suture and lateral margins. Head with front convex, deeply punctate, densely clothed with appressed pale pubescence, long erect hairs numerous; interantennal area broadly concave; vertex coarsely, densely punctate; antennae a little longer than body, very sparsely gray pubes- cent beneath, long erect hairs numerous on basal segments, third segment longer than scape, fourth subequal to third, fifth shorter than fourth. Pronotum Vot. XXXIX] CHEMSAK AND LINSLEY: MECAS 163 broader than long, sides subparallel; punctures moderately coarse, dense, calluses absent; pubescence dense, appressed, lateral bands broad, yellowish, median band narrower, usually whitish, remainder of surface finely pubescent, long, erect hairs numerous; prosternum densely pubescent; meso- and metasternum densely pubescent, rather coarsely punctate at sides, metepisternum yellow pubes- cent over posterior half. Elytra over twice as long as broad; punctures coarse, close, becoming finer toward apex; pubescence between longitudinal bands fine, appressed, with longer erect hairs numerously interspersed; apices rounded. Legs finely, densely pubescent; tarsal claws with teeth subequal in length. Abdomen densely pubescent, narrowly yellow at sides of apical sternites; last sternite deeply impressed for its entire length. Length, 6.5—-8 mm. FEMALE. Form and size similar. Antennae about as long as body. Abdomen with last sternite shallowly impressed near apex. Length, 6.5-8 mm. TYPE LOCALITY. Western States and Texas. RANGE. Southeastern United States to New Mexico. FLIGHT PERIOD. March to July. Host PLANts. Unknown. One specimen was collected on Colubrina texensis (Rhamnaceae) in Texas, but it is very unlikely that this shrub is a host. REMARKS. The absence of pronotal calluses and the distinctive pubescent bands make this species easily recognizable. Breuning (1955) states that the head, pronotum, and elytra are densely and very finely punctate but this is an illusion produced by the pubescence. Actually, they are coarsely punctate, the elytra less so apically. MATERIAL EXAMINED. Twenty-one specimens from South Carolina, Alabama, Texas, and New Mexico. Mecas confusa Chemsak and Linsley, new species. (Figure 8.) Mecas inornata (not Say), BLANCHARD, 1887, Ent. Amer., vol. 3, p. 86; Horn, 1888, Trans. Amer. Ent. Soc., vol. 15, p. 301; Lenc and HAmirton, 1896, Trans. Amer. Ent. Soc., vol. 23, p. 152; BratcHiEy, 1910, Coleoptera—in Indiana, p. 1090. Mate. Form moderate sized, subparallel; color black, body densely clothed with thick, grayish, recumbent pubescence which obscures the surface. Head with front convex, finely densely punctate, darker suberect hairs short, about half as long as second antennal segment; interantennal area very shallowly con- cave; vertex sparsely punctate, large punctures well separated; antennae about as long as body, scape finely gray pubescent, remaining segments to ninth gray pubescent beneath, third segment longer than scape, fourth shorter than third, fifth shorter than first, remaining segments gradually decreasing in length. Pronotum broader than long, sides rounded, base impressed; disk convex, calluses absent; large deep punctures irregular, well separated, each [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 164 Se Sa PS Se == = SS a ee a ee é. sley, Mecas confusa Chemsak and Lin FIGURE 8. VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 165 puncture bearing a long erect hair; pro-, meso- and metasterna densely clothed with recumbent pubescence which obscures the surface. Elytra less than 2% times as long as broad; punctures coarse, close, linearly arranged, becoming obsolete at apex; recumbent pubescence completely obscuring surface, base with numerous rather short suberect hairs, these becoming shorter and recurved toward apex; apices obliquely subtruncate. Legs very densely pubescent; tarsal claws with inner tooth almost as long as outer. Abdomen very densely pubes- cent; last sternite impressed for its entire length. Length, 10-13 mm. FEMALE. Similar in form and size. Abdomen with last sternite linearly impressed, apex broadly V-shaped. Length, 10-14 mm. MATERIAL EXAMINED. Holotype male (California Academy of Sciences) from Luling, Gonzales Co., Texas, V-3-53 (B. J. Adelson). Allotype from Gonzales, Gonzales Co., Texas, V-2-53 (M. Wasbauer). Paratypes as follows: 2 64, same data as holotype; 1 2,1 2, same data as allotype; 1 2, Ft. Sam Houston, Texas (C. Grant); 1 ?, Corpus Christi, Texas, VI-28-42 (E. S. Ross) ; 2 64,1 2, Palmetto State Park, Gonzales Co., Texas, IV-11-53 (M. Wasbauer), V-10-53 (B. J. Adelson); 1 6, Lee Co., Texas, May (R. Oertel); 1 2, Hidalgo Co., Texas, VI-22-33 (S. Bromley). Other material, not paratypical includes: 1 2, Texas (F. Blanchard collection); 1 2°, Lee Co., Texas, June; 1 2, 49 mi. N. of Raymondville, Kenedy Co., Texas, VI-30-61 (R. L. Westcott); 1 4,5 22, Lake Texoma, 2 mi. E. of Willis, Oklahoma, June, July, 1965 (R. M. Bohart) ; Que elark Co,, Kansas, June (¥. H. Snow). This species closely resembles M. cineracea but may be separated by its larger size, denser overall pubescence, shorter erect hairs on the front of the head, and by the structure of the tarsal claws. In M. confusa the inner tooth of the claws is elongate and almost as long as the outer one; in M. cineracea the tooth is very small. The two species are sympatric, at least in parts of Texas, but it is not now known whether they infest the same or different host plants. Mecas femoralis (Haldeman). Phytoecia femoralis HALDEMAN, 1847, Trans. American Philos. Soc., vol. (2)10, p. 59. Mecas femoralis, LeContE, 1852, Jour. Acad. Philadelphia vol. (2)2, p. 155; Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; Lenc and Hamirton, 1896, Trans. American Ent. Soc., vol. 23, pp. 152, 153; Casey, 1913, Memoirs on the Coleoptera, vol. 4, p. 360 (in.) ; BreuNtING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 143. Mate. Size small, subparallel; color black, femora reddish; pubescence grayish. Head with front convex, densely punctate, densely clothed with ap- pressed pubescence and numerous suberect, dark hairs; vertex rather coarsely, closely punctate, densely pubescent; antennae about as long as body, basal seg- ments sparsely gray pubescent beneath, long, erect hairs sparse, third segment longer than first, fourth shorter than third, fifth shorter than fourth. Pronotum about as long as broad; sides almost subparallel; disk convex, rather coarsely, 166 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. closely punctate; pubescence dense, short, appressed, long, erect hairs numerous; prosternum densely pubescent; meso- and metasternum densely clothed with re- cumbent pubescence, sides more coarsely punctate. Elytra about 2’ times as long as broad; punctures coarse, dense, becoming finer toward apex; pubescence dense, appressed, with longer erect hairs numerously interspersed; apices rounded. Legs moderately densely pubescent; tarsal claws with inner tooth very short. Abdomen densely pubescent; last sternite shallowly impressed over most of its length. Length, 6-8 mm. FEMALE. Very similar in size and shape. Abdomen with last sternite im- pressed over apical one-half. Length, 6-8 mm. TYPE LOCALITY. Not given. RANGE. Southeastern United States. FLIGHT PERIOD. May to July. Host PLANTS. Unknown. Remarks. Mecas femoralis can be recognized by its small size, rather uniform pubescence, lack of pronotal calluses, and reddish femora. This species appears to be rare in collections. The eleven specimens we have seen vary very little. MATERIAL EXAMINED. Eleven individuals from North Carolina to Florida have been seen. Mecas cineracea Casey. Mecas cineracea CAsry, 1913, Memoirs on the Coleoptera, vol. 4, p. 360; Vocr, 1949, Pan- Pacific Ent., vol. 25, p. 184. (record) Saperda cineracea, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 139. Saperda bicallosa BREUNING (not Martin), 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 140. Mate. Form rather small, parallel; color black, body densely clothed with gray recumbent pubescence. Head with front convex, appressed pubescence obscuring punctures, longer erect hairs numerous; interantennal area almost plane; vertex moderately coarsely, separately punctate; antennae about as long as body, basal segments finely gray pubescent beneath, long erect hairs decreasing in number toward apex, third segment longer than first, fourth shorter than third, fifth a little shorter than first, remaining segments gradually de- creasing in length. Pronotum broader than long, sides almost parallel, shal- lowly impressed at base; disk convex, without calluses, punctures coarse, separated; appressed pubescence obscuring surface, long erect hairs numerous; prosternum densely pubescent; meso- and metasternum densely pubescent, coarsely punctured at sides. Elytra about 2% times as long as broad; punctures coarse, contiguous, becoming finer toward apex; pubescence obscuring surface, long suberect hairs numerous near base; apices obliquely truncate. Legs very densely pubescent; tarsal claws with inner tooth small. Abdomen densely VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 167 pubescent; last sternite shallowly impressed for its entire length. Length, 6-10 mm. FEMALE. Similar in form and size. Antennae shorter than body. Abdomen with last sternite linearly impressed. Length, 7-11 mm. TYPE LOCALITY. Harris Co., Texas. RANGE. Southeastern United States to Arizona and Colorado and _ north- eastern Mexico. FLIGHT PERIOD. April to August. Host pLants. Helenium microcephalum, Baileya multiradiata. Remarks. This species may be recognized by its small size, lack of pronotal calluses, uniform coloration and pubescence, and by the small inner tooth of the tarsal claws. It was incorrectly transferred to Saperda by Breuning (1955). A series of specimens from western New Mexico and Arizona have thicker pubescence than the Texas examples. However, there appears to be a gradient in this character from east to west as is apparent in the material at hand. MATERIAL EXAMINED. More than 200 specimens from South Carolina to Florida, to Arizona and Colorado. Mecas cinerea (Newman). (Figure 9.) Saperda cinerea NEWMAN, 1840, Entomologist vol. 1, p. 13. Mecas cinerea, GAHAN, 1888, Trans. American Ent. Soc., vol. 15, p. 300. Mecas senescens BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 203; CAsEy, 1913, Memoirs on the Coleoptera, vol. 4, p. 360. Mecas rubripes BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 203; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 140, 143. New synonymy. Mecas rubripes morpha callosoreducta BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 140, 143. New synonymy. Mecas inornata GAHAN, 1888 (not Say), Trans. Amer. Ent. Soc. vol. 15, p. 300. Mate. Form moderate sized, elongate, slender, parallel; color black, femora and/or elytra often reddish, pronotum occasionally with a median longitudinal reddish band; pubescence gray to yellowish, dense, appressed, long erect hairs fairly numerous. Head with front convex, finely separately punctate; inter- antennal area slightly concave; vertex deeply, separately punctate; pubes- cence dense, obscuring surface, long erect hairs moderately numerous; an- tennae slightly longer than body, segments gray pubescent beneath, basal segments with a few long erect hairs beneath, third segment longer than first, fourth subequal to third, fifth equal to first, remaining segments gradually decreasing in length, eleventh subacute at apex. Pronotum slightly broader than long, cylindrical; disk with two small glabrous calluses on each side of basal median elongate callus; punctures moderately coarse, separated at middle and subconfluent at sides; appressed pubescence obscuring surface, “(UBUUMIN) Dasau29 SspIaPyY JO 9dUGIINIIO UMOUY “6 AMNSIA ‘uolsstwed Aq poonpoiday ‘“osvotyD Jo AyisiaAtuy) ay} Aq /¢E6T IYSIUAdOO deur aseg NOILD3FO8d V3IE¥-IWNDI S,3NNOB YNOLNOD 100d 0001». su2LIROM oor [Proc. 4TH SrEr. EIN oe CALIFORNIA ACADEMY OF SCIENCES 168 VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 169 denser at middle and forming a vague longitudinal band, long erect hairs fairly sparse, shorter erect hairs more numerous; prosternum densely pubescent; meso- and metasternum densely pubescent, rather densely punctate at sides. Elytra usually about 3 times as long as broad; punctures coarse and contiguous basally, becoming finer and sparser toward apex; pubescence short, dense, appressed, long erect hairs numerous basally, becoming shorter and suberect toward apex; apices rounded. Legs moderately densely pubescent; tarsal claws with inner tooth slightly shorter than outer. Abdomen densely pubescent; last sternite rather shallowly impressed for its entire length. Length, 8-11 mm. FEMALE. Form similar. Antennae about as long as elytra. Pronotum more transverse, distinctly broader than long. Abdomen with last sternite linearly impressed. Length, 8-13 mm. TYPE LOCALITY. Of ‘cinerea,’ Mexico; ‘senescens,’ Puebla; ‘rubripes,’ Mex- ico; ‘callosoreducta,’ Mexico. RANGE. San Luis Potosi to Morelos and Nayarit (fig. 9). FLIGHT PERIOD. June to September. REMARKS. The number of calluses on the pronotum, elongate body form, and the length of the antennae distinguish this species from the other Mexican species of Mecas. The body and antennal length, and different tarsal claws separate it from M. pergrata. It is difficult to detect a definite geographical variational trend from the material available for study. The more northern individuals are all black while most of the southern specimens possess reddish femora and often, reddish elytra. Two individuals from Nayarit have a reddish longitudinal band on the pronotum and yellowish epipleurae. MATERIAL EXAMINED. MEXICO. San Luis Potosi: 2 646, 17 miles W. of Xilitla, 4700 ft., VII-22-54 (Univ. Kansas Mex. Exped.); Hidalgo: 1 4,2 92, 7 miles NE. of Zimapan, VIII-15-58 (H. F. Howden); Distrito Federal: 5 64, 7 22, Temascaltepec, 1931 (G. B. Hinton); 1 ¢, 2 22, Real de Arriba, Temascaltepec, VII-32 (H. E. Hinton), VII-33 (Hinton and Usinger); 2 4, 5 22, Tejupilco, Temascaltepec, VI-VII-33 (Hinton and Usinger); Puebla: 1 2,15 miles S. of Puebla, 6200 ft., IX-6-57 (H. A. Scullen); 1 4, Atlixco, VII-23-56 (R. & K. Dreisbach); Mexico: 14,12, 3 miles N. of Valle de Bravo, VI-28-29-65 (G. H. Nelson); Morelos: 7 é 6, Cuernavaca VII-6-38, VII-29-61, 7000 ft. (R. & K. Dreisbach); 1 ¢, 12, Cuernavaca-Acapulco Road, VIII-22-36 (Ball & Stone); 1 ¢, 4 miles E. of Cuernavaca, 6000 ft., VI-25-59 (H. E. Evans); 1 6, 7 km. E. of Cuernavaca, 5700 ft., VIII-11-62 (Evans); Nayarit: 1 6,1 2, La Mesa de Nayarit, VII-21-55 (B. Malkin). Mecas pergrata (Say). Saperda pergrata SAy, 1824, Jour. Acad. Philadelphia, vol. 3, p. 407; HaLpEMAN, 1847, Trans. American Philos. Soc., vol. (2)10, p. 55; LEConTE, 1859, Complete Writings of Thomas Say, vol. 2, p. 190. 170 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Stenostola pergrata, HALDEMAN, 1847, Proc. American Philos. Soc., vol. 4, p. 373; LEConreE, 1852, Jour. Acad. Nat. Sci. Philadelphia, vol. (2)2, p. 154. Mecas pergrata, Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; Lenc and Hamiton, 1896, Trans. American Ent. Soc., vol. 23, pp. 152, 153; BLarcHLry, 1910, Coleoptera—in Indiana, pp. 1090, 1091; CAsry, 1913, Memoirs on the Coleoptera, vol. 4, p. 361; CRAIGHEAD, 1923, Dom. Canada Agr. Bull., vol. 27, p. 138; Knut, 1946, Ohio Bio. Survey, Bull. vol. 39, pp. 274, 275, pl. 22, fig. 86; BrEUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 140, 144, fig. 1. Mecas pergrata morpha semiruficollis BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 140, 145. New synonymy. Stenostola gentilis LECONTE, 1852, Jour. Acad. Nat. Sci. Philadelphia, vol. (2)2, p. 154. Mecas discovittata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 140, 143. New synonymy. MALE. Form moderate sized, parallel; color black, femora pale reddish, elytra occasionally partly reddish; pubescence dense, short, recumbent, grayish. Head with front convex, punctures rather fine, well separated; pubescence dense, appressed, long dark erect hairs numerously interspersed; appressed pubescence thicker around eyes; vertex rather densely punctate; antennae shorter than body, segments gray pubescent beneath, outer segments annulate, third segment longer than first, fourth subequal to first, remaining segments gradually decreasing in length. Pronotum broader than long, sides slightly rounded; disk convex, four glabrous calluses present in addition to median callus; punctures rather sparse, scattered; apex and base usually with a narrow band of dense yellowish pubescence, remaining surface partially obscured, long, erect hairs numerously interspersed; prosternum densely pubescent; meso- and metasternum densely pubescent, densely punctate at sides. Scutellum densely clothed with yellowish recumbent pubescence. Elytra about 2’ times as long as broad; punctures rather coarse, contiguous at base, becoming finer toward apex; pubescence short, re- cumbent, partially obscuring surface, longer suberect hairs numerous, suture and lateral margins narrowly clothed with dense, yellowish, appressed pubescence; apices rounded. Legs finely, densely pubescent; femora reddish; tarsal claws with inner tooth small. Abdomen densely pubescent; last sternite shallowly im- pressed over most of its length. Length, 6-11 mm. FEMALE. Similar in form and size. Antennae extending to about second abdominal segment. Abdomen with last sternite linearly impressed. Length, 6-12 mm. TypeE LocaLity. Of ‘pergrata,’ Platte River, Nebraska; ‘gentilis,’ Missouri Territory; ‘semiruficollis, Texas; ‘discovittata,’ Colorado. RANGE. Great Plains to southeastern United States, New Mexico, and north- eastern Mexico. FLiGHT PERIOD. April to July. Host pLAnts. Aster (roots), Helianthus. Remarks. The five glabrous spots of the pronotum, reddish femora, and the Vor. XXXIX] CHEMSAK AND LINSLEY: MECAS 171 densely pubescent lines on the suture and lateral margins of the elytra will readily distinguish this species. In certain parts of the range, the elytra tend to be reddish down the disk and frequently the pronotum is also partially reddish. Hasits. According to Craighead (1923) larvae feed in the stems of Aster and down into the roots, completely hollowing the latter. Subsequently that portion of the stem of the plant breaks off at the surface of the ground. Small heaps of frass are exuded about the base of the plant. Only one larva is found in each stem. Mecas cirrosa Chemsak and Linsley, new species. Mate. Form moderate sized, robust; color black; pubescence dense, gray, recumbent. Head with front convex, rather finely, irregularly punctate; inter- antennal area rather deeply impressed; vertex coarsely, separately punctate; pubescence dense, appressed, long erect hairs numerous, pale and dark colored; antennae slightly shorter than body, segments robust, somewhat flattened, all segments gray pubescent beneath, dark brown above, segments from second densely fringed with long curved hairs beneath, third segment slightly longer than first, fourth a little shorter than third, fifth shorter than first, remaining segments gradually decreasing in length. Pronotum broader than long, sides broadly rounded; disk convex, each side with a small glabrous, premedian callus, narrow median callus extending almost length of disk; punctures moderately coarse, irregular, subconfluent; pubescence fine, appressed, long, pale, erect hairs numerous; prosternum densely pubescent; meso- and metasternum densely pubescent, minutely punctate. Elytra more than 2% times as long as broad; basal punctures moderately coarse, well separated, punctures becoming fine and quite sparse toward apex; pubescence fine, dense, appressed, basal margin with a few long, erect, pale hairs, remainder of surface with very short, suberect, dark hairs; apices rounded. Legs densely pubescent; tarsal claws with inner tooth short, obtuse, lobe-like. Abdomen densely pubescent; last sternite deeply impressed. Length, 13 mm. FEMALE. Form similar. Antennae shorter than body, curved hairs less numerous beneath. Abdomen with last sternite linearly impressed. Length, 10.5 mm. . TYPE MATERIAL. Holotype male (California Academy of Sciences) from 5 miles N. of Guanajuato, Guanajuato, Mexico, VII-25-54 (E. I. Schlinger). The specimen that we consider as the female of this species differs in several respects. The pubescence appears thicker and the antennae have a much sparser fringe. This individual is from Km. 320, near Hacienda Balvanera, Queretero, Mexico, VIT-13-55. Remarks. Although the tarsal claws are uniquely different from other 172 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. species of Mecas, we consider them to be bifid and the other morphological char- acteristics are similar enough to place ‘cirrosa’ in the genus. Mecas humeralis Chemsak and Linsley, new species. Mate. Form moderate sized, parallel; color black; pubescence dense, grayish and brownish. Head with front convex, rather finely, separately punctate; interantennal area impressed; vertex rather finely, separately punc- tate; pubescence dense, appressed, long, dark, erect hairs numerous; antennae about as long as body, outer segments slightly flattened, all segments gray pubescent beneath, dark brown above, long erect hairs sparse on basal segments, third segment much longer than first, fourth a little shorter than third, fifth equal to first, remaining segments gradually decreasing in length. Pronotum slightly broader than long, sides broadly rounded; disk convex, each side with a large glabrous callus before middle, median callus long, rather broad; punctures moderately coarse, subconfluent; pubescence dense, appressed, long erect hairs numerous; prosternum densely pubescent; meso- and metasternum densely pubescent, finely densely punctate at sides. Elytra about 3 times as long as broad; basal punctures coarse, contiguous, becoming finer and sparser toward apex; pubescence dense, recumbent, gray on disk, humeri glabrous, epipleurae and sides with a broad band of brown pubescence extending from under humeri almost to apex, long erect hairs abundant at base, becoming shorter and recurved toward apex; apices rounded. Legs densely pubescent; tarsal claws with inner tooth short, blunt, lobe-like. Abdomen densely pubescent, first three sternites with a narrow glabrous band at base; last sternite deeply impressed. Length, 13 mm. FEMALE. Form similar. Antennae about as long as body. Legs with tarsal claws having the short inner tooth slightly acute. Abdomen lacking glabrous lines on sternites, last sternite linearly impressed. Length, 10.5 mm. TYPE MATERIAL. Holotype male (California Academy of Sciences) from El Molino, Jalisco, Mexico, VII-10-56 (R. & K. Dreisbach); allotype from Guadalajara, Jalisco, VII-24-51 (P. D. Hurd). REMARKS. This species has tarsal claws similar to those of M. cirrosa. The different antennae and antennal pubescence will readily separate the two species. The glabrous humeri also make M. humeralis distinctive. Mecas bicallosa Martin. Mecas bicallosa Martin, 1924, Ent. News, vol. 35, p. 244; Hatcu, 1971, Univ. Washington Publs. Biol., vol. 16, p. 155. Saperda bicallosa, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 139, 140. Mate. Form moderate sized, parallel, rather robust; color black, body densely clothed with short, appressed, grayish pubescence. Head with front VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 173 convex, pubescence obscuring surface, long erect hairs very numerous; inter- antennal area plane, vertex deeply punctate; antennae extending to about third abdominal segment, segments through fourth gray pubescent, dark at apices, basal segments with numerous long, suberect hairs, segments from third with long hairs beneath, these decreasing in number toward apex, third segment longer than first, fourth subequal to first, remaining segments short, subequal in length. Pronotum broader than long, sides rounded; disk convex, with two glabrous calluses at middle and a smaller median one behind middle; punctures rather fine, deep, separated; pubescence obscuring surface, very long erect hairs numerous; prosternum densely pubescent, front coxal cavities narrowly open behind; meso- and metasternum densely clothed with recumbent and_sub- depressed pubescence. Elytra more than twice as long as broad; punctures at base coarse, dense, becoming finer toward apex; pubescence obscuring surface, long, suberect hairs abundant over basal half; apices rounded, often vaguely, obtusely toothed. Legs very densely pubescent; tarsal claws with inner tooth very small, short. Abdomen densely pubescent; last sternite shallowly, rather broadly impressed. Length, 10-13 mm. Femate. Form similar. Antennae slightly shorter. Abdomen with last sternite narrowly linearly impressed, apex shallowly concave. Length, 10- >) ma. Type LocaLity. Martins Springs, Lassen Co., California. RANGE. Washington to northern Baja California, to Colorado. FLIGHT PERIOD. April to August. Host PLaAnts. Artemisia tridentata. Remarks. The bicallused pronotum and abbreviated distal antennal seg- ments characterize this species. Breuning (1955) incorrectly synonymized M. bicallosa with M. cineracea Casey and transferred both to Saperda. MATERIAL EXAMINED. A total of 127 specimens were examined from Washington, Oregon, California, Nevada, Idaho, Utah, Colorado and Baja California. Mecas ambigena Bates. Mecas ambigenus BATES, 1881-1885, Biologia Centrali-Americana, Coleoptera, vol. 5, pp. 203, 426; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 142, 147. Mecas pseudambigena BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 142, 147. New synonymy. Mate. Form rather small, short, parallel; color black; pubescence dense, rather coarse, brownish, appressed, long dark erect hairs numerous. Head with front convex, finely densely punctate; interantennal area concave; vertex distinctly, separately punctate; pubescence short, appressed, front with numerous very long erect black hairs; antennae extending to about apical of elytra, 174 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. segments to tenth gray pubescent beneath, segments from fourth narrowly annulate at base, basal segments with a moderate number of suberect hairs beneath, scape robust, a little shorter than third segment, fourth subequal to first, remaining segments decreasing in length. Pronotum broader than long, sides broadly rounded; disk convex with a glabrous callus on each side before middle, elongate median basal callus often vague or absent; punctures coarse, confluent; pubescence short, appressed with numerous long erect hairs inter- spersed; prosternum densely pubescent; meso- and metasternum densely pubes- cent, finely densely punctate. Elytra less than 2% times as long as broad; punctures coarse, contiguous, becoming finer toward apex; pubescence mod- erately densely appressed, suture narrowly lined with dense yellow-brown pubescence, long erect hairs numerous over basal ’2, shorter and suberect toward apex; apices rounded. Legs densely pubescent; tarsal claws with inner tooth smaller than outer. Abdomen densely pubescent, finely punctate; last sternite deeply impressed for about ™ its length. Length, 9 mm. FEMALE. Similar in form and size. Antennae extending to about apical % of elytra. Abdomen with last sternite lightly linearly impressed. Length, 10 mm. Type LocaLity. Of ‘ambigena,’ Mexico; ‘pseudambigena,’ Mexico. RANGE. Durango to Mexico. FLIGHT PERIOD. June and July. Remarks. The shorter form and short antennae will distinguish M. ambigena from other species of Mecas. MATERIAL EXAMINED. MEXICO. Durango: 1 ¢, 25 miles W. of Durango, 7,500 ft., VI-23-64 (J. E. Martin); Zacatecas: 2 64, 4 miles W. of Monte Escobido, 7,800 ft., VII-19-20-54 (R. H. Brewer); Mexico: 1 ¢&, Toluca (Wickham). Mecas menthae Chemsak and Linsley, new species. (Figures 10, 11.) Mecas marginella, LINSLEY, KNULL, and STATHAM (not LeConte), 1961, Amer. Mus. Nov., nNOwZ20505 pas2- Mate. Form moderate sized, subparallel; color black; pubescence dense, short, appressed, grayish to gray-brown, long erect dark hairs numerous. Head with front convex, rather finely, separately punctate; vertex deeply, separately punctate; pubescence dense, short, appressed, antennal tubercles dark pubescent above, long erect hairs numerous on front and vertex; antennae slightly longer than elytra, segments to tenth gray pubescent beneath, segments from fifth narrowly pale annulate at base, long erect hairs fairly numerous beneath on basal segments, third segment longer than first, fourth shorter than third but longer than first, fifth equal to first, remaining segments gradually decreasing in length. Pronotum broader than long, sides broadly rounded; disk convex, each side of middle with a rather large, glabrous callus, middle with an elongate VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 175 = SRN RSs “ if} f We i Ss SoS = ~~ sas SAS So Ficure 10. Mecas menthae Chemsak and Linsley, 92. [Proc. 4TH Serr. CALIFORNIA ACADEMY OF SCIENCES 176 ‘AQ[SUIT pue YyesuloyD avyjuawm spray JO 9dUAIINIDIO UMOUY ‘TI Fano ‘uotsstuiad Aq paonpoiday ‘oseotyD JO AjysIoAtUA, 9Yy} Aq LEG] J4stIAdoo dew aseg = 3ONsIDNO7 153m sol s! We a oa ——= NOILD3°O¥8d VY3YNV-1VND3 S/3NNOE YNOLNOD 1004 O00) Su2L3MOTIM OOF ooc oor 001 Re s711n09¢ ooe 001 aqvos VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 177 callus toward base; punctures rather fine, separated; pubescence short, appressed, obscuring surface, long erect hairs numerous; prosternum densely pubescent; meso- and metasternum densely pubescent, finely densely punctate at sides. Scutellum densely clothed with appressed pubescence. Elytra more than 2% times as long as broad; punctures coarse, contiguous to about apical %, very fine at apex; pubescence dense, short appressed, lateral margins and suture with a narrow band of appressed pubescence, long erect hairs numerous basally, shorter toward apex; apices rounded. Legs densely pubescent; tarsal claws with inner tooth slightly shorter than outer. Abdomen densely pubescent; last sternite deeply impressed for about * of its length. Length, 8-12 mm. FEMALE. Similar in form, slightly more robust. Antennae about as long as elytra. Abdomen with last sternite linearly impressed. Length, 9-13 mm. MATERIAL EXAMINED. Holotype male, allotype (California Academy of Sciences) and 99 paratypes (60 males, 39 females) from 8 miles W. of El Palmito, Sinaloa, Mexico, VII-19, 24, 29-64, VIII-5-64 on Agastache (J. A. Chemsak, J. A. Powell, H. F. Howden). Additional material not paratypical assignable to this species includes: 9 6 6,5 2 2, 6.5 miles E. of Potrerillos, Hwy. 30, Sinaloa, VIII-20-21-64 (E. 1. Schlinger, P. Rauch); 2 ¢ 4, 20 miles E. of Villa Union, Sinaloa, VIII-20-64 (P. Rauch); 1 2, 63 miles E. of Jct. Hwy. 15 & 40 on Hwy. 40, Mexico, VIII-28-64 (D. C. & K. A. Rentz, J. A. Grant); 1 ¢, El Pichon, Nayarit, Mexico, VI-25-63 (J. Doyen); 1 4, Southwestern Research Station, Chiricahua Mts., Arizona, VIII-24-58 (P. D. Hurd); 1 6,1 2°, Yank’s Spring, Sycamore Canyon, Tumacacori Mts., Santa Cruz Co., Arizona, VII-28- 65 (H. B. Leech); 1 ¢, Oak Creek Canyon, 12 miles S. of Sedona, Arizona, VII-18-57 (C. W. O’Brien); 1 2, Swift Trail, between Ladybug Saddle and Shannon Park, Pinaleno Mountains, Graham County, Arizona, VI-27-58 (J. M. & S. N. Burns). Other specimens from Mexico tentatively assigned to M. menthae; 1 2, 2 miles S. of Tlaquepaque, Jalisco, VII-11-53 (C. & P. Vaurie) ; 1 2,5 km. W. of Zacapu, Michoacan, VII-13-51 (H. E. Evans); 1 2, Real de Arriba, Distrito Federal, VII-32 (H. E. Hinton); 1 6, 2 2%, Temescaltepec, Distrito Federal, 1931 (G. B. Hinton); 1 2, Tejupilco, Temescaltepec, VI-33 (H. E. Hinton, R. L. Usinger). This species differs from M. ambigena by the much longer antennae, finer pubescence, and much less coarsely punctate pronotum. The type series was collected during the day from the apical leaves of the mint Agastache. The adults were found resting in the curve of the smaller top leaves of the plant. Numerous individuals were mating and this plant is prob- ably the larval host. Mecas cana (Newman). Saperda cana NewMan, 1840, Entomologist, vol. 1, p. 12; LeConrer, 1852, Jour. Acad. Nat. Sci. Philadelphia, vol. (2)2, p. 164. 178 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Mecas cana, GAHAN, 1888, Trans. American Ent. Soc., vol. 15, p. 300; Lenc and Hamirton, 1896, Trans. American Ent. Soc., vol. 23, p. 152; Casey, 1913, Memoirs on the Coleoptera, vol. 4, p. 360; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 148. MALE. Form moderate sized, subparallel; color black, body densely clothed with gray recumbent pubescence. Head with front convex, pubescence obscuring punctures, longer, dark, suberect hairs numerous; interantennal area almost plane; vertex finely, separately punctate; antennae slightly shorter than body, segments to sixth rather sparsely grey pubescent beneath, long erect hairs de- creasing in number toward apex; third segment longer than scape, fourth shorter than third but longer than first, remaining segments gradually decreasing in length. Pronotum broader than long, sides rounded to subparallel; basal and apical margins narrowly margined; disk convex, each side with a flat glabrous callus before middle, middle usually with a vague linear callus near base; punctures moderately coarse, rather sparse, partially obscured by pubes- cence; longer erect hairs pale with dark setae interspersed mostly at sides; prosternum densely pubescent; meso- and metasternum densely pubescent; metasternum deeply rather densely punctate at sides. Elytra over 2% times longer than broad; punctures moderately coarse, well separated, becoming finer and sparser toward apex; pubescence obscuring surface, longer suberect hairs numerous; apices obliquely subtruncate. Legs densely pubescent; tarsal claws with inner tooth small. Abdomen occasionally reddish, densely pubescent; last sternite moderately impressed for its entire length. Length, 9-15 mm. FEMALE. Form similar. Antennae shorter than body. Abdomen with last sternite linearly impressed. Length, 10-16 mm. TYPE Locatity. St. John’s Bluff, Florida. RANGE. Southeastern United States to Texas, northeastern Mexico, and South Dakota. REMARKS. This species may be separated from M. confusa by the short inner tooth of the tarsal claws. The pronotal calluses readily distinguish it from M. cineracea and the elongate antennal segments from M. bicallosa. Two subspecies can be recognized. Mecas cana cana (Newman). Saperda cana NEWMAN, 1840, Entomologist, vol 1, p. 12; LeConre, 1852, Jour. Acad. Nat. Sci. Philadelphia, vol. (2)2, p. 164. Mecas cana, GAHAN, 1888, Trans. American Ent. Soc., vol. 15, p. 300; Casry, 1913, Memoirs on the Coleoptera, vol. 4, p. 360; BREUNING, 1955, Mem. Soc. Roy. Nat. Belgique, vol. 27, p. 148. Body densely grayish pubescent, elytra narrowly clothed at suture and lateral margins with bands of appressed pubescence. Length, 10—-12.5 mm. TypE LocALity. St. John’s Bluff, Florida. RANGE. Florida. VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 179 FLIGHT PERIOD. April to October. Host pLrants. Ambrosia, Flaveria linearis. REMARKS. This subspecies appears to be restricted to Florida, primarily the southern portion. Although Breuning (1955) states that the type of M. c. cana appears to be lost, it is in the collection of the British Museum (Natural History). Mecas cana saturnina (LeConte). Stenostola saturnina LECONTE, 1859, Smithsonian Contr. Knowledge, vol. 11, p. 21. Mecas saturnina, GAHAN, 1888, Trans. American Ent. Soc. vol. 15, p. 300; Horn, 1888, Trans. American Ent. Soc., vol. 15, p. 301; Brreuntnc, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 146; Wirson, 1960, Comm. Inst. Biol. Control Tech. Comm. vol. 1, p. 62; StrmpE and WARWICK, 1962, Anim. Behaviour, vol. 10, p. 112 (habits). Mecas inornata (not Say), Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; KNULL, 1946, Ohio Biol. Surv. Bull. vol. 39, p. 274; Dmton and Ditton, 1961, Man. Common Beetles of Eastern North America, p. 652, pl. 65, no. 17. Mecas brevicollis CAsEY, 1913, Memoirs on the Coleoptera, vol. 4, p. 362. Similar in form and size to typical subspecies. Pubescence uniformly gray or yellowish, elytra without pubescent bands on margins and suture. Length, 9-16 mm. TYPE LocaALity. Of ‘saturnina, Kansas; ‘brevicollis, WKansas. RANGE. Alabama to northeastern Mexico to South Dakota. FLIGHT PERIOD. May to August. Host pLants. Ambrosia, Xanthium, Helianthus, Gaillarda. Adults have also been taken on Prosopis and Salvia in Texas. INCERTAE SEDIS Mecas albovitticollis Breuning. Mecas albovitticollis BrEuNtNG, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 146. Mecas (?)———————————(?) Bates, 1885, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 427. “Antennae a little longer than the body. Lower lobes of the eyes obviously longer than the cheeks. Head and pronotum densely and very finely punctate. Pronotum transverse. Elytra slightly truncate at the apices, densely and finely punctate. “Black covered with light gray pubescence. Pronotum ornated with three straight, longitudinal, discal, whitish bands. Scutellum having whitish pubescence. Antennae faced with deep brown pubescence. “Length: 10 mm; width: 2% mm. “Type: a male from Mexico in the British Museum.” We have been unable to place this species on the basis of the above original description and were unable to locate the type at the British Museum. However, we believe that this species was based on the single specimen cited by Bates in the Biologia Centrali-Americana. Since this reference stated: “One example 180 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. of a distinct species, in bad condition, doubtfully belonging to this genus.’’, we will follow Bates and exclude it from the genus Mecas. Mecas marmorata Gahan. Mecas marmorata GAHAN, 1892, Trans. Ent. Soc. London. 1892, p. 268, pl. 12, fig. 7; BREUNING, 1955, Mem. Soc. Roy. Belgique, vol. 27, pp. 140, 152. Mate. Form elongate, slender; color black, pronotum and elytra suffused with pale reddish brown, antennal segments 2 to 10 reddish; pubescence dense, yellowish, appressed, arranged in patches on elytra, forming a longitudinal band down middle of pronotum. Pronotum cylindrical, as long as broad; mesosternal process slightly broadened. Elytra more than 3 times longer than broad; punctures coarse, irregular; apices produced, dentate. Legs short; mesotibiae with sinus; tarsal claws with inner tooth smaller than outer. Abdomen with last sternite deeply impressed. TYPE LOCALITY. Guerrero, Mexico (not restricted further). The elongate body, apically produced elytra, and cylindrical pronotum appear sufficient to exclude this species from Mecas. Since we have examined only the type in the British Museum (Natural History) and one other male from 17 miles south of Puebla, Puebla, IX-6-57 (H. A. Scullen) we have not attempted to clarify the generic status of ‘marmorata’ at this time. ACKNOWLEDGMENTS This study was made as part of a National Science Foundation sponsored monograph of North American Cerambycidae (Grant GB-31120-X). Although a large portion of the material we examined came from the California Academy of Sciences and the California Insect Survey, Berkeley, we wish to gratefully acknowledge the following institutions for the loan of specimens: American Museum of Natural History; Canadian National Collection, Ottawa; Cornell University; Field Museum of Natural History, Chicago; Los Angeles County Museum; Museum of Comparative Zoology, Harvard University; Instituto Tecnologico y de Estudios Superiores de Monterrey, Monterrey, Mexico; Ohio State University; Oregon State University; United States National Museum; University of California, Davis and Riverside, University of Kansas; collections of G. H. Nelson and R. L. Westcott. We are deeply appreciative to Celeste Green for preparing the illustrations and to the authorities of the British Museum (Natural History) for permitting us to examine type specimens in their collection. LITERATURE CITED BaArerc, W. J. 1921. A girdler on artichokes and other little-known insect pests. Journal of Economic Entomology, vol. 14, pp. 99-100. VoL. XX XIX] CHEMSAK AND LINSLEY: MECAS 181 Bates, H. W. 1881-1885. Biologia Centrali-Americana, Insecta, Coleoptera, Lamiidae, vol. 5, pp. 153- 224, pls. xii-xv, pp. 334-436, pls. xxii-xxv. [Mecas, pp. 203-205.] BEUTENMULLER, W. 1896. Food-habits of North American Cerambycidae. Journal of the New York Ento- mological Society, vol. 4, pp. 73-81. Brake, S. F. 1951. Compositae. Jn: Kearney, T. H. and R. H. Peebles, Arizona Flora, pp. 829-971. University of California Press. BLANCHARD, F. 1887. Notes on Coleoptera. Entomologia Americana, vol. 3, pp. 85-88. BLATCHLEY, W. W. 1910. On the Coleoptera known to occur in Indiana. Indiana Department of Geology and Natural Resources, Bulletin 1, 1386 pp. BREUNING, S. 1952. Revision einiger Gattungen aus der Gruppe der Saperdini Muls. Entomologische Arbeiten aus der Museum G. Frey, vol. 3, pp. 107-213. 1955. Revision du genre Mecas LeConte. Memoires de la Société Royale d’Entomologie de Belgique, vol. 27, pp. 138-152, 1 fig. 1967. Mecas. Catalogue des Lamiaires du Monde, vol. 10, pp. 830-832. CasEy, T. L. 1913. Mecas LeC. Memoirs on the Coleoptera, vol. 4, pp. 360-363. CuHEmMsAK, J. A., AND E. G. LINSLEY 1965. A revised key to the species of Elytroleptus with notes on variation and distri- bution. The Pan-Pacific Entomologist, vol. 41, pp. 193-199. CRAIGHEAD, F. C. 1923. North American cerambycid larvae. Dominion of Canada, Department of Agriculture Bulletin No. 27, new series, 238 pp., 44 pls. Diton, E. S. anp L. S. Ditton 1961. A manual of the common beetles of Eastern North America. Row, Peterson and Company. 884 pp. GaHaNn, C. J. 1888. Notes on some types of North American Cerambycidae in the British Museum. Transactions of the American Entomological Society, vol. 15, pp. 299-300. 1892. Additions to the Longicornia of Mexico and Central America, with notes on some previously-recorded species. Transactions of the Entomological Society of London. 1892, pp. 255-274. Gimour, E. F. 1962. Synopsis of the tribe Aerenicini. Revista de Biologia Tropical, vol. 10, pp. 123-147. HALDEMAN, S. S. 1847. Material toward a history of the Coleoptera Longicornia of the United States. Transactions of the American Philosophical Society, vol. (2)10, pp. 27-66. 1847. Corrections and additions to his paper on the Longicornia of the United States. Proceedings of the American Philosophical Society, vol. 4, pp. 371-376. Hatcu, M. H. 1971. The beetles of the Pacific Northwest. Part V. University of Washington Publi- cations in Biology, vol. 16, 662 pp. 182 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Horn, G. H. 1878. Notes on some genera of Cerambycidae of the United States. Transactions of the American Entomological Society, vol. 7, pp. 41—50. 1888. Additional notes. Transactions of the American Entomological Society, vol. 14, pp. 300-301. Kincssury, J. M. 1967. Poisonous plants of the United States and Canada. Prentice-Hall, New Jersey. xiii + 626 pp., 130 figs. Knut, J. N. 1946. The long-horned beetles of Ohio. Ohio Biological Survey Bulletin 39, pp. 133- 354, 29 pls. LAcorDAIRrE, J. T. 1872. Genera des coléoptéres. . ., vol. 9, no. 2, pp. 411-930. LEConrmE, J. L. 1852. An attempt to classify the longicorn Coleoptera of the part of America north of Mexico. Journal of the Academy of Natural Sciences of Philadelphia, vol. (2)2, pp. 139-178. 1859a. The Coleoptera of Kansas and Eastern New Mexico. Smithsonian Contributions to Knowledge, no. 11, pp. 1-58. 1859b. The complete writings of Thomas Say on the entomology of North America. 2 vols. New York. 1873a. New species of North American Coleoptera. Part II. Smithsonian Miscel- laneous Collections, No. 264, pp. 169-240. 1873b. Classification of the Coleoptera of North America. Part II. Smithsonian Miscellaneous Collections, No. 265, pp. 279-348. LEConTE, J. L., anp G. H. Horn 1883. Classification of the Coleoptera of North America. Smithsonian Miscellaneous Collections, no. 507, xxxviii +- 567 pp. Lenc, C. W. ano J. HAMILTON 1896. Synopsis of the Cerambycidae of North America. Part III, the Lamiinae. Trans- actions of the American Entomological Society, vol. 23, pp. 101-178. Linsey, E. G. 1962. Synopsis of the genus Elytroleptus Dugés. Folia Entomologica Mexicana, no. 3, pp. 1-13. Linstey, E. G., J. N. Knut, Ano M. STATHAM 1961. A list of Cerambycidae from the Chiricahua Mountain area, Cochise County, Arizona. American Museum Novitates, no. 2050, pp. 1-34, 24 figs. Martin, J. O. 1924. Studies in the genus Mecas. Entomological News, vol. 35, pp. 244-245. NEwmMan, E. 1840. Entomological notes. Entomologist, vol. 1, pp. 1-16. Norp, J. C., AND F. B. KnicHT 1970. Saperda inornata Say, 1824 (Insecta, Coleoptera): Proposed use of the plenary powers to designate a neotype to stabilize the nomenclature. Bulletin of Zoo- logical Nomenclature, vol. 27, pp. 123-128. Rey, C. V. 1880. Food habits of the longicorn beetles or wood borers. American Entomologist, vol. 3, pp. 237-239, 270-271. VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 183 Save) Tr. 1824. Descriptions of coleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long. Journal of the Academy of Natural Sciences of Philadelphia, vol. 3, pp. 139-216. STRIDE, G. O., AND E. P. WArRwIckK 1962. Ovipositional girdling in a North American cerambycid beetle, Mecas saturnina. Animal Behaviour, vol. 10, pp. 112-117. THomson, J. 1864. Systema cerambycidarum on expose de tous les genres compris dans la famille des Cerambycides et familles limitrophes. Memoires des Société Royale des Sciences de Liege, vol. 19, pp. 1-540. 1868. Physis recueil d’Histoire Naturelle, vol. 2, pp. 6-208. TOWNSEND, C. H. T. 1884. Notes on some Coleoptera taken in South Louisiana. Psyche, vol. 4, pp. 219-222. Voct, G. B. 1949. Notes on Cerambycidae from the lower Rio Grande Valley, Texas. The Pan- Pacific Entomologist, vol. 25, pp. 137-144, 175-184. Witson, F. 1960. A review of the biological control of insects and weeds in Australia and Australian New Guinea. Commonwealth Institute of Biological Control, Technical Com- munication no. 1, pp. 1-102. ° Li : wi wall i" ae rf a A ai \ oy GH) WwW shé iy ’ Oba ~* PROCEEDINGS OF THE Weoatls Hale, wage, CALIFORNIA ACADEMY OF §S FOURTH SERIES Vol. XXXIX, No. 13, pp. 185-256; 26 figs.; 7 tables. July 9, 1973 COMPARATIVE REVISION OF SCOMBEROIDES, OLIGOPLITES, PARONA, AND HYPACANTHAUS WITH COMMENTS ON THE PHYLOGENETIC POSITION OF CAMPOGRAMMA (PISCES: CARANGIDAE)' = William F. Smith-Vaniz? and Jon C. Staiger Rosenstiel School of Marine and Atmospheric Science, University of Miami 10 Rickenbacker Causeway, Miami, Florida 33149 TABLE OF CONTENTS LVDS CYST ce SS ee RE ner eee WO OE 186 ee een aE DEY Hoe rants ek ei Oe oe Te Tae ee 186 Mrrseercrseanur MeCnOUS . te a ee 187 Semmsmscumucrores Lacepede = 2 toe oe ee eee 190 Mevarorspecies\GL scomberoides — 193 Neomberoides commersonianus Lacépede ——-__-___ 194 ReaMICROIGeS COld ((CUVIEL)" 2225. ee 199 RUCHOIOeS VSN |(CHOrSKal) Se a ee 205 tie ROTC SHLOUM\ OC UNIGE)) oe A. 2 ee a ae eee eee 209 DamuUerOres. SPINOSUS (SMITROV )) 22 ee ee ee eee Dil ene AOU pTILes Gili o= Se Sa 2 2 AUS Vie a ee 213 evarospecies. 0: Oligoplitess =! ee 217 Oicoplites réjuigens Gilbert: and Starks — 27 Mapes SGiICNS (EOC) ey F2RN0 i) Fes 3 ee ee 218 1 Contribution number 1624 from the Rosenstiel School of Marine and Atmospheric Science, University of Miami. 2Present address: The Philadelphia Academy of Natural Sciences, Nineteenth and the Parkway, Philadelphia, Pa. 19103. [185] 186 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Olicoplites-saurus (Block and Schneider). Eee 221 Oligopliesraitus,( Gunther), ..2 4 32% 2". 5 J ee eee 222 Oligoplites pelometa (Cuvier) .....<-. eee 225 Genus Parone Bete. 2 tS ee 225 Porona signata (Jenyns) ...- Eee 227 Genus Aypacanthus Ratinesque\._..__.__ Eee 228 Hy pacanthus ana (Linnaeus) Eee 230 Relationships and Zoogeography .....___.._.___.._ ____.__ _ = aaa Zou Phylogenetic position of Campogramma Regan __.---._--------_--------------- 244 SUMMMATY = 32s ee ee Se i A Oe ee ee 247 Nominal species of Scomberoides and Oligoplites ____----_________ 248 Literature Cited. 2-2 ee eee 250 INTRODUCTION This study evolved as a result of our efforts to unscramble the confused synonymies of Indo-Pacific carangids of the genus Scomberoides. The initial effort led us to review the species of Oligoplites, a New World genus closely related to Scomberoides, last treated by Schultz (1945). It soon became obvious that our paper would not be complete unless we included coverage of several monotypic Atlantic genera that are superficially similar to the ‘leatherjackets’ of American waters. Comparison of these three carangid genera is especially desirable because they, together with Oligoplites, are the only ones, of the 18 Atlantic genera that we currently recognize, which do not occur on both sides of the Atlantic. The combinations of characters exhibited by these genera have impressed us with the artificiality of the present subfamilial partitioning of the Carangidae. In discussing relationships of these genera it is necessary to em- phasize some of the inadequacies of the present subfamilial classification of the Carangidae. ACKNOWLEDGMENTS We wish to thank the following individuals who assisted us in obtaining data for our study, either through hospitality at their institutions or by providing data on, or loans of, specimens (abbreviations refer to institutions and are ex- plained in the methods section): Donn E. Rosen, AMNH; John R. Paxton, AMS; N. B. Marshall and Alwyne C. Wheeler, BMNH; William N. Eschmeyer and staff, CAS; M. L. Bauchot, MNHN; Carl L. Hubbs and Richard H. Rosen- blatt, SIO; Warren C. Freihofer, SU; Frederick H. Berry, TABL; Boyd W. Walker, UCLA; C. Richard Robins, UMML; Victor G. Springer, Martin F. Gomon and Edgar E. Gramblin, USNM; Jérgen G. Nielsen, ZMC; R. J. McKay, WAM. Vor. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 187 Special thanks are due J. R. Paxton and R. J. McKay for providing identi- fications and locality data for collections of Scomberoides housed in their re- spective institutions, thus confirming our tentative zoogeographical conclusions. Tomio Iwamoto examined MNHWN types of Scomberoides (subsequently re- examined by us) that provided much of the impetus for initiation of the present paper. Additional help came from various sources. James E. Bohlke and William N. Eschmeyer facilitated our study by making literature available. Phillip C. Heemstra called to our attention several pertinent references. William H. Hulet and Francisco J. Palacio assisted in translating Italian and Portuguese literature. Grady W. Reinert illustrated the specimens shown in figure 1. The manuscript was typed by Mrs. Kathie Jeffries. Work space at the Southeast Fisheries Center Miami Laboratory, National Marine Fishery Service, where much of the material cited in this paper is housed, was generously provided by the Director, Mr. Harvey R. Bullis. Frederick H. Berry and C. Richard Robins reviewed the manuscript and offered suggestions for improvement. Major collections of Scomberoides critical to our study were obtained in Ceylon under Smithsonian Foreign Currency Grant 3818, C. Richard Robins and Frederick H. Berry, principal investigators. Field work in Ceylon was greatly facilitated by the cooperation of Mr. A. S. Mendis, Deputy Director (Research), Department of Fisheries, Ceylon. The second author was able to pursue this study through support of the National Geographic Society-Univer- sity of Miami Deep-Sea Biology Program (University of Miami research account 8852R), Gilbert L. Voss, Principal Investigator. Partial support was also pro- vided by a grant from the Tai Ping Foundation to Smith-Vaniz for study of Indian Ocean Carangidae. CHARACTERS AND METHODS Osteological terminology generally follows that of Suzuki (1962); a specific exception is the “postmaxillary process” which is here more appropriately termed a premaxillary spur since it is a posterior projection on the premaxilla that braces the maxilla. A basibranchial is defined (Nelson, 1969, p. 480) as the median part of the gill arch endoskeleton that lies between successively paired arch-elements. The number of the basibranchial refers to the paired arch-element behind it. Rayless predorsal interneurals are termed predorsal bones (Smith and Bailey, 1961). All carangids have at least 3 predorsal bones; counting poste- tiorly, the fourth element, whether supporting a dorsal-fin spine or not, has a well developed spinelike projection on its anterodorsal margin. This structure, commonly termed a procumbent spine, may occasionally protrude through the 188 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. skin. In the genera treated in this study, all anterior dorsal pterygiophores support a single fin spine. Thus, a shortening of the dorsal-fin base through loss of anterior spines results in a concomitant increase in number of predorsal bones. The first fin-ray (spine) is often greatly reduced, depressed posteriorly, and can easily be overlooked. In carangids there is a tendency for the number of external spines to decrease in large specimens due to absorption, as evidenced from examination of radiographs. In this study large specimens were not x- rayed or dissected; thus the frequency of specimens with the least number of dorsal spines may be slightly lower than indicated in tables 2 and 4. The last dorsal- and anal-fin rays are split to the base, have a single pteryg- iophore, and were counted as one ray. The first caudal vertebra has a well developed haemal spine which artic- ulates with the first anal pterygiophore; the urostyle is regarded as the terminal caudal vertebra. Counts of outer row dentary teeth in Scomberoides and Oligoplites are lateral counts (one side only). Counts were made on the dentary that appeared to have the most teeth. Broken teeth were counted but empty tooth sockets and teeth that had not attained a functional position in the outer row were excluded. Symphyseal dentary canines were also excluded in the case of Scomberoides. Pectoral-fin ray counts are lateral counts (one side only). The dorsalmost fin-ray element is spinelike and was counted separately. Gillraker counts are lateral counts and were usually made on the right side of each fish. Only gillrakers on the lateral side of the first arch were counted. The gillraker at the angle of the upper and lower limb has the major portion of its base on the ceratobranchial bone and was included in the count of the lower limb. Rudimentary gillrakers (with the width of raker greater than its length) were not included in the counts. All measurements are straight line (point to point) measurements. Mea- surements smaller than 120 mm. were made with dial calipers. Measurements given in the material examined sections are fork length (FL), unless otherwise stated, and have been rounded off to the nearest millimeter. In large fishes with relatively rigid caudal fins, such as carangids, the end of the hypural plate is often difficult to determine precisely. In addition to being easier to determine than standard length, fork length is more familiar to fishery biologists, who should have the greatest need to identify carangids. The following measurements were used in this study: Fork length (FL). Snout tip to tip of shortest median caudal-fin ray. Standard length (SL). Snout tip to posterior end of hypural bones (caudal base). Head length (HL). Snout tip to posterior margin of fleshy opercular flap. Snout-postorbit length. Snout tip to anterior margin of posterior adipose eyelid. VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 189 Snout length. Snout tip to posterior margin of anterior adipose eyelid. Upper jaw length. Snout tip to posterior margin of maxilla. Body depth Dz to Ag. Origin of terminal dorsal spine to origin of terminal anal spine. Height of fin lobes. Origin of fin to tip of longest segmented ray. Pectoral-fin length. Origin of spine to tip of longest ray. Abbreviations of institutions cited are as follows: AMNH American Museum of Natural History, New York City AMS Australian Museum, Sydney BMNH British Museum (Natural History), London CAS California Academy of Sciences, San Francisco FMNH Field Museum of Natural History, Chicago MNHN Muséum National d’Histoire Naturelle, Paris NFIS Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt SIO Scripps Institution of Oceanography, University of California, La Jolla SU Stanford University, Division of Systematic Biology (collection trans- ferred to CAS) TABL _ Southeast Fisheries Research Center, National Marine Fishery Service, Miami (formerly Tropical Atlantic Biological Laboratory) UCLA University of California at Los Angeles UMML University of Miami, Rosenstiel School of Marine and Atmospheric Science, Miami USNM National Museum of Natural History, Division of Fishes, Washington, DPC. ZMC Zoological Museum, Copenhagen WAM Western Australian Museum, Perth. Uncataloged California Academy of Sciences specimens collected under the auspices of the George Vanderbilt Foundation (GVF) are reported under their register or station numbers. Most collections from Ceylon are uncataloged and listed by station numbers. Abbreviations refer to the following collectors listed in chronological order of their visits to Ceylon: F. J. Schwartz; W. F. Smith- Vaniz; P. C. Heemstra; C. C. Koenig. Localities in material examined sections are abbreviated. Summary state- ments of material examined include only specimens examined by the authors. 190 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Scomberoides Lacépéde, 1802 Scomberoides LACEPEDE, 1802, p. 50 (type-species: Scomberoides commersonianus LACEPEDE, 1802, by subsequent designation of JORDAN, 1917, p. 60). Orcynus RAFINESQUE, 1815, p. 84 (nomen nudum; substitute name for Scomberoides LACEPEDE, 1802). Chorinemus Cuvier in Cuvier and Valenciennes, 1831, p. 367 (type-species: Scomberoides commersonianus LACEPEDE, 1802, by subsequent designation of JORDAN, 1917, p. 137). Rhaphiolepis FOWLER, 1905, p. 59 (as a subgenus of Scomberoides; type-species: Chorinemus tol Cuvier in Cuvier and Valenciennes, 1831, by original designation). Eleria JORDAN and SEALE, 1905, p. 774 (type-species: Eleria philippina JoRDAN and SEALE, 1905 = Chorinemus tala Cuvier in Cuvier and Valenciennes, 1831, by monotypy). Palaeoscomber SmirRNov, 1936, p. 49-59 (type-species: Palaeoscomber spinosus SMIRNOV, 1936, by original designation). The carangid genus Scomberoides comprises four recent and one fossil species of tropical and subtropical marine fishes restricted to the Indian and western and central Pacific oceans. All recent species attain at least 45 cm. fork length. Scomberoides commersonianus, the largest species, occasionally exceeds 100 cm. and is a valued game fish. In some areas S. commersonianus supports a seasonal fishery of considerable importance. Major catches are obtained from drift nets set at depths of 15 to 18 m. in waters surrounding offshore islands (James, 1967). Depending on the region, scomberoides are considered excellent food fishes (Smith, 1949, p. 224) or they are mostly salted and not generally esteemed (Chan, 1968, p. 82). Halstead et al. (1972) give a detailed description of the venom apparatus of Scomberoides lysan. Venom glands are associated with the first seven dorsal- fin spines and the first two anal spines. The anal spines, rather than the dorsal spines, are the most venomous. The anal spines are equipped with a frictional locking device which resembles that found in the fin spines of catfishes and balistid triggerfishes. Scomberoides lysan is capable of inflicting painful stings. The nearly identical morphology of these spines in the other species of Scom- beroides and Oligoplites (see comments under O. saurus) suggest that all species of both genera are venomous. The osteology of Scomberoides has received inadequate attention. The only species studied in detail (under the name Chorinemus orientalis) is Scomberoides lysan (Suzuki, 1962). The only known fossil species of Scomberoides was orig- inally described as belonging to a new genus, Palaeoscomber, of Scombridae (Smirnov, 1936) and was subsequently redescribed as a species of Oligoplites (Danil’chenko, 1960). The nomenclature of Scomberoides species has been greatly confused by a plethora of nominal species inadequately described by early workers. Differ- ences ascribed to nominal species have frequently been based on unrecognized ontogenetic changes. Changes in dentition associated with growth led Jordan VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 191 TasleE 1. Nominal species of Scomberoides recognized in selected publications with our present allocation. Authors Species Present Study commersonianus tala lysan tol Smith (1970) lysan — tolooparah tala Williams (1958) lysan — sanctipetri — Roxas and Agco (1941) lysan tala tolooparah tol Weber and lysan tala tolooparah tol De Beaufort (1931) sanctipetri Wakiya (1924) lysan _ sanctipetri formosanus orientalis moadetta Day (1878) lysan tala sanctipetri moadetta toloo and Seale (1905, p. 774) to describe a new genus and species based on juveniles of S. tala. The failure of recent workers to critically read original descriptions and examine types adequately has resulted in the consistent misapplication of the name Scomberoides lysan (Forskal). Because of the confused nomenclature of Scomberoides species, we present in table 1 a list of the species recognized in several major papers with our present allocation of them. An annotated list of the nominal species of Scomberoides and Oligoplites is given in the appendix. NOMENCLATURE. Considerable confusion exists as to whether Scomberoides Lacépéde, 1802 was validly described and thereby has priority over Chorinemus Cuvier, 1831. Williams (1958) briefly discussed the problem and rejected Scomberoides, as did Weber and De Beaufort (1931) and Smith (1970). Re- jection of Scomberoides as a valid generic name by several recent workers is probably attributable to the comments of Barnard (1927, p. 562) who credits the rejected name to Lacépéde (1800), in which only a figure is given with the caption ‘“Scomberoide commersonnien.” Jordan (1917, p. 60) rightly credits Scomberoides to Lacépéde (1802), in which ‘‘Les Scomberoides” was latinized to accommodate the description of three new species. Caranx and Trachinotus, widely accepted carangid generic names, originate from the same work and were described in the same manner as Scomberoides. Williams (1958, p. 416) states that “the use of Chorinemus has been common and widespread since 1833 whilst Scomberoides has been restricted (except Wakiya, 1924) to American authors of the last 75 years.”’ He further states “should Scomberoides be proved to have priority over Chorinemus C.V..... : then a case exists for submission to the International Commission of Zoological Nomenclature for a suspension of the Rules, in that strict application of the Rules will clearly result in greater confusion than uniformity.” In their review 192 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. of the Philippine Carangidae Roxas and Agco (1941) used Scomberoides as did Oshima (1925) and McCulloch (1929). Except for Rofen (1963), American authors of the last 75 years, the most recent of which are Gosline and Brock (1965), have consistently used Scomberoides instead of Chorinemus. Although Williams apparently attaches little importance to this, it should be noted that American authors are responsible for a large portion of the literature treating these fishes. Confusion alse exists as to what constitutes the type-species of the genus. Several workers have recognized Scomberoides noelii as the type-species. Lacépede (1802) described three species in his new genus, listing them on page 50 with brief diagnoses (descriptions expanded on following pages) in the fol- lowing order: (1) Scomberoides noelii; (2) S. commersonianus; (3) S. saltator. He did not, however, specifically designate a type-species. The first author to subsequently designate a type-species for Scomberoides in accordance with Article 69a of the International Code of Zoological Nomenclature was Jordan (1917, p. 60), who selected S. commersonianus. The published illustration of S. commersonianus (as “Scomberoide commersonnien”) (Lacépéde, 1800 pl. 20, Fig. 3) is sufficiently accurate for positive identification. The possession of needlelike scales does distinguish the type-species of Rhaph- iolepis from its congeners. However, the transition from oval-shaped to needle- like scales is obvious in Scomberoides (see figure 3), and we do not believe that recognition of subgenera in such a small group of closely related species is justified. The sole character mentioned in the original description to distinguish Eleria from Scomberoides was based on unrecognized ontogenetic changes in dentition (see discussion under S. tala). Reasons for referral of Palaeoscomber to the synonymy of Scomberoides are discussed under S. spinosus. DeEscrIPTION. Body moderately to strongly compressed. Adipose eyelids mod- erately to poorly developed. Body, except head, covered with embedded scales, broadly lanceolate to needlelike. Lateral-line scales not developed into scutes. Lateral line without side branches, straight except for slight curve over pectoral fin. Anterior rim of shoulder girdle smooth, without fleshy papillae or deep groove near isthmus. Premaxillae not protractile, connected anteriorly to snout at midline by a wide fleshy bridge. Swimbladder strongly bifid posteriorly, length of bifurcated portion of swimbladder equal to or greater than undivided part. Posterior dorsal- and anal-fin rays consisting of semidetached finlets, distal fourth to half of ray not connected by interradial membrane (unattached portion of rays increasing with growth); ultimate and penultimate rays more closely spaced than adjoining rays, fully connected by interradial membrane; none of dorsal- or anal-fin rays produced into long filaments. Pelvic fins rel- atively short, depressible into shallow abdominal grooves. Pectoral fins short, not falcate, 53 to 71 percent head length in adults. Maxilla extends posteriorly Vout. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 193 from middle of eye to well behind orbital rim, 43 to 65 percent head length, in adults. Predorsal bones 3, rarely 4 (4 in one of 60 specimens); dorsal-fin rays VI or VII + I, 19-21; total dorsal-fin rays 27-29. Anal-fin rays II + I, 16-20. Pectoral-fin rays I, 16-18. Pelvic-fin rays I, 5. Vertebrae 10 + 16 = 26; epipleural ribs 9 or 10. Branchiostegal rays 3 + 5 (epihyal + ceratohyal). Upper gillrakers 0-8; lower gillrakers 7-20; total gillrakers 8-27. Fronto-supraoccipital crest low; temporal crest extending forward to edge of cranium slightly in front of middle of orbit; preorbital region short. Epiotics broadly united along midline of cranium internally. Zygapophyses of exoccipital joined beneath foramen magnum. Parasphenoid not expanded into a broad, flattened plate posteriorly. Myodome with broad, posterior opening. Basi- occipital without a pair of lateral processes on ventral surface. Ascending process of premaxilla short and triangular; maxilla long and slender, rather closely attached to premaxilla. Premaxillary spur absent. Small supramaxilla present. Interosseous space between dorsal arm of dentary and upper margin of articular greatly reduced. Well developed teeth on dentary, premaxilla, palatine, and vomer; two distinct rows of dentary teeth (see discussion under Zoogeography and Relationships Section); symphyseal premaxillary and dentary teeth of juveniles (except S. /ysan) robust and caniniform. Mesopterygoids covered with firmly ankylosed, minute, granular teeth. Pharyngeals not noticeably enlarged, covered with sharp, pointed teeth. Well developed suborbital shelf present; lower and posterior suborbitals not greatly expanded posteriorly. Basibranchials 1-2 with a pair of tooth plates, third basibranchial with two or more irregular tooth plates. Lower limb of posttemporal not noticeably short or thickened. Postcoracoid process absent. Interpelvic keel well developed; post pelvic process of each side coalesced entire length, not forming an apical fork. Inferior vertebral foramina absent. First anal pterygiophore and haemal spine of first caudal vertebra firmly attached to form a strong strut. Anal pterygiophore expanded anterolaterally to form roof over first two anal spines; first two anal spines articulated proximally with each other; second anal spine asymmetrical, with deep groove on one side of anterior face. Caudal skeleton with 2 epurals and 2 pairs of uroneurals. Key To Recent Spectres oF Scomberoides Color pattern will readily separate most specimens of the four species. In large individuals and especially in small specimens that are not freshly caught the spots may fade or completely disappear; this is particularly true of fish market specimens. aeeGalirakerses) tom is dorsal-tine lobe) unttonmly plemented = 2 1b. Gillrakers 21 to 27; distal half of dorsal-fin lobe abruptly and heavily pigmented __ 3 2a. Large oval blotches above or touching lateral line; upper jaw extends well beyond posterior margin of eye, especially in adults (fig. 2a); snout 21 to 26 (mean 23.4) 194 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. percent HL; dentary teeth of inner and outer rows subequal in adults; specimen fig- UL ell yp oS cat ven ae ee 2 Et ie ee ee S. commersonianus 2b. Vertically elongate blotches intersecting lateral line; upper jaw extends slightly beyond posterior margin of eye (fig. 2b); snout 26 to 30 (mean 28.2) percent HL; inner row dentary teeth distinctly larger than outer row teeth in adults; specimen figure 1b —__ S. tala 3a. Double series of 6 to 8 dusky, roundish blotches above and below lateral line, occasionally connected by narrow isthmus; scales lanceolate (fig. 3b) ; specimen figure 1c ___ S. lysan ¢ 3b. Oval or vertically oblong blotches, the first 4 or 5 intersecting lateral line; scales slender, needlelike -(fig) Sd); specimen figure Id) 22 == Eee Ss Hal! Scomberoides commersonianus Lacépéde. (Figures la, 2a, 3a, 4a, 5—7, 14c, 16b, 19, 23b, 25b; tables 2-3.) Scomberoides commersonianus LACEPEDE, 1802, p. 50 (type locality Madagascar). Scomber madagascariensis SHAW, 1803, p. 590, pl. 85 (type locality Madagascar). Chorinemus exoletus EHRENBERG in Cuvier and Valenciennes, 1831, p. 379 (278) (based solely on an unpublished drawing by Ehrenberg of a fish from Lohaia, Red Sea). Chorinemus delicatulus RICHARDSON, 1846, p. 269 (based solely on a drawing in the Reeves Collection of Chinese fish drawings). Chorinemus leucopthalmus RicHARDSON, 1846, p. 269 (based solely on a drawing in the Reeves Collection of Chinese fish drawings). NOMENCLATURE. The figure of S. commersonianus (as “Scombéroide com- mersonnien”) and original description are sufficiently accurate for positive identification, and the name has consistently been applied to the correct species, although usually in synonymy (see nomenclature section under S. lysan). Al- though in the original description S. commersonianus was spelled with a double “n,”’ we prefer the emended spelling with a single “‘n,’’ which is correct. The description of Chorinemus exoletus refers to a deep-bodied species with a long upper jaw. This together with the stated type locality is sufficient to confidently identify it as conspecific with S. commersonianus. Whitehead (1969) has published the original drawings upon which the de- scriptions of Chorinemus delicatulus and C. leucopthalmus are based. These drawings and that of Scomber madagascariensis are reasonably diagnostic and are undoubtedly based on S. commersonianus. Although the large oval blotches characteristic of S. commersonianus are not shown on the sides of Chorinemus delicatulus and C. leucopthalmus, they rapidly fade and disappear in dead specimens. DescripTION. Characters given in the generic description are not repeated here. Frequency distributions of selected meristic characters are given in table 2. Dorsal-fin rays VI-VII + I, 19-21; anal-fin rays II + I, 16-19; pectoral- fin rays, I, 17-19. Gillrakers 0-3 + 7-12 = 8-15. Upper jaw extending well beyond posterior margin of eye, especially in adults (fig. 2a). Scales broadly lanceolate (fig. 3a). Inner and outer row of dentary teeth subequal in adults (fig. 4a); dentary typically with one or two pair of symphyseal canines, at least VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 195 Ficure 1. Scomberoides species: a, S. commersonianus, TABL 107366, 468 mm. FL; b, S. tala, TABL 107365, 464 mm. FL; c, S. lysan, TABL 107364, 523 mm. IGS Ch, ual TABL 107363, 450 mm. FL. [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 196 16 Ch Caen OKamemmamer CL 9 CemeS come ce me 6 = pa Aa Eg 104 CEG SANS Gk OG AG. AS UDsK] ONC S0 ae en en oe Cee at a SI Oy OC UN e kee fae —— DIDI OSS Oe erie eee eee i i eee OL Lb SSieasole alin (x “SIUDIUOSHOUINLOD x INC 4E M6 Se we BE be Ie OW ON rN A OL Si i wy Al je O 6 § sotedg SISAeA (ED [BIOL LES. Ce ai on a ce OS Oa sey eS Cg JIE an ae! 2 ps 101 SVC Os aes Ss Sy SS WO Bo Y CG Wy We G@ Wo = S32 = UDSK] eee Se pee toe C2 Ol SS Sap e: L GUS ORS a TO: Oia DIDI ae meee he Bol) Dee OC SOT. T OS © 3 nee eee STC) COSTELLO SMOULLOD sl 2 OW St awl el @ i OW @ & 2 See NGS ci sie 6 Ol Ste” Cw eco) oles RO sataedg SIOYVY [[ID JoMmo'T SIIYeY [ED 1toddq G 89 ce = ae S02 16 ov vy T 68 G 104 =a L £8 s = 0°02 v6 8 £8 € 98 8 UDSK] aaa 9 08 € ce 6°61 06 9 el IT v8 9 DIDI = ¢ TZ eT (4 L161 L8 T £9 (BG £8 v SNUDIMOSAAUULOD 0Z 61 SI LT OT x N 14 02 61 TAS SrA satadg SABY PoyUIUI5IS SABY Pa}UaUISISG soutds 'q sAby Ul_-[eUYy SsABY UlA-[esiog "$a19agS SaplO1aqwuorg U2 S4azIDADYI IYSiMaU {Oo UOoLNQrysYp KoUaNbatg *7 AAV, VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 197 Cc d Ficure 2. Upper jaw development of adult Scomberoides species: a, S. commersonianus, TABL 107192, 355 mm. FL; b, S. tala, TABL 107712, 351 mm. FL; c, S. lysan, TABL 107364, 412 mm. FL; d, S. tol, TABL 107714, 410 mm. FL. Maxilla shown in black. in young; young with dentary teeth of outer row numerous and considerably more closely spaced than inner row teeth. Proportional measurements apply only to specimens longer than 150 mm. FL, and are expressed as percent FL unless otherwise stated. Depth (origin Dz to Ag) 25.7-36.2; height dorsal-fin lobe 14.4-19.8; height anal-fin lobe 13.5- 18.5; upper jaw 55.9-64.6 percent head length; snout 20.6—26.0 percent head length. PIGMENTATION. Sides with 5 to 8 large oval blotches above or touching lateral line, first two may intersect lateral line. Dorsal and anal fins uniformly pigmented. Pectoral fin of adults frequently with a dusky blotch ventrally. In life, body gray to silvery white ventrally, dusky green or bluish dorsally; 198 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TABLE 3. Comparison of several proportional measurements in Scomberoides species expressed as hundredths of head length. Upper Jaw Length Snout Length <20 cm. = = Species Range FL N Range x Range % commersonianus 82-198 19 59-64 60.1 21-26 23.6 tala 71-199 23 53-61 58.2 26-30 28.3 lysan 45-189 20 51-56 53.9 28-31 29.0 tol 44-181 17 47-51 48.6 30-33 Sue3 Upper Jaw Length Snout Length re pper J g g Species Range FL N Range X Range x commersonianus 224-943 64 56-65 60.6 21-26 23.4 tala 201-571 58 54-61 57.8 26-30 28.2 lysan 201-585 65 50-54 51.6 28-32 30.0 tol 209-468 70 43-49 46.3 29-33 Silks lateral blotches plumbeous gray. Large individuals often golden, especially ventrally. Annunziata and Cedrone (1972, p. 61) present a color photograph of a large specimen from Madagascar. DISTRIBUTION. Widely distributed throughout the Indian Ocean and Indo- Australian Archipelago. The species has also been reliably reported from the Red Sea and Persian Gulf. Scomberoides commersonianus appears to be re- stricted to neritic waters and does not occur east of the New Guinea-Solomon Islands region. Records of the species (as S. /ysan) from Tahiti and other central Pacific islands are based on misidentifications. MATERIAL (96 specimens, 82—943 mm. FL, from 53 collections). KENYA: Mombasa Fish Market, TABL uncataloged Anton Bruun cruise 9 station, FT-2 (1: 398). WEST PAKISTAN: off Karachi, TABL uncataloged (3: 236-295). INDIA: Gulf of Kutch, TABL uncataloged (1: 203); Bombay, TABL 107190 (1: 384), 107350 (1: 158); Palk Strait, TABL 107292 (4: 127-288), TABL uncataloged FHB 66-36 (1: 766); Porto Novo, TABL 107303 (1: 198), TABL uncataloged (1: 190). CEYLON: TABL 107192 FJS 69-41 (9: 327-422), FJS 69-45 (6: 195-244), FJS 69-64 (1: 474), FJS 69-50 (1: 400), TABL 107366 S-V 69-77 (3: 303-476), S-V 69-86 (2: 224-254), S-V 69-126 (1: 335), S-V 69-151 (2: 754-762), PCH 69-183 (1: 617), PCH 69-185 (7: 123-158), TABL 107715 PCH 69-201 (3: 398-443), PCH 69-218 (10: 130-168), PCH 69-267 (1: 503), PCH 69-305 (1: 913), CCK 69-1 (1: 843), CCK 69-44 (1: 797), CCK 69-53 (1: 475), CCK 69-60 (7: 234-349), CCK 69-97 (2: 160- 218). GULF OF THAILAND: GVF station 57-45 (2: 355-391), GVF station 57-92 (1: 520), GVF station 57-83 (2: 559-579), GVF station 60-113 (2: VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 199 849-943), GVF station 60-146 (2: 727-729), GVF-1548 (1: 133), GVF-1557 (2: 186-197), GVF-2195 (1: 93, cleared and stained), GVF-2513 (2: 727-739), GVF-2546 (1: 291), GVF-2547 (6: 243-612). HONG KONG: GVF station HiK-111 (1: 829). PHILIPPINES: Luzon, USNM 72190 (1: 172), 83447 (1: 241), 168322 (1: 309), SU 20341 (2: 82-116); Samar, USNM 149810 (1: 129); Iloilo, USNM 72189 (1: 129), 149812 (1: 152); Palawan, USNM 149813 (1: 128), 149815 (1: 151). NORTH BORNEO: Sandakan, USNM 149814 (1: 172), 168323 (1: 234). JAVA: Djakarta, TABL 107300 (1: 266). AUSTRALIA: Western Australia, Exmouth Gulf, Onslow, WAM P.2772 (1: 302), P.15487 (1: 175); Port Hedland, WAM P.180 (1: 329); Broome, WAM P.555 (1: 95); Northern Territory, Charles Point, AMS IA.7781 (1: 164); Port Darwin, AMS 1.9778 (1: 300), IA.4375 (1: 280); Darwin, AMS 1B.3168 (1: 420), IB.3171 (1: 85); Bedwell Point, AMS IA.7678 (1: 356); Gulf of Carpentaria, Groote Eylandt, USNM 173954 (4 of 7: 240-345); 17°22’S., 149°45’E., AMS T.15557-118 (5: 87-172) ; Queensland, Burdekin River, USNM 47835 (1: 275), AMS A.18299-300 (2: 285-615); Lindeman Island, AMS IA.6279 (1: 297); Hayman Island, AMS IA.6104 (1: 460); New South Wales, Port Jackson, USNM 47910 (1: 357). NEW GUINEA: AMS A.12694 (1: 300); Port Moresby, AMS 1.13357 (1: 565). Scomberoides tala (Cuvier). (Figures 1b, 2b, 3c, 4b, 5-7, 15b, 17, 19, 24b; tables 2-3.) Chorinemus tala CUVIER in Cuvier and Valenciennes, 1831, p. 377 (type locality Malabar). ?Chorinemus toloo Cuvier in Cuvier and Valenciennes, 1831, p. 377 (description based on “toloo parah” RUSSELL, 1803, p. 29, pl. 137, Vizagapatam). Eleria philippina JorpAN and SEALE, 1905, p. 744, fig. 1 (type locality Negros Island, Philippines). Chorinemus hainanensis Cuu and CHENG, 1958, p. 317, fig. 2 (type localities Sanya, Kanchium, Kwonghoi, and Chinglan, China). NOMENCLATURE. In the Muséum National d’Histoire Naturelle there are two collections (A. 6587 and A. 6588) from Malabar, collected by Belanger, which are labeled syntypes of Chorinemus tala Cuvier. A. 6588, here designated lectotype of C. tala, is a single specimen 298 mm. SL. A. 6587 consists of two specimens of S. tol Cuvier. In his treatment of the western Indian Ocean species of Scomberoides, Smith (1970, p. 221) referred S. tol to the synonymy of S. tala. This erroneous action was based on his examination of one of the specimens of A.6587. Smith apparently was unaware of the existence of other material labeled as types of S. tala, and fortunately did not designate a lectotype. Although Weber and De Beaufort (1931, p. 282) and Smith (1970, p. 219) considered the species insufficiently described for positive identification, the following statements given in the original description of Chorinemus tala leave no doubt as to its specific identity: (1) “Chorinemus tala differs [from the 200 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 3. Scales of Scomberoides species (from area between anal-fin origin and lateral line): a, S. commersonianus, TABL 107715, 398 mm. FL; b, S. lysan, TABL 107364, 412 mm. FL: c, S. tala, TABL 107712, 370 mm. FL: d, S. tol, TABL 107714, 410 emma Hees seale: 20.0 mm. preceding species, C. commersonianus | by a smaller gape, a maxillary that hardly extends beyond the posterior border of the orbit. (2) The maxillary is, in addition, covered by the suborbital and only shows its posterior third; it is enlarged more posteriorly and is cut off more squarely. (3) The teeth are stronger in proportion. (4) The curve of the snout is a little less straight .. . , but the shape of the body, the scales, the lateral line and the number of rays are as in the preceding species.” Statement 4 and the fact that S. tala was compared with S. commersonianus, suggest that the name S. tala applies to a deep-bodied species with high fin lobes. Scomberoides tol and S. lysan are both relatively slender species (fig. 6) and have shorter fin lobes (fig. 7) than do the other two species of Scomberoides. Statements 1 and 4 exclude S. tol, in which the maxilla does not reach the pos- terior border of the orbit and the scales are needle-shaped (fig. 3d). Statement 3 excludes S. lysan and S. tol, in which the teeth in both jaws are relatively small in adults and applies best to the species here recognized as S. tala, which has the largest teeth (fig. 4b). Statements 1 and 2 are sufficient to exclude S. commersonianus (compare figs. 2a and 2b). The fact that Cuvier recognized S. commersonianus (and several of its synonyms) and described as new, from Malabar specimens, examples of the three remaining valid species of Scomberoides leaves no doubt that he was able VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 201 10 mm Ficure 4. Dentary teeth of adult Scomberoides species: a, S. commersonianus; b, S. tala, c, S. lysan, d, S. tol. Data as in figure 2. Teeth of inner row shown in black. to distinguish them. It is obvious from his description of S. fala that he would not have confused it with S. tol. At some time type labels or MNHN catalog numbers must have become inadvertently mixed. Since the description of S. tol was based in part on specimens from Malabar, the two specimens of that species labeled as types of S. tala are here designated putative paralectotypes of Scom- beroides tol (Cuvier). Russell’s poor figure of ‘“toloo parah,” upon which the original description of Chorinemus toloo is based, cannot be positively identified as S. tala, although it most closely resembles that species. Weber and De Beaufort (1931) included S. toloo in their synonymy of Scomberoides tala; the figure of Chorinemus toloo published in Day’s Fishes of India is clearly referable to S. tala. Jordan and Seale distinguished their new genus and species, Eleria philip pina, from Scomberoides solely on the basis of strong symphyseal canines on the lower jaw. They apparently were unaware that such canines are typically present in the young of S. tala. The description and figure of Chorinemus hainanensis leave no doubt as to the identity of this nominal species. In their description, Chu and Cheng em- phasized the absence of spots on side of body. Like a number of authors, they were not cognizant of the fact that these spots frequently fade completely. DEScRIPTION. Characters given in the generic description are not repeated here. Frequency distributions of selected meristic characters are given in table 2. Dorsal-fin rays VI-VII + I, 19-21; anal-fin rays IJ + I, 17-19; pectoral- 202 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 110 100 COMMERSONIANUS 90 TALA LYSAN TOL 80 => => 70 == S60 za LJ —J ae