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B R AR 1 ES SMITHSONIAN “INSTITUTION, -«i tf\ ^ . €i\ -?(• I ?AR I ES^’SMITHSONIAN^ INSTITUTION NOIXfUliSNI NVIN0SHillMs‘”s3 I avaaiT. — — to — '^ - S5^^\ “ NUMBER 236 JANUARY 10, 1973 3 % TWO ANT GENERA NEW TO THE UNITED STATES (Hymenoptera: Formicidae) By Roy R. Snelling CONTRIBUTIONS IN SCENCE NATURAL HISTORY MUSEUM LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- able footnotes must be typed on separate sheets. Several of one kind may be placed on a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing with Latin American subjects. Summaries in other languages are not required but are strongly recommended. Summaries will be published at the end of the paper. (8) A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor TWO ANT GENERA NEW TO THE UNITED STATES (Hymenoptera: Formicidae)' By Roy R. Snelling^ Abstract: The two genera Rogeria (Myrmicinae) and Acropyga (Formicinae) are recorded from the United States. Rogeria is repre- sented by two new species: R. creightoni in Texas and R. huachucana in Arizona. Also present in Arizona is the new species Acropyga epedana. These records are believed to be natural northern termini of the ranges of these genera, rather than accidental introductions. The types of the new species are deposited in the Natural History Museum of Los Angeles County. In this paper the myrmicine genus Rogeria and the formicine genus Acropyga are recorded for the first time from the United States. Two species of Rogeria, one in Texas and one in Arizona, and one species of Acropyga in Arizona are described below as new species. Neither of these generic additions would seem to be the result of introduction within historical times; this seems especially true of the two Arizona species. Both are found at the southern end of the Huachuca Mountains where introduction is not likely. Further, this area is known to be the northern terminus for many Mexican species of Neotropical affinities. The Texas species may be the result of an introduction but, if so, I have been unable to correlate it with any previously described species. More probably this, too, is a largely Mexican species distributed along the tropical east coast of Mexico, with its northern terminus in the lower Rio Grande Valley, a well-known distribution pattern. Genus ROGERIA Emery The genus Rogeria consists of two dozen poorly known species, all Neo- tropical. Most of these species have been described within the past 50 years, from one or a few specimens. The small size and cryptic habits of these ants have mitigated against the accumulation of large series in collections. The taxonomy of the genus is confused, and only in recent years has it consisted of more than the bare descriptions of isolated new species. The works of Kempf (1961, 1962, 1963, 1964, 1965) have greatly assisted in understanding this genus. Until the present paper, the northernmost species were recorded from Honduras on the mainland, but several are known from Cuba. In addition, there are at least two undescribed species in Mexico which I have seen. Among the ant genera recorded from the United States, Rogeria may be recognized by the following combination of worker characteristics: Myrmicinae; worker monomorphic; antennae twelve-segmented, the last three forming a 'Review Committee for this Contribution Charles L. Hogue George C. Wheeler Thomas J. Zavortink ^Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, California 90007 1 2 Contributions in Science No. 236 robust club longer than the remainder of the flagellum; tibiae II and III without apical spurs; propodeum armed; thoracic dorsum without defined sutures and distinctly convex in profile; anterior peduncle of petiole well defined from node or not. Of the genera previously recorded from the United States, Macromischa is most closely related to Rogeria. Curiously, Macromischa, also of Neotropical origin, is represented in this country by one species in southern Texas and one in southern Arizona, as is true of Rogeria. Rogeria may be separated from Macro- mischa by the larger antennal club (shorter than remainder of flagellum in Macromischa), much smaller eyes (more than 50 facets in Macromischa), presence of an antero-inferior pronotal angle, strongly sculptured head and thorax, smaller size and terricolous habitat {Macromischa is arboreal). Rogeria creightoni, new species Figure 1 DIAGNOSIS. Eyes small, composed of 8-12 pigmented facets; head and thorax coarsely reticulose; pronotum abruptly descending from mesonotum and separated from it by transverse carinula; propodeum with transverse welt basal- ly; petiolar peduncle subopaque, reticulo-punctate ventrally, without ventral crest but with distinct anterior tooth. WORKER (Holotype). Measurements: HE 0.61; HW 0.55; SL 0.41; PW 0.40; WL 0.68; TL 2.43 mm. Head as shown in figure 1, Cl 90. Mandibles smooth and shiny, with scat- tered fine piligerous punctures; cutting margin quinquedentate, apical tooth longest, remaining teeth sharply triangular. Median lobe of clypeus elevated, narrow, without median carinula but with sharp lateral carinulae, sharply trun- cate and perpendicular anteriorly in profile; lateral lobes with several costae. Frontal lobes moderately expanded, covering antennal insertions, maximum width about 1/3 HW. Scape short, ending well below occipital margin, SI 75. Flagellum 1 1 -segmented, the three-segmented club much longer than remain- der of flagellum, median segments clearly broader than long. Eyes small, con- sisting of 8-12 facets, removed from mandibular insertion by about 1.2 times their maximum diameter. Front and sides of head with fine, sharp, widely spaced longitudinal rugulae, forming reticula on occiput and genae; interspaces smooth and shiny; gula shiny, with sparse oblique rugulae. Thorax as shown in figure 1. Promesonotum with sharp humeral angles and separated from pronotal neck by a coarse, irregular rugule; dorsum rugoso- reticulate, interspaces smooth and shiny; sides shiny, with coarse, irregular rugu- lae; pronotal collar irregularly finely rugulose, interspaces densely punctate and dull; inferior pronotal angle defined but neither acute nor subdentate; lower pronotal margin transparent and slightly reflexed. Meso- and metapleurae shiny, coarsely and irregularly longitudinally rugulose. Propodeal dorsum strongly sloping, basally with distinct transverse welt, coarsely reticulose basal- ly, transversely rugulose between bases of spines, posterior face nearly vertical. 1973 Two Ant Genera New to the United States 3 Figure 1. Upper: Rogeria creightoni, worker, frontal view of head, lateral view of head, thorax and abdomen. Middle: R. huachucana, frontal view of head, lateral view of head, thorax and abdomen. Lower: Acropyga epedana, worker, frontal view of head, lateral view of head, thorax and petiole, outline of petiole in posterior view (pubescence omitted). Figures by Ruth Ann DeNicola. 4 Contributions in Science No. 236 smooth and shiny; spines about twice as long as their width at base, longer than distance between them. Tibiae II and III without evident apical spurs. Petiolar and postpetiolar profiles as in figure 1 . Petiole distinctly peduncu- late, anterior peduncle longer than height of node, laterally marginate; peduncle closely punctate, slightly shiny; node slightly shiny, densely punctate and ob- scurely transversely rugulose anteriorly, shinier, more coarsely, longitudinally, rugulose posteriorly; dull, closely punctate laterally and ventrally, with low longitudinal carina, ending anteriorly in a low, oblique, flattened tooth. Post petiole, from above, a little broader than long, node smooth and shiny, with sparse fine piligerous punctures. Caster smooth and shiny, with sparse fine piligerous punctures. Body and appendages with numerous well-spaced fully erect coarse hairs; head and gaster, in addition, with numerous finer, shorter appressed to suberect hairs. Color ferruginous, head and gaster a little darker. TYPE MATERIAL. Holotype and one paratype worker: La Feria, Camer- on County, TEXAS, 16 Feb. 1971 (W.S. Creighton). Holotype in LACM, para- type in collection of W. S. Creighton. VARIATION. The paratype is nearly identical to the holotype, with the fol- lowing measurements: HL 0.61; HW 0.53; SL 0.41; PW 0.38; WL 0.63; TL 2.34 mm; Cl 87; SI 77. DISCUSSION. The type specimens were taken from the yard of the Creighton residence in a now highly disturbed habitat, once mesquite-acacia savannah. This species is not closely allied to the following. Among the Caribbean species, it is evidently most like R. scabra Weber from Cuba, especially in de- tails of petiolar and postpetiolar sculpturation. It differs from R. scabra in the more declivitous pronotum, broader propodeal spines and the smooth and shiny posterior face of the propodeum. In Kempfs key (1963) to the South American species it goes to the vicinity of R. pellecta Kempf, but differs by the longer spines, distinct humeral angles and many other characters. The well-defined pet- iolar node will serve to separate this ant from the Irogera series of species. ETYMOLOGY. This species is dedicated to my friend and colleague, Wil- liam S. Creighton, who generously turned his specimens over to me and permit- ted deposition of the type in the LACM. Rogeria huachucana, new species Figure 1 DIAGNOSIS. Small species; body compact; eyes minute, with about four facets in greatest diameter; no erect hairs present on cephalic and thoracic dor- sa; cephalic rugulae fine and obscure; thoracic dorsum without evident rugulae; epinotal spine short, dentiform. WORKER (Holotype), Measurements: HL 0.60; HW 05.0; SL 0.40; PW 0.35; WL 0.65; TL 2.33 mm. Two Ant Genera New to the United States 5 1973 ! Head as shown in figure 1, Cl 83. Mandibles smooth and shiny, with scat- j tered fine piligerous punctures; cutting margin quinquedentate, apical tooth largest, remaining teeth low, basal tooth a little smaller. Median lobe of clypeus elevated, narrow, with fine median carinula, strongly truncate anteriorly, per- pendicular; lateral borders very weakly carinulate; lateral lobes without costae. Frontal lobes moderately expanded, covering antennal insertions, maximum width about 1/3 HW. Antennal scape short, ending well below occipital margin, SI 80. Flagellum 1 1 -segmented, the three-segmented apical club much longer than remainder of flagellum, median segments about twice broader than long. Eyes minute, consisting of about eight facets, removed from mandibular inser- tion by twice their maximum length. Front and sides of head with low, fine, ir- regular rugulae, spaces between rugulae equal to, or exceeding, width of rugu- lae; interspaces roughened and with scattered shallow punctures; occiput with a few obscure, basically transverse rugulae in middle; gula with a few coarser, transverse rugulae, shinier than front. Thorax as shown in figure 1. Promesonotum without humeral angles, gent- ly convex in both directions, inferior pronotal angle subdentate; surface with very fine, low widely spaced rugulae, interspaces roughened and slightly shiny, with scattered fine, obscure punctures, anterior face and neck of pronotum aru- gulose, closely punctate. Meso- and metapleura arugulose, finely and densely punctate. Propodeum, above lateral teeth, mostly sloping, but with narrow trans- verse basal area, abruptly sloping toward petiolar insertion below teeth; basal area densely punctate, without rugulae, sloping area with fine transverse rugu- lae, surface shinier than that of basal area; lateral teeth short, triangular, shorter than their basal width, separated by about two times their length. Legs shiny, finely and superficially punctate; tibiae II and III without evident apical spurs. Petiolar and postpetiolar profiles as in figure 1. Petiole distinctly peduncu- late; anterior peduncle a little longer than height of node, closely and finely punctate above, laterally marginate, sides and venter shiny, sparsely and finely punctate; venter with short longitudinal carina, ending anteriorly in a short tooth; node shiny, finely and sparsely punctate. Postpetiole a little broader than long, shiny, finely and sparsely punctate. Gaster smooth and shiny, with sparse fine piligerous punctures. Front and occiput of head, thoracic dorsum, petiole, postpetiole and first gastric segment with scattered fine appressed hairs. Clypeus with about six long erect hairs on disc; mandibles with a few short erect hairs; gula with about three short erect hairs on each side; coxae with a number of erect hairs; gastric seg- ments beyond first with scattered long erect hairs. Color yellowish ferruginous, head a little darker, legs a little lighter. TYPE MATERIAL. Holotype and two paratype workers; 8.1 mi. SE of Sunnyside, elev. 5950 feet, Cochise County, ARIZONA, 22 Aug. 1971 (R. R. Snelling, No. 71-41); 1 paratype worker, Huachuca Mts., Cochise Co., ARIZO- NA, elev. 5850-5900 feet, T24S, R20E, sec. 4, 2 Sept. 1972 (R. R. Snelling, No. 6 Contributions in Science No. 236 72-58). Holotype and two paratypes in LACM, one paratype in collection of W. S. Creighton. VARIATION. The four specimens are so very similar in size that there is nearly no evident variation. The following measurements exhibit the slight ranges presently known: HL 0.60-0.63; HW 0.50-0.53; SL 0.40; PW 0.35-0.36; WL 0.53-0.66; TL 2.27-2.33 mm; Cl 83-85; SI 75-80. DISCUSSION. The 1971 series was taken from beneath a large, deeply imbedded stone on a south-facing slope at the southern end of the Huachuca Mountains, in oak-juniper woodland. The single 1972 specimen was taken in a similar area, beneath a small stone which also concealed a colony of Cremato- gaster browni Buren. This species may be readily separated from the few known North American species by its small size, reduced eyes and lack of erect hairs on much of the body. It appears, to judge from the description, to be most like R. bruchi Sant- schi of Argentina, to which it will run in Kempfs key to the South American species (1963). ETYMOLOGY. Named for the Huachuca Mountains of southern Arizona whence the types were collected. Genus ACROPYGA Roger Ants of the genus Acropyga are small to minute, yellowish species which, in the field may be confused with Brachymyrmex, another genus of formicines of similar habits. Among the known genera of ants in the conterminous United States, Acropyga may be recognized by the following combination of characters: Formicinae; antennae 10-segmented (8-11 in exotic species); eyes lateral, small to minute, situated closer to mandibular articulation than occiput; ocelli absent; posterior clypeal margin contiguous with rim of antennal socket; mandibles slender, quadri- or tridentate, cutting margin oblique, often continuous with basal margin; maxillary palpi two-segmented; labial palpi three-segmented; promesonotal and metanotal sutures present; propodeum oblique; petiole high, compressed. According to the most recent treatment of the New World species Weber (1944) there are twenty-nine nominate forms in the Neotropical Region. Other species in the Ethiopian and Australasian Regions are assigned to Acro- pyga. A total of four subgenera (Acropyga s. str., Atopodon, Malacomyrma and Rhizomyrma) have been named, of which only Rhizomyrma has been re- corded from the New World. The subgenera are purportedly distinguished from one another by mandibular characters (shape and dentition) and the number of antennal segments. However, the entire range of characters is covered by the New World Rhizomyrma. Because I have not had the opportunity to examine material of the Old World species in sufficient detail, no formal synonymy is pro- posed at this time, even though I have no faith in the validity of any of these sub- generic names as they currently are characterized. The species of Acropyga are largely, if not entirely, subterranean in their 1973 Two Ant Genera New to the United States 7 habits and collections are usually from Berlese samples. Several of the Neotropi- cal species are of agricultural importance, because they tend and disperse coc- coids which feed on the roots of coffee plants. According to Weber (1944) the association is a true symbiotic one. He opined that the New World species may all be obligate coccoidophiles. For a fuller discussion of the relationship of Acro- pyga to coccoids see Weber’s paper. Acropyga epedana, new species Figure 1 DIAGNOSIS. Mandible slender, cutting margin oblique and quadridentate; eye minute, consisting of one or two facets; antennae 10-segmented, scape sur- passing occipital margin, terminal segment longer than preceding four seg- ments; pubescence and hairs abundant on all body surfaces. WORKER (Holotype), Measurements: HL 0.50; HW 0.50; SL 0.37; PW 0.03; WL 0.53; TL 1.75 mm. Head, in frontal view, as wide as long. Cl 100; widest at about eye level, sides nearly straight; occiput slightly concave in middle; in lateral view, thickest above eye level, occiput compressed, eyes at lower one-fourth of side. Mandi- bles slender, cutting margin strongly oblique, with three large triangular teeth and a minute tooth at basal angle. Apical margin of clypeus evenly slightly con- vex; clypeus in profile with apical third abruptly sloping to margin. Scape reach- ing a little beyond occipital margin, SI 74; median flagellar segments distinctly broader than long, apical segment large, longer than preceding four segments combined. Thorax short and robust, PW 0.67 x WL. Pronotal face declivitous; meso- metanotum flattened, evenly curved from pronotum to propodeum. Propodeum entirely oblique, without distinct basal face, surface flat. Legs stout; tarsal seg- ments flattened. Petiolar scale, in profile, compressed, fully erect, crest thin; in frontal view, crest very slightly concave. Integument smooth and shiny on head and thorax; head with abundant seti- gerous punctures; coarser and more conspicuous on lower half; gaster shiny, in- tegument roughened and with obscure coarse piligerous punctures. Head with abundant appressed and subappressed short, coarse pubescence on sides and beneath; front and occiput with much pubescence fully erect; with scattered long erect hairs, numerous on clypeus. Thorax and propodeum with sparse subappressed to fully erect coarse pubescence and scattered long, erect coarse hairs. Gaster with abundant pubescence, subappressed to erect on ter- gites, more closely appressed laterally, appressed on venter; with numerous long, fully erect hairs. Scape pubescence abundant, reclinate to suberect, coarse. Legs similar, coarse hairs abundant on tibiae; tarsal segments with coarse seta- like hairs and a coarse seta on apical corners of segments. Color uniformly brownish yellow, mandibles transparent, cutting margin piceous. 8 Contributions in Science No. 236 FEMALE. Unknown. MALE. Unknown. TYPE MATERIAL. Holotype and three paratypes, 3.1 mi. W of summit of Montezuma Pass, 5800 ft. elev., Huachuca Mts., Cochise Co., ARIZONA, 14- VIII-1969 (R. R. Snelling, No. 69-307), all deposited in LACM. DISCUSSION. The type specimens were under a stone also concealing a colony of Paratrechina sp., in an area of oak-juniper woodland. The species is probably wholly hypogaeic. In the key by Weber (1944) A. epedana will run to A. mesonotalis Weber from Haiti. The mesonotum of that species is distinctly elevated and differen- tiated from the remainder of the thorax in profile; in A. epedana the mesonotum is nearly flat and continuous with the metanotum in profile. Geographically, the nearest described species is A. exsanguis (Wheeler) based on three specimens from Jalapa, Mexico. That species has tridentate mandibles, 8-9 segmented an- tennae and lacks erect hairs over most of the body. The Honduran A. wheeler i Mann also has tridentate mandibles, shorter scapes which Just attain the occipi- tal margin, the antennae are nine-segmented with the last segment almost as long as the preceding three and the propodeum with a distinct basal face. The Panamanian species, A. panamensis Weber, has a distinctly transverse mandibu- lar cutting margin, short scapes which do not attain the occipital margin and the head is longer than broad. In A. goeldii Forel, the remaining recorded Central American species, the eyes consist of several facets, the mesonotum is high and strongly arched and the propodeum has a distinct, though short, basal face. ETYMOLOGY. Gr., epedanos, weak or feeble, so-called because of the thin, collapsible integument. Literature Cited Kempf, W. W. 1961. Remarks on the ant genus “Irogera” Emery, with the description of a new species. Rev. Brasil. Biol. 21:435-441. 1962. Miscellaneous studies on Neotropical ants. II. Studia Entomol. 5:1-38. 1963. Additions to the Neotropical ant genus “Rogeria” Emery, with a key to the hitherto recorded South American species. Rev. Brasil. Biol. 23:189-196. 1964. Miscellaneous studies on Neotropical ants. III. Studia Entomol. 7:45-71. 1965. Nota preliminar sobre algumas formigas Neotropicas, descritas por Frederick Smith. Rev. Brasil. Biol. 25:181-186. Weber, N. A. 1944. The Neotropical coccid tending ants of the genus Acropyga Roger. Ann. Entomol. Soc. Amer. 37:89-122. Accepted for publication August 7, 1972 Printed in Los Angeles, California by Anderson, Ritchie and Simon on Simpson Lee Recovery Text f .5-0 7. 7J C:ilu9 NUMBER 237 JANUARY 19, 1973 A NEW BUTTERFLYFISH OF THE GENUS CHAETODON AND A NEW ANGELFISH OF THE GENUS CENTROPYGE FROM EASTER ISLAND ' By John E. Randall and David K. Caldwell |i 1 ! CONTRIBUTIONS IN SCIENCE NATURAL HISTORY MUSEUM • LOS ANGELES COUNTY A NEW BUTTERFLYFISH OF THE GENUS CHAETODON AND A NEW ANGEEFISH OF THE GENUS CENTROPYGE FROM EASTER ISEANDi By John E. Randall^ and David K. Caldwell^ Abstract: The chaetodontid fish Chaetodon lit ns Randall and Caldwell is described from Easter Island, the only known locality. It is brown with no dark markings; the dorsal and anal fins are narrowly edged with white except the spinous portion of the anal which is almost entirely white; the caudal is pale posteri- orly. The pomacanthid Centropyge Iwtnnuitna Randall and Caldwell is also described from Easter Island. It is brown, with the head, nape, abdomen, and caudal peduncle orange-yellow; a black spot appears posteriorly on the opercle and a blue ring nearly encircles the eye; a blue margin is visible on the soft portion of the anal fin, with blue markings posteriorly in the dorsal and anal fins. This species was collected subsequently in the Pitcairn Group, Rapa, and the Austral Islands. In his “Los Peces de la Islade Pascua,” de Buen (1963) listed the 40 species of fishes known from Easter Island at that time. Among them was one species of butterflyfish, Eorcipiger longirostris, now recognized as E. flavissimus Jordan and McGregor (Randall and Caldwell, 1 970). No angelfishes were listed. Three large collections of fishes made in recent years at Easter Island have increased the number of species of fishes known from the island to 107. The first was a single large rotenone station made inshore at Anakena Cove by Ramsey Parks and the crew of the yacht “Chiriqui” on October 1, 1958. This collection was held at UCLA in the care of Boyd W. Walker until 1960 and was then transferred to the Natural History Museum of Los Angeles County (LACM). Extensive collections were made during the Canadian Medical Expedition to Easter Island in 1964 and 1965 by Ian E. Efford, Jack A. Mathias and associates. These fishes are deposited mainly at the University of British Columbia (BC). The third collection was made by Randall, Gerald R. Allen, and Bruce A. Baker in January and February, 1969, with support of a grant from the National Geographic Society. Except for certain types, these speci- mens are at the Bernice P. Bishop Museum (BPBM), Honolulu. Thanks are due Robert J. Lavenberg and Norman J. Wilimovsky for the loan of LACM and BC specimens, respectively, and for field data. These collections contained additional specimens of Eorcipiger flavissi- 1 Review Committee for this Contribution James Bohlke William N. Eschmeyer Robert J. Lavenberg Camm C. Swift ^Bernice P. Bishop Museum, Honolulu, Hawaii ^Communication Sciences Laboratory, University of Florida, Gainesville, Florida 1 Contributions in Science No. 237 |t 2 jl mus. They also included specimens of a new chaetodontid fish of the genus j Chaetodon and a new pomacanthid fish of the genus Centropyge which are the I subject of the present paper. We are also grateful to Dr. Herbert R. Axelrod of ij TFH Publications for the color plate of the two new species of fishes. I In addition to the Bishop Museum, Natural History Museum of Los i Angeles County, and the University of British Columbia, paratypes have been j' deposited at the British Museum (Natural History) [BM(NH)], California i Academy of Sciences (CAS), Museum National d’Histoire Naturelle (MNHN), I and the National Museum of Natural History (USNM). ! Chaetodon litus Randall and Caldwell, new species j Figures 1, 2, Table 1 i Holotype: BPBM 6659, 96.5 mm standard length (SL), female, Easter Island, I Anakena Cove, 6-10 ft., Chemfish, J. E. Randall and G. R. Allen, January j 31, 1969. Paratypes: LACM 6560-1, 19(85-1 18.5 mm SL), Easter Island, Anakena Cove, east side, 100 yd. NE of sand beach, boulder bottom with brown algae (75%) and Litlio- thamnium (15%) cover, 0-15 ft., water temperature 69° F, rotenone, R. Parks and crew of the ketch “Chiriqui,” January 1, 1958; BC 65-403, 1(107.5 mm SL), Easter Island, Hanga Roa, rocky pool, 3-5 m, rotenone and spear, I. E. Efford and J. A. Mathias, December 23, 1964; BC 65-407, 1(112 mm SL), Easter Island, 200 m N of Hanga Roa, I. E. Efford and J. A. Mathias, December 23-24, 1964; BC 65-429, 2(107 — 108 mm SL), Easter Island, Rano Raraku on south coast, rotenone, I. E. Efford and J. A. Mathias, January 8, 1965; BC 65-430, 2(100.5— 107 mm SL), Easter Island, Rano Raraku area, 3-8 m, rotenone and spear, I. E. Efford and J. A. Mathias, January 8, 1965; BC 65-434, 1(93 mm SL), Easter Island, 200 m N of Hanga Roa, I. E. Efford and J. A. Mathias, January 13, 1965; BC 65-440, 2(44.5 — 96 mm SL), Anakena Cove, tide pools, 0.5-8 m, rotenone, I. E. Efford and J. A. Mathias, January 15, 1965; BC 65-446, 2( 1 10— 122.5 mm SL), Anakena Cove, 7-8 m, rotenone, I. E. Efford and J. A. Mathias, January 16, 1965; BC 65-451, 1(93.5 mm SL), Easter Island, off Hanga Roa, 1-5 m, rotenone, I. E. Efford and J. A. Mathias, February 2, 1965; BC 65-455, 2(39.5 —40 mm SL), Easter Island, Hanga Roa, subtidal, rotenone, I. E. Efford and J. A. Mathias, February 5, 1965; BC 65-457, 1(1 10.5 mm SL), Easter Island, Hanga Roa, subtidal, rotenone, I. E. Efford and J. A. Mathias, February 5 or 6, 1965; BC 65-458, 2(42— 111 mm SL), Easter Island, Hanga Roa, rotenone, I. E. Efford and J. A. Mathias, February 6, 1965; BPBM 6655, 2(99 — 107 mm SL), Easter Island, Hanga Piko boat channel, 6 ft., spear, G. R. Allen, January 18, 1969; BPBM 6660, 1(107 mm SL), Easter Island, Hanga Piko boat channel, 10 ft., spear, J. E. Randall, January 20, 1969; BPBM 6662, 4(36-40 mm SL), Easter Island, tide pools between Hanga Roa and Hanga Piko, 1-3 ft., J. E. Randall and G. R. Allen, January 25, 1969; BPBM 10438, 1(87 mm SL), same data as holotype; USNM 208218, 1( 100.5 mm SL), same data as holotype; BPBM 6661, 2(40 — 1 15 mm SL), Easter Island, wreck between Hanga Roa and Hanga Piko, 15 ft., spear and Chemfish, J. E. Randall and B. A. Baker, January 27, 1969; BPBM 6658, 1(107 mm SL), Easter Island, Mataveri O Tai, 25 ft., Chemfish and spear, J. E. Randall and G. R. Allen, February 2, 1969; BPBM 6663, 1(39.8 mm SL), Easter Island, off Ahu Akapu on west coast, 70 ft., Chemfish, J. E. Randall and B. A. Baker, February 3, 1969; BM (NH) 1972.10.5.2, 1(41.7 mm SL), same data as preceding; BPBM 6657, 1(113 mm SL), Easter Island, off Ahu Akapu, 80 ft., Chemfish, J. E. Randall and 1973 New Easter Island Fishes 3 G. R. Allen, February 5, 1969; BPBM 6664, 1(39 mm SL), Easter Island, tide pools j between Hanga Roa and Hanga Piko, 1-3 ft., Chemfish, J. E. Randall and G. R. Allen, February 6, 1969; BPBM 6656, 1(97.2 mm SL), Easter Island, off Motu Tautara, 60 ft., Chemfish, J. E. Randall and G. R. Allen, February 7, 1969. | Diagnosis: A Chaetodon, as defined by Ahl (1923), with dorsal rays XIII (rarely | XIV), 22 to 25; anal rays III, 1 8 to 2 1 ; pectoral rays 15 or 16; lateral-line scales | 37 to 43; snout slightly produced, its length 2.6 to 2.8 in head; body depth 1.6 ; to 1.8 in SL; brown, the centers of the scales paler than edges, with no black | markings on head or body; margins of dorsal and anal fins white, the spinous portion of the anal almost completely white; a crescentic hyaline zone pos- ; teriorly in caudal fin. | Description (counts in parentheses apply to 39 paratypes unless otherwise specified): Dorsal rays XIII, 25 (22 to 25, except two XIV, 23, modally 23 or 24); anal rays III, 20 (18 to 21, modally 19, one with 18 and one with 21); Table 1 Proportional Measurements of Type Specimens of Chaetodon litiis expressed as Thousandths of SL. HOLOTYPE PARATYPES BPBM USNM BPBM BPBM BPBM 6659 208218 6656 10438 6663 Standard length (mm) 96.5 100.5 97.2 87.0 39.8 Greatest depth of body 593 616 592 610 553 Width of body at gill opening 145 151 139 147 153 Head length 297 290 305 301 354 Snout length 110 110 1 17 109 125 Diameter of eye 85 88 89 95 119 Postorbital length of head 132 123 130 126 134 Bony interorbital width 90 90 92 95 113 Length of upper jaw 75 75 74 71 93 Least depth of caudal peduncle 102 96 100 103 113 Length of caudal peduncle 50 50 56 45 42 Snout to original of dorsal fin 383 375 380 379 410 Snout to origin of pelvic fins 409 403 423 407 459 Length of caudal fin 197 204 203 230 248 Length of pectoral fins 255 274 267 276 280 Length of pelvic fins 279 280 276 284 352 Length of pelvic spine 197 224 211 225 276 Length of dorsal fin base 806 807 772 815 691 Length of first dorsal spine 69 89 80 85 131 Length of second dorsal spine 135 139 164 172 280 Length of third dorsal spine 180 177 200 203 321 Length of last dorsal spine 181 174 189 202 228 Length of longest dorsal soft ray 196 209 207 237 227 Length of anal fin base 383 384 364 402 370 Length of first anal spine 109 101 121 124 155 Length of 2nd anal spine 238 246 220 230 250 Length of third anal spine 228 263 231 233 253 Length of longest anal soft ray 209 204 210 230 248 4 Contributions in Science No. 237 I pectoral rays 16(15 or 16, modally 15, except one abnormal specimen with 12 I and another with 13 rays on one side; normally upper two and lower-most unbranched); pelvic rays 1,5; principal caudal rays 17 (15 branched); pored I lateral-line scales 37 to 43 (modally 39 and 40); vertical scale rows from upper I end of gill opening to caudal base about 50; scales above lateral line to origin of dorsal fin 13; scales below lateral line to origin of anal fin 26; circum- peduncular scales 27; gill rakers 18 (4 specimens) branchiostegal rays 6; vertebrae 24 (34 specimens). Body deep, the maximum depth 1 .6 to 1 .8 in SL, and compressed, the width behind gill opening 3.6 to 4.3 in depth; head (measured from front of upper lip to end of opercular membrane) 2.8 to 3.5 in SL (head relatively long in juveniles); snout slightly produced, its length to front of upper lip 2.6 to 2.8 in head; interorbital space convex, the bony width 3.1 to 3.3 in head; caudal peduncle deeper than long, the least depth 2.9 to 3.15 in head, the length (measured horizontally from rear base of anal fin to caudal base) 2.3 to 2.55 in least depth. Mouth small, oblique, the lower jaw projecting; maxillary reaching a vertical below nostrils. Teeth very close-set, slender, and elongate (the length about one-fifth eye diameter in adults), in six rows at front of upper jaw and nine at front of lower. A small bony prominence at front of snout aligned with anterior and posterior nostrils, distance separating these knobs contained about four times in eye diameter of adults. A slight, fleshy prominence at corner of mouth below posterior end of maxillary. Anterior and posterior nostrils separated by a distance about equal to opening of anterior nostril; a well-developed fleshy rim on anterior nostril, broadening to a slight flap dorsoposteriorly; posterior nostril slightly elongate horizontally, with a low fleshy rim anteriorly. Margins of opercular bones smooth except for a few small serrae or crenulations on the broadly rounded angle of the preopercle. Scales finely ctenoid. Head completely scaled except for a narrow region at front of snout including the pair of bony prominences mentioned above; scales on head and thorax notably smaller than on body, those on opercle in about 8 or 9 irregular near-vertical rows (as counted near ventral edge). Small scales reaching nearly to margins of dorsal and anal fins except anterior spinous portions; scales on caudal fin extending about two-thirds to three-fourths the distance to posterior margin; paired fins scaled only basally. Lateral line following curved contour of back, ending beneath base of about third-from-last dorsal ray. Origin of dorsal fin above upper edge of preopercular margin; first dorsal spine about half length of second, the second spine about 1 .2 to 1 .3 in third (of adults); fifth and sixth dorsal spines the longest, about 1.4 in head; last dorsal spine very slightly shorter than third. Longest dorsal soft ray (about the tenth) approximately equal to longest dorsal spine. Interspinous membranes of anterior portion of dorsal fin deeply incised. Figure 1. Juvenile of Chaetodon litas Randall and Caldwell, paratype, BPBM 6662, 36 mm SL, Easter Island. Anal spines notably longer and stouter than first three dorsal spines, first about equal to snout length, second about twice as long as first, and third subequal to second. Longest anal soft ray (about the sixth) slightly shorter than second and third anal spines. Caudal fin varying from slightly rounded to slightly emarginate, its length 1 .3 to 1 .5 in head. Pectoral fins somewhat pointed, slightly shorter than head (1.05 to 1.25 in head length), not reaching as far posteriorly as pelvic fins. Origin of pelvic fins posterior to base of pectorals; filamentous tip of first pelvic ray reaching to or beyond anus; pelvic spine 1 .2 to 1 .5 in head. ' ■"^1 <:C'- . a=’^;V.; ®;i ^!»s^i^^^;aa^^wfis^ ■fe !• ’J^®v\f '^V ^^•^^^^^V,;^^-• Fig. 3. Centropyge hotumatua Randall and Caldwell, holotype, BPBM 6665, 67.6 mm standard length, Easter Island. Fig. 2. Chaetodon litus Randall and Caldwell, holotype, BPBM 6659, 96.5 mm standard length, Easter Island. 6 Contributions in Science No. 237 Color in alcohol brown, centers of scales paler than edges; dorsal and anal fins brown with a narrow whitish margin except spinous portion of anal fin almost entirely whitish; dorsal fin with a dark brown submarginal line (brown of anal fin slightly darker than dorsal fin and submarginal dark brown not visible or only faintly discernible); caudal fin brown with a pale crescentic area posteriorly in fin (nearly one-third length of fin in broadest central portion); pectoral fins pale, except scaled basal portion brown; pelvic fins dark brown, except spine pale. In life this fish is colored almost the same as in preservative; the contrast between the darker brown edges of the scales of the body and the paler centers is greater, and the white edges of the dorsal and anal fins are more vivid in life. Remarks: The closest relative of Chaetodon litus is an undescribed species from Pitcairn and Rapa which is black on its anterior half and abruptly bright yellow on the posterior half (Burgess and Randall, MS). Of the described species in Chaetodon, C. litus most resembles C. daedalma Jordan and Fowler (1902) in color; however, daedalma is clearly distinct in having 16 anal rays, short posterior spines in the dorsal fin, a more rounded caudal fin, and larger scales on the opercle. C. litus was collected only at Easter Island; it was taken from the shallows to depths of 80 feet. The young were found in tidepools. Randall (1970) wrote that G. R. Allen once observed it picking at the bodies of other fishes, hence it appears to be a part-time cleaner, as reported for a few other species of Chaetodon. Named litus, a greek adjective, meaning plain or simple, in reference to its plain color — very much in contrast to the striking color patterns of most butterflyfishes. Centropyge hotumatua Randall and Caldwell, new species Figures 3, 4, Table 2 Holotype: BPBM 6665, 67.6 mm SL, male, Easter Island, offshore from Ahu Akapu on the western side, 70 ft., bottom mainly coral and coral rock with brown algae, Chemfish, J. E. Randall and B. A. Baker; February 3, 1969. Paratypes: BPBM 10439, 1(64.0 mm SL), same data as holotype; MNHN 1970-1 16, 1(53.5 mm SL), off Ahu Akapu, 80 ft., Chemfish, J. E. Randall and G. R. Allen, February 5, 1969; USNM 205521, 1(64.8 mm SL), same data as preceding; CAS 13390, 1(45.2 mm SL), off Motu Tautara on west side of island, 125 ft., sand and rock bottom with ledges, Chemfish, J. E. Randall and B. A. Baker, February 12, 1969; BPBM 133 14, 2(27.8 — 36.8 mm SL), Oeno, Pitcairn Group, north side of atoll, reef in 40-60 ft., rotenone, J. E. Randall and crew of the schooner “Westward,” December 18, 1970; BPBM 13326, 2(22.6 — 27.5 mm SL), Pitcairn, off Down Rope on southeast side of island, reef in 90 to 100 ft., rotenone, J. E. Randall, D. B. Cannoy, and S. R. Christian, December 23, 1970; LACM 32817-1, 2(23.8—57.5 mm SL), Pitcairn, off Christian’s Point, 40-60 ft., rotenone, J. E. Randall, D. B. Cannoy, and S. R. Christian, December 28, 1970; BM(NH) 1972.10.5.1, 2(23.0— 5 1.0 mm SL), 1973 New Easter Island Fishes 7 same data as preceding; BPBM 13312, 5(24.9-61.8 mm SL), Pitcairn, off McCoy on south side of island, reef in 75-85 ft., rotenone, J. E. Randall, D. B. Cannoy, J. R. ! Haywood, J. D. Bryant, and S. R. Christian, January 4, 1971; BPBM 13315, 3(24.5- 46.2 mm SL), Pitcairn, patch reef off Gannet Ridge on north side of island, 130-145 vi ft., rotenone, J. E. Randall, D. B. Cannoy, J. R. Haywood, R. R. Costello, J. D. Bryant, and C. R. Christian, January 6, 1971; BPBM 12275, 3(32.0-48.0 mm SL), C Ducie, Pitcairn Group, off southwest side of atoll; reef in 100 ft., rotenone, J. E. II Randall, D. B. Cannoy, R. R. Costello, S. R. Christian and R. N. McNair, January II 15, 1971; BPBM 13012, 1(60.6 mm SL), Rapa, reef at entrance to Haurei Bay, 50 ft., spear, J. E. Randall, February 15, 1971; BPBM 12793, 1(33.0 mm SL), Raivavae, Austral Islands, outside barrier reef, south of Motu Haa, 120 ft., spear, D. B. Cannoy, February 25, 1971. Diagnosis: A Centropyge, as defined by Fraser-Brunner (1933), with the l|l head, thorax, abdomen, and caudal peduncle pale (yellow-orange in life) in 1} contrast to the dark brown median portion of body, a black spot posteriorly on opercle associated with small blue markings, and a blue ring around all but 3 anterior edge of eye. , ; Description (counts in parentheses apply to Easter Island paratypes): Dorsal rays XIV, 17 (one with 18) (last ray branched to base); anal rays 111,18 (last 2 rays close at base); pectoral rays 17 (one with 16) (upper 2 and lowermost unbranched); pelvic rays 1,5; branched caudal rays 15 (upper and lowermost ■ unbranched rays of fin not reaching posterior margin); pored scales of lateral line 36 (34 to 37); vertical scale rows from upper end of gill opening to caudal base 45 (43 to 46); scales above lateral line to origin of dorsal fin 7 (one with 6); scales below lateral line to origin of anal fin 19 (18 to 20); circumpeduncular scales 20 (20 to 22) ; gill rakers 6+17 (one with 6+16) (raker at angle included in lower-limb count); branchiostegal rays 6; upper teeth 70 (62 to 68); lower teeth 68 (53 to 60). Body deep, the maximum depth 1.8 to 1.9 in SL, and compressed, the width behind gill opening 2.6 to 3 in depth; head (measured from front of upper lip to end of opercular membrane) 3 to 3.35 in SL; snout 2.65 to 2.95 in head; eye 2.55 to 3.1 in head; interorbital space slightly convex, the bony width 3.2 to 3.65 in head; caudal peduncle deeper than long, the least depth 2.2 to 2.5 in head, the length (measured horizontally from rear base of anal fin to caudal base) 1.5 to 1.6 in least depth. Mouth small, terminal, the gape horizontal; maxillary reaching slightly posterior to a vertical at anterior nostril; upper lip nearly as broad (vertically at front) as lower, the upper lip height almost half eye diameter. Teeth slender and elongate, close-set, flexible, tricuspid (the central cusp notably longer and broader than the lateral ones), in about three well-spaced rows in jaws, the teeth of the outer row the largest; no teeth on roof of mouth; tongue short and broadly rounded. Gill membranes narrowly attached to isthmus; 5 (1 to 3) fleshy papillae in mid-ventral line between chin and isthmus; anterior and posterior nostrils separated by a space about half diameter of posterior nostril; posterior nostril slightly larger than anterior, separated from eye by a space about equal to its 8 Contributions in Science No. 237 Table 2 Proportional Measurements of Type Specimens of Centropyge hotumatua expressed as Thousandths of SL. HOLOTYPE PARATYPES BPBM 6665 BPBM 10439 USNM 205521 CAS 13390 MNHN 1970-1 16 Standard length (mm) 67.3 64.0 64.8 45.2 53.5 Greatest depth of body 557 532 542 538 555 Width of body behind gill opening 195 202 186 192 187 Head length 327 325 298 304 328 Snout length 118 122 111 109 111 Diameter of eye 106 108 102 118 117 Postorbital length of head 106 105 102 106 1 10 Bony interorbital width 91 89 93 94 90 Least depth of caudal peduncle 132 129 133 133 136 Length of caudal peduncle 82 86 85 88 — Snout to origin of dorsal fin 405 405 381 388 384 Snout to origin of pelvic fins 375 378 367 380 360 Length of caudal fin 297 300 278 293 295 Length of pectoral fin 302 313 303 308 328 Length of pelvic fin 363 314 360 377 341 Length of pelvic spine 182 181 191 185 — Length of dorsal fin base 712 680 701 700 — Length of first dorsal spine 62.5 70 69 67 72 Length of second dorsal spine 89 109 107 1 12 1 16 Length of third dorsal spine 145 141 139 148 146 Length of last dorsal spine 201 200 201 208 188 Length of longest dorsal soft ray 221 219 218 204 216 Length of anal fin base 408 400 415 410 — Length of first anal spine 113 125 136 111 114 Length of second anal spine 190 172 200 175 175 Length of third anal spine 231 228 234 225 215 Length of longest anal soft ray 263 259 255 265 247 diameter; anterior nostril with a membraneous edge and an elevated posterior flap. A stout spine at corner of preopercle, its length varying from about three-fourths eye diameter to full eye diameter; 12 (10 to 13) small spines along upper margin of preopercle and one nearly half as long as spine at angle on lower edge of preopercle, usually contiguous to spine at angle; 2 or 3 moderate spines on interopercle; a few small spines on subopercle; opercle with 2 broad flat spines, one near upper end of gill opening and the other at about the level of lower edge of eye; preorbital with 4 to 7 spines along ventral edge. Scales coarsely ctenoid (up to 32 ctenii on margins), the exposed portion ridged; scales extending more than halfway to margin of dorsal and anal fins (nearly to margin in middle of fins); scales on basal third of caudal fin, with small scales extending out on rays for another third of length of fin; paired fins scaled only basally. Head almost completely scaled. 1973 New Easter Island Fishes 9 ^ Lateral line steeply arched, ending at about base of fourth-from-last dorsal ray. i Origin of dorsal fin above upper end of gill opening; first dorsal spine ' about two-thirds length of second dorsal spine; second spine about three-fourths . length of third spine; last four spines subequal, the last contained 1.45 to 1.7 times in head length; longest dorsal soft ray (about the eighth) 1.35 to 1.5 in head length; membranes of anterior portion of dorsal fin deeply incised, while those between posterior spines only slightly indented; a small cirrus projecting from membrane just posterior to tips of dorsal spines. The three anal spines slightly longer than their dorsal counterparts, the third spine 1.25 to 1.5 in head length; longest anal soft ray (about the eighth) 1.15 to 1.3 in head length. Caudal fin nearly as long as head, its posterior border varying from near- truncate with the lobes slightly produced, particularly the upper (holotype), to slightly rounded. Pectoral fins slightly pointed, about equal to head length, reaching to level of origin of anal fin or slightly beyond. Origin of pelvic fins slightly posterior to base of pectorals; first soft ray of pelvic fins prolonged, reaching well beyond origin of anal fin (varying from base of second anal spine to base of second anal soft ray); pelvic spine about half length of pelvic fin. Color in alcohol: body dark brown posterior to a line passing from origin of dorsal fin slightly posterior to pectoral base and thence to origin of anal fin; body anterior to this line, and head, pale; caudal peduncle pale (in the smallest paratype the dark brown zone is narrower, beginning at an approximate line from base of fifth dorsal spine to base of third anal spine and ending posterior to a line from base of ninth dorsal soft ray to base of twelfth anal soft ray); upper posterior portion of opercle and opercular membrane above lower opercular spine dark brown; some dark pigment on shoulder region at upper end of gill opening, largely covered by opercle; posterior two-thirds of eye with an irregular dark brown edge which is broadest posteriorly (about one-fifth eye diameter at its widest point), some pigment extending downward onto upper posterior part of eye; a small bilobed brown spot on preopercular margin on left side (more pigment on three of the paratypes at this location, particularly the smallest); centro-basal region of dorsal and anal fins, where covered with ciliated scales like the body, dark brown; rest of these fins brown, paler distally, with a trace of irregular horizontal banding distally, posterior to eleventh dorsal spine, and a large dark brown (almost black) spot on outer part area between 7th to 13th dorsal soft rays (within this large spot there are elongate blotches of denser pigmentation); some dark pigment on outer part of 14th and 15th soft dorsal rays; cirri at tips of dorsal spines and some of adjacent mem- branes blackish; caudal fin dusky, most of pigment on rays; paired fins pale; membraneous flap lying beneath and slightly above spine at corner of pre- opercle brownish. In life the pale areas are bright orange-yellow, this color extending along Contributions in Science No. 237 10 Figure 4. Transforming prejuvenile and juvenile of Centropyge hotumatua Randall and Caldwell, paratypes, BPBM 13326, 22.6 and 27.5 mm SL, Pitcairn. anterior and outer part of dorsal and anal fins, where largely masked by brown, and onto upper and lower edges of caudal fin; rim of posterior two-thirds of orbit bright blue (a zone of black on upper part of eye below the blue edge); some blue markings around black spot at upper posterior portion of opercle; short blue lines in large dark spot posteriorly in dorsal fin, mostly parallel to rays, those in outer part of spot longer; a blue line on outer part of 14th and 15th rays; first half of soft portion of anal fin with a bright blue margin, followed by a series of bright blue lines, each along outer part of next six rays; last four rays of fin without blue markings. Remarks: Eighteen valid species of Centropyge are presently known, including hotumatua but not Holacanthus multifasciatus Smith and Radcliffe which Fraser-Brunner (1933) placed in Centropyge. The latter species warrants 1973 New Easter Island Fishes 11 a genus by itself (Randall, MS). In addition, Randall has collected three other forms of Centropyge in Oceania which await description (Randall and Burgess, ^ in progress). All of the species of Centropyge are very similar in body form and counts, but they are distinctive in color pattern. It is principally in color that hotumatua is separable from the other species of the genus. C. hotumatua was first collected at Easter Island in 1969, and the descrip- tion was prepared from specimens taken then. It was believed that this angelfish was another native species among the high percentage of endemic fishes known for the island (Randall, 1970). However, it was collected in 1970-71 at islands of the Pitcairn Group, Rapa, and Raivavae in the Austral Islands. It was also seen at Tubuai at 100 to 125 feet. It was not observed, however, at Rurutu in the Australs nor at Rarotonga in the Cook Islands. This species was collected on coral or rock bottoms in the depth range of 40 to 145 feet. Named for Hotumatua, the legendary Polynesian chieftain who first colonized Easter Island. Literature Cited Ahl, E. 1923. Zur Kenntnis der Knochenfischfamilie Chaetodontidae insbesondere der Unterfamilie Chaetodontinae. Arch. Natg. Berlin 89(5): 1-205. De Buen, F. 1963. Los Peces de la Isla de Pascua. Bol. Soc. Biol. Concepcion 35-56: 3-80. Fraser- Brunner, A. 1933. A revision of the chaetodont fishes of the subfamily Pomacanthinae. Proc. Zool. Soc. London, pp. 543-599. Jordan, D. S., and H. W. Fowler. 1902. A review of the Chaetodontidae and related families of fishes found in the waters of Japan. Proc. U.S. Nat. Mus. 25:5 13-563. Randall, J. E. 1970. Easter Island an ichthyological expedition. Oceans 3(3): 48-59. AND D. K. Caldwell. 1970. Clarification of the species of the butterflyfish genus Forcipiger. Copeia 4: 727-731. Accepted for publication December 29, 1972 NUMBER 238 JANUARY 29, 1973 Soq.'JZ L sQg THE ANT GENUS CONOMYRMA IN THE UNITED STATES (Hymenoptera: Formicidae) By Roy R. Swelling CONTRIBUTIONS IN SCIENCE NATURAL HISTORY MUSEUM • LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- able footnotes must be typed on separate sheets. Several of one kind may be placed on a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing with Latin American subjects. Summaries in other languages are not required but are strongly recommended. Summaries will be published at the end of the paper. (8) A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor j j r ! THE ANT GENUS COlStOMYRMA IN THE UNITED STATES I (HYMENOPTERA: FORMICIDAE)* j| I By Roy R. Snelling2 i Abstract: The generic name Conomyrma Ford is recognized as a valid genus and all United States species formerly assigned to I Dorymyrmex are placed in Conomyrma. The subgenus Biconomyrma - Conomyrma (N. SYN.). The specific and varietal names applied to i North American forms are considered and the following species are i| recognized: bicolor (Wheeler), flavopecta (M. Smith) and insana j (Buckley) (= pyramicus of North American literature = flavus Mc- Cook = nigra Pergande = antillana Ford = smithi Cole = brunnea of I North American literature = wheeleri Kusnezov, all N. SYN.). i j Conomyrma was established by Forel (1913) as a subgenus of Dormyrmex. i Santschi (1922) designated Prenolepis pyr arnica Roger, one of the originally i included species, as the type species of Conomyrma; Donisthorpe (1943) desig- nated the same species as the type. The subgeneric name was rejected by Creighton (1950) on the grounds that Conomyrma was coextensive with Dory- myrmex, once those species lacking a propodeal tooth were removed to Arauco- myrmex, as was done by Gallardo (1916). Kusnezov (1952) restudied the problem and proposed to divide Dorymyr- mex into two genera, Dorymyrmex and Conomyrma. Within Dorymyrmex five subgenera, including Araucomyrmex, were recognized. Conomyrma was divided into two subgenera: Biconomyrma and Conomyrma, s. str. The genus Cono- myrma was separated from Dorymyrmex in the worker caste by the lack of a psammophore and in the female caste by the presence of a single, versus two, cubital cells in the forewing. Eisner (1957) noted that the proventriculi of the workers of Dorymyrmex and Conomyrma were different. Larvae were described by G. C. and J. Wheeler (1951) for “Dorymyrmex pyramicus” {-Conomyrma pyramica, sensu Kusnezov) and “Araucomyrmex tener” and differences be- tween them were noted. The two were separated in the key by means of the posterior projection: a postero-ventrally directed cone in pyramicus and a knob in tener. Kusnezov (1952) failed to designate type species for the new subgenera proposed, but corrected his oversight in a subsequent paper (1959). In this latter paper he proposed elevation of several of the subgenera recognized in 1952 to full generic status. Thus, Biconomyrma was elevated to generic level. 'Review Committee for this Contribution Arthur C. Cole, Jr. Charles L. Hogue George C. Wheeler ^Entomology Section, Natural History Museum of Los Angeles County, Los Angeles, California 90007 1 2 Contributions in Science No. 238 Figure 1-4, profile of head and thorax of: 1, Conomyrma pyramicus (Brazil); 2, C. insana '(Texas); 3, C. flavopectus (Florida); 4, Dorymyrmex sp. (Chile). Figures 5-6, underside of head of: 5, C. bicolor (Arizona); 6, Dorymyrmex sp. (Chile). Figures 7-8, forewing of female: 7, C. insana (Texas); 8, C. pyramicus (after Kusnezov, 1952). Figures by Ruth A. DeNicola. 1973 The Ant Genus CONOMYRMA in the United States 3 The separation of Biconomyrma from Conomyrma depends upon slight differences in wing venation of the female and thoracic profile of the worker. Since one species (flavopectus M. Sm.) possesses worker characteristics of Conomyrma, s. str., and female characteristics of Biconomyrma, it is obvious that the attempted segregation of Conomyrma species into subgenera is un- tenable. Hence, Biconomyrma - Conomyrma (NEW SYNONYMY). Conomyr- ma, as a genus, is sufficiently defensible in the characteristic wing venation of the female, the reduced psammophore, and structure of the proventriculus in the worker that it may be recognized as a genus apart from Dorymyrmex. I differ with Kusnezov and prior authors with respect to their use of the word psammophore. These workers claim that a psammophore is present in ants now assigned to Dorymyrmex but absent in those placed in Conomyrma. I believe that the setalike hairs on the ventral head surface must be considered a psammophore. In those ants which truly lack a psammophore, the hairs present are short, usually irregular in length, and randomly distributed. A psam- mophore is said to be present when there is a discrete group of elongate hairs, uniform in length, arranged in a definite pattern, the result of which is the pres- ence of a discrete “basket” on the cephalic venter. These hairs are typically flattened and distinctly curved or even curled. Such is the case in both Cono- myrma (Fig. 5) and Dorymyrmex (Fig. 6). The only appreciable difference is that in the latter genus the hairs extend forward over the oral cavity; they are half, or more, as long as the head is wide. In Conomyrma the hairs are quite short, much less than half the head width, and end far short of the oral cavity. Dorymyrmex, with its related subgenera and/or genera, was restricted to South America by Kusnezov (1952). Conomyrma, together with Biconomyrma, was distributed from Argentina and Chile north to the United States and the Caribbean. Kusnezov (1952) listed the species in Conomyrma- Biconomyrma. Half of these are South American species and have no direct bearing on our problems, hence are not considered. Roger (1863) described Prenolepis pyramica from a single Specimen from Corrientes, State of Bahia, Brazil. The name was transferred to Dorymyrmex by Mayr (1866). By 1900 pyramicus was assumed to range from Argentina to the southern United States and over the entire Caribbean area. It had acquired a number of varieties and subspecies and had also received a wholly different identity. Wheeler (1902) recognized that Formica insana Buckley, 1866, described from central Texas, belonged to Dorymyrmex and stated that insana was undoubtedly a synonym of pyramicus. Buckley’s insana was uniformly black or brownish black; Roger stated that pyramicus possessed a yellowish red head and thorax and brownish gaster. It may be seen from this, then, that pyramicus as described by Roger agreed closely with the description of the new variety, bicolor, of Wheeler (1906). In 1912 Emery listed four subspecies and five varieties of pyramicus throughout its range. The first attempt to classify our forms was that of Creighton who considered that within the United States there existed the species pyramicus, represented by three subspecies: 4 Contributions in Science No. 238 pyramicus pyramicus (Roger, 1863) = insana Buckley, 1866 = flavus McCook, 1879 = nigra Pergande, 1895 = smithi Cole, 1936 pyramicus bicolor Wheeler, 1906 pyramicus flavopectus M. Smith, 1944 Creighton’s most important contribution was that morphological characters were used, for the first time, in differentiating our forms. Kusnezov (1952) removed these taxa from Dorymyrmex to Conomyrma, dividing them between his two subgenera in the following manner: subg. Biconomyrma bicolor (Wheeler) brunnea (Forel) wheeleri Kusnezov subg. Conomyrma flavopectus (M. Smith) pyr arnica (Roger) All these were treated as species occurring in the United States; the new species wheeleri was added from Tucson, Arizona, and brunnea, originally described from Argentina was tentatively thought by Kusnezov to occur here. Since these five were divided between subgenera established on morphological characters, it follows that the species were separable into two groups and that bicolor was thus severed from pyramicus. It is evident in reading Kusnezov’s paper that his concept of pyramicus was based on material from South America and that the inclusion of the United States and Mexico in its range was based largely on the literature. Presumably, too, pyramicus in this interpretation continued to carry the various synonyms assigned to it by Creighton. The inclusion of brunnea as a part of our fauna was based on a series of specimens from Colorado Springs, Colorado, and determined as that form by Wheeler; Kusnezov wisely accepted this determination with reservation. As indicated above, I do not consider the subgenus Biconomyrma worthy of recognition since 'it is based on minor characters, but these characters are use- ful in separating species. The workers of flavopecta and pyramica both possess a mesonotum which in profile slopes evenly into the mesopropodeal suture, there being no abrupt declivity behind. In bicolor, brunnea, and wheeleri, the meso- notum in profile is abruptly declivitous behind, often descending vertically, or nearly so, into the mesopropodeal suture. Based on Brazilian material, pyramica is a bicolored ant, as noted above, and always seems to possess a pair of moder- ately long, fully erect hairs on the pronotal dorsum. The forewing of the females has a characteristic venation (Fig. 8), as noted and figured by Kusnezov (1952). No known North American form possesses the mesonotal and venation charac- ters of Conomyrma, s. str. of Kusnezov. Although Kusnezov assigned flavopecta 1973 The Ant Genus CONOMYRMA in the United States 5 to Conomyrma, s. str., the venation of the female forewing is the same as that of bicolor and insana (Fig. 7). No specimens fulfilling the criteria here estab- lished for pyramicus have been seen from North America and I feel this name should be removed from our lists. The ant long referred to pyramicus in North American literature has a meso- notum which is sharply declivitous behind; the venation of the forewing of the female is that which Kusnezov attributed to his subgenus Biconomyrma and the insect is uniformly brownish to blackish, the lower part of the head lighter. This ant clearly is not the same entity as the South American pyramicus. The earliest available name for this ant is insana. The record for brunnea from Colo- rado, cited above, must also be referred to insana. The synonymy for this name is as follows: insana (Buckley, 1866) = pyramicus, sensu Wheeler, 1902; Creighton, 1950, etc., not of Roger, 1863 = flavus McCook, 1879. NEW SYNONYMY. = nigra Pergande, 1895. NEW SYNONYMY. = antiliana Forel, 1911. NEW SYNONYMY. = smithi Cole, 1936. NEW SYNONYMY. = brunnea, Kusnezov, 1952. Misidentification = Kusnezov, 1952. NEW SYNONYMY. Kusnezov’s wheeleri is known only from the two type specimens from Tucson, Arizona. They should be in his collection at the Instituto Miguel Lillo, Tucuman, but efforts to locate them have not been successful. There is nothing in the description to indicate that this ant is anything other than insana, a com- mon ant in the Tucson area. I am sure Kusnezov described it solely because it seemed to belong to his Biconomyrma dind could not, therefore, by “pyramicus.” Because bicolor is consistent in its color pattern and because it is broadly sympatric with insana and does not intergrade with it, I agree with Cole (1957) that this species must be accorded specific recognition. The form of the meso- notum and the wing venation of the female ally bicolor with insana. The prono- tum lacks erect hairs and this, together with the shape of the mesonotum, will separate it from the true pyramicus of South America. Literature Cited Buckley, S. B. 1866. Descriptions of new species of North American Formicidae. Proc. Ent. Soc. Phila. 6:152-172. Cole, A. C., Jr. 1957. Notes on western ants. J. N. Y. Entomol. Soc. 65:129-131. Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. 104:1-585. Donisthorpe, H. 1943. A list of the type-species of the genera and subgenera of the Formicidae. Ann. Mag. Nat. Hist. (ser. 11) 10:617-737. Eisner, T. 1957. A comparative morphological study of the proventriculus of ants. Bull. Mus. Comp. Zool. 116:439-490. 6 Contributions in Science No. 238 Emery, C. 1912. In Wytsman, P., Genera Insectorum. Dolichoderinae. Ease. 137:1-50. Forel, a. 1913. Formicides du Congo beige recoltes par M. M. Bequaert, Luja, etc. Rev. Zool. africaine 2:306-351. Gallardo, A. 1916. Las hormigas de la Republica Argentina. Subfamilia Dolichoderi- nas. Anal. Mus. Nac., Buenos Aires 28:1-130. Kusnezov, N. 1952. El estado real del grupo Dorymyrmex Mayr. Acta Zool. Lilloana 10:427-448. 1959. Die Dolichoderinen-Gattungen von Siid-Amerika. Zool. Anz. 162:38-51. Mayr, G. 1866. Myrmecologische Beitrage. Sitz. Akad. Wiss. Wien. 53:484-517. Roger, J. 1863. Die neu aufgefUhrten Gattungen und Arten meines Formiciden Ver- zeichnisses. Berl. Ent. Zeits. 7:131-214. Santschi, F. 1922. Myrmecines, Dolichoderines et autres formicides neotropiques. Bull. Soc. Sci. Nat. Laussane 54:345-378. Wheeler, G. C., and J. Wheeler. 1951. The ant larvae of the subfamily Dolichoderinae. Proc. Entomol. Soc. Wash. 53:169-210. Wheeler, W. M. 1902. A consideration of S. B. Buckley’s “North American Formicidae.” Trans. Texas Acad. Sci. 4:1-15. 1906. The ants of the Grand Canon. Bull. Amer. Mus. Nat. Hist. 22:329-345. Accepted for publication September 26, 1972 S a j . j CjlL 9^2 NUMBER 239 FEBRUARY 28, 1973 A NEW SPECIES OF THE GENUS LEPIDOPHYMA (REPTILIA; XANTUSIIDAE) FROM GUATEMALA By Robert L. Bezy I CONTRIBUTIONS IN SCICNCE s:, > > ■ . ■ : ^ LOS ANGELES COUNTY A NEW SPECIES OF THE GENUS LEPIDOPHYMA (REPTILIA; XANTUSIIDAE) FROM GUATEMALA' By Robert L. Bezy^ Abstract: A new species, Lepidophyma mayae, is described from eight specimens from the El Peteh-Alta Verapaz lowlands of Guatemala. At its type locality, L. mayae is sympatric with L. flavi- maculatum, from which it differs in several aspects of scalation and color pattern. Studies of the systematics of the lizards of the genus Lepidophyma indicate that eight specimens from the El Peten-Alta Verapaz lowlands of Guatemala represent a previously unnamed species. This new species is described here in j advance of the completion of a review of the genus. | I thank Dr. William E. Duellman for loan of specimens from the Museum of ! Natural History, University of Kansas (KU), and for generously making avail- | able field notes and a color slide of the type material. I am also grateful to Miss Alice G. C. Grandison and Dr. E. N. Arnold of the British Museum (Natural His- tory), London, Dr. Jean Guibe of the Museum National d’Histoire Naturelle, Paris, and Dr. Eugen Kramer of the Museum d’Histoire naturelle, Basle, (MHNB) for permission to examine specimens in their care and for the hospi- tality they extended during my visit to their respective institutions. I thank Dr. and Mrs. Jerry Nilsson for the financial support which enabled me to travel to these museums. Lepidophyma mayae, new species Figures 1-3 Holotype. — KU 59554, an adult female collected near Chinaja, elev. 140 m, Depto. Alta Verapaz, Guatemala by William E. Duellman and John Wellman on 24 June 1960. Paratypes.—¥A] 55863, 59556, 59558-9; LACM 75194-5 (formerly KU 59553 and 59557, respectively) all topotypes; MHNB 3751 from “Vera Paz,” Guatemala. Diagnosis. — Lepidophyma mayae differs from L. gaigeae, radula, dontomasi, and sylvaticum (all included by Smith, 1942, in the genus Gaigeia) in having conspicuously enlarged tail whorls separated dorsally by 4-5 interwhorls (vs. poorly differentiated tail whorls separated dorsally by 2-3 interwhorls); from •Review Committee for this Contribution Roy W. McDiarmid John R. Meyer John W. Wright ^Associate Curator of Herpetology, Natural History Museum of Los Angeles County, Los Angeles, California 90007 1 1973 A New Species of LEPIDOPHYMA 2 Figure 1. Dorsal and ventral views of the holotype (KU 59554) of Lepidophyma mayae, new species. L. smithii and occulor in having 29-35 (vs. 16-27) total femoral pores^ and 33- 46 (vs. 16-24) tubercle rows along side of the body between axilla and groin; from L. micropholis in having 4. 0-5.0 (vs. 5. 5-7.0) dorsal scales between para- vertebral rows of tubercles and 38-43 (vs. 57-68) gulars between the fold and the second pair of infralabials; from L. tuxtlae in having 29-35 (vs. 20-29) femoral pores, 35-38 (vs. 38-42) ventrals, including preanals, and 1-2 (vs. 4-9) temporals between seventh supralabial and postocular, total of both sides; from L. paja- panensis in having 1-2 (vs. 6-10) temporals and 23-25 (vs. 26-30) fourth toe lamellae; from L. flavimaculatum in having 33-46 (vs. 25-35 for all examined, 25-29 for specimens from Guatemala) tubercle rows. Description of Holotype. — Measurements (in mm): snout-vent length, 64.4; tail length, 92.1 (of which 17.6 is regenerated); head length, 15.3; head width, 9.1; head depth, 7.0; eye diameter, 2.6; fourth toe length, 8.3. Scales on dorsal surface of head. — Rostral, pentagonal, broader than high, scarcely visible from above; nasals, rectangular, meeting on a short middorsal suture; frontonasal, octagonal, in contact with the pentagonal median which separates the pentagonal prefrontals; frontals, hexagonal, meeting on a long middorsal suture, and contacting the median and prefrontals anteriorly, and the parietal and interparietals posteriorly; parietals, pentagonal, separated by an elongate, hexagonal interparietal in the posterior third of which the parietal ^Definitions of meristic characters given here are followed throughout the paper and terminology for scalation is largely that of Savage (1963). 3 Contributions In Science No. 239 Figure 2. Paratype of Lepidophyma mayae, new species (KU 55863, snout-vent length, 50 mm). From a Kodachrome slide taken by William E. Duellman. “eye” is faintly visible; postparietals, hexagonal, meeting on a long middorsal suture, but lacking anomalous postparietal sutures. Scales on right lateral surface of head. — Rostral followed by seven supra- labials; first, fifth and sixth supralabials higher than the others; fifth supralabial entering the orbit; nostril bordered by first supralabial, nasal, and postnasal; anterior loreal higher than postnasal and separating it from the prefrontal; posterior loreal large, only slightly longer than high; most of the scales of the orbit indistinct; upper preocular, high and narrow; lower preocular, rectangular, and in contact with upper preocular, posterior loreal, and fourth and fifth supra- labials; pupil, round; supraoculars, long, narrow, and indistinct; two postoculars, both rectangular, except that the upper one has a long thin dorsal extension between the orbit and the upper first temporals; lower first temporal (=enlarged pretympanic) approximately equal in size to the upper first temporal; a large second temporal, in contact with the third temporal, postparietal, parietal, upper first temporal, and 11 pretympanic scales; third temporal, one-third the size of the upper second temporal; seven enlarged auriculars along the anterior ear mar- gin; 65 granular pretympanics in the area anterior to' the ear opening, postero- ventral to the enlarged temporals, and dorsal to the last supralabial and the angle of the jaw. Scales on ventral surface of head. — Mental followed by four pairs of enlarged infralabials, the first two pairs larger and each with a midventral suture; third infralabials smaller and separated by the granular gulars (pregulars); the fourth infralabials, one-fourth the size of the third and followed on each side by five granular infralabials; 36 granular gulars (includes pregulars) between the ears, 42 between gular fold and midventral contact of second infralabials. Scales on body. — Two rows of enlarged, trihedral, paravertebral tubercles, separated by five rows of middorsal granular scales; scales of paravertebral rows heterogeneous with some of the enlarged trihedral tubercles separated by smaller more granular scales; 32 enlarged tubercles in right paravertebral row from above axilla to above groin; row of scales immediately dorsal to each of the two paravertebral rows, slightly enlarged (one to two times the size of other mid- dorsal granules) and weakly keeled; 188 middorsal granular scales between 1973 A New Species of LEPIDOPHYMA 4 occiput and rump; two to three rows of granular scales bordering the paraverte- brals laterally, and separating them from the enlarged, nearly homogeneous, trihedral tubercles that cover the sides of the body in the area above the ven- trals and between the axilla and groin; 40 vertical rows of enlarged tubercles along the right side of the body between axilla and groin; ten longitudinal rows of large, square, ventral plates at midbody; outer row of ventrals, smallest and only 15 scales long; 38 transverse rows of ventral scales (including preanals) between gular fold and vent. Scales on appendages. — Forelimbs covered by subequal, small, conical tu- bercles; fourth finger with 18 undivided subdigital lamellae; dorsal surface of hind limb with enlarged, trihedral tubercles, interspaced with smaller, more granular scales; ventral surface of hind limb with small, smooth scales; 17 fem- oral pores on each leg; the femoral pores are “undeveloped” and consist of shal- low pits with little evidence of glandular secretion; fourth toe with 25 subdigital lamellae of which eight are entire (i.e., lack a midventral suture); tail with rings of enlarged trihedral tubercles (whorls) separated by rings of smaller, weakly trihedral scales (interwhorls); four interwhorls dorsally (between each whorl), with two incomplete and two complete across midventral line. Color and color pattern. — Dorsal surface of head, dark brown, darkest in the interorbital and parietal area; tan parietal spot on posterior two thirds of inter- parietal; labials, dark brown to black, sutures marked with pale yellow-tan; gular region mottled with pale yellow-tan, dark brown, and tan; dorsum of body obscurely mottled with dark brown to black and lighter tan-brown; a row of small (ca. three scales in diameter) pale tan, round spots separated by larger (ca. five scales long) dark brown to black rectangular spots, along outer (lateral) side of row of paravertebral tubercles; side of body, dark brown with many small (ca. two scales) pale, dirty yellow flecks, becoming more numerous nearer the venter; venter, pale yellow-tan with numerous small dark brown spots; tail with a check- ered pattern due to the alternation of the light and dark spots on the whorls with those on the interwhorls; light spots on the interwhorls, nearly continuous on middorsal and midventral lines. Variation. — The hypodigm (Ic?, 49, 3 juveniles) of L. mayae varies as fol- lows: femoral pores, 29 (32.6 ± 0.8) 35; tubercle rows, axilla to groin, 33 (38.3 ± 1.3) 46; scales between paravertebral tubercle rows, 4 (4.2 ± 0.1) 5; dorsal caudal interwhorls, 4 (4.1 ±0.1) 5; ventral caudal interwhorls 2 (2.1 ± 0.1) 3; median prefrontal present in all; temporals, 1 (1.9 ± 0.1) 2; gulars, 38 (40.6 ± 1.0) 43; middorsal scales, occiput to rump, 162 (170.0 ± 3.4) 188; large paravertebral tubercles (right row) from above axilla to above groin, 21 (28.4 ± 2.2) 42; ventrals, 35 (36.3 ± 0.4) 38; fourth toe lamellae, 23 (23.6 ± 0.3) 25; divided fourth toe lamellae, 10 (12.9 ± 0.7) 16. The color patterns of the paratypes are similar to that of the holotype (Fig. 3). Duellman (field notes, 1960) recorded that in life KU 55863 had dull brown spots, lips barred with cream, and a dull reddish brown iris. The sex ratio of the type series of L. mayae was published earlier under “species novum” (Bezy 1972:20). 5 Contributions In Science No. 239 I 1973 A New Species of LEPIDOPHYMA 6 Distribution and Ecology. — Lepidophyma mayae is presently known only from the type locality and “Vera Paz,” Guatemala. Duellman (1963) published descriptions and photographs of the rain forest at Chinaja and notes on the habi- tat, activity, color patterns, and sizes of the specimens of Lepidophyma collected there. Duellman (field notes, 1960) recorded that all seven specimens of L. mayae collected were found in the forest; two were active by day, three were taken under logs, and two under rocks. The specimen from “Vera Paz” was collected by Gustav Bernoulli (Muller 1878a) and may be from either the lowlands (probably between Chisec and Coban) or the highlands (probably Coban; see Miiller 1878b and Stuart 1948:9). Etymology. — The species is named for the Maya of El Peten. Discussion. — Nearly a century ago Miiller (1878a) called attention to a spec- imen of an unknown species of Lepidophyma from “Vera Paz,” Guatemala, pointing out its differences from the specimens of L. smithii for which he had earlier (1877) inadvertantly published his manuscript name Akleistops guate- malensis. In 1971 I visited Basle to examine this specimen (MHNB 3751 from “Vera Paz”) and the “type series” of A. guatemalensis (MHNB 3748-50, 3752, 8039-40 from Mazatenango). Comparisons with data from 570 specimens (in- cluding all holotypes) of all nominal taxa of the genus Lepidophyma indicated that: (1) Akleistops guatemalensis Muller (1877) is a “synonym” of Lepido- phyma smithii Bocourt (1876) and (2) the specimen from “Vera Paz” (MHNB 3751) represents L. mayae, the species described in the present paper. Duellman (1963) reported a series of Lepidophyma f. flavimaculatum col- lected at Chinaja and 15 km NW Chinaja near the Peten-Alta Verapaz border in Guatemala, noting that there was considerable variation in color pattern among the specimens included. Examination of this series (Fig. 3) suggested to me that two species are present. The two specimens from 15 km NW of Chinaja (KU 55864 and 59552) and one from Chinaja (KU 59555) are assignable to L. flavimaculatum, while seven from Chinaja (KU 55863, 59554, 59556, 59558-59; LACM 75194-95) represent L. mayae. The specimens of L. mayae were initially segregated from those of L. flavimaculatum by their more numerous lateral tubercles (33-46 vs. 26-27) arranged in less discrete rows. When thus segregated, the L. mayae also differ from the L. flavimaculatum of this series in having fewer femoral pores (30-35 vs. 38-44), temporals (1-2 vs. 6-7), gulars (38-43 vs. 44-49), fourth toe lamellae (23-25 vs. 27-28), and a less distinctly spotted dorsum and a more punctate venter (Fig. 3) Of the other nine species of the genus, L. mayae is most similar to tuxtlae, pajapanensis, and flavimaculatum. The question of whether or not L. mayae Figure 3. Dorsal and ventral views of a series of Lepidophyma flavimaculatum (two speci- mens on the left) and L. mayae, new species (four specimens on the right). From left to right: KU 55864 (from 15 km NW Chinaja, El Pete'n, Guatemala); KU 59555, 59554; LACM 75194; KU 59558, 55863 (all from Chinaja, Alta Verapaz, Guatemala). 7 Contributions In Science No. 239 should be considered specifically distinct from all three of these is greatly simpli- fied by the fact that it is sympatric with L. flavimaculatum to which it is most similar (see Diagnosis, above). My decision to consider it specifically distinct from the allopatric L. tuxtlae and L. pajapanensis is somewhat more arbitrary, made after consideration of (1) the morphological differences between known pairs of sympatric species {L. tuxtlae and pajapanensis in Veracruz, L. tuxtlae and flavimaculatum in Chiapas, L. gaigeae and flavimaculatum in Queretaro, and L. flavimaculatum and mayae in Guatemala) and (2) the geographical varia- tion in the wide-ranging species, L. smithii and L. flavimaculatum. Resumen Una nueva especie es descrita de ocho ejemplos de las tierras bajas en El Peten y Alta Verapaz, Guatemala. A la localidad tipica, L. mayae es simpatrico con L. flavimaculatum, del cual se distingue por escamacion y coloracion. Literature Cited Bezy R. L. 1972. Karyotypic variation and evolution of the lizards in the family Xantu- siidae. Contrib. Sci. 227:1-29. Bocourt, M. 1876. Note sur quelques reptiles de I’lsthme de Tehuantepec (Mexique) donnes par M. Sumichrast au Muse'um. J. de ZooL, Paris 5:386-41 1. Duellman, W. E. 1963. Amphibians and reptiles of the rainforests of southern El Pete'n, Guatemala. Univ. Kans. Publ. Mus. Nat. Hist. 15:205-249. Muller, F. 1877. Mittheilungen aus der herpetologischen Sammlung des Basler Museums, I. Ueber einige seltene und neue Reptilien aus Guatemala. Verb. Naturforsch. Ges. Basel 6:390-411. 1878a. Katalog der im Museum und Universitatskabinet zu Basel aufgestellten Amphibien und Reptilien nebst Anmerkungen. Ibid., 6:559-709. 1878b. Dr. Gustav Bernoulli. Gestorben den 18. Mai 1878 in S. Franzisco. Ibid., 6:710-736. Savage, J. M. 1963. Studies on the lizard family Xantusiidae IV. The genera. Los Angeles Co. Mus., Contrib. Sci. 71:1-38. Smith, H. M. 1942. Mexican herpetological miscellany. Proc. U.S. Nat. Mus. 92:349-395. Stuart, L. C. 1948. The amphibians and reptiles of Alta Verapaz Guatemala. Misc. Publ. Mus. Zool. Univ. Mich. 69:1-109. Accepted for publication September 26, 1972 ^ /• 73 W f NUMBER 240 FEBRUARY 28, 1973 A NEW SPECIES OF ATELOPUS (ANURA, BUFONIDAE) FROM NORTHEASTERN SOUTH AMERICA By Roy W. McDiarmid MiiiSiiiiiiiii mi CONTRIBUTIONS IN SCICNCE LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. ( 3 ) Footnotes should be avoided if possible. Acknowledgments as footnotes will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- able footnotes must be typed on separate sheets. Several of one kind may be placed on a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing with Latin American subjects. Summaries in other languages are not required but are strongly recommended. Summaries will be published at the end of the paper. (8) A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor A NEW SPECIES OE A TELOPUS (ANURA, BUFONIDAE) FROM NORTHEASTERN SOUTH AMERICAi By Roy W. McDiarmid^ Abstract: A new species of bufonid frog, Atelopus ver- miculatus, is described from a series of adults and juveniles from French Guiana. The new species has an internal tympanum and middle ear and in this respect differs from all species of Atelopus Qxctpi flavescens and spiimarius, from which it is easily distin- guished by its small size and distinctive coloration. The amphibian fauna of northern South America is poorly understood. New and little known species frequently are collected in biologically unworked areas of this vast tropical region. On three different occasions Dr. Philip A. Silverstone visited northern South America and made important collections of amphibians and reptiles incidental to his studies of dendrobatid frogs. Some of the new material has been described recently by Silverstone (1971), Trueb (1971), and Brame and Wake (1972). Among the specimens collected by Silverstone in a remote area of French Guiana was a series of an undescribed species of Atelopus. It is ironic that this new species of Atelopus should be found less than 100 km south of the type locality of Atelopus flavescens Dumeril and Bibron 1841, the oldest taxon in the genus. With reference to the dorsal color pattern I propose that this frog be called Atelopus vermiculatus, new species Figures 1 and 2 Holotype: LACM 42060; an adult female (Figs. 1 and 2) collected by Philip A. Silverstone along a forest trail paralleling the lower Riviere Matarony, a tributary of the Fleuve Approuague, French Guiana during the day, 29 July 1968; 35 m elevation. Paratypes: LACM 42049-59, 42061-63, 42070-79 are topoparatypes collected on 24-30 July and 17-18 August, 1968; LACM 42064-66 from forested hills near Crique Ipoucin on 5-6 August, 1968, between 8-180 m; LACM 42067, 42069 from forest near ORSTOM camp at Sant Tortue, on the Fleuve Approuague at its confluence with the Crique Tortue on 8 and 1 1 August, 1968, 75 m; LACM 42068 from along a trail between Fleuve Ap- 1 Review Committee for this Contribution W. Ronald Heyer Jay M. Savage John W. Wright ^Research Associate, Section of Herpetology, Natural History Museum of Los Angeles County; and Department of Biology, University of South Florida, Tampa, Florida 33620 1 2 Contributions in Science No. 240 Figure 1. Holotype of Atelopus vermiculatus, new species (LACM 42060). Left, dorsal view; right, ventral view. prouague and summit of Montagnes Tortue on 9 August, 1968, 100 m. All specimens were collected by Philip A. Silverstone. Diagnosis: Atelopus vermiculatus is a relatively small species (snout-vent lengths for adult males and females are 19.3-22.1 mm, x = 21.27, N = 4 and 26.8-31.3 mm, x= 28.96, N= 15, respectively) that is easily distinguished from all species of Atelopus, except flavescens and spumarius, by the presence of a well-developed internal tympanum, annulus tympanicus and middle ear. It differs from A. flavescens and A. spumarius in its smaller size and distinctive pattern and coloration. In addition, Atelopus vermiculatus can be defined on the basis of the following characters: presence of ostia pharyngea and vocal slits in males; the absence of dorsolateral rows of warts and obvious glandular areas on the sides of the head, over the eyes, in the paratoid region and dorsally and laterally on the back, sides and limbs; subacuminate and protruding snout; relatively smooth skin, lacking coni apicali, moderately webbed foot, modal webbing formula (from Savage and Heyer, 1967) 1 0 — 1 \\ Vi — IVi III W4 — 3 Vi IV 3 Vi — V; and dorsal color pattern of yellowish, greenish, or tan vermiculations on a dark brown background with a rose pink wash on the belly and thighs. Description of Holotype: Adult female (Figs. 1 and 2); head narrower than body; head slightly longer than wide; snout subacuminate in dorsal view, tip slightly rounded; snout protruding over lower lip, acute in lateral profile; canthus rostralis rounded to slightly angular; loreal region flat; lips not flared; nostrils elliptical, directed laterally; nostrils closer to tip of snout than to eye; snout moderately long, distance from eye to nostril slightly greater than diameter of eye; interorbital width slightly greater than length of upper eyelid; 1973 New Species of Atelopus 3 Figure 2. Holotype of Atelopus vermiculatus, new species. Ventral view of right hand and foot. outer margin of upper eyelid not noticeably thickened; surface of head concave between nostril, flat to slightly rounded between eyes; external tympanum absent; supratympanic (paratoid) ridge well developed; pretympanic ridge moderately developed; tongue moderate in size, elliptical, posterior three- fourths free; two patches of pigment on anterior lateral third of tongue; choanae ovoid, not visible when roof of mouth viewed from directly below; ostia pharyngea large. Skin relatively smooth, lacking dorsolateral row of warts; skin covered with tiny scattered verrucae visible microscopically; outlines of suprascapulae, neural ridge of vertebrae, anterior and posterior portions of coccyx and dorsolateral expanses of sacral diapophyses visible through skin on dorsal surface; postero ventral quarter of belly and medioventral surface of thighs (rump patch) wrinkled; anal opening on low protuberance, directed posteriorly, opening slightly above midlevel of thighs; anal opening covered dorsally by anal flap consisting of one large and two smaller ventrally directed papillae; series of anal furrows radiating from opening, best developed ventral to opening; palmar tubercle single, smooth; thenar tubercle indistinct; subarticular 4 Contributions in Science No. 240 Figure 3. Variation in dorsal pattern of selected adult female paratypes of Atelopus vermiculatus, new species (from left to right, LACM 42076, SV-30.6 mm; 42065, 31.3 mm; 42068, 27.8 mm; 42064, 30.7 mm). tubercles few, low and smooth; finger tips rounded, slightly swollen, slightly broader than penultimate joint; webbing between fingers fleshy, reduced; webbing formula for hand 1 C2 — 2C2 II 2 — 31/2 III 3C2 —21/2 IV; no tarsal fold; sole of foot smooth; inner metatarsal tubercle reduced, nearly continuous with subarticular tubercle of first toe; outer metatarsal tubercle small, oval; other metatarsal tubercles absent; subarticular tubercles low, oval, indistinct; supernumerary tubercles absent; toe tips rounded, not or very slightly expanded; webbing fleshy; webbing formula for foot 10 — 1 ll V2 — 21^ III m —31/2 IV 31/2 — m V. Measurements and Proportions: All measurements are in millimeters: snout-vent length (SV) 29.5; tibia length (TL) 13.2; foot length (FL) 9.9; head length (HL) 8.9; head width (HW) 8.6; snout length (SL) 4.4; eye diameter 2.5; interorbital width 3.7; distance from eye to nostril 3.0; sacral width 7.9; TL/SV 0.45; FL/SV 0.34; HL/SV 0.30; HW/SV 0.29; HW/HL 0.97; SL/HL 0.49. Coloration of Holotype (in alcohol): Dorsal surface of body and limbs dark brown with extensive tan vermiculation. Vermiculate pattern most con- centrated on head, least on sides of head and sides of body. Pale brown area between eye and upper lip. Poorly defined tan blotch over the suprascapulae. Posterior surface of thighs brown with scattered, irregular tan specks. Ventral surfaces dirty tan to light brown, darkest in pectoral-gular area. Posterior section of belly and medioventral surface of thighs creamy white. Ventral surface of hands tan; ventral surface of feet light brown; tubercles lighter tan to cream. Variation in Paratypes: There are 14 adult female, 4 adult male and 12 immature (sex undetermined) paratypes. Except for details of coloration, the adult females are similar to the holotype. The males, however, are considerably smaller, have moderately developed brown dorsolateral nuptial pads on the thumbs and proportionately thicker upper forelimbs than the females. They have the same general variation in color and pattern found in the females. The immatures range from 14.6 to 25.9 SV and have the same basic pattern and color as the adults. There is no obvious indication in this series of ontogenetic change in color, pattern or proportions. The dorsal pattern of the females 1973 New Species of Atelopus 5 varies from broadly vermiculate (LACM 42065) to nearly uniform light brownish tan (LACM 42064); most of the females have a fine vermiculate pattern similar to that of the holotype. This variation in dorsal pattern is illustrated by selected individuals (Fig. 3). There is considerable variation in ventral pigmentation in adult females. Generally the throat and anterior half of the body is washed, finely speckled, or vermiculated with various shades and intensities of brown over dirty white; all have a creamy white, grayish white or dirty white patch on the posterior part of the belly and ventral surfaces of the thighs. This light area may include most of the belly (extending to the pectoral area) and entire ventral surface of the limbs or may be restricted to a small area on the posterior eighth of the belly and central area of the thighs; most have some brown speckling on the lateral edges of the ventral surface of the thighs and on the ventral surface of the forelegs. The ventral surfaces of the manus are light; a few specimens have a brownish wash on the palms. All have a brown central solar patch; the toes are basically light; there is brown color between the subarticular tubercles in a few individuals. Three of the four males have a typical vermiculate dorsal pattern. The fourth (LACM 42066) is irregularly marked with tan and brown; traces of the vermiculate pattern are evident on the head, sides and limbs. Ventrally the males are lighter than the females; as compared to the females, the brown pigmentation is reduced on the chin and anterior belly and consists of only a few small light brown irregular lines or faint scattered speckling. All immatures are marked with the typical vermiculate pattern. The ground color in one specimen (LACM 42067) is faint and the vermiculation, while still present, is gray both on the limbs and dorsal surface. Ventrally the immatures have the same range in pattern and coloration as described for the adults. In life the dorsal coloration for all specimens is about the same as described above except that the vermiculate pattern is brighter, sometimes greenish golden (comparison of coloration of preserved animals and Kodachrome slides). The females and largest immatures have a striking rose pink ventral wash, brightest on the belly and thighs; obscured to various degrees on the ventral surfaces of chin, chest, and limbs by brown coloration and pattern. The rump patches are much lighter and only have a faint rose pink coloration. Dr. Silverstone provided the following color notes made in the field from living specimens LACM 42049 (PAS 4187) immature, 20.1 mm snout-vent length and LACM 42050 (PAS 4188) adult female, 27.9 mm SV: “Iris black with gold ring around pupil; dorsum dull dark brown with very faint light yellowish reticulum on back and limbs; extreme posterior belly orange pink; ventral surface of thighs, anterior part of posterior belly and axillae rose pink; rest of venter dull medium brown with slight rose pink tinge.” Color notes from LACM 42064 (PAS 4404) adult female, 30.7 mm SV and LACM 42065 (PAS 4405) adult female, 3 1.3 mm SV are: “Back and head in one specimen (LACM 42064) dull greenish with purplish areas dorsolaterally; limbs above mixed purple and green; the other specimen (LACM 42065) is dark purple- 6 Contributions in Science No. 240 brown above, with dull green reticulum on entire dorsal surface; venter on both specimens dull purple, brighter than back, lightest and brightest on limbs; posterior belly and part of right side of belly bright rose pink; on one specimen (LACM 42065) tiny black spotting on throat, breast, edges of limbs, and edges of belly.” Measurements and proportions for the paratypes are contained in Table I. Specimens were categorized as adult female (eggs present), adult male (enlarged testes and nuptial pads present), immature (no eggs, gonads not enlarged, no nuptial pads). No attempt was made to sex the immatures. There is well-defined size dimorphism between males and females; the adult females are much larger than adult males. The data also suggest that proportionately males have a slightly larger head than females. Table I Range and Mean of Measurements and Proportions of Paratypes of Atelopus vermiculatus Male Female Immature (N = 4) (N=14) (N=12) SV* 19.3 - 22.1 1 (21.27) 26.8 - 31.3 (28.92) 14.6 - 25.9 (18.68) TL* 9.2 - 10.4 (9.82) 12.2 - 13.9 (13.03) 6.5 - 11.5 (8.45) FL* 6.2 - 8.7 (7.37) 9.1 - 11.1 (9.99) 4.9 - 8.8 (6.27) HL* 6.4 - 7.2 (6.90) 8.0 - 9.5 (8.67) 5.1 - 7.6 (6.01) HW* 6.1 - 6.9 (6.60) 7.7 - 9.0 (8.31) 4.9 - 7.4 (5.77) SL* 3.2 - 3.5 (3.37) 3.9 - 4.6 (4.29) 2.5 - 3.8 (3.04) TL/SV 0.45- 0.48 (0.461) 0.44- 0.48 (0.450) 0.43- 0.49 (0.452) FL/SV 0.32- 0.39 (0.345) 0.32- 0.38 (0.344) 0.30- 0.38 (0.336) HL/SV 0.32- 0.33 (0.324) 0.29- 0.31 (0.298) 0.29- 0.36 (0.323) HW/SV 0.30- 0.32 (0.310) 0.27- 0.30 (0.286) 0.28- 0.34 (0.310) HW/HL 0.94- 0.97 (0.956) 0.91- 0.99 (0.957) 0.89- 1.05 (0.962) SL/HL 0.48- 0.50 (0.489) 0.44- 0.54 (0.495) 0.43- 0.57 (0.505) *SV = snout-vent length; TL = tibia length; FL = foot length; HL = head length; HW = head width; SL = snout length; all measurements are in mm. Osteology: All information was obtained from a cleared and stained adult female (LACM 42075), 28 mm SV. Only those osteological traits that have been demonstrated to have interspecific variation are discussed (see McDiarmid, 1971). This specimen was included in the analysis and definition of the genus Atelopus presented elsewhere (McDiarmid, 1971). The following osteological points are of interest: nasals separated medially, fused to underlying chondrocranium; occipital grooves in frontoparietal covered; wings of vomer well developed; anterior end of the parasphenoid pointed, slightly overlapping sphenethmoid; no anterior projection on squamo- sal; quandratojugal well developed, overlapping maxillary; lateral head of posterior end of prootic reduced; atlas completely fused with first trunk vertebra; neural arches ornately adorned, without connection between dorsal ornamentation on consecutive vertebrae; transverse processes of trunk vertebrae about equal in length with irregular, scalloped edges; anterior edges 1973 New Species of Atelopus 7 Figure 4. Map of the Fleuve Approuague Drainage System in French Guiana show- ing the locality records of Atelopus vermiculatus: 1-near Crique Ipoucin; 2-trail from Fleuve Approuague to Montagues Tortue; 3 -near confluence of Crique Tortue and Fleuve Approuague; 4-lower Riviere Matarony. of sacral diapophyses irregular; coccyx with bicondylar articulation and lateral flange extending about half its length; sternum of pectoral girdle well developed and ossified; prepollex a single distinct element about the same length as metacarpal 1; prepollex and metacarpal 1 fused at proximal end; phalangeal formula 1 -2-3-3 for manus and 2-2-3-4-3 for foot. Habitat and Distribution: All specimens of Atelopus vermiculatus were collected during the day between 0800 and 1800 from 24 July to 18 August, 8 Contributions in Science No. 240 Figure 5. Map of northeastern South America showing locality records of Atelopus vermiculatus (triangles) and Atelopus flavescens (solid circles). 1968. When collected the frogs were moving slowly (making short jumps or walking) on the forest floor. The area around the Bruynzeel lumber camp on the lower Riviere Matarony (Fig. 4) consists of low forested hills; the hills are well drained but there are swampy creeks at their bases. The forest is relatively undisturbed. Some large, buttressed trees, greater than 25 m high, are present but the forest consists primarily of palms and some smaller trees. The undergrowth is not very dense and walking is easy. The undergrowth shrubs are head high or less; a species of Dracontium (Araceae) is common. There is a heavy cover of dead leaves on the forest floor. Some of the common trees are the spiny palm Astrocaryum, Lecythis davisii (Lecythidaceae), Clusia sp. (Guttiferae), Ster- culia sp. (Sterculiaceae), Apeiba sp. (Tiliaceae), and Parkia sp. (Leguminosae). The other three localities are farther up stream along tributaries of the Fleuve Approuague (Fig. 4). Apparently there is no tidal influence in the creek at these localities. The forest in this area is undisturbed and similar in stature to 1973 New Species of Atelopus 9 that near the Bruynzeel camp. The spiny palm Astrocaryum is common here also, together with Attalea spectabilis (Palmae), species of Sterculia and Caryocar (Caryocaraceae). At present, A. vermiculatus is known only from the drainage system of the Fleuve Approuague. Additional collecting in the basins of the Riviere Compte to the northwest and the Rio Oiapoque to the south may reveal its presence in those river systems. The closely related species A. jiavescens has been collected east of the Fleuve Approuague at several localities in the region drained by the Crique Grand Inini, a tributary of the Maroni Riviere (MCZ 43502-08). Atelopus jiavescens also is known from Cayenne (type locality) and several localities in Surinam and Guyana to the east (Boulenger 1882; Van Lidth de Jeude 1904) as well as from the Brazilian territory of Amapa (LACM 42045-48), and the state of Para (KU 129954-60) to the south (Fig. 5). Ecology: It is interesting that mature females outnumbered males nearly 4 to 1 . The larger size of the females might lead one to conclude that a collecting bias for larger frogs influenced the sample. While this might in part account for the difference, the relatively large sample of immature specimens suggests no such bias. A more reasonable explanation may lie in the habitats sampled. McDiarmid (1971) suggested that male Atelopus spend much of the breeding season along streams while the females often are found on hillsides and in the forest some distance from the streams. Silverstone collected most of the specimens along trails leading through the forest and away from the rivers and streams. Thus, it appears that most of the collecting was done in areas where females might be more common than males. In this sense A. venniculatus apparently partitions the habitat according to sex. This same situation has been recorded for A. varius in Costa Rica (McDiarmid, 1971) and A. cruciger in Venezuela (Sexton, 1958). All large females collected contain ovarian eggs. No recently spent females were found. The largest males were reproductively active and have moderately developed nuptial pads and enlarged testes. It appears that these frogs were about to begin breeding. All other species of Atelopus for which data are available breed in streams (McDiarmid, 1971). Streams in seasonal tropical forests are subject to tremendous fluctuations in water level during the rainy season. Amphibians that breed in streams during the rainy season experience high mortality to eggs and early larval stages as a result of high water and movement of rocks and debris on the stream bottom during torrential storms. As a result, there has been strong selection to concentrate breeding activity in the dry season in some stream breeding frogs (i.e., Smilisca sordida, James, 1944; Duellman and Trueb, 1966). The breeding sites and larvae of A. vermiculatus are unknown. However, it is assumed that this species is also a stream breeder. The timing of breeding activity in A. vermiculatus apparently corresponds to the seasonal distribution of rain in the area. April and May are the wettest months (>450 mm/month) with a gradual drying through June, July, and August; the main dry season (< 40 mm/month) occurs in September and October. The fact that all large females and males collected in July and 10 Contributions in Science No. 240 Figure 6. Dorsal and ventral views of an adult female Atelopus flavescens (LACM 42046, SV-35.9) and an adult female Atelopus vermiculatus, new species (LACM 42056, SV-28.2). August are reproductively active and that September and October are the driest months supports my contention that Atelopus vermiculatus carries on breeding activity during the dry season. The streams are slower and quieter during this period, and there is less chance of periodic flooding that could result in high mortality to the eggs and larvae. Remarks: One of the major problems encountered in understanding relationships among species of Atelopus is the lack of well-defined species groups. McDiarmid (1971) referred to long-legged and short-legged species of Atelopus and mentioned some morphological traits associated with some of the species in each group. In some respects (i.e. habitus) A. vermiculatus would be associated with the long-legged species, as represented by A. spurrelli and 1973 New Species of Atelopus 1 1 A. varius. However, A. vermiculatus has an internal tympanum and middle ear and in this respect is' most closely related to A. flavescens and A. spumariiis (including A. pulcher, Rivero, 1968). These three species form a natural group and are distinct from all other species of Atelopus that have been studied. Rivero (1968) included Atelopus flavescens and A. spumarius in the same group and suggested the two species probably were remnants of a once- continuous population. At that time A. flavescens was known only from French Guiana (Dumeril and Bibron, 1841), British Guiana and “Interior of Brazil” (Boulenger, 1882), and Surinam (Van Lidth de Jeude, 1904). Literature reports of A. flavescens from near Quito, Ecuador (Cornalia, 1849) and from Central Brazil (Miranda-Ribeiro, 1926) are in error. Atelopus spumarius is known from eastern Peru and Ecuador. Collection of specimens from the Amazon Basin (Eig. 5) indicates a much wider range for A. flavescens and lends support to Rivero’s suggestion of a once-continuous population. Additional material from the western parts of the Amazon Basin and subsequent detailed analysis of the species might indicate that A .flavescens and A . spumarius are conspecific. Atelopus vermiculatus apparently was derived from A. flavescens and subsequently isolated in the drainage system of the Fleuve Approuague in French Guiana. A morphometric comparison of A. vermiculatus and four samples of A. flavescens, using the proportions listed in Table I, was of little use in distinguishing the two species, primarily because of the interpopulational variation in A. flavescens. Thus, the two species are most easily distinguished on the basis of the striking differences in size and coloration. Adult A. flavescens are considerably larger (mature male SV 24.2-32.2, x = 28.9, N= 15; mature female SV 29.5-40.3, x = 34.3, N= 10) than adult A. vermiculatus (mature male SV 19.3-22.1, x=21.3, N = 4; mature female SV 26.8-31.3, x = 28.9, N= 14). The marked differences in coloration between the two species are illustrated in Figure 6. Specimens Examined Atelopus flavescens: Guyana (British Guiana): Kartabo (AMNH 21332-34, 53304-05, 13528-29); Kalacoon (AMNH 70997-71001 -f- 6); Kangaruma (AMNH 39725). Surinam: Brownsberg (AMNH 77449). French Guiana: Degrade Leonce, Petite Inini Riviere (MCZ 43502); on trail between Sophie and La Greve (MCZ 43503-07); on trail between Dorlin and Sophie (MCZ 43508). Brazil: Territorio de Amapa, Serado Navio, Serra de Veado, hills east of mine (LACM 42045-48); Para, Sudam Floral Reserve, 74 km SE Santarem (KU 129954- 60). Acknowledgments Philip A. Silverstone collected the type series and provided detailed field notes and Kodachrome slides of the new species. The following individuals and institutions provided comparative material for the study: William E. Duellman and Martha L. Crump, Museum of Natural History, University of Kansas (KU); Ernest E. Williams and Benjamin Shreve, Museum of Comparative 12 Contributions in Science No. 240 Zoology, Harvard University (MCZ); John W. Wright, Natural History,- Museum of Los Angeles County, Los Angeles (LACM); and Richard G. Zweifel and Charles W. Myers, American Museum of Natural History, New \ York (AMNH). Mercedes McDiarmid and John Wright read an earlier draft | of this paper and made several helpful suggestions. Jim Feigl photographed ' the specimens of Atelopus and Betty Ingalls illustrated Figure 2. I would like } to thank all of these people for their help. | Resumen j Una nueva especie de rana, Atelopus vermiculatus, se describe basada i en una serie de adultos y juveniles colectados en Guiana Frances. La nueva i! especie tiene un timpano interno y oido medio y, en este respecto, se distingue de todas otras especies de Atelopus con excepcion de A. flavescens y spumar- ius, de cuales puede ser distinguida facifmente por el tamaho pequeho y la coloracion distintiva. Literature Cited Boulenger, G. a. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. Second Edition. British Museum, London. ' 503 pp., plates 1-30. Brame, a. H., Jr., and D. B. Wake. 1971. New species of salamanders (Genus Bolitoglossa) from Colombia, Ecuador and Panama. Contrib. Sci. 219:1-34. CoRNALiA, E. 1849. Vertebratorum synopsis in Museo Mediolanense extantium quae per novam orbem Cajetanus Osculati collegit annis 1846-47-48. Milano. . 16 pp., 1 plate. j Duellman, W. E., and L. Trueb. 1966. Neotropical hylid frogs, genus Smilisca. 1 Univ. Kans. Publ., Mus. Nat. Hist. 17(7):28 1-375. | Dumeril, a. M. C., and G. Bibron. 1841. Erpetologie generale ou histoire naturelle : complete des reptiles. Paris. Vol. 8, 792 pp. | James, M. S. 1944. Notes on the breeding of Hyla nigripes in Costa Rica. Copeia i 1944(3):147-148. McDiarmid, R. W. 1971. Comparative morphology and evolution of frogs of the neotropical genera Atelopus, Dendrophryniscus, Melanophrynisciis, and Oreoplirynella. Bull. Los Angeles Co. Mus. Nat. Hist., Sci. 12:1-66. Miranda-Ribeiro, a. 1926. Notas para servirem ao estudo dos Gymnobatrachios (Anura) Brasileiros. Archivos Mus. Nac. Rio de Janeiro 27:1-227, pis. 1-22, figs. 1-110. Rivero, J. A. 1968. More on the Atelopus (Amphibia, Salientia) from western South America. Caribbean J. Sci. 8(1 and 2): 19-29. Savage, J. M., and W. R. Heyer. 1967. Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central America. Beitr. Neotrop. Eauna 5:111-131. Sexton, O. 1958. Observations on the life history of a Venezuelan frog, Atelopus cruciger. Acta Biol. Venezuelica 2(2 1):235-242. Silverstone, P. a. 1971. Status of certain frogs of the genus Colostethus, with descriptions of new species. Los Angeles Co. Mus., Contrib. Sci. 215:1-8. Trueb, L. 1971. Phylogenetic relationships of certain neotropical toads with the description of a new genus (Anura: Bufonidae). Los Angeles Co. Mus., Contrib. Sci. 216:1-40. Van Lidth de Jeude, T. W. 1904. Reptiles and Batrachians from Surinam. Notes Lyden Mus. 25:83-94, Plate 7. Accepted for publication September 7, 1972 Printed in Los Aneeles L'alifnrni'n AnH<»rcrin NUMBER 241 MARCH 20, 1973 Z 7-^ A LATE MIOCENE RECORD OF LEPAS LINNAEUS (CIRRIPEDIA, LEPADIDAE) FROM SOUTHERN CALIFORNIA By Victor A. Zullo CONTRIBUTIONS IN SCICNCE NATURAL HISTORY MUSEUM LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ; | ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not ' be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- able footnotes must be typed on separate sheets. Several of one kind may be placed on a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing with Latin American subjects. Summaries in other languages are not required but are strongly recommended. Summaries will be published at the end of the paper. (8) A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor A LATE MIOCENE RECORD OF LEPAS LINNAEUS (CIRRIPEDIA, LEPADIDAE) FROM SOUTHERN CALIFORNIA' By Victor A. Zullo^ Abstract; Scuta of one species, and scuta and carinae of a second species of Lepas Linnaeus (Cirripedia, Lepadidae) are locally abundant in late Miocene fish-bearing diatomaceous shales of the Yorba Member, Puente Formation in Los Angeles County, California. These fossils, preserved primarily as impressions on bedding planes, constitute the second recorded fossil occurrence of Lepadomorph (‘goose’) barnacles from the eastern margins of the Pacific basin. The majority of scuta are similar to those of the extant species L. (Dosima) fascicularis Spengler, but are regarded as a new species, L. (D.) latiscutis. The remainder, with associated carinae, resemble those of the extant species L. (Lepas) anserifera Linnaeus. The pre- vious record, from the late Pliocene San Diego Formation in southern California, was of a single scutum similar to that of the extant spe- cies L. (L.) pectinata Spengler. Introduction Although balanomorph (‘acorn’) barnacles are common in rocks of Cenozoic age on the Pacific coast of North America, lepadomorphs (‘goose’ barnacles) are rarely encountered. The single previous record (Zullo, 1969:4) was of a solitary scutum resembling that of the extant cosmopolitan species Lepas (Lepas) pectinata Spengler from the late Pliocene San Diego Formation, south- ern California. The material described herein adds a pre-Pliocene record for two species to the southern California Cenozoic fauna. The family Lepadidae has a meager fossil record that is limited to remains assigned to the genus Lepas. Extant members of the genus, comprising seven species and two varieties in three subgenera, are pelagic, and several are vir- tually cosmopolitan in their distribution. Most species attach to floating objects such as seaweed and drifting wood, but L. fascicularis Spengler can construct its own float by enlargement of its stalk after initially attaching to floating mate- rials. The scarcity of fossil lepads is attributable to their pelagic habit and to the fragility of their preservable hard parts. Withers (1953) in his monographic study of Tertiary cirripeds recognized the following extinct species, ranging in age from middle Eocene to Pliocene: Lepas (Lepas) stenzeli Withers (1953), middle Eocene, Weches Formation, Claiborne Group, Texas. 'Review Committee for this Contribution J. Wyatt Durham William A. Newman Edward C. Wilson ^Research Associate in Invertebrate Paleontology, Natural History Museum of Los Angeles County; and University of North Carolina at Wilmington, Wilmington, North Carolina 28401. 1 2 Contributions in Science No. 241 L. (L.) reflexa Withers (1953), upper Eocene (Bartonian), Lower Barton Beds, England. L. (L.) aquitanica Fischer (1886), lower Miocene (Aquitanian-Burdigalian), France. L. (L.) orbignyi Fischer (1886), lower Miocene (Burdigalian), France. ’ L. (L.)harringtoni Laws (1948), lower Miocene (Altonian), Southland Series, New Zealand. L. (L.) pritchardi Hall (1902), lower Miocene (Janjukian), Australia. L. (L.) mallandriniana Seguenza (1876), upper Miocene (Tortonian), Messina, Sicily. L. (Dosima) rovasendai De Alessandri (1895), lower and middle Miocene (Aquitanian — Helvetian), northern Italy. L. (D.) delicatula Withers (1953), Pliocene (Plaisancian), Coralline Crag, England. Identification of extant specimens based solely on capitular plate (i.e., shell) morphology can be difficult. Shape, ornamentation and structure vary markedly from individual to individual, and in many cases ultimate identifica- tion rests on characteristics of the soft body. Many of the above-named fossil species are based on single plate types, and only two are known from terga, scuta and carinae. Given the variation in plate morphology and the widespread occurrence of extant species, it is quite likely that our knowledge of diversity I within Lepas during the Cenozoic is less than that suggested by the list of nom- 1 inate species presented above. I Among the new California Miocene specimens, the subgenus Lepas is repre- | sented by numerous scuta and two carinae that are not sufficiently diagnostic to | identify with or distinguish from known species. The subgenus Dosima Gray is | represented by numerous scuta and a fragmentary impression of what might be 1 a tergum. Scuta have been described for only two of the three known species of i Dosima, and the new specimens are readily distinguished from these. The late Pliocene species L. delicatula is known only from two incomplete carinae. How- ever, the shape and thickness of these carinae appear to preclude their associa- tion with the type of scutum represented in the California Miocene. This conclu- i sion, supported by the disparity in ages of the two occurrences, has prompted | my decision to regard the California Dosima as a distinct species. Locality The new specimens are from diatomaceous shale of the Yorba member of the Puente Formation. The locality (no. 1209, Natural History Museum of Los Angeles County, Invertebrate Paleontology collection) is at longitude 117° 54' 41" W, latitude 33° 59' 28" N, Vi mi west of the intersection of Los Palacios and Fullerton roads, Rowland Heights, City of Industry, USGS, La Habra 7.5 min topographic quadrangle, 1950 edition, Los Angeles County, California. Age and Paleoecology The Rowland Heights locality, notable for abundant marine fish remains, has yielded few invertebrate megafossils. Age determination, based upon corre- Late Miocene Record of LEPAS 3 I 1973 i lation with Foraminifera-bearing rocks at similar stratigraphic levels in the I eastern Puente Hills indicates a late Mohnian (late Miocene) age (Durham and i Yerkes, 1964). Preservation of the fragile and readily scattered plates of Lepas I appears to have been facilitated by unusual depositional circumstances sug- gested by the fossils and sediments of the Yorba Member. The abundance of pelagic remains, the scarcity of benthic invertebrates, the excellent preservation exhibited by the fish, and the kinds of foraminifers and fish present suggest a low energy, low productivity environment at bathyal depths. Presumably the lepads, floating in surface waters, sank to the bottom where they remained in a relatively undisturbed state until burial. Systematics Suborder Lepadomorpha Pilsbry, 1916 Family Lepadidae Darwin, 1851 Genus Lepas Linnaeus, 1758 Subgenus Dosima Gray, 1825 Lepas (Dosima) latiscutis, new species Figures 1-3 Diagnosis. Scutum subquadrate, broader than tall, with fine radial striae, and lacking both an apico-umbonal ridge and growth below the short, straight basal ridge. Description. Scutum thin, papery, subquadrate in shape, usually broader than high; occludent margin nearly straight at base, becoming broadly convex towards rounded apex; tergolateral rnargin produced, straight to slightly con- cave in upper half, broadly convex in lower half; basal margin short, straight, thickened and inflexed; apico-umbonal ridge lacking; growth ridges broad and prominent, crossed by distinct but not prominent radial striae. Remarks. The specimens available consist of a dense aggregation of scuta. Most of these are preserved as impressions, but a few retain original shell mate- rial. No carinae, and only a fragment of what might be a tergum were observed (Fig. 1). The density of aggregation and the selectivity in plate types preserved suggests post depositional concentration, possibly coupled with selective win- nowing of the smaller plates during settlement through the water column. The scutum of L. latiscutis differs from that of L. fascicularis Spengler (Figs. 7, 12) in its broader proportions, its blunt apex, its straight basal margin, the development of external radial striation, and the lack of growth below the basal ridge. The scutum of the Miocene species L. rovasendai (Fig. 11) is higher than wide, with indistinct growth ridges, an angular apex, and a distinct apico- umbonal ridge. The carina of the late Pliocene species L. delicatula (Figs. 8-10) is not readily associated with the type of scutum exhibited by L. latiscutis. As the carina of L. delicatula is less angulate than that of L. fascicularis, and as the lower pait of the tergolateral margin of the scutum tends to parallel the lower margin of the carina, then the tergolateral margin of the scutum of L. delicatula should be less angulate and its lower half should more nearly parallel the occlu- dent margin. The tergolateral margin of the scutum of L. latiscutis is similar in 4 Contributions in Science No. 241 Figures 1-3. Lepas (Dosima) latiscutis, new species. (1) scutum and partial impression of a plate suggestive of a tergum, /, height of scutum, 15 mm; (2) scuta on bedding plane of diatomaceous shale, width of block in photograph, 55 mm; (3) scutum, holotype, length of basal margin, 1 1 mm. Figures 4-6. Lepas (Lepas) sp. . (4) partial carina, length 8 mm; (5) scutum?, height, 14 mm; (6) scuta and partial carina, c, length of carina, 9 mm. Late Miocene Record of LEPAS 5 1973 Figures 7, 12. Lepas fascicularis Spengler, showing extreme variations in plate shape. (7) ‘typical’ form, from Pilsbry, 1907, U.S. Nat. Mus. Bull. 60, pi. 9, fig. 6, capitular height, 20 mm; (12) rarer broad form, from Broch, 1959, Conseil Intern. Explor. de la Mer, Zoo- plankton 83, fig. 5a, capitular height, 33 mm. Figures 8-10. Lepas delicatula Withers, from Withers (1953:pl. 58, figs. 13-14). (8) basal part of Carina, greatest length, 6.8 mm; (9) apical part of carina (incomplete), greatest length, 8.8 mm; (10) outer view of basal part of carina shown in fig. 8. Figure 11. Lepas rovasendai De Alessandri, from Withers (1953, pi. 58, fig. 4a), height, 4.5 mm. 6 Contributions in Science No. 241 shape to that of L. fascicularis, suggesting that its carina was more angulate than that known for L. delicatula. Also, the carinae of L. delicatula are heavily calcified, and it is unlikely that associated scuta would be as delicate as those representing L. latiscutis. Type disposition. The type and only known lot from the Yorba Member of the Puente Formation is deposited in the Natural History Museum of Los Angeles County, Invertebrate Paleontology collection. Subgenus LET AS Linnaeus, 1758 Lepas (Lepas) sp. Figures 4-6 Description. Scutum subtriangular, taller than broad; umbo basal, apex acute, basal margin straight; exterior ornamented by growth lines and faint radial striae; apico-umbonal ridge indistinct, close to occludent margin. Carina short, convex, broadest in lower third, attenuated apically; basal part unknown. ^ Remarks. The specimens referred to Lepas (Lepas) sp. are poorly preserved, : but definitely represent a species other than L. latiscutis. The two species were ’ not observed to co-occur on the same block of shale, and the density of plates of Lepas sp. was considerably less than that of L. latiscutis. Among extant spe- cies, these specimens most closely approximate L. anserifera Linnaeus. Acknowledgments I wish to thank Dr. Edward C. Wilson of the Natural History Museum of Los Angeles County for providing the specimens described herein, and for in- formation regarding the locality. Support for this study was provided by the Department of Marine Sciences Research, University of North Carolina at Wilmington. Literature Cited De Alessandri, G. 1895. Contribuzione alio studio dei Cirripedi fossili d’ltalia. Boll. Soc. geol. Ital. 13:234-314, pis. 3-5. Durham, D. L., and R. F. Yerkes. 1964. Geology and oil resources of the eastern Puente Hills area, southern California. U.S. Geol. Surv. Prof. Paper 420-B: 1-62. Fischer, P. 1886. Sur deux especes de Lepas fossiles du Miocene des environs de Bor- deaux. Act. Soc. Linn. Bordeaux 40:189-192, pi. 4. Hall, T. S. 1902. New or little known fossils from the Tertiaries of Victoria. Proc. Roy. Soc. Victoria (new series) 15:80-85, pi. 11. Laws C. R. 1948. A new fossil cirripede from New Zealand Miocene beds. Trans. Roy. Soc. New Zealand 77: 151-152, text figs. 1-2. Seguenza G. 1876. Ricerche paleontologiche intorno ai Cirripedi Terziarii della provincia di Messina, pt. 2, Lepadidi. Atti Accad. pontaniana 10:265-481, pis. a-b, 1-10. Withers T. H. 1953. Catalogue of fossil Cirripedia in the Department of Geology, vol. HI. Tertiary. Brit. Mus. (Nat. Hist.), xv + 396 pp., 64 pis. ZuLLO, V. A. 1969. Thoracic Cirripedia of the San Diego Formation, San Diego County, California. Los Angeles Co. Mus., Contrib. Sci. 159:1-25, figs. 1-77. Accepted for publication September 26, 1972 Printed in Los Angeles, California by Anderson, Ritchie and Simon on Simpson Lee Recovery Text NUMBER 242 JUNE 4, 1973 MIDWATER FISHES FROM THE GULF OF CALIFORNIA AND THE ADJACENT EASTERN TROPICAL PACIFIC By Gary D. Brewer CONTRIBUTIONS IN SCIENCE HI NATURAL HISTORY MUSEUM LOS ANGELES COUNTY MIDWATER FISHES FROM THE GULF OF CALIFORNIA AND THE ADJACENT EASTERN TROPICAL PACIFICi By Gary D. Brewer^ Abstract: This faunal study is based on over 56,000 specimens, representing at least 113 species distributed in 49 families, including Alepocephalidae, Apogonidae, Astronestidae, Balistidae, Bathylagidae, Bothidae, Branchiostegidae, Bregma- cerotidae, Carangidae, Cetomimidae, Chiasmodontidae, Chau- liodontidae, Coryphaenidae, Cynoglossidae, Diodontidae, Eche- neidae, Engraulidae, Exocoetidae, Gonostomatidae, Holocen- tridae, Idiacanthidae, Kyphosidae, Linophrynidae, Macrouridae, Melamphaeidae, Melanocetidae, Melanostomiatidae, Moridae, Myctophidae, Nemichthyidae, Neoscopelidae, Nomidae, Ogcoce- phalidae, Oneirodidae, Ophidiidae, Paralepididae, Polynemi- dae, Scopelarchidae, Scorpaenidae, Scyliorhinidae, Serrivo- meridae, Scombridae, Sternoptychidae, Stomiatidae, Stroma- teidae, Thaumatichthyidae, Trachipteridae, Trichiuridae, and Zoarcidae. Specimens were taken by 10-ft Isaccs-Kidd Midwater jj Trawl during the University of Southern California R/V Velero |; IV cruises in the Gulf of California and the adjacent eastern i| tropical Pacific in 1967 and 1970. Approximately 90 percent of the total number of specimens belonged to two families, Myc- tophidae and Gonostomatidae, and the majority of these to only ij two species, Triphotiirus mexicanus and Cyclothone acclinidens, ! respectively. i Several endemic species {Avocettina bowersi, Bathophilus | filifer, Chauliodiis barbatiis, Lestidiops pacificiim, Benthosema ‘ panamense, Diaplius pacificiis, Bregmaceros bathy master, Bro- j tidoides emmelas, Melamphaes laeviceps, Melamphaes macro- I cephalus, Oneirodes luetkeni, Melanocetus ferox, Borophryne j apogon) are adapted, exclusively to the unique hydrological | features (i.e. the extensive oxygen minimum layer) present in the eastern tropical Pacific. Furthermore, the tropical eastern Pacific is a center of origin and distribution for a number of fishes which occur along the west coasts of North and South America, including Vincigiierrki lucetia, Idiacanthus antros- tomus, Diogenichthys laternatiis, Lampanyctiis parvicauda, Melamphaes acanthomas, Scopeloberyx robustas, and Scopelo- gadas mizolepis bispinosus. A comparison of trawl data from inside the Gulf of Cali- fornia with those data from along the coast of Mexico indicate the Gulf to be a limiting environment for a number of eastern Pacific fishes. 1 Review Committee for this Contribution Robert J. Lavenberg H. Geoffrey Moser Basil G. Nafpaktitis ^Department of Biological Sciences, Allan Hancock Foundation, University of Southern California, Los Angeles, Calif. 90007 1 1973 Gulf of California Midwater Fishes 2 INTRODUCTION For a number of years, members of the Department of Biology and Allan Flancock Foundation of the University of Southern California, with support from the National Science Foundation, have been sampling the midwater fauna of the eastern Pacific Ocean aboard the R/V Velero IV. As a part of this program, special cruises to the Gulf of California and to the waters off the west coast of Mexico (the Middle American Trench) were made in November, 1967 and January, 1970, respectively. The purpose of these cruises was to survey the midwater fauna of these areas using a 10-ft Isaacs-Kidd Midwater Trawl (IKMWT) (Isaacs and Kidd, 1953). Almost no deep sea collecting had been undertaken in the eastern tropical Pacific before 1891, when the Albatross made its historic voyage from the Galapagos Islands to the tip of Baja California. Garman’s description (1899) of the midwater and benthic fishes collected during that cruise included 180 species, of which 85% were described as new to science. Townsend and Nichols (1925) listed deep sea fishes from Albatross stations on the outer coast of Baja California and from two stations at the mouth of the Gulf of California. Parr (1931) described midwater fishes taken during the Pawnee Expedition from the west coast of Central America and Mexico. Mesopelagic fishes from off the coast of Peru were reported by Morrow (1957), and from the Peru-Chile Trench by Bussing (1965). Laven- berg and Fitch (1966) made 25 tows with a midwater trawl in the Gulf of California, and Robison (1972) has recently discussed the distribution of 39 species of midwater fishes in the Gulf of California. Several other reports of fishes from the Gulf of California and the eastern tropical Pacific have been published, but generally, these are con- cerned with epipelagic and near-shore species (Meek and Hildebrand, 1923- 28; Breder, 1936; Brock, 1938; Fowler, 1944; Seal, 1940-46; Ricker, 1959; Clemens and Nowell, 1963). Other reports have dealt with only a particular group of midwater fishes, such as those by Beebe and Vander Pyl (1944) and Alverson (1961) on the family Myctophidae. More recently, data from the wide-ranging EASTROPAC cruises were used by Moser and Ahlstrom (1970) and Ahlstrom (1971), to describe development in members of the family Myctophidae and the kinds and abundance of fish larvae in the eastern tropical Pacific, respectively. The eastern tropical Pacific is a hydrographically unique region, largely independent of the major Pacific Ocean gyre systems, with water masses peculiar to it. This uniqueness is reflected in the prominent endemic element of the fauna. A comprehensive survey of the midwater ichthyofauna is not available for a large portion of the eastern tropical Pacific. This study is an attempt to partially fill that void. Furthermore, a knowledge of the systematics and distribution of the eastern tropical Pacific ichthyofauna is essential for an understanding of the origin and relationships of a number of species which range into the Transi- tional Waters off the west coasts of North and South America. 3 Contributions in Science No. 242 This report is intended to 1) give an annotated list of the fishes taken by the R/V Velero IV from two cruises in the Gulf of California and the adjacent eastern tropical Pacific; and 2) to correlate the physico-chemical environment with the midwater fish distribution in the tropical eastern Pacific and areas adjacent to it. !i I fl MATERIALS AND METHODS i Cruise 922, to the Gulf of California by the R/V Velero IV, was under-,, j taken in November, 1967. A total of 35 trawls were made with a 10-ft IKMWT.I i An additional 26 IKMWT’s were taken during cruise 1061, in January, 1970, from off Acapulco, Mexico, to Cabo San Lucas, Baja California. Together, the collections from the two cruises represent a continuous series of stations in the eastern Pacific and the Gulf of California between 16° N, 100° W and | 29° N, 113° W. Station data and a map of the study area are shown in Table 1 and Figure 1, respectively. Station data are listed in sequence, the southernmost stations from cruise 1061 are given first. Included in the data are the date of each trawl and the location when the net started down and when it reached the j surface after fishing. Time, in Pacific Standard Time, is listed when the net started down. Bottom depths were taken from fathometer readings and are j listed in meters. The maximum depth of each trawl was determined from | Benthos time-depth recorder charts. Since the net was without a closing device and fished continuously while in the water, this period is listed as total a fishing time. The period in which the net fished at the approximate desired depth is listed as time at depth. An attempt was made to specifically identify every fish taken. However, this proved impossible because of a number of specimens belonging to poorly \ known families and genera, some larval forms, and damaged individuals. ! When specific determination could not be made, individuals were placed in a higher taxonomic category and the reasons briefly explained in the annota- | tions. A number of specimens in the collection undoubtly belong to undescribed species. These individuals have been so designated and have become available to specialists on the various groups. The listing of orders and families in the annotated list largely follows the phylogenetic sequence adopted by Greenwood et al. (1966), with genera and species in each family listed alphabetically. Positive station numbers follow each species name, and these are followed, in parenthesis, by the number of individuals captured at each station. The size ranges are listed to the nearest millimeter. In all cases, sizes of specimens are in standard length. A plus sign following a measurement indicates that a posterior portion of the animal was missing. Maps, showing the locations of capture are included for the more significant captures only. Since the net was open and fishing continuously at all depths, a number of epipelagic forms were captured. All fishes taken are reported here, but in the discussion major emphasis is placed on the midwater fishes. Gulf of California Midwater Fishes 4 ; 1973 Table 1 Station Data Cruise 1061 Station Date S Lat. N Location End Long. W Hour Bottom Depth Wire Out Max. Trawl Depth Time At Depth Total Fish. Time tart Long. W Lat. N 13727 1-12-70 16° 16' 100° 08' Dip net station 13728 1-12-70 16° 16' 100° 08' 16°17' 100° 14' 22:48 3290 1220 435 1:00 2:02 13729 1-13-70 16° 17' 100° 14' 16°18' 100° 21' 01:05 3290 2750 1100 1:00 2:50 13730 1-13-70 16°33' 100°36' 16°39' 100°59' 04:22 3840 3970 1290 3:00 5:52 13736 1-14-70 17°05' 101°52' 17°01' 102° 07' 08:28 2750 3970 1240 3:00 5:49 13737 1-14-70 17°01' 102°07' 17°08' 102° 18' 14:28 4300 2750 900 2:00 4:22 13738 1-14-70 17°08' 102° 18' 17° 10' 102°22' 19:00 4080 915 250 1:00 1:36 13739 1-15-70 17047' 103°20' 17°54' 103°36' 04:47 3020 3970 1320 3:00 5:57 13740 1-15-70 17°54' 103°36' 18°00' 103°49' 10:51 3200 3360 1050 2:00 4:24 13741 1-15-70 18°00' 103°49' 18°00' 104°01' 15:28 2840 2750 875 2:00 3:59 13748 1-17-70 19°05' 104° 10' 19°05' 105° 13' 13:40 1560 3360 1050 2:00 3:50 13749 1-17-70 19°05' 105°13' 19° 14' 105°32' 17:40 1460 1525 410 1:43 2:30 13750 1-17-70 19014' 105°32' 19°24' 105°45' 20:20 3290 3660 1200 3:00 5:30 13751 1-18-70 19° 24' 105°51' 19°39' 105°51' 02:10 3020 3050 950 3:00 5:04 13757 1-18-70 19°53' 105°59' 20°05' 106° 06' 19:40 2510 3050 940 3:00 5:03 13758 1-19-70 20° 05' 106° 06' 20°21' 106° 10' 00:55 2560 3050 950 3:00 5:07 13759 1-19-70 20°21' 106° 10' 20°40' 106°24' 06:07 2270 3660 1280 3:00 6:08 13760 1-19-70 21°04' 106°21' 21°14' 106°32' 15:37 4210 3060 980 3:00 5:07 13761 1-19-70 21° 19' 106°32' 21°19' 106° 39' 20:50 3660 1220 375 1:30 2:14 13762 1-19-70 21°19' 106°39' 21°30' 106°48' 23:20 3300 3360 1090 2:00 5:30 13763 1-20-70 21°30' 106°48' 21039' 106°53' 04:04 2750 3050 1015 2:00 4:06 13771 1-21-70 22°21' 108° 12' 22°25' 108°29' 15:15 2560 3050 940 2:23 4:22 13772 1-21-70 22° 25' 108°29' 22°31' 108°45' 19:49 2710 3360 1040 2:30 4:43 13773 1-22-70 22°50' 109°09' 22°35' 109°36' 06:10 1970 3660 1275 2:53 5:40 13777 1-22-70 22°34' 109° 29' 22°36' 109°54' 20:30 2800 3970 1150 3:00 5:52 13778 1-23-70 22°36' 109°54' 22°37' 110°11' 02:33 2710 3360 1000 3:00 5:16 13779 1-23-70 22°37' 110° 11' 22°41' 110° 28' 08:01 2560 3050 875 3:00 5:04 Cruise 922 11731 11-07-67 21°26' 106° 43' 21°15' 106°37' 22:35 2790 1070 325 2:00 3:13 11732 11-08-67 21°15' 106°37' 21°02' 106° 39' 03:00 3660 1980 625 2:00 3:50 11733 11-08-67 21°02' 106°39' 20°35' 106° 17' 07:05 4160 3960 1500 4:00 8:15 11734 11-08-67 20°35' 106° 17' 20° 49' 106°24' 15:25 3294 1070 400 2:00 2:57 11735 11-08-67 20°49' 106°24' 21°00' 106°30' 18:35 3360 1980 650 2:00 4:00 11748 11-11-67 21039' 106°58' 22°00' 107°19' 13:20 2820 3960 1675 4:00 8:16 11749 11-11-67 22°00' 107° 19' 22°51' 108° 14' 21:53 2790 1370 450 2:00 3:02 11750 11-12-67 22°51' 108° 14' 22° 24' 108°01' 14:28 2560 3960 1370 4:00 8:38 11751 11-12-67 22024' 108°01' 23°03' 108° 17' 23:16 2880 3960 1500 4:00 8:36 11752 11-13-67 23°03' 108° 17' 22°33' 107°58' 07:47 2870 2290 730 3:00 5:06 11764 11-15-67 23°13' 108°09' 23°32' 108° 24' 10:15 2470 3960 1500 4:00 8:30 11765 11-15-67 23°32' 108°24' 23°42' 108°33' 18:52 2470 2290 730 2:00 4:16 11766 11-15-67 23°42' 108°33' 24°01' 108°52' 23:14 2650 3960 1500 4:00 8:26 11767 11-16-67 24°01' 108°52' 24°23' 109° 10' 08:12 2560 3960 1500 4:00 7:13 11768 11-16-67 24023' 109° 10' 24° 04' 108°54' 15:38 2560 3960 1500 4:00 7:39 11781 11-22-67 24°58' 110° 12' 25° 10' 110° 14' 00:57 1120 1980 550 2:00 3:15 11782 1 1-22-67 25° 10' 110° 14' 25°28' 110° 12' 04:22 1900 3960 1500 4:00 6:54 11783 11-22-67 25°31' 110°09' 25°36' 110° 09' 11:25 2050 1070 400 2:00 2:48 11784 11-22-67 25°36' 110°09' 25°49' 110° 10' 14:20 2020 2290 680 2:00 3:33 11785 11-22-67 25°49' 110° 10' 25°26' 109°58' 18:05 1920 3960 1500 4:00 7:02 11786 11-23-67 25°26' 109°58' 25°03' 110°23' 01:20 2200 3960 1500 2:45 5:50 11798 11-25-67 26°02' 110°23' 26° 19' 110°42' 14:27 1830 3960 1500 4:00 7:03 11799 11-25-67 26° 10' 110° 42' 26° 34' 110°40' 21:45 2280 3050 950 4:00 7:10 11800 11-26-67 26°34' 110° 40' 26°44' 110°41' 05:07 1280 1980 550 2:00 3:20 11801 11-26-67 26°44' 110°41' 26°59' 110°58' 08:35 1280 2590 800 3:00 4:52 11802 11-26-67 26°59' 110°58' 27° 11' 111°17' 13:40 1460 3960 1500 4:00 7:02 11803 11-26-67 27°11' 111°17' 27° 14' 111°22' 20:52 1700 305 100 1:00 1:10 11804 11-26-67 27° 14' 111°22' 27°33' 111°52' 22:15 1920 3960 1500 5:20 8:25 11818 11-28-67 27°27' Mechanical failure, all specimens lost 11819 11-29-67 27°43' 111°56' 27°56' 112° 10' 18:14 1560 3050 950 3:35 2130 660 0:25 6:06 11820 11-30-67 27°56' 112°11' 28° 17' 112°28' 00:30 600 1370 400 4:30 6:22 11821 11-30-67 28°33' 112°49' 28°46' 113°06' 11:15 732 2130 630 4:10 5:31 11822 11-30-67 28°46' 113°06' 28°35' 112°52' 16:53 1280 1530 450 3:00 4:00 11823 11-30-67 28°35' 112°52' 29° 03' 113°21' 21:00 915 2750 850 8:17 9:37 11839 12-03-67 27°08' 111°37' 27°01' 111°32' 10:25 1875 3360 1100 1:23 3:20 5 Contributions in Science No. 242 All material has been deposited at the Natural History Museum of Los Angeles County. ANNOTATED LIST OF FISHES SQUALIFORMES Scyliorhinidae Galeus piperatus Springer and Wagner, 1966 11821 (1) 283 mm. Figure 1. Midwater trawl stations occupied in the Gulf of California and the eastern tropical Pacific during November 1967 (squares) and January 1970 (circles). 1973 Gulf of California Midwater Fishes 6 ANGUILLIFORMES Leptocephali 13729 (11) 65-226 mm; 13730 (2) 103-112 mm; 13737 (1) 120 mm; 13738 (2) 62-106 mm; 13739 (4) 85-124 mm; 13741 (2) 73-99 mm; 13748 (12) 84-180 mm; 13749 (11) 49-191 mm; 13750 (10) 82-119 mm; 13751 (11) 74-182 mm; 13758 (25) 73-120 mm; 13759 (22) 93-165 mm; 13760 (12) 53-108 mm; 13762 (10) 52-123 mm; 13763 (10) 90-189 mm; 13771 (8) 80-165 mm; 13772 (16) 50-180 mm; 13777 (1) 80 mm; 13779 (2) 77-120 mm; 11734 (49) 62-175 mm. On the basis of pigmentation, body length and depth, and the shape and size of the head and jaws, at least eight species of leptocephali are represented in the collection. Serrivomeridae Serrivomer sector G2Lvm?Lr\, 1899 (Fig. 2) 13729 (1) 394 mm; 13730 (2) 410-526 mm; 13736 (3) 398-525 mm; 13737 (5) 340-410 mm; 13739 (4) 351-508 mm; 13740 (8) 410-605 mm; 13741 (1) 300 mm; 13748 (7) 340-410 mm; 13751 (3) 448-540 mm; 13757 (7) 404-473 mm; 13758 (6) 460-490 mm; 13759 (5) 344-558 mm; 13760 (2) 380-458 mm; 13761 (1) 325 mm; 13762 (6) 441-561 mm; 13763 (4) 400-603 mm; 13771 (4) 425-485 mm; 13772 (3) 420-470 mm; 13773 (1) 422 mm; 13777 (3) 443-574 mm; 13778 (3) 470-530 mm; 13779 (10) 425-520 mm; 11733 (8) 362-600 mm; 11748 (2) 425-558 mm; 11750 (4) 432-504 mm; 11751 (5) 412-584 mm; 11752 (1) 420 mm; 11764 (4) 448-H-537 mm; 11766 (3) 419-590 mm; 1 1767 (5) 339-553 mm; 1 1768 (1) 606 mm; 1 1782 (2) 405-575 mm; 1 1784 (1) 150-f mm; 1 1785 (6) 420-586 mm; 1 1786 (2) 480-540 mm; 11802 (3) 480-546 mm; 11804 (5) 465-570 mm; 11819 (1) 490 mm; 1 1839 (1) 554 mm. Nemichthyidae Avocettina howersi (Garman, 1899) (Fig. 2) 13738 (2) 240-386-f mm; 13748 (1) 323 -f mm; 13757 (2) 249-f-455 mm; 13758 (2) 347-390-hmm; 13760(3) 346-420 mm; 1376 1 ( 1) 365 -f- mm; 1 1766 ( 1 ) 420 mm; 11799 (1) 420 mm; 11802 (1) 441 mm; 11804 (1) 458 mm; 1 1819 (3) 453-515 mm. Avocettina infans (Gunther, 1878) (Fig. 2) 13773 (2) 455-525 mm; 13778 ( 1) 552 mm; 1 1750 (1) 612 mm; 11751 (1) 490+ mm; 11765 (1) 612 mm; 11766 (1) 714 mm; 11767 (1) 645 mm; 11802 (1) 554 mm. It is with some hesitation that I have assigned specific names to the above specimens of Avocettina. The taxonomic limits of the genus are not well understood, and much confusion exists in the literature. Part of the confusion arises because the long, slender bodies of the animals are easily broken, and after regeneration of new caudal fin rays, it is difficult to determine how much of the tail has been lost. Thus, at times, meristic data are of little value. The problem is further confounded by the apparent extreme intraspecific variation in such characters as number of fin rays, number of vertebrae, and length of snout. In their revision of the family, Roule and Bertin (1929) suggested that Nemichthys infans (Gunther), Labichthys elongatus Gill and Ryder, Labich- thys gilli Bean, and Labichthys bowersi Garman were synonymous with Avocettina infans (Gunther). This suggestion was based on the extreme variability of several characters in “Dana” specimens from both Atlantic and Pacific Oceans. They were hesitant to include bowersi in the synonmy, but 7 Contributions in Science No. 242 only because Carman’s dorsal ray count of 252 was low compared to over 300 rays for other described species. All other morphometric and meristic data of bowersi, they felt, were within the range found for A. infans. In the present specimens a broad range of morphometric and meristic characters was similarly found. Two groups were finally established on the basis of cephalic sensory pore arrangement. In A. bowersi, the anteriormost supraorbital pore lies in front of the posterior nare, while in A. infans all of the supraorbital pores are posterior to the nares. The ranges of counts and measurements were found to vary for the two species as follows: Dorsal rays Lateral line pores Diameter of eye into postorbital distance (distance from posterior margin of the eye to the opercular opening) Dorsal-fin origin Anal-fin origin beneath A. infans 303-340 183-195 2.7-3.T 0.0-0. 5 eye diameters behind the posterior end of the pectoral fin base 24-34 dorsal-fin ray A. bowersi 202-287 143-183 3.8-4.4 1. 5-2.0 eye diameters behind the posterior end of the pectoral fin base 16-24 dorsal-fin ray Environmental influences undoubtedly play an important role in causing variation in morphometric and meristic characters in these eels, but the degree of geographical variation for each species is as yet, unknown. Clearly, a more detailed study with a larger sample size is needed. Nemiclithys scolopacens Richardson, 1848 (Fig. 3) 13751 (2) 293-856 mm; 13759 (2) 867-924 mm; 13760 (1) 795 mm; 13772 (2) 300- 785 mm; 13777 (1) 310 mm; 13779 (3) 880-989 mm; 11766 (1) 795+ mm; 11767 (1) 853 mm; 1 1768 (2) 835-1 147 mm; 11784 (1) 529 mm; 1 1785 (1) 882 mm; 11786 (2) 651-760+ mm; 1 1798 (1) 755 mm; 1 1799 (2) 535-970 mm; 11801 (1) 880 mm; 1 1802 (1) 626+ mm; 11803 (1) 556 mm; 11804 (4) 389-960 mm; 1 1819 (2) 715 + - 748 mm; 11820 (1) 605 mm. CLUPEIFORMES Engraulidae Engraulidae — unidentified larvae 13760 (2) 13 mm. SALMONIFORMES Bathylagidae Bathylagus nigrigenys Parr, 1931 (Fig. 3) 13728 (1) 36 mm; 13729 (8) 25-44 mm; 13730 (2) 31-64 mm; 13736 (2) 51-52 mm; 13737 (6) 22-87 mm; 13738 (5) 28-79 mm; 13740 (25) 21-89 mm; 13741 (24) 19- 74 mm; 13748 (6) 21-40 mm; 13750 (3) 37-70 mm; 13751 (6) 25-71 mm; 13757 (12) 24-73 mm; 13758 (27) 20-73 mm; 13759 (5) 25-53 mm; 13760 (33) 25-85 mm; 13761 (6) 23-63 mm; 13763 (29) 24-79 mm; 13771 (17) 28-69 mm; 13772 (1 1) 1973 Gulf of California Midwater Fishes 8 Figure 2. Locations of capture of A) Serrivomer sector, B) Avocettina bowersi, and C) Avocettina infans indicated by blackened squares and circles. 9 Contributions in Science No. 242 »■ 26- 69 mm; 1 1731 (1) 58 mm; 1 1732 (5) 23-31 mm; 11733 (10) 15-82 mm; 11748 (6) ll 14-65 mm; 1 1749 (2) 63-74 mm; 1 1750 (9) 17-83 mm; 11751 (3) 19-50 mm; 1 1764 if (2) 52-53 mm; 11766 (1) 58 mm; 11767 (1) 80 mm; 11782 (1) 25 mm; 11802 (1) 63 mm. | According to larval surveys by Ahlstrom (1971), this species is common if throughout the eastern tropical Pacific north of 5° S. In the present study, t B. nigrigenys was taken only occasionally within the Gulf, where it is largely replaced by the more abundant Leuroglossus stilbius stilbius (this supports . earlier distributional data by Lavenberg and Fitch, 1966). Furthermore, while i L. stilbius stilbius is taken off the outer Baja California coast, B. nigrigenys is apparently replaced by the closely related B. wesethi (Wisner, 1962; Berry and Perkins, 1966). Bat by lag us pacific us Gilbert, 1890 I 11750 (1) 155 mm. This is the first record of Bathylagus pacificus from the Gulf of California, 1 and the southernmost record. It was previously recorded from the Gulf of |i Alaska to the Pacific side of northern Baja California (Fitch and Lavenberg, ' 1968). , I Bathylagus wesethi Bolin, 1939 13779 (4) 24-59 mm. Leuroglossus stilbius stilbius Gilbert, 1890 (Fig. 3) 11764 (3) 27-36 mm; 11765 (1) 26 mm; 1 1767 (8) 21-56 mm; 1 1768 (16) 15-47 mm; ■ 11781 (10) 21-80 mm; 1 1782 (3) 26-32 mm; 11784 (6) 31-74 mm; 1 1785 (5) 20-34 ' 1 mm; 1 1786 (6) 24-30 mm; 1 1798 (37) 24-67 mm; 1 1799 (4) 25-50 mm; 1 1800 (129) 27- 79 mm; 11801 (90) 24-79 mm; 11802 (21) 21-92 mm; 11803 (441) 23-65 mm; 1 1804 (21) 23-55 mm; 11819 (23) 22-74 mm; 11820 (632) 25-96 mm; 11821 (51) II 31-51 mm; 1 1839 (29) 21-91 mm. , Borodulina (1968) has recognized three subspecies of Leuroglossus, L. | stilbius schmidti from the Bering Sea, L. stilbius stilbius from the north eastern Pacific, and L. stilbius urotranus from the Peru-Chile Trench. A single < specimen of L. stilibus stilbius reported from the Gulf of Panama (Borodulina, 1968) is of questionable identity. In the present study, 26 IKMWT’s outside the Gulf of California, along the coast of Mexico failed to yield representatives of this form. Gonostomatidae Cyclothone accliniclens Garman, 1899 (Fig. 4) 13728 (499) 21-51 mm; 13729 (229) 27-47 mm; 13730 (176) 26-50 mm; 13736 (130) 27-56 mm; 13737 (564) 20-53 mm; 13738 (444) 25-49 mm; 13739 (321) 28-54 mm; 13740 (614) 26-48 mm; 13741 (462) 17-51 mm; 13748 (628) 20-53 mm; 13749 (138) 26-50 mm; 13750 (205) 20-51 mm; 13751 (1846) 20-55 mm; 13757 (2556) 20-53 mm; 13758 (1962) 17-54 mm; 13759 (1302) 20-48 mm; 13760 (734) 18-51 mm; 13761 (930) 23-55 mm; 13762 (261) 20-48 mm; 13763 (903) 20-50 mm; 13771 (879) 18-52 mm; 13772 (1189) 16-49 mm; 13773 (454) 22-53 mm; 13777 (78) 10-55 mm; 13778 (458) 24-54 mm; 13779 (653) 17-48 mm; 1 1733 (1) 35 mm. This was the second most abundant fish taken in the survey. A total of 18,606 individuals were collected at 27 stations. Surprisingly, not a single 1973 Gulf of California Midwater Fishes 10 Figure 3. Locations of capture of A) Nemichthys scolopaceiis, B) Bathylagiis nigrigenys, and C) Leuroglossiis stilbiiis stilbiiis indicated by blackened squares and circles. Contributions in Science No. 242 1 1 individual was taken north of 22°50'N in the Gulf of California. This phenomenon is discussed in a later section. || Cyclotlione pallida Brauer, 1902 (Fig. 4) ( 13729 (1) 62 mm; 13730 (2) 50-56 mm; 13737 (4) 53-65 mm; 13739 (6) 50-64 mm; | 13740 (2) 45-57 mnr, 13741 (2) 42-45 mm; 13748 (2) 45-55 mm; 13750 (2) 54-61 mm; [ 13751 (2) 58-61 mm; 13759 (3) 55-59 mm; 1 3762 (4) 38-64 mm; 13763 (2) 5 1-55 mm; I! 13772 (8) 50-59 mm; 13773 (3) 53-59 mm; 13778 (7) 44-60 mm; 13779 (2) 54-58 mm. | Cyclotlione signata Garman, 1899 (Fig. 4) y 13748 (1) 19 mm; 13751 (1) 21 mm; 13757 (1) 28 mm; 13758 (2) 24-33 mm; 13760 I (1) 26 mm; 13762 (1) 16 mm; 13763 (1) 18 mm; 13779 (1) 21 mm. I Diplophos taenia Gunther, 1873 (Fig. 5) ||] 13763 (2) 36 mm; 13773 (1) 84 mm; 13777 (1) 44 mm; 13778 (1) 78 mm; 11731 (3) I 36-105 mm; 11732 (6) 86-106 mm; 11734 (12) 58-102 mm; 11748 (3) 68-99 mm; | 11750 (1) 119 mm; 11752 (4)95-115 mm; 11786 (1) 93 mm. | Until the taxonomic status of this species is thoroughly reviewed, the | specific identification of these individuals must remain tentative. Johnson (1970) suggested that three species of Diplophos described from the Pacific, | D. pacificus Gunther, D. orientalis Matsubara, and D. proximus Parr, are | synonymous with the Atlantic form D. taenia. In an earlier study by Grey I (1960), Pacific specimens were placed in D. taenia, but later (1964), she I questioned this synonomy because of lower photophore counts in the Pacific | specimens. 1 The range of photophore counts for the present specimens are followed, ? in parentheses, by counts by Grey (1964) for the Atlantic D. taenia: BR 11 (12); IV 39-41 (47); VAV 14-16 (16-17); AC 41-44+ 2 (46-47 + 2-3); IC 97- 102 (111-113); OA 77-84 (71). Body proportions of these Pacific specimens are not significantly different from those listed for the Atlantic D. taenia. FZ/zc/gz/c/T/V/ ///ccr/rt (Garman, 1 899) (Fig. 5) ! 13728 (52) 9-36 mm; 13729 (27) 1 1-35 mm; 13730 (6) 18-41 mm; 13736 (17) 15- 45 mm; 13737 (9) 22-51 mm; 13738 (23) 7-35 mm; 13739 (94) 22-46 mm; 13740 (5) j 15-36 mm; 13741 (11) 12-30 mm; 13748 (19) 15-30 mm; 13749 (10) 12-31 mm; ! 13750 (23) 9-34 mm; 1375 1 (18) 12-36 mm; 13757 (7) 22-39 mm; 13758 (38) 9- 46 mm; 13759 (13) 12-46 mm; 13760 (9) 12-30 mm; 13761 (2) 11-54 mm; 13762 (21) 8-33 mm; 13763 (23) 10-48 mm; 13771 (24) 14-44 mm; 13772 (3) 13-19 mm; 13773 (24) 19-28 mm; 13777 (12) 20-44 mm; 13778 (10) 12-39 mm; 13779 (32) 13- 46 mm; 1 1732 (1) 12 mm; 1 1733 (201) 1 1-38 mm; 1 1734 (150) 12-43 mm; 1 1735 (331) 12-38 mm; 11748 (148) 15-41 mm; 1 1751 (135) 12-46 mm; 11752 (90) 11- 33 mm; 1 1764 (38) 14-41 mm; 1 1765 (2) 13-29 mm; 1 1766 (1) 20 mm; 1 1767 (19) 18-31 mm; 11768 (18) 12-27 mm; 11781 (23) 12-34 mm; 11782 (18) 16-46 mm; 11783 (82) 12-34 mm; 1 1784 (23) 12-30 mm; 11785 (7) 30-41 mm; 11786 (52) 14- 52 mm; 11798 (31) 14-41 mm; 1 1799 (59) 18-51 mm; 11801 (42) 13-34 mm; 11802 (37) 1 1803 (44) 13-53 mm; 11804 (40) 12-48 mm; 11820 (1) 37 mm; 11839 (59) mm. Vinciguerria liicetia was described by Ahlstrom and Counts (1958) as “the most ubiquitous and abundant species of fish in plankton collections from the eastern Pacific.” Over 2000 individuals were collected at 52 stations in the present study. • • *00 ♦/ 1973 Gulf of California Midwater Fishes 12 Figure 4. Locations of capture of A) Cyclothone acclinidens, B) Cyclothone pallida, and C) Cyclothone signata indicated by blackened squares and circles. 13 Contributions in Science No. 242 Sternoptychidae Argyropelecus lychnus (Garman, 1899) (Fig. 5) 13773 (1) 32 mm; 13777 (1) 50 mm; 13778 (1) 50 mm; 11766 (1) 35 mm; 11781 (1) 34 mm; 1 1783 (6) 1 1-16 mm; 1 1784 (1) 13 mm; 1 1820 (5) 35-64 mm. According to the latest revision of the family by Baird (1971), only two species of hatchetfishes occur in the eastern tropical Pacific, Argyropelecus lychnus and Sternoptyx obscura Garman. Astronesthidae Borostomias panamensis Regan and Trewavas, 1930 11782 (1) 220 mm. Borostomias panamensis was originally described from the Gulf of Panama, but has been taken as far north as Point Conception (Berry and Perkins, 1966). Melanostomiatidae Bathophilus filifer Garman, 1899 (Fig. 6) 13729 (5) 21-54 mm; 13730 (1) 56 mm; 13737 (2) 62-63 mm; 13738 (5) 35-82 mm; 13739 (5) 21-82 mm; 13740 (4) 27-75 mm; 13741 (22) 61-79 mm; 13748 (4) 21-76 mm; 13749 (4) 15-73 mm; 13750 (3) 66-86 mm; 13751 (9) 56-86 mm; 13757 (1) 68 mm; 13758 (3) 44-60 mm; 13759 (3) 71-78 mm; 13760 (1) 73 mm; 13763 (2) 65- 69 mm; 13772 (2) 60-75 mm; 1 173 1 (2) 69 mm; 1 1732 (2) 29-30 mm; 1 1733 (3) 29-68 mm; 11734 (4) 43-67 mm; 1 1735 (1) 58 mm; 11750 (7) 63-85 mm; 11752 (2) 56-66 mm; 1 1764 (1) 76 mm; 1 1766 (4) 63-83 mm; 1 1767 (1) 82 mm; 1 1782 (1) 67 mm. Bathophilus sp. 1 1766 (1) 83 mm. This specimen is damaged and specific identification cannot be made with certainty. Chauliodontidae Chauliodus barbatus Garman, 1899 13741 (1) 40 mm; 13751 (1) 198 mm. Chauliodus macouni Bean, 1890 11802 (1) 188 mm. Morrow, in his review (1964) of the genus, describes the above species as the only two occurring in the eastern Pacific. C. macouni is frequently taken in the north eastern Pacific and was previously known from the Bering Sea to off northern Baja California (Berry and Perkins, 1966). C. barbatus was originally described from the Gulf of Panama (13°01'N) and was recorded in the Peru-Chile Trench by Bussing (1965). According to Morrow (1961), C. barbatus “probably ranges to central Mexico.” The northernmost station that yielded C. barbatus 13751 (19°24'N). Stomiatidae Stomias atriventer Garman, 1899 (Fig. 6) 13740 (1) 29 mm; 13741 (1) 124 mm; 13750 (3) 29-35 mm; 13757 (2) 44-56 mm; 13758 (5) 76-145 mm; 13759 (10) 89-162 mm; 13760 (3) 96-125 mm; 13771 (13) 97-211 mm; 13772 (20) 87-202 mm; 13773 (9) 43-183 mm; 13777 (1) 140 mm; 13778 (19) 30-172 mm; 13779 (88) 56-210 mm; 11731 (1) 181 mm; 1 1732 (2) 38- 1973 Gulf of California Midwater Fishes 14 Figure 5. Locations of capture of A) Diplophos taenia, B) Vinciguerria lucetia, and C) Argyropelecus lychnus indicated by blackened squares and circles. q 11748 (1) 72 mm; 11751 (1) 48 mm; 11782 (1) 29 mm. i These specimens strongly resemble Bajacalifornia drakei (Beebe) in most | morphometric and meristic data as listed by Parr (1937) and Bussing (1965). j However, the snout length (in percent of body length) of the present specimens 1 1 (13.1) is longer than previously listed (10.5 and 10.9, respectively). Also, the ; ' number of gill rakers in the present specimens (19) is low compared to the f | range (24-26) listed by the above authors. The maxillary reaches to below j the anterior rim of the orbit in the present individuals, while in the specimens listed by Parr (1937) and Bussing (1965), the maxillary reaches to below the middle of the eye. Barbantus curvifrons (Roule and Angel, 1931) '>■ 11767 (1) 107 mm. I This specimen is in good condition and is remarkably similar to sped- ;; mens described by Parr (1960) from the Atlantic and Indian Oceans. ■ Counts for the Velcro specimen are followed by ranges given by Parr (1960): Dorsal 15 (15); anal 16 (15-17); pectoral 25 (20-24); ventral 8 (7-8); C transverse scale rows from lateral line to dorsal 8 (6-8), and from lateral line | to anal 8 (6-8); gillrakers 5+14 (4-5 + 14). Body proportions (in percent of I body length) for the Velero specimen are followed in parentheses by those listed by Parr (1960) for a 1 15 mm specimen from the Indian Ocean: Head 27.5 (28.7); snout 6.0 (5.6); orbit 9.8 (9.8); upper jaw 13.5 (14.4); lower jaw 1973 Gulf of California Midwater Fishes 16 Figure 6. Locations of capture of A) Bathophiliis filifer, B) Stomias atriventer, and C) Idicanthus antrostomus indicated by blackened squares and circles. 17 Contributions in Science No. 242 15.1 (14.9); interorbital 4.6 (4.5); snout to dorsal 58.3 (62.6); snout to anal 68.6 (68.9); base of pectoral 4.6 (4.8); longest pectoral ray 9.6 (9.6); greatest depth 20.5 (21.7); longest gillraker 3.7 (3.5). The bony, horizontal spines projecting laterally on each side at the tip of the lower Jaw, which characterize the genus, are largely worn off in the present specimen. Holthyrnia macrops Maul, 1957 1 1766 (1) 63 mm; 1 1799 (2) 56-68 mm. Holtbyrnia melanocephala (Vaillant, 1888) 13758 (1) 38 mm. The specific identification of this young alepocephalid is questionable because the light organs are not fully developed. Normichthys campbelli Lavenberg, 1965 13740 (1) 86 mm; 13771 (2) 81-98 mm. This species was previously known from the Santa Catalina Basin, off southern California (Lavenberg, 1965). Pellisolus faciUis Parr, 1951 13737 (1) 98 mm; 13739 (3) 41-100 mm; 13740 (1) 27 mm; 13759 (1) 22 mm; 11764 (1) 34 mm; 1 1733 (1) 47 mm; 1 1766 (1) 59 mm. Talismania bifurcata (Parr, 1951) 13737 (1) 27 mm; 13738 (1) 38 mm; 13741 (1) 50 mm; 13779 (1) 89 mm; 11782 (1) 208 mm; 1 1798 (1) 28 mm; 1 1839 (1) 28 mm. Alepocephalidae — unidentified 13730 (2) 35-80 mm. The larger specimen is in fair condition and is characterized by the presence of two supramaxillaries; the absence of scales (no scale pockets are evident); the origin of the dorsal fin being well in advance of the anal fin origin; the maxillary reaching well beyond the posterior margin of the eye; the teeth in the jaws being pluriserial; the head being about 36% of standard length; and by lacking a shoulder organ. Counts for the same specimen are as follows: Dorsal 15-16; anal 11; ventral 8; pectoral 6; gillrakers on first arch 13; pyloric caeca 7. Based on keys by Parr (1937, 1951, 1952), the combination of the above characters prevent any generic designation. Alepocephalidae — unidentified 13730 (1) 15 mm; 13757 ( 1) 24 mm; 13777 (1) 15 mm; 1 1767 (1) 56 mm. These individuals are small or badly damaged. Paralepididae Lestidiops pacificum (Parr, 1931) (Fig. 7) 13736 (1) 127 mm; 13737 (1) 125 mm; 13738 (1) 29 mm; 13740 (2) 83-87 mm; 13741 (1) 68 mm; 13771 (1) 139 mm; 11731 (1) 76 mm; 11733 (4) 78-167 mm; 11734 (1) 86 mm; 1 1750 (1) 96 mm. 1973 Gulf of California Midwater Fishes 18 Figure 7. Locations of capture of A) Bajacalifornia bnrragei, B) Lestidiops pacificum, and C) Scopelarchoides nicholsi indicated by blackened squares and circles. 19 Contributions in Science No. 242 Scopelarchidae Scopelarchoides nicholsi Parr, 193 1 (Fig. 7) 11731 (1) 99 mm; 11733 (2) 34-46 mm; 11735 (1) 102 mm; 11750 (3) 25-104 mm; 11752 (1) 96 mm; 11764 (5) 90-106 mm; 11766 (1) 101 mm; 11768 (1) 105 mm; 1 1781 (1) 83 mm; 1 1786 (2) 69-99 mm. Myctophidae Benthosema panamense (Taning, 1932) (Fig. 8) 13736 (3) 15-19 mm; 11765 (1) 14 mm; 11768 (1) 10 mm; 11782 (1) 18 mm; 11783 (35) 12-22 mm; 11784 (10) 12-22 mm; 11786 (4) 19-23 mm; 11798 (6) 14-19 mm; 1 1799 (3) 12-16 mm; 1 1800 (8) 13-17 mm; 1 1801 (12) 14-42 mm; 1 1802 (29) 14-36 mm; 11803 (71) 11-47 mm; 11819 (11) 11-36 mm; 11820 (21) 12-41 mm; 11821 (9) 16-37 mm; 11839 (38) 10-34 mm. Benthosema panamense is endemic to the tropical eastern Pacific. Diaphiis pacificus Parr, 1931 (Fig. 8) 13728 (44) 5-28 mm; 13729 (1) 26 mm; 13730 (58) 17-30 mm; 13736 (27) 16-30 mm; 13737 (29) 22-32 mm; 13738 (36) 9-30 mm; 13739 (34) 7-31 mm; 13740 (83) 9-32 mm; 13741 (58) 9-29 mm; 13748 (43) 13-32 mm; 13749 (10) 11-31 mm; 13750 (4) 10- 11 mm; 13751 (10) 11-32 mm; 13758 (15) 11-34 mm; 13759 (17) 25-29 mm; 13760 (18) 9-33 mm; 13761 (6) 9-31 mm; 13762 (2) 16-29 mm; 13763 (26) 12-31 mm; 13771 (10) 21-31 mm; 13772 (8) 10-29 mm; 13773 (18) 19-30 mm; 13777 (5) 25-28 mm; 13778 (3) 20-33 mm; 13779 (1) 17 mm; 1 1731 (5) 13-32 mm; 11732 (68) 13-43 mm; 1 1733 (68) 13-33 mm; 11734 (485) 9-34 mm; 1 1735 (8) 13-30 mm; 11748 (5) 15-28 mm; 1 1749 (5) 9-31 mm; 11750 (30) 11-31 mm; 11751 (42) 10-31 mm; 1 1752 (61) 9-35 mm; 11764 (4) 21-26 mm; 11765 (1) 16 mm; 11766 (15) 16-32 mm; 11767 (9) 9-22 mm; 11768 (6) 10-32 mm; 11782 (1) 32 mm; 11783 (22) 13-34 mm; 11784 (2) 14-15 mm; 11786 (4) 17-35 mm; 11800 (1) 31 mm; 11801 (3) 23-24 mm; 1 1839 (2) 14-21 mm. Diaphus pacificus is endemic to the eastern tropical Pacific. Diogenichthys laternatus (Carman, 1899) (Fig. 8) 13728 (1) 16 mm; 13729 (3) 9-10 mm; 13730 (4) 20-24 mm; 13736 (10) 10-23 mm; 13737 (5) 12-17 mm; 13738 (15) 11-24 mm; 13739 (3) 13-32 mm; 13740 (48) 9- 25 mm; 13741 (2) 16-26 mm; 13748 (4) 18-25 mm; 13749 (4) 13-20 mm; 13751 (58) 16-25 mm; 13757 (3) 21-24 mm; 13758 (1) 24 mm; 13760 (11) 12-26 mm; 13761 (4) 22-24 mm; 13762 (5) 10-28 mm; 13763 (25) 8-24 mm; 13771 (58) 8-25 mm; 13772 (18) 18-25 mm; 13773 (60) 15-27 mm; 13777 (15) 21-24 mm; 13778 (24) 15-25 mm; 13779 (24) 18-26 mm; 11731 (133) 12-26 mm; 11732 (166) 10-27 mm; 11733 (1 17) 10-26 mm; 11734 (120) 10-25 mm; 11735 (74) 10-23 mm; 11748 (95) 11- 26 mm; 11749 (24) 15-24 mm; 11750 (71) 9-28 mm; 11751 (63) 10-28 mm; 11752 (115) 10-21 mm; 1 1764 (57) 13-25 mm; 11765 (49) 11-27 mm; 11766 (125) 11-26 mm; 11767 (136) 11-28 mm; 11768 (109) 13-26 mm; 11781 (44) 13-27 mm; 1 1782 (38) 12-27 mm; 11783 (1) 26 mm; 11784 (66) 14-26 mm; 11785 (22) 16- 24 mm; 1 1786 (86) 13-25 mm; 1 1798 (51) 13-27 mm; 11799 (13) 14-26 mm; 11800 (112) 15-30 mm; 11801 (206) 14-28 mm; 11802 (74) 12-28 mm; 11803 (33) 13- 26 mm; 11804 (23) 16-26 mm; 11819 (8) 9-24 mm; 11820 (3) 21-25 mm; 11839 (288) 13-27 mm. This was the second most commonly taken myctophid and the third most numerous species of any kind collected during the two cruises. A total of 2827 individuals were taken at 55 stations. Gonichthys tenuiculus (Carman, 1899) 13773 (2) 42 mm; 13778 (1) 50 mm; 13779 (3) 39-44 mm; 1 1764 (1) 14 mm; 1 1781 (1)31 mm; 11798 (1) 46 mm. 1973 Gulf of California Midwater Fishes 20 Figure 8. Locations of capture of A) Benthosema pamimense, B) Diaplius pacificiis, and C) Diogenichthys laternatiis indicated by blackened squares and circles. 21 Contributions in Science No. 242 Hygophum atratiim (Garman, 1899) (Fig. 9) 13740 (1) 49 mm; 13748 (1) 43 mm; 13760 (1) 49 mm; 13761 (1) 50 mm; 13761 (5) 35-60 mm; 13763 (2) 38-48 mm; 13771 (2) 48-50 mm; 13772 (1) 45 mm; 13773 (5) 24-48 mm; 13778 32-50 mm; 13779 (4) 37-53 mm; 11731 (1) 46 mm; 11732 (1) 17 mm; 11733 (4) 15-42 mm; 11734 (11) 22-48 mm; 11748 (1) 20 mm; 11751 (6) . 13-48 mm; 1 1752 (72) 17-60 mm; 1 1764 (4) 24-45 mm; 1 1765 (1) 43 mm; 11767 (2) | 17-36 mm; 11781 (2) 20-23 mm; 11785 (1) 32 mm; 11799 (1) 56 mm; 11800 (1) 51 mm; 11801 (3) 40-47 mm; 11802 (1) 48 mm; 11803 (1) 46 mm; 11804 (17) 12- 5 1 mm. Lampanyctiis idostigma Parr, 1931 (Fig. 9) 13750 (1) 58 mm; 13751 (1) 38 mm; 13757 (7) 30-66 mm; 13759 (3) 55-60 mm; 13761 (5) 35-60 mm; 13762 (13) 27-75 mm; 13771 (22) 31-80 mm; 13778 (119) 26-82 mm; 13779 (191) 30-86 mm; 11731 (13) 18-57 mm; 11733 (11) 21-49 mm; 11734 (62) 18-62 mm; 11735 (6) 27-44 mm; 11748 (10) 21-60 mm; 11749 (7) 21-33 mm; 11750 (45) 24-76 mm; 11751 (7) 24-60 mm; 1 1752 (57) 19-58 mm; 11764 (18) 21-66 mm; 11765 (5) 23-54 mm; 11766 (47) 19-79 mm; 11767 (20) 48-79 mm; 11768 (18) 42-73 mm; 11781 (7) 23-60 mm; 11782 (5) 32-82 mm; 11784 (1) 23 mm; 1 1786 (2) 20-53 mm. Lampanyctus parvicauda Parr, 1931 (Fig. 9) 13728 (6) 29-54 mm; 13729 (4) 24-76 mm; 13730 (84) 21-83 mm; 13736 (102) 23-90 mm; 13737 (77) 26-70 mm; 13738 (16) 24-80 mm; 13739 (40) 30-87 mm; 13740 (23) 20-90 mm; 13741 (13) 27-66 mm; 13748 (2) 20-80 mm; 13749 (3) 27- 62 mm; 13750 (29) 25-97 mm; 13751 (20) 22-75 mm; 13757 (16) 44-93 mm; 13758 (2) 24-68 mm; 13759 (14) 26-95 mm; 13760 (10) 60-83 mm; 13761 (1) 66 mm; 13762 (8) 26-90 mm; 13763 (4) 78-94 mm; 13771 (2) 32-95 mm; 13772 (3) 75-92 mm; 13773 (21) 25-45 mm; 13777 (49) 17-96 mm; 13778 (5) 22-95 mm; 13779 (39) 21-113 mm; 11731 (2) 27-89 mm; 1 1733 (75) 21-93 mm; 11734 (20) 32-90 mm; 1 1735 (7) 22-88 mm; 11748 (242) 23-89 mm; 11749 (14) 23-75 mm; 11750 (68) 21-92 mm; 11764 (25) 22-79 mm; 11765 (2) 63-75 mm; 11766 (82) 20-101 mm; 11767 (14) 23-85 mm; 11768 (1) 91 mm; 11781 (1) 28 mm; 11782 (3) 23-29 mm; 11784 (1) 92 mm; 1 1786 (3) 22-34 mm; 11800 (1)73 mm; 11804 (1) 84 mm. Lampanyctus sp. 13740 (3) 25-31 mm; 13757 (1) 13 mm; 11749 (2) 25-28 mm. These damaged specimens can not be identified to species. Bolinichthys longipes (Brauer, 1906) 13777 (1) 21 mm. Myctophum aurolatematum Garman, 1899 (Fig. 10) 13728 (3) 21-58 mm; 13729 (6) 26-77 mm; 13730 (1) 73 mm; 13736 (19) 19-74 mm; 13737 (1) 45 mm; 13738 (1) 25 mm; 13739 (2) 23-44 mm; 13740 (3) 21-22 mm; 13741 (1) 70 mm; 13743 (1) 75 mm; 13748 (2) 21 mm; 13751 (1) 23 mm; 13758 (1) 22 mm; 13771 (2) 67-77 mm; 13772 (1) 21 mm; 13773 (5) 32-36 mm; 1 1732 (1) 51 mm; 1 1733 (2) 20 + -25 mm; 11748 (6) 26-80 mm; 1 1750 (1) 22 mm; 11751 (1) 19 mm. Myctophum sp. 13773 (1) 24 mm. This specimen is badly damaged. Taaningichthys bathyphilus Taning, 1928 1 1733 (1) 62 mm; 1 1767 (1) 57 mm. O ♦ o ♦ 1973 Gulf of California Midwater Fishes 22 Figure 9. Locations of capture of A) Hygophiini atratiun, B) Lampanyctus idos- tigma, and C) Lcinipanyctus parvicciuda indicated by blackened squares and circles. 23 Contributions in Science No. 242 Triphot urns mexicanus GiVoQYi, 1890 (Fig. 10) 13760 (3) 42 mm; 13762 (1) 40 mm; 13763 (7) 25-49 mm; 13771 (36) 28-57 mm; 13772 (6) 40-56 mm; 13773 (10) 19-64 mm; 13777 (3) 43-57 mm; 13778 (20) 28- 60 mm; 13779 (262) 14-69 mm; 11783 (14) 15-20 mm; 11822 (3) 25-44 mm; 11823 (18) 18-46 mm. Over 23,000 specimens of T. mexicanus were captured at a single trawl station in the Gulf; unfortunately labels indicating the exact station number have subsequently been lost. Probably the population of T. mexicanus in the Gulf is enormous. Such aggregations have been noted previously by Lavenberg and Fitch (1966). According to Moser and Ahlstrom (1970), T. mexicanus has been con- fused with the closely related T. oculeus (Garman). The former ranges throughout the Transitional Water of the eastern north Pacific and into the Gulf of California while T. oculeus ranges from off Panama to Peru. The southernmost station that yielded T. mexicanus was 13760 (21°N). Myctophidae — unidentified 13729 (2) 6 mm; 13738 (10) 6-8 mm; 13739 (7) 7-8 mm; 13750 (3) 5-10 mm; 13758 (6) 7-11 mm; 13759 (1) 21 mm; 11733 (3) 19-20 mm; 11750 (2) 11-15 mm; 11751 (10) 11-27 mm. These myctophids were either damaged or too small to identify. Neoscopelidae Scopelengys tristis Alcock, 1892 (Fig. 10) 13736 (2) 136-175 mm; 13739 (1) 148 mm; 13740 (3) 140-167 mm; 13748 (1) 139 mm; 13751 (3) 135-150 mm; 13757 (6) 98-139 mm; 13758 (1) 155 mm; 13760 (1) 140 mm; 13763 (3) 112-157 mm; 13771 (2) 89-133 mm; 11748 (1) 84 mm; 11751 (1) 161 mm; 11768 (1) 61 mm; 11786 (3) 113-146 mm; 11798 (1) 150 mm; 11799 (4) 104-142 mm; 1 1802 (5) 104-175 mm; 1 1804 (1) 158 mm; 11819 (5) 1 12-160 mm. CETOMIMIFORMES Cetomimidae Gyrinomimus bruuni Rofen, 1957 13729 (1) 71 mm; 13739 (1) 101 mm; 11748 (2) 72-104 mm; 11768 (1) 199 mm. This entire family is poorly known and in need of revision. The counts and measurements of the present specimens fit descriptions of G. bruuni, which was described from off Kenya. Cetomimus sp. 13737 (1) 86 mm; 1 1786 (1) 127 mm. These specimens probably belong to an undescribed species. Cetomimidae — unidentified 11733 (1) 92 mm; 11764(1) 102 mm. These two bizarre individuals can not be placed in any of the known genera of the family. They are characterized by a long, curved upper jaw that projects well beyond the slightly curved lower jaw. Their description will appear in a forthcoming paper by Gerald Citek, of the University of Southern California. 1973 Gulf of California Midwater Fishes 24 Figure 10. Locations of capture of A) Myctophiim aurolaternatum, B) Triphoturiis mexicanus, and C) Scopelengys tristiis indicated by blackened squares and circles. 25 Contributions in Science No. 242 !? ' LOPHIIFORMES Ogcocephalidae Ogcocephalidae — unidentified 11800(1) 32 mm. ' Melanocetidae ^ Melanocetus ferox Regan, 1926 11748 (1) 114 mm. This species has so far been reported only from the eastern tropical Pacific. This and the following ceratioids were identified by Theodore Pietsch of the University of Southern California. Melanocetus johnsoni Gunther, 1864 13729 (1) 15 mm; 13730 (1) 20 mm; 13737 (1) 20 mm; 13739 (1) 18 mm; 13751 (1) 24 mm; 13758 (1) 20 mm; 13759 (1) 24 mm; 13760 (2) 19 mm; 13761 (2) 19 mm; (1) 17 mm; 13772 (2) 20 mm; 13779 (1) 34 mm; 1 1764 (1) 22 mm. | I Oneirodidae Dolopichthys pullatus Regan and Trewavas, 1932 11764 (1) 48 mm. Bertelsen (1951) synonymized D. pullatus with Dolopichthys longicornus, but recent work by Pietsch (1972b) based on larger sample size has shown this species to be distinct. Oneirodes luetkeni (RegSin, 1925) 13740 (2) 45-60 mm; 13759 (1) 15 mm. One i rode s sp. 13750(1) 24 mm. This individual cannot be assigned to any known species until further work on the genus clarifies existing taxonomic problems. Oneirodidae — unidentified 11767 (1) 11 mm. According to Pietsch (personal communication), this individual belongs to an undescribed genus. Thaumatichthyidae Thaumatichthys pagidostomus Smith and Radcliff, 1912 13737 (1) 22 mm. Thaumatichthys pagidostomus has been taken in the Atlantic and Indian Oceans (Bertelsen, 1951), and this is the first record of the species from the Pacific. Centrophrynidae Centrophryne spinulosa Regan and Trewavas, 1932 13730 (2) 168-209 mm; 13737 (1) 16 mm. The 168 mm female taken at station 13730 had a male Melanocetus johnsoni attached to its upper lip, and was the subject of a paper by Pietsch and Nafpaktitis (1971). This monotypic family was recently reviewed by Pietsch (1972a). 1973 Gulf of California Midwater Fishes 26 Linophrynidae Borophryne apogon Regan, 1925 (Fig. 1 1) 13729 (2) 13 mm; 13736 (1) 19 mm; 13737 (1) 15 mm; 13739 (1) 36 mm; 13740 (1) 35 mm; 13741 (1) 14 mm; 11751 (1) 14 mm; 1 1767 ( 1) 33 mm; 1 1768 (2) 16-19 mm; 11783 (1) 18 mm; 1 1786 (1) 13 mm; 11798 (8) 13-110 mm; 11799 (1) 13 mm; 11802 (8) 12-36 mm; 1 1803 (1) 15 mm; 1 1804 (5) 18-25 mm. This species is apparently endemic to the tropical eastern Pacific. GADIFORMES Bregmacerotidae Bregmaceros atlanticus Goode and Bean, 1886 (Fig. 1 1) 13730 (1) 43 mm; 13739 (1) 22 mm; 13748 (1) 48 mm; 13749 (6) 18-51 mm; 13761 (2) 38 mm; 13771 (3) 24-26 mm; 1 1732 (2) 45-49 mm; 11733 (3) 38-49 mm; 11734 (9) 22-52 mm; 11750 (6) 24-56 mm; 11751 (2) 47-49 mm; 11752 (4) 45-50 mm; 1 1764 (1) 56 mm; 1 1766 (1) 47 mm; 1 1802 (1) 50 mm. Bregmaceros bathy master iordan and Bollman, 1890 13749 (2) 36-42 mm; 1 1734 (2) 34 mm. Bregmaceros spp. 13759 (2) 15 mm; 13760 (1) 13 mm; 13763 (1) 17 mm; 13758 (6) 8-1 1 mm. There are at least three forms represented by these unidentified larvae and juveniles. The latest revision of the bregmacerotids by D’Ancona and Cavinato (1965) lists three species of Bregmaceros {B. atlanticus, B. bat hy master, and B. macclellandii Thompson) as occurring in the eastern tropical Pacific, all of which were taken in the Gulf of Panama. B bathymaster was subsequently taken in the Gulf of California (Lavenberg and Fitch, 1966; Robison, 1972), and Ahlstrom (1971) has found five kinds of Bregmaceros larvae in the eastern tropical Pacific. The key to Bregmaceros by D’Ancona and Cavinato relies heavily on longitudinal scale counts, a difficult character in these small, often poorly preserved fish. Many of the counts and measurements overlap considerably among the various species, making specific identification difficult. In the present specimens, B. atlanticus has been distinguished from B. bathymaster by vertebral counts, taken from radiographs. Moridae Microlepidium vereciindum (Jordan and Cramer, 1896) 13759 (3) 63-82 mm; 1 1750 (1) 72 mm. Microlepidium grandiceps Garman (1899), from the eastern tropical Pacific, has been found to be synonymous with M. verecundum (Fitch and Barker, 1972). Ophidiidae Brotidoides emmelas (Gilbert, 1890) 13771 (1) 67 mm; 11732 (1) 46 mm; 11734 (1) 51 mm; 1 1735 (1) 43 mm; 11749 (2) 42-48 mm; 1 175 1 (1) 43 mm; 11765 (2) 51-55 mm. Ophidiidae — unidentified 13728 (1) 10 mm; 13739 (2) 14-44 mm; 13750 (2) 35-41 mm; 13761 (1) 16 mm; 13763 (1) 28 mm; 13772 (1) 38 mm; 1 1786 (1) 36 mm. 27 Contributions in Science No. 242 1973 Gulf of California Midwater Fishes 28 I Zoarcidae i Melanostigma pammelas GiVoQri, 1893 (Fig. 11) I 13758 (1) 96 mm; 11767 (1) 77 mm; 11798 (4) 60-83 mm; 11802 (4) 57-89 mm; I 11804 (5)40-113 mm; 11819(2) 105-115 mm; 11839 (2) 26-88 mm. i Following Bussing (1965), these specimens were identified on the basis ! of vertebral and fin-ray counts (specimens were X-rayed). McAllister and I Rees (1964) in their revision of the genus, proposed an antitropical distribution ! for the three then known species of Melanostigma and suggested a means of i dispersal for the group. The subsequent discovery of a fourth species, M. I bathium Bussing (1965), in the Peru-Chili Trench, between 33° and 46°S, i supports the hypothesis of McAllister and Rees (1964) regarding the existence j of a tropical barrier. The capture of M. pammelas at 20° N (station 13758) is the southernmost I record of this species. i I Macrouridae j Coelorhynchus scaphopsis GWhQVi, 1890 13771 (1) 97 mm; 13778 (1) 81 mm; 11748 (1) 91 mm; 11819 (1) 69 mm. ATHERINIFORMES j Exocoetidae Exocoetus monocirrhiis KiohdiYdson, 1846 13737 (1) 36 mm. This specimen was taken by dip net. Hyporhamphus iinifasciatus (Ranzani, 1842) 13727 (9) 18-84 mm; 13728 (1) 48 mm; 13730 (7) 33-65 mm; 13741 (1) 31 mm. BERYCIFORMES Melamphaeidae Melamphaes acanthomus Ebeling, 1962 (Fig. 12) 13729 (2) 70-76 mm; 13737 (2) 29-83 mm; 13738 (2) 21-31 mm; 13740 (1) 99 mm; 13741 (1) 81 mm; 13748 (5) 66-84 mm; 13749 (2) 25-87 mm; 13751 (7) 75-96 mm; 13757 (2) 51 mm; 13758 (12) 78-105 mm; 13759 (3) 74-86 mm; 13760 (9) 74-90 mm; 13762 (2) 91-107 mm; 13763 (7) 75-97 mm; 13771 (8) 85-98 mm; 13772 (6) 56-105 mm; 13773 (1) 42 mm; 13777 (1) 67 mm; 13778 (3) 88-115 mm; 13779 (2) 77-87 mm; 1 1733 (2) 65-93 mm; 1 1750 (2) 42-88 mm; 1 1751 (3) 74-93 mm; 1 1767 (5) 70-92 mm; 11768 (1) 99 mm; 1 1785 (1 1) 63-99 mm; 1 1786 (1) 95 mm; 11798 (5) 75-84 mm; 11799 (7) 73-89 mm; 11802 (10) 73-101 mm; 11804 (8) 79-89 mm; 11819 (8) 71-94 mm; 11839 (3) 80-96 mm. Melamphaes laeviceps Ebeling, 1962 13750 (1) 84 mm; 13751 (1) 27 mm; 13773 (1) 93 mm. Melamphaes maerocephalus Parr, 1931 (Fig. 12) 13728 (1) 45 mm; 13729 (7) 24-1 12 mm; 13736 (4) 26-98 mm; 13737 (7) 27-101 mm; 13739 (10) 26-97 mm; 13740 (1) 42 mm; 13741 (3) 42-98 mm; 13748 (3) 96-100 mm; 13750 (6) 27-102 mm; 13751 (4) 23-77 mm; 13757 (1) 106 mm; 13758 (2) 56-82 mm; 13759 (3) 40-99 mm; 13772 (8) 68-104 mm; 13773 (4) 76-108 mm; 11732 (2) 58- 68 mm; 1 1733 (6) 54-102 mm; 1 1734 (3) 41-79 mm; 11735 (2) 58-66 mm; 11748 (11) 28-94 mm; 11750 (9) 42-94 mm; 1 1751 (8) 46-88 mm; 11752 (2) 66-92 mm; 11766 (6) 54-95 mm; 11767 (3) 89-106 mm; 1 1768 (2) 70-83 mm; 1 1782 (3) 77- 95 mm; 1 1786 (6) 52-91 mm; 1 1798 (1) 70 mm; 1 1802 (1) 103 mm. 29 Contributions in Science No. 242 Figure 12. Locations of capture of A) Melamphaes acanthomiis, B) Melampluies macrocephalus, and C) Melamphaes spinifer indicated by blackened squares and circles. 1973 Gulf of California Midwater Fishes 30 Melamphaes spinifer Ebeling, 1962 (Fig. 12) 13736 (1) 25 mm; 13738 (2) 30-39 mm; 13739 (1) 35 mm; 13740 (1) 31 mm; 13750 (4) 26-41 mm; 13751 (3) 23-48 mm; 13757 (1) 24 mm; 13759 (1) 44 mm; 13760 (1) 62 mm; 13763 (2) 57-64 mm; 13772 (5) 33-56 mm; 13773 (1) 59 mm; 11732 (4) 26-48 mm; 1 1733 (3) 22-55 mm; 1 1748 (5) 37-66 mm; 1 1749 (1) 21 mm; 1 1750 (3) 17 + -43 mm; 1 1751 (3) 21-49 mm; 1 1764 (3) 44-53 mm; 1 1765 (2) 49-55 mm; 11766 (4) 20-50 mm; 1 1768 (1) 26 mm; 1 1784 (1) 54 mm. Melamphaes spp. 13739 (5) 16-17 mm; 13771 (1) 18 mm. These juveniles could not be identified specifically. Poromitra crassiceps {Gunther, 1878) (Fig. 13) 13729 (1) 71 mm; 13737 (1) 60 mm; 13739 (1) 123 mm; 13740 (7) 69-131 mm; 13751 (2) 48-55 mm; 13758 (1) 46 mm; 13759 (4) 67-120 mm; 13762 (3) 39-122 mm; 13763 (1) 106 mm; 13772 (2) 63-65 mm; 13773 (1) 71 mm; 13778 (2) 99-126 mm; 1 1750 (8) 78-121 mm; 1 1766 (1) 57 mm; 1 1767 (2) 66-88 mm. Poromitra sp. 13730 (1) 61 mm; 13739 (1) 65 mm; 1 1733 (1) 75 mm; 11748 (1) 64 mm; 11750 (1) 62 mm; 1 1751 ( 1) 64 mm. These fish represent an undescribed species of Poromitra. Their small eye, low dorsal fin-ray count (I II- 10), and low gillraker count (23-24) dis- tinguish them from the known species of this genus. According to Ebeling (personal communication), this species ranges from the eastern tropical Pacific through the west Pacific and the Indian Ocean. Scopeloheryx rohiistus {Gunther, 1887) (Fig. 13) 13729 (2) 41-46 mm; 13730 (7) 29-64 mm; 13736 (6) 31-61 mm; 13737 (10) 24-60 mm; 13739 (17) 30-57 mm; 13750 (7) 35-69 mm; 13751 (4) 30-44 mm; 13759 (4) 30-46 mm; 13762 (2) 42-60 mm; 13773 (20) 32-66 mm; 13777 (9) 28-58 mm; 13778 (14) 35-52 mm; 11733 (2) 31-53 mm; 11748 (6) 48-64 mm; 11750 (5) 41-56 mm; 11751 (13) 34-66 mm; 11764 (26) 38-67 mm; 11766 (24) 36-62 mm; 11767 (3) 48-58 mm; 1 1768 (8) 40-65 mm; 1 1782 (8) 45-68 mm; 1 1785 (4) 48-61 mm; 1 1786 (15) 49-59 mm; 1 1798 (3) 54-60 mm; 1 1799 (2) 49-55 mm; 1 1802 (1) 63 mm. Scopelogadus mizoloepis hispinosus (Gilbert, 1890) (Fig. 13) 13728 (18) 12-60 mm; 13729 (15) 18-51 mm; 13730 (3) 30-50 mm; 13736 (10) 20- 60 mm; 13737 (11) 21-55 mm; 13738 (32) 16-51 mm; 13739 (19) 10-54 mm; 13740 (168) 23-65 mm; 13741 (83) 17-55 mm; 13748 (33) 15 mm; 13749 (9) 37-53 mm; 13750 (19) 19-59 mm; 13751 (75) 14-54 mm; 13757 (96) 10-50 mm; 13758 (99) 8-57 mm; 13759 (71) 25-62 mm; 13760 (263) 13-67 mm; 13761 (61) 11-52 mm; 13762 (14) 10-63 mm; 13763 (96) 9-68 mm; 13771 (24) 28-50 mm; 13772 (10) 16-57 mm; 13773 (9) 14-43 mm; 13777 (1) 37 mm; 13778 (5) 33-55 mm; 13779 (18) 28-63 mm; 1 1731 (38) 11-63 mm; 1 1732 (38) 10-56 mm; 1 1733 (57) 20-60 mm; 1 1734 (12) 18-44 mm; 1 1735 (38) 18-56 mm; 1 1748 (15) 29-54 mm; 11749 (7) 30-56 mm; 1 1750 (51) 25-57 mm; 1 1751 (16) 11-59 mm; 11752 (139) 1 1-58 mm; 11764 (2) 28-32 mm; 1 1765 (8) 31-57 mm; 1 1766 (5) 17-40 mm; 1 1767 (10) 26-50 mm; 11768 (5) 29-52 mm; 1 1781 (5) 34-63 mm; 11782 (7) 26,-52 mm; 11784 (1) 40 mm; 11785 (2) 37-39 mm; 1 1798 (1) 44 mm; 1 1799 (2) 35-37 mm; 11800 (8) 26-49 mm; 1 1801 (1) 49 mm; 11819 (8) 22-34 mm. Holocentridae Holocentriis suborhitalis Gill, 1863 13727 (12) 17-40 mm; 13731 (6) 14-34 mm. 31 Contributions in Science No. 242 Figure 13. Locations of capture of A) Poromitra crassiceps, B) Scopeloberyx rohustus, and C) Scopelogadiis mizolopis bispinosiis indicated by blackened squares and circles. 1973 Gulf of California Midwater Fishes 32 LAMPRIDIFORMES Trachipteridae Tnichipteriis sp. 13738 (1) 1 1 mm. SCORPAENIFORMES Scorpaenidae Sebastes sinensis Gilbert, 1890 1 1822 (14) 46-128 mm; 1 1823 (22) 47-148 mm. Sebastes sp. 11823 (1) 160 mm. This specimen represents an undescribed species (Lo-chai Chen, personal communication). Scorpaena sp. 13741 (1)21 mm. Scorpaenidae-unidentified 13738 (13) 3-8 mm; 13739 (1) 5 mm; 13740 (2) 9-13 mm; 13750 (2) 9-13 mm. PERCIEORMES Apogonidae Apogon dovii Giinther, 1861 13737 (1) 20 mm. This individual was identified by a key in Breder (1936), but because of its small size, the specific identification is not certain. Branchiostegidae CcinlolatHns prince ps (Jenyns, 1842) 13751 (1) 16 mm; 13758 (2) 14-18 mm; 13762 (1) 18 mm. Echeneidae Phtheirichthys I i neat ns (Menzies, 1791) 13743 (1) 5rmm. Remora remora (Linnaeus, 1758) 13731 (1) 65 mm. Carangidae Caranx canimis Gunther, 1869 13727 (7) 56-72 mm; 13728 (1) 55 mm; 13738 (1) 52 mm; 13748 (1) 82 mm. Decapterus hypodns Gill, 1862 13751 (1) 97 mm. Selene brevoortii (Gill, 1863) 13736 (1) 18 mm. The identification of this juvenile must remain tentative until comparative material is obtained. Coryphaenidae Coryphaena hippnrns Linnaeus, 1758 13727 (2) 91 mm. Kyphosidae Kyphosus elegans (Peters, 1869) 13727 (3) 55-70 mm. 33 Contributions in Science No. 242 Figure 14. Locations of capture of A) Chios moclon niger, and B) Kali nornmni indicated by blackened squares and circles. 1973 Gulf of California Midwater Fishes 34 Polynemidae Polydactyliis opercniaris Gill, 1863 13761 (1)41 mm. Chiasmodontidae Cliiasmoclon niger Johnson, 1863 (Fig. 14) 13730 (1) 105 mm; 13737 (1) 74 mm; 13739 (2) 78-1 19 mm; 13748 (1) 103 mm; 13750 (1) 1 16 mm; 13773 (1) 39 mm; 1 1734 (2) 43-44 mm; 1 1748 (1) 44 mm; 1 1750 (2) 39-89 mm. This genus is in need of revision, but these specimens fit Norman’s ( 1929) key for C. niger. Kali nornumi (Parr, 1931) (Fig. 14) 13729 (2) 64-139 mm; 13737 (1) 142 mm; 13740 (1) 142 mm; 13741 (2) 56-74 mm; 13748 (1) 125 mm; 13757 (1) 139 mm; 13758 (1) 136 mm; 13759 (1) 133 mm; 13760 (1) 160 mm; 13761 (1) 132 mm; 13772 (1) 136 mm; 13779 (1) 76 mm. Trichiuridae Lepidopiis xantnsi Goode and Bean, 1896 1 1820 ( 1 ) 314 mm. Scombridae Scombridae — unidentified 13731 (47) 7-17 mm. Nomeidae Cubiceps carinatns Nichols and Murphy, 1944 13728 (6) 6-18 mm; 13729 (1) 16 mm; 13736 (1) 77 mm; 13738 (1) 20 mm; 13741 (1) 1 1 mm; 13748 (1) 11 mm; 13759 (1) 98 mm; 13760 (1) 17 mm; 13773 (2) 64-68 mm; 13778 (1) 24 mm. These fish were identified with the help of the key to the stromateoid fishes by Haedrich and Horn (1969). The species was originally described from the Panama Bight by Nichols and Murphy ( 1944). Stromateidae Peprilus spp. 13730 ( 1) 13 mm; 13736 (1) 23 mm; 13739 (1) 6 mm; 13751 (1) 1 1 mm; 13760 (1) 13 mm; 13761 (1)5 mm. According to Horn (1970), three species of Peprilus occur in all or part of the study area, but these young fish are too small to identify. PLEURONECTIFORMES Bothidae Citharichthys sp. — larvae 13739 (1) 9 mm; 13761 (1) 12 mm. Etropus crossatiis Jordan and Gilbert, 1881 13728 (2) 13 mm; 13729 (2) 10-24 mm; 13749 (4) 13-23 mm; 13750 (2) 16-20 mm; 13751 (1) 13 mm; 13759 (1) 22 mm; 13762 (2) 15-16 mm; 13763 (1) 13 mm. Bothidae — unidentified 13748 (1) 13 mm; 13751 (1) 10 mm. 35 Contributions in Science No. 242 I Cynoglossidae Symphurus spp. 13741 (1) 19 mm; 13751 (1) 23 mm; 13758 (1) 20 mm; 13759 (3) 17 mm. Two species of Symphurus are represented here. One form has a large ;j midlongitudinal, darkly pigmented area, the other has small scattered areas of pigmentation. ^ TETRAODONTIFORMES Balistidae Canthiclerniis rotufuhis {de Proce, 1822) 13727 (4) 35-1 1 1 mm. Alutera scripta (Osbeck, 1765) 13731 (1) 156 mm. Diodontidae Diodofi holacanthus Linnaeus, 1758 13743 (3) 60-82 mm. Unidentified 13727 (3) 13-15 mm; 13728 (14) 4-18 mm; 13729 (6) 4-14 mm; 13736 (2) 17-21 mm; 13738 (9) 8-13 mm; 13739 (9) 4-8 mm; 13748(1) 16 mm; 13749 (1) 8 mm; 13750 (6) 7-28 mm; 13758 (26) 4-22 mm; 13759 (6) 9-25 mm; 13761 (7) 4-10 mm; 13762 (3) 4-22 mm; 13763 (2) 8-10 mm; 13771 (1) 78 mm; 11766(1) 14 mm; 1 1767 (1) 35 mm. At least a dozen forms are represented in these larvae and damaged individuals. SUMMARY OF FISHES Over 56,000 individual fish were taken during cruises of the R/V Velero /F to the Gulf of California and the eastern tropical Pacific. These specimens belong to at least 14 orders, 49 families, 82 genera, and 113 species. The exact number of species cannot be determined until a number of larval forms are identified. A single species of the family Myctophidae made up a substantial portion of the total number of specimens. Over 23,000 individuals of Triphoturus mexicanus were taken at a single IKMWT station. The gonostomatid, Cyclotiwne acclinidens, contributed over 18,000 individuals, making it the second most numerous species taken. The myctophid, Diogenichthys laterna- tus, and the gonostomatid, Vinciguerria lucetia were also quite numerous, with over 2,900 and 2,100 specimens, respectively. The most frequently captured species were Diogenichthys laternatus (55 stations), Vinciguerria lucetia (52 stations), the melamphaeid Scopelogadus mizolepis bispinosus (50 stations), and the myctophid Diaphus pacificus (47 stations). The most numerous species, Triphoturus mexicanus and Cyclotiwne acclinidens, were captured in only 12 and 27 trawls, respectively. At least 26 species were not taken in more than one IKMWT, and of these, 24 species were represented by single specimens. The greatest number of species identi- fied from one trawl was 30, taken at station 1 375 1 . Gulf of California Midwater Fishes 36 1973 The most speciose orders were the Salmoniformes, with 39 species distributed in 15 families and 32 genera, and the Perciformes, with 13 families, 18 genera, and 21 species. The families with the greatest number of species were the Myctophidae, with 10 genera and 11 species; the Alepocephalidae (including Searsidae), with 7 genera and 9 species; and the Melamphaeidae, with 4 genera and 8 species. Two families, the Myctophidae and the Gonos- tomatidae, included approximately 90% of all individuals taken. Table 2 provides a summary of fishes collected. The number of positive trawl stations and the number of individuals are listed for each species. Further- more, a comparison is made of the number of occurrences and the number of specimens between positive stations within the Gulf of California (north of 23 °N), and stations south of the Gulf. This table will be considered in the discussion section in reference to the Gulf as a limiting environment. DISCUSSION In recent years it has become common practice for biologists to relate water masses, as defined by temperature-salinity curves, to the distribution of pelagic organisms. Since the studies by Bieri (1959), McGowan (1960), Brinton (1962), and Ebeling (1962), it has been popular to superimpose the distribution of pelagic organisms over water mass boundaries. Although water masses are not the final answer to distributional problems, as pointed out recently by Backus et al. (1970), the fact remains that the distributions of many pelagic species are best understood when considered in relation to temperature-salinity and oxygen profiles. In the following distributional analysis, water masses are used to categorize the broad distributional trends of the midwater fishes collected during the survey. The water mass characteristics of the eastern tropical Pacific remain largely independent of those in adjacent regions. The area, lying in the con- cavity formed by North, Central, and South American coastlines, is protected from the major Pacific gyral systems by its geographic position. The conserva- tive and non-conservative properties of the eastern tropical Pacific waters reflect this isolation when compared to adjacent regions. The subsurface waters of the entire eastern tropical Pacific, north of the equator, and in the Gulf of California, are relatively uniform and result from the slow horizontal spread of water from the Equatorial Undercurrent. The great depths in the southern and northern sections of the eastern tropical Pacific are filled with Intermediate Water of Antarctic and North Pacific origin, respectively. Probably the outstanding feature of the subsurface waters off the coast of Mexico and in the Gulf of California is the presence of an oxygen minimum layer. This layer forms a zone as thick as 1200 m off Mexico, where oxygen content falls below 1.0 ml/1. Oxygen concentrations below 0.1 ml/1 are present vertically through several hundred meters (Griffiths, 1968). Several midwater species taken in the survey are apparently endemic to the eastern tropical Pacific. These include Avocettina bowersi, Bathophilus 37 Contributions in Science No. 242 Table 2 Summary of fishes collected in the Gulf of California and the eastern tropical Pacific by the R/V Velero Gulf of California Pacific Ocean South of Gulf of California Total No. of No. of No. of No. of No. of No. of Species occurrences specimens occurrences specimens occurrences specimens Scyliorhinidae Galeus piperatus 1 1 0 0 1 1 Leptocephali 0 0 20 221 20 221 Serrivomeridae Serrivomer sector 13 35 26 108 39 143 Nemichthyidae Avocettina bower si 5 7 6 11 11 18 Avocettina infans 4 4 4 5 8 9 Nemichthys scolopaceus 14 21 6 11 20 32 Engraulidae unidentified 0 0 1 2 1 2 Bathylagidae Bathylagus nigrigenys 5 6 26 264 31 270 Bathylagus pacificus 0 0 1 1 1 1 Bathylagus wesethi 0 0 1 4 1 4 Leuroglossus stilbius stilbius 20 1536 0 0 20 1536 Gonostomatidae Cyclothone acclinidens 0 0 27 18,616 27 18,616 Cyclothone pallida 0 0 16 52 16 52 Cyclothone signata 0 0 8 9 8 9 Diplophos taenia 2 5 9 30 11 35 Vinciguerria lucetia 20 686 32 1498 52 2184 Sternoptychidae Argyropelecus lychnus 5 14 3 3 8 17 Astronesthidae Borostomias panamensis 1 1 0 0 1 1 Melanostomiatidae Bathophilus filifer 5 9 23 95 28 104 Bathophilus sp. 1 1 0 0 1 1 Chauliodontidae Chauliodus bar bat us 0 0 2 2 2 2 1973 Gulf of California Midwater Fishes 38 Gulf of California Pacific Ocean South of Gulf of California Total Species No. of No. of occurrences specimens No. of No. of occurrences specimens No. of No. of occurrences specimens Chauliodus macouni 1 1 0 0 1 1 Stomiatidae Stomias atriventer 20 188 20 189 40 377 Idiacanthidae Idiacanthus antrostomus 0 0 17 29 17 29 Alepocephalidae Bajacaliforn ia burragei 6 16 3 3 9 19 iia jaca liforn ia sp. 1 1 2 3 3 Barbantus curvifrons 1 1 0 0 1 1 Holtbyrnia macrops 2 3 0 0 2 3 Holtbyrnia melanocephala 0 0 1 1 1 1 Normichthys campbelli 0 0 2 3 2 3 Pellisolus facillis 3 3 4 6 6 9 Tal is mania bifur cat a 3 3 4 4 7 7 unidentified 1 1 4 5 5 6 Paralepididae Lestidiops pacificum 0 0 10 14 10 14 Scope larch idae Scope larchoides nicholsi 6 11 4 7 10 18 Myctophidae Benthosema panamense 16 260 1 3 17 263 Diaphus pacificus 13 131 34 1282 47 1413 Diogenichthys laternatus 22 1658 33 1269 55 2927 Gonich thys tenuiculus 3 3 3 6 6 9 Hygophum atratum 12 106 16 47 28 153 Lampanyctus idostigma 10 180 17 523 27 703 Lampanyctus parvicauda 11 134 33 1021 44 1155 Lampanyctus sp. 0 0 3 6 3 6 Bolinicthys longipes 0 0 1 1 1 1 39 Contributions in Science No. 242 Gulf of California Pacific Ocean South of Gulf of California Total Species No. of No. of occurrences specimens No. of No. of occurrences specimens No. of No. of occurrences specimens Myctophum aurolaternatum 0 0 21 61 21 61 Myctophum sp. 0 0 1 1 1 1 Taaningichthys bathyphilus 1 1 1 1 2 2 Triphoturus mexicanus 3 23,000 9 348 12 23,000 unidentified 0 0 9 44 9 44 Neoscopelidae Scopelengys tristus 7 20 12 25 19 45 Cetomimidae Gyrinomimus bruuni 1 1 3 4 4 5 Cetomimus sp. 1 1 1 1 2 2 unidentified 1 1 1 1 2 2 Ogcocephalidae unidentified 1 1 0 0 1 1 Melanocetidae Melanocetus ferox 0 0 1 1 1 1 Melanocetus johnsoni 1 1 12 15 13 16 Oneirodidae Dolopichthys pullatus 1 1 0 0 1 1 Oneirodes luetkeni 0 0 2 3 2 3 Oneirodes sp. 0 0 1 1 1 1 unidentified 1 1 0 0 1 1 Thaumatichthyidae Thaumatichthys pagidostomus 0 0 2 3 2 3 Linophrynidae Borophryne apogon 9 28 7 8 16 36 Bregmacerot idae Bregmaceros atlanticus 4 7 11 36 15 43 Bregmaceros bathymaster 0 0 2 4 2 4 Bregmaceros spp. 0 0 4 10 4 10 Moridae Microlepidium verecundum 0 0 2 4 2 4 Ophidiidae Brotuloides emmelas 1 2 6 7 7 9 unidentified 1 1 6 8 7 9 1973 Gulf of California Midwater Fishes 40 Gulf of California Pacific Ocean South of Gulf of California Total Species No. of No. of occurrences specimens No. of No. of occurrences specimens No. of No. of occurrences specimens Zoarcidae Melanostigma pammelas 6 18 1 1 7 19 Macrouridae Coelorhynchus scaphopsis 1 1 3 3 4 4 Exocoetidae Exocoetus monocirrhus 0 0 1 1 1 1 Hyporhamphus unifasciatus 0 0 4 18 4 18 Melamphaeidae Melamphaes acan thomus 10 59 23 85 33 144 Melamphaes laeviceps 0 0 3 3 3 3 Melamphaes macrocephalus 8 24 22 105 30 129 Melamphaes spin if er 5 11 18 42 23 53 Melamphaes spp. 0 0 2 6 2 6 Poromitra crassiceps 2 3 13 34 15 37 Poromitra sp. 0 0 6 6 6 6 Scopeloberyx robust us 10 94 16 128 26 222 Scopelogadus mizoloepis bispinosus 15 204 35 1534 50 1738 Holocentridae Holocentrus suborbitalis 0 0 2 18 2 18 Trachipteridae Trachipterus sp. 0 0 1 1 1 1 Scorpaenidae Sebastes sinensis 2 36 0 0 2 36 Sebastes sp. 1 1 0 0 1 1 Scorpaena sp. 0 0 1 1 1 1 unidentified 0 0 4 18 4 18 Apogonidae Apogon dovii 0 0 1 1 1 1 Branchiostegidae Caulolatilus princeps 0 0 3 4 3 4 Echeneidae Phtheirichthys lineatus 0 0 1 1 1 1 Remora remora 0 0 1 1 1 1 41 Contributions IN Science No. 242 Pacific Ocean South of Gulf of California Gulf of Califonia Total No. of No. of No. of No. of No. of No. of Species occurrences specimens occurrences specimens occurrences specimens Carangidae Caranx caninus Decapterus 0 0 4 10 4 10 hypodus Selene 0 0 1 1 1 1 brevoortii 0 0 1 1 1 1 Coryphaenidae Coryphaena hippurus 0 0 1 2 1 2 Kyphosidae Kyphosus elegans 0 0 1 3 1 3 Polynemidae Polydactylus opercularis 0 0 1 1 1 1 Chiasmodontidae Chiasmodon niger 0 0 9 12 9 12 Kali normani 0 0 12 14 12 14 Trichiuridae Lepidopus xantusi 1 1 0 0 1 1 Scombridae unidentified 0 0 1 47 1 47 Nomeidae Cubiceps carinatus 0 0 10 16 10 16 Stromateidae Peprilus spp. 0 0 6 6 6 6 Bothidae Citharichthys sp. Etropus 0 0 2 2 2 2 cr os sot us 0 0 8 15 8 15 unidentified 0 0 2 2 2 2 Cynoglossidae Symphurus spp. 0 0 4 6 4 6 Balistidae Canthidermis rotundas Alutera 0 0 1 4 1 4 scrip ta 0 0 1 1 1 1 Diodontidae Diodon holacanthus 0 0 1 3 1 3 Unidentified 2 2 15 96 17 98 Gulf of California Midwater Fishes 42 1973 ' filifer, Chauliodus barbatus, Lestidiops pacificum, Benthosema panamense, Diaphus pacificus, Bregmaceros bathymaster, Brotuloides emmelas, Melcim- phaes laeviceps, Melamphaes macrocephcdus, and the ceratioids, Oneirodes ' luetkeni, Melcmocetiis ferox, and Borophryne apogon. From the north flows the California Current carrying relatively cool, low salinity water which is gradually warmed and mixed with subtropical water as it approaches the tip of the Baja Peninsula. Oceanographers have classically considered the region around the tip of Baja California to be the southern limit of Transitional Water, a derivative of Subarctic Water (Sverdrup et al., 1942). It is not mere coincidence, then, that the southern range limits of several northern species have been found in this area. The capture of two bathylagids, Bathylagus wesethi and Bathylagus pacificus, and the zoarcid Melanostigma pammelas, in the mouth of the Gulf, suggests the influence of California Current water in this region. California Current water may at times enter the Gulf (Griffiths, 1968), and this may explain the presence of another northern species, Chauliodus macouni, some distance into the Gulf itself. The eastern tropical Pacific is the center of origin and distribution of a number of species that are common in the Transitional Water Mass along the west coast of North America. Those species able to adapt to the cooler, less saline Transitional Water may have achieved their northern ranges by being carried by the subsurface counter current flowing northward along the coast. The following fish, taken in the present survey, range into the Transi- tional Water of the California Current system: Vinciguerria lucetia, A rgy ro- pe/ecus lychnus, Borostomias panamensis, Stomias atriventer, Idiacanthus antrostomus, Bajacalifornia burragei, Talismania bifurcata, Holtbyrnia melanocephala, Normichthys campbelli, Pellisolus facilis, Scopelarchoides nicholsi, Diogenichthys laternatus, Gonichthys teiniiculus, Hygophum atratum, Lampanyctus parvicauda, Melamphaes acanthomas, Poromitra crassiceps, Scopeloberyx robustus, Scopelogadus mizolepis bispinosus. Kali normani, and Lepidopus xantusi. A system of water masses and currents, analagous to those in the north- east Pacific, exists along the west coast of South America. In this system, water of subantarctic origin flows through a transitional region where it is gradually warmed and mixed with subtropical water as it turns westward and becomes the South Equatorial Current. A tongue of warm, high salinity water, of low oxygen content indicates southward movement of subsurface equatorial water along the coast of Chile (Craddock and Mead, 1970). It is tempting to relate this southern extension of equatorial water to the presence of tropical species taken along the coast of South America (Bussing, 1965; Craddock and Mead, 1970). These species include Vinciguerria lucetia, Bathylagus nigrigenys, Pellisolus facilus, Idia- canthus antrostomus, Diogenichthys laternatus, Lampanyctus parvicauda, Melamphaes acanthomas, Melamphaes spinifer Scopeloberyx robustus, and Scopelogadus mizolepis bispinosus. As mentioned earlier, several of these species have been taken quite far north along the west coast of North 43 Contributions in Science No. 242 America. The northern and southern range limits of some of these forms seem to be far removed from their breeding area based on the paucity of juvenile fish in these areas, in which case they may very well represent expatriate populations. The number of widely distributed species continues to increase as mid- water trawl data from all parts of the world are published. Several species from the, present survey have been shown to have circumtropical or world- wide distributions. These include Serrivomer sector, Nemichthys scolopaceus, A vocettina infans, Cyclothone acclinidens, Cyclothone pallida, Diplophos taenia, Bolinichthys longipes, Taaningichthys bathyphilus, Scopelengys tristns, Holtbyrnia macrops, Bregmaceros atlanticus, Thaumatichthys pagidostonms, Dolopichthys pullatus, Melanocetus johnsoni, Centrophryne spinulosa, and Chiasmodon niger. A comparison of the Gulf of California midwater ichthyofauna with that of the adjacent Pacific is of interest. The faunas of the Gulf and outer coast of Baja California are indeed distinct (Berry and Perkins, 1966; present data). The differences may be attributed to water mass characteristics and currents in the two regions. However, faunal differences in the midwaters between the Gulf and the adjacent tropical Pacific are difficult to explain because the depths of the Gulf are filled with water which has identical characteristics with that of the entire eastern tropical Pacific. Furthermore, there is no shallow sill restricting fish movement into the Gulf. Nevertheless, some type of barrier is apparently present, as the following discussion will illustrate. Not a single individual of the genus Cyclothone was taken in the Gulf of California. This is remarkable because members of the genus are among the most abundant fishes in the world (Fitch and Lavenberg, 1968), and Robison (1972) captured them throughout the Gulf in summer and fall cruises of the Stanford Oceanographic Expeditions. Cyclothone spp. were taken at each of the 26 stations of cruise 106 1 (in November) south of the Gulf and across the Gulf entrance — a total of over 1 8,000 individuals and as many as 2,500 at a single station (Fig. 4). During cruise 922 (in January), only one individual of the genus was captured, despite the fact that several trawls were taken in the same area, at similar depths, across the Gulf entrance. Garman ( 1 899) recorded C. acclinidens in the Gulf at 27° N, but none were taken by Lavenberg and Fitch (1966) in their midwater trawl survey in March and April. It is difficult to explain this phenomonon on the basis of available data. The answer may, in part, lie in the poorly known fluctuating surface and subsurface currents in the Gulf. The movement of water in and out of the Gulf has been pictured by Roden (1962) as an outflow of water at the surface and an inflow at greater depths during the winter, with the reverse occurring during the summer. The life habits of Cyclothone [i.e. restricted vertical migration (De Witt, 1 972) and the timing of their life stages] may be out of phase with the fluctuating currents, which could exclude them from the Gulf Gulf of California Midwater Fishes 44 1973 during part of the year. In any case, more data are needed before any con- clusions can be drawn concerning this problem. A number of species appear restricted to waters south of the Gulf, or to waters south of the southernmost one-third of the Gulf. Nineteen midwater species captured in waters south of the Gulf were not taken in the Gulf north of 23 °N (Table 2). Only 7 midwater species taken in the Gulf were not captured at stations south of 23 °N. Twenty-five trawls taken within the Gulf resulted in over 28,000 individuals and an average of 13 species per trawl, while 35 trawls south of 23 °N also took over 28,000 individuals but with an average of 22 species per trawl. It must be realized that some 23,000 of the Gulf specimens were represented by a single species (Triphoturus mexicaniis) taken in a single trawl. These findings, which agree with those of Robison (1972), indicate a relatively depauperate midwater ichthyofauna in the Gulf as compared to that in waters to the south. The impoverishment of the Gulf midwater ichthyofauna can perhaps best be considered in relationship to the Gulf as a limiting environment. Despite the fact that the subsurface waters in the Gulf are similar to the eastern tropical Pacific waters, broadly fluctuating surface salinities and temperatures are present in the Gulf when compared to surface waters to the south (Roden, 1962; Wyrtki, 1967; Griffiths, 1968). Fluctuating surface waters combined with the remoteness of the Gulf (at higher and higher latitudes) to open ocean currents may, in part, explain the Gulf as a limiting environment to the epipelagic larvae of some midwater fishes. In conclusion, the Gulf of California and eastern tropical Pacific remain largely independent of the major current and water mass systems in adjacent areas. This independence is reflected in some unique hydrographical con- ditions and results in biotic features which will remain among the most intriguing of the world’s oceans. ACKNOWLEDGMENTS I thank Basil Nafpaktitis for his constructive and critical review of the manuscript, and Theodore Pietsch for his helpful suggestions and identifi- cation of all ceratioid material. Several other persons helped to initially sort and identify much of the material, and I thank Gerald Citek, Brent Davy, Robert Lavenberg, Richard McGinnis, Mary Nafpaktitis, Jerry Neumann, William O’Day, and Catherine Rainwater for their efforts. Resumen Este estudio faunistico esta basado sobre 56,000 especimenes, represen- tando de menos 1 13 especies las cuales fueron capturadas en una red Isaacs- Kidd Midwater Trawl de 10 pies de tamano, durante los cruceros del R/V Velero IV, University of Southern California, en el Golfo de California y el 45 Contributions in Science adyacente Pacffico oriental tropical durante 1967 y 1970. Aproximadamente*i 90% del mimero total de especimenes pertenecen a dos familias, Myctophidae m y Gonostomatidae, y la mayoria de estos a solo dos especies, Triphoturus |i mexicanus y Cyclothone acclinidens, respectivamente. ? La hidrografia del Pacffico oriental tropical es considerada en relacion | a su ictiofauna endemica de aguas de profundidad media y el papel del | Pacffico oriental tropical como un centro de origen y distribucion de peces, a lo largo de las costas occidentales de Norte y Sud America es discutida. Una comparacion do la fauna do Golfo de California con aquella a lo largo de las costa de Mexico se considera brevemente. i Literature Cited Ahlstrom, E. H. 1971. Kinds and abundance of fish larvae in the eastern tropical ! Pacific, based on collections made on EASTROPAC I. Fish. Bull. 69: 3-77. , AND R. C. Counts. 1958. Development and distribution of Vinciguerria | lucetia and related species in the eastern tropical Pacific. U.S. Fish Wildl. Serv., Fish. Bull. 58: 363-416. ' Alverson, F. G. 1961. Daylight surface occurrence of myctophid fishes off the i coast of Central America. Pac. Sci. 15: 483. Backus, R. H., J. E. Craddock, R. L. Haedrich, and D. L. Shores. 1970. The ; distribution of mesopelagic fishes in the equatorial and western north Atlantic Ocean. J. Mar. Res. 28: 179-201. Baird, R. C. 1971. The systematics, distribution, and zoogeography of the marine hatchetfishes (family Sternoptychidae). Bull. Mus. Comp. Zool., Harvard 142: 1-128. Barlow, G. W. 1961. Causes and significance of morphological variation in fishes. Syst. Zool. 10: 105-117. Beebe, W., and M. Vander Pyl. 1944. Eastern Pacific expeditions of the New York Zoological Society. XXXIII. Pacific Myctophidae (Fishes). Zoologica, N.Y. 29(9): 59-95. Berry, F. H., and H. C. Perkins. 1966. Survey of the pelagic fishes of the California Current area. U.S. Fish Wildl. Serv., Fish. Bull. 65(3): 625-682. Bertelsen, E. 1951. The ceratioid fishes, ontogeny, taxonomy, distribution and biology. Dana Rep. 39. 276 pp. Bieri, R. 1959. The distribution of the planktonic Chaetognatha in the Pacific and their relationship to the water masses. Limnol. Oceanogr. 4: 1-28. Borodulina, O. D. 1968. Taxonomy and distribution of the genus Leuroglossus (Bathylagidae, Pisces). Vopr. Ichthyol. 8: 1-10. Breder, C. M. 1936. Heterosomata to Pediculata from Panama to Lower California. Scientific results of the second oceanographic expedition of the Pawnee 1926. Bull. Bing. Oceanogr. Coll. 2(3): 1-56. Brinton, E. 1962. The distribution of the Pacific euphausiids. Bull. Scripps Inst. Oceanogr. 8: 5 1-270. Brock, V. C. 1938. Notes on the ranges of fishes from Lower California and the west coast of Mexico. Copeia 1938: 128-131. Bussing, W. A. 1965. Studies of the midwater fishes of the Peru-Chile Trench. In G. A. Llano (ed.). Biology of the Antarctic Seas II. Antarctic Res. Ser. 6. Nat. Acad. Sci., Nat. Res. Counc. Publ. 1297, pp. 185-227. Clemens, H. B., and J. C. Nowell. 1963. Fishes collected in the eastern Pacific during tuna cruises, 1952-1959. Calif. Fish and Game 49(4): 240-264. li 1973 Gulf of California Midwater Fishes 46 I Craddock, J. E., and G. E. Mead. 1970. Midwater fishes from the eastern South ^ Pacific Ocean. Scientific results of the southeast Pacific expedition. Anton j Bruun Rep. 3. 46 pp. i D’Ancona, U., and G. Cavinato. 1965. The fishes of the family Bregmacerotidae. Dana Rep. 64. 92 pp. DeWitt, E. a. 1972. Bathymetric distributions of two common deep sea fishes, Cyclothone acclinidens and C. signata, off southern California. Copeia 1972; 88-92. Ebeling, a. W. 1962. Melamphaidae I. Systematics and zoogeography of the species in the bathypelagic fish genus Melcunphaes Gunther. Dana Rep. 58. 164 pp. Eitch, J. E., and L. W. Barker. 1972. The fish family Moridae in the eastern North Pacific with notes on morid otoliths, caudal skeletons, and the fossil record. Nat. Mar. Fish. Serv., Fish. Bull. 70(3): 565-584. Fitch, J. E., and R. J. Lavenberg. 1968. Deep-water teleostean fishes of California. Univ. Calif. Press, Berkeley. 155 pp. Fowler, H. W. 1944. Results of the 5th George Vanderbilt Expedition (1941). Acad. Nat. Sci. Phila. Monogr. 6: 57-475. Garman, S. 1899. Reports on the exploration off the west coasts of Mexico, Central, and South America, and off the Galapagos Islands ... by the . . . Albatross. Mem. Mus. Comp. Zool., Harvard 24. 43 1 pp. Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Amer. Mus. Nat. Hist. Bull. 13(4): 341-455. Grey, M. 1960. A preliminary review of the family Gonostomatidae, with a key to- the genera and the description of a new species from the tropical Pacific. Bull. Mus. Comp. Zool., Harvard 122(2): 57-125. 1964. Family Gonostomiatidae. In fishes of the Western North Atlantic. Mem. Sears Found. Mar. Res. 1, pt. 4, pp. 78-240. Griffiths, R. C. 1968. Physical, chemical, and biological oceanography of the entrance to the Gulf of California, spring of 1960. U.S. Fish and Wildl. Serv. Spec. Sci. Rep., Fisheries 573. 47 pp. Haedrich, R. L., and M. Horn. 1969. A key to stromateioid fishes. Woods Hole Oceanogr. Inst., Tech. Rep. Ref. No. 69-70. Horn, M. 1970. Systematics and biology of the stromateid fishes of the genus Peprilus. Mus. Comp. Zool., Harvard 140 (5): 165-261. Isaacs, J. D., and L. W. Kidd. 1953. Isaacs-Kidd midwater trawl. Scripps Inst. Oceanogr., Ref. 53-3. 21 pp. Johnson, R. K. 1970. A new species of Diplophos (Salmoniformes: Gonostomatidae) from the western Pacific. Copeia 1970: 437-443. Lavenberg, R. J. 1965. A new species of searsiid fish, Nonnichthys camphelli, from the eastern North Pacific Ocean. Bull. So. Calif. Acad. Sci. 64(1): 22-26. , and a. W. Ebeling. 1967. Distribution of midwater fishes among deep water basins of the southern California shelf. In Proceedings of the Symposium on the Biology of the California Islands. R. N. Philbrick (ed.), pp. 185-201. , AND J. E. Fitch. 1966. Annotated list of fishes collected by midwater trawl in the Gulf of California, March-April 1964. Calif. Fish and Game 52(2): 92-1 10. McAllister, D. E., and E. I. S. Rees. 1964. A revision of the eelpout genus Melanostigmci with a new genus and with comments on Maynea. Bull. Mat. Mus. Canada 199: 85-109. McGowan, J. A. 1960. The relationship of the distribution of the planktonic worm, Poeohiiis meseres Heath, to the water masses of the North Pacific. Deep-sea Res. 6: 125-139. Meek, S. E., and S. F. Hildebrand. 1923-28. The marine fishes of Panama. Field Mus. Nat. Hist. Publ., Zool. Ser. 15, parts 1, 2, 3. 1045 pp. 47 Contributions in Science No. 242 Morrow, J. E. 1957. Studies in ichthyology and oceanography off coastal Peru. Mid-depth fishes of the Yale South American Expedition. Bull. Bing. Oceanogr Coll. 16(2): 56-71. 1961. Taxonomy of the deep sea fishes of the genus Cluiuliodiis. Bull. Mus. Comp. Zool., Harvard 25(9); 249-294. 1964. Family Chauliodontidae. In Fishes of the Western North Atlantic. Mem. Sears Found. Mar. Res. 1, pt. 4, pp. 274-289. Moser, H. G., and E. H. Ahlstrom. 1970. Development of lanternfishes (family Myctophidae) in the California Current. Part I. Species with narrow-eyed larvae. Bull. Los Angeles Co. Mus. Nat. Hist. 7. 145 pp. Norman, J. R. 1929. The teleostean fishes of the family Chiasmodontidae. Ann. Mag. Nat. Hist., Ser. 10, 3: 529-544. Nichols, J. T., and R. C. Murphy. 1944. A collection of fishes from the Panama Bight, Pacific Ocean. Bull. Amer. Mus. Nat. Hist. 83(4): 221-259. Parr, A. E. 1931. Deepsea fishes from the western coast of North America. Bull. Bing. Oceanogr. Coll. 2(4). 53 pp. 1937. Concluding report on fishes. Bull Bing. Oceanogr. Coll. 3(7). 79 pp. 1951. Preliminary revision of the Alepocephalidae, with the introduction of a new family, Searsidae. Amer. Mus. Novitates, 1531.21 pp. 1952. Revision of the species currently referred to Alepocephalns, Hali- sauriceps, Batliytroctes and Bajacalifornia, with introduction of two new genera. Bull. Mus. Comp. Zool., Harvard 107(4): 255-269. 1960. The fishes of the family Searsidae. Dana Rep. 51. 108 pp. PiETSCH, T. W. 1972a. A review of the monotypic deep-sea anglerfish family Centrophrynidae: Taxonomy, distribution, and osteology. Copeia 1: 17-47. 1972b. Ergebnisse der Forschungsreisen des FES “Walther Herwig” nach Siidamerika. XIX. Systematics and distribution of Ceratioid Fishes of the genus Dolopichthys (Family Oneirodidae), with the Description of a new species. Arch. Fisch Wiss. XXII (Heft 1): 1-28. , AND B. C. Nafpaktitis. 1971. A male Melanocetiis johnsoni attached to a female Centrophryne spiniilosa (Pisces: Ceratioidea). Copeia 2: 322-324. Robison, F. H. 1972. Distribution of the midwater fishes of the Gulf of California. Copeia 3: 448-461. Roden, G. I. 1962. Oceanographic aspects of the Gulf of California. In T. H. Van Andel and G. G. Shor (eds.). Marine Geology of the Gulf of California. Amer. Assoc. Petroleum Geol. pp. 30-58. Ricker, K. E. 1959. Mexican shore and pelagic fishes collected from Acapulco to Cape San Lucas during the 1957 cruise of the Marijean. Mus. Contrib. 3, Inst. Fish., Univ. Brit. Columbia. 18 pp. Route, L., and L. Bertin. 1929. Les poissons apodes appartenant au sous-ordre des nemichthydiformes. Dana Rep. 4. 1 13 pp. Seal, A. 1940-46. Report on fishes from the Allan Hancock Expedition in the California Academy of Sciences. Allan Hancock Found. Publ., Univ. So. Calif. 9: 1-46. SvERDURP, H. U., M. W. Johnson, and R. H. Fleming. 1942. The Oceans. Prentice- Hall, New York. 1087 pp. Townsend, C. H., and J. T. Nichols. 1925. Deep sea fishes of the Albatross Lower California Expedition. Bull. Amer. Mus. Nat. Hist. 52(1): 1-20. WiSNER, R. L. 1962. Midwater trawl surveys. In oceanographic studies during operation Wigmam. Limnol. and Oceanogr. Supp. 7. Wyrtki, K. 1967. Circulation and water masses in the eastern equatorial Pacific Ocean. J. Oceanol. and Limnol. 1(2): 1 17-147. Accepted for publication April 5, 1973 Printed in Los Angeles, California by Anderson, Ritchie and Simon on Simpson Lee RecoveruBL 5-0 l .13 C 2 /- ^ ^ ^ NUMBER 243 JUNE 14, 1973 THE SYSTEMATICS OF GASTEROSTEUS ACULEATUS (PISCES: GASTEROSTEIDAE) IN CENTRAL AND SOUTHERN CALIFORNIA By Stephen T. Ross t, CONTRIBUTIONS IN SCICNCE p , I LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- able footnotes must be typed on separate sheets. Several of one kind may be placed on a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing with Latin American subjects. Summaries in other languages are not required but are strongly recommended. Summaries will be published at the end of the paper. ( 8 ) A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor THE SYSTEMATICS OF GASTEROSTEUS ACULEATUS (PISCES: GASTEROSTEIDAE) IN CENTRAL AND SOUTHERN CALIFORNIAi By Stephen T. Ross^ Abstract: Morphological, breeding, and chromosome studies of the three principally allopatric forms of G. aculeatus: the anad- romous G. a. aculeatus and the freshwater G. a. microcephalus and G. a. williamsoni are presented. Emphasis is primarily on the two latter forms. Combined population samples of the williamsoni form are morphologically distinct from combined samples of microce- phalus with 96% joint nonoverlap of character index values. Recip- rocal crosses between G. a. williamsoni and G. a. microcephalus do not conclusively demonstrate postmating isolating mechanisms, although hybrid crosses show only 69% fertilization compared to 90% for the controls. There are no statistically significant differ- ences in mortality between control and test crosses for either em- bryos or larvae. Positive identification of laboratory reared F| hy- brids by plate number, spine length, or by a character index is not possible due to dominance of G. a. microcephalus. The value of plate counts in the determination of wild F| hybrids is thus ques- tionable. G. a. aculeatus, G. a. microcephalus, and G. a. william- soni all have a diploid chromosome number of 42. Retention of the subspecific nomenclature is favored. Introduction The threespine stickleback, Gasterosteus aculeatus Linnaeus, displays complex patterns of variation throughout its distribution. Populations of Gasterosteus on the Pacific Coast of North America have recently been sub- jected to considerable research by Hagen (1967), McPhail (1969), Miller and Hubbs (1969), Narver (1969), Hagen and McPhail (1970), and Hagen and Gil- bertson (1972). Hagen and McPhail discussed the extreme variability of many populations and presented and reviewed evidence calling for specific recogni- tion of G. trachurus and G. leiurus. In contrast, Miller and Hubbs (1969) sup- ported retention of the subspecies G. a. aculeatus, G. a. microcephalus, and G.a. williamsoni, as suggested by Jordan and Hubbs (1925). Most of the problems discussed by Hagen and McPhail pertain to stickleback populations in the Pacific Northwest. For the purpose of this investigation, centered in central and southern California, the nomenclature proposed by Jordan and Hubbs (1925) will be followed. However, as stated by Hagen and McPhail (1970), “A revision of the nomenclature may be necessary once the data are in hand.” This study attempts to clarify certain relationships of the central and southern California populations of sticklebacks. 'Review Committee eor this Contribution Carl L. Hubbs Robert J. Lavenberg Joseph S. Nelson 2Dept. of Biology, University of South Florida, Tampa, Florida 33620 1 2 Contributions in Science No. 243 Nomenclature and Distribution This section discusses the literature dealing with the systematics of G. a. williamsoni in southern California. First locality records from an area are cited, but additional records from the same area are omitted unless they conflict with the original identifications or contain additional information. References to G. a. microcephalus are restricted to major papers. A listing of the earlier liter- ature is given by Eigenmann (1886: 245) and Evermann and Clark (1931). Gasterosteus aculeatus williamsoni Girard Gasterosteus williamsoni Girard, 1854: 133. (original description; type locality, Williamson’s Pass, determined by Miller (1960) to be Soledad Canyon of the Santa Clara River, Los Angeles County, California). Eucalia williamsoni, Jordan and Copeland, 1876: 141. (identification doubtful as to genus; Williamson’s Pass, California). Gasterosteus williamsoni, Gilbert, 1893. (possible subspecific recogni- tion; San Bernardino County, California). Jordan and Evermann, 1896: 750. (characters; abundant in southern California). Gasterosteus williamsoni williamsoni, Rutter, 1896: 245. (survey of Pacific slope populations; “True williamsonE from Santa Ana basin at Col- ton, Riverside and San Bernardino, San Bernardino County, California. Also eight type specimens from Williamson’s Pass, Santa Clara River). Gasterosteus santaeannae. Regan, 1909: 345. (original description; type locality, Santa Ana River, Colton, California). Gasterosteus cataphractus, Snyder, 1913: 17. (characters; examination of Rutter’s (1896) data). Gasterosteus aculeatus williamsoni, Jordan and Hubbs, 1925: 200. (syn- onymy — cataphractus - aculeatus; santaeannae - williamsoni; Santa Ana drainage in southern California). Hubbs, 1929: 4. (plate II, figure 2). Miller, 1939: 168. (recorded from San Juan Creek and Arroyo Trabuco, Orange County, California). Miller, 1960: 350. (characters; San Juan Creek and Ar- royo Trabuco samples do not represent unarmored form, also recorded from Alamo Creek and Huasna River in Santa Maria River drainage, Santa Bar- bara County, California). Miller, 1961: 388. (Los Angeles River, San Gabriel River and Mohave River (introduced)). Miller and Hubbs, 1969: 52. (charac- ters, plate counts for southern California populations). Distribution. Santa Clara River in Soledad Canyon, Santa Ana^, San GabrieP, Los Angeles^, and Mohave rivers, and Alamo Creek and the Huasna River (Santa Maria River drainage). Miller and Hubbs (1969) reported G. a. microcephalus x williamsoni intergrades introduced in the Mohave River, and G. a. microcephalus introduced into Alamo Creek and the Huasna River. ^Probably extirpated between 1942-1961 (Miller, 1969) 1973 Systematics of GASTEROSTEUS ACULEATUS 3 Gasterosteus aculeatus microcephalus Girard Gasterosteus microcephalus Girard, 1854; 133. (original description; type locality, Four Creek (Kaweah River), San Joaquin Valley, California). Eigen- mann, 1886: 245. (characters; distribution from Bering Strait to Todos Santos Bay). Jordan, 1894: 141. (San Luis Obispo County). Gasterosteus williamsoni microcephalus, Rutter, 1896: 245. (characters; list of synonyms; distribution from Unalaska to Santa Clara River in southern California). Jordan and Evermann, 1896: 751. (characters; distribution, Alaska to Todos Santos Bay). Gasterosteus cataphractus, Jordan and Gilbert, 1899: 443. (intergrades into fully plated form and thus lacks characters of a subspecies). Evermann and Goldsborough, 1906: 273. (distribution in Alaska). Snyder, 1913: 71. (distribution: streams tributary to Monterey Bay). Gasterosteus aculeatus, Regan, 1909: 435. (characters; synonymy of European and American forms). Gasterosteus aculeatus microcephalus, Jordan and Hubbs, 1925: 202. (distribution: California, Aleutian Islands and Japan). Miller and Hubbs, 1969. (characters; plate counts for Pacific Coast populations). Gasterosteus aculeatus, Hagen, 1967: 1637. (characters and isolating mechanisms of British Columbia populations). Gasterosteus leiurus, Hagen and McPhail, 1970: 147. (discussion of isolating mechanisms). Distribution. Freshwater of Pacific Coast of North America from Bering Strait (Eigenmann, 1886) to Arroyo del Rosario, Baja California (Hubbs, in Follett, 1960), with a hiatus in its range between the Santa Clara River and San Juan Creek (Miller and Hubbs, 1969: 63). Material Examined The localities, numbers of specimens, dates of collection, and museum numbers are given below (see also Fig. 1). Material without museum numbers is at California State College, Fullerton. Santa Cruz County: San Lorenzo River (5), 10 July, 1957, California Academy of Sciences 26738; San Lorenzo River (10), 3 April, 1969, Los Angeles County Museum, LACM 30905-1. Monterey County: Carmel River (30), 17 June, 1966, LACM 30889-1; Big Sur River (15), 19 June, 1966, LACM 30907-1; San Antonio River, Hunter- Liggett Military Reservation, (25), 25 February, 1968; Nacimiento River, below Nacimiento Dam (11), 25 February, 1968. San Luis Obispo County: Toro Creek (12), 2 July, 1968, LACM 30916-1; Morro Creek (30), 26 Decem- ber, 1967, LACM 30888-1; San Luis Obispo Creek, in San Luis Obispo, (25), 26 December, 1967, LACM 30909-1; San Luis Obispo Creek (11), 21 January, 1968, LACM 30927-1; San Luis Obispo Creek (23), 25 February, 1968, LACM 30923-1; San Luis Obispo Creek (25), 27 June, 1968, LACM 30894-1; Price Creek (29), 22 June, 1968, LACM 30938-1; Arroyo Grande Creek (30), 20 Aug- ust, 1967, LACM 30904-1; Alamo Creek (10), 9 December, 1967, LACM 30897- 1; Alamo Creek (17), 25 February, 1968, LACM 30932-1. Santa Barbara 4 Contributions in Science No. 243 County: Salsipuedes Creek (27), 20 August, 1967, LACM 30911-1. Ventura County: Ventura River at Foster Park (30), 15 September, 1967; Piru Creek, below Piru Dam (5), 15 September, 1967, LACM 30908-1; Santa Clara River near the Los Angeles- Ventura County line (25), 8 July, 1968, LACM 30937-1. Los Angeles County: Santa Clara River in Soledad Canyon, west of Bee Can- yon (44), 15 September, 1967, LACM 30892-1; Santa Clara River, Soledad Canyon (25), 27 June, 1968, LACM 30925-1. San Bernardino County: Mohave River, at Las Flores junction. Summit Valley (30), 18 July, 1966, LACM 30890-1. Orange County: Arroyo Trabuco, near Mission San Juan Capistrano (30), 15 January, 1968. Experimental studies were conducted with fish from Soledad Canyon of the Santa Clara River, Los Angeles County (the type locality of G. a. william- soni), and San Luis Obispo Creek, San Luis Obispo County. Also, a male G. a. aculeatus from Scott Creek, Santa Cruz County, was utilized in breeding studies and individuals of G. a. aculeatus from the San Lorenzo River, Santa Cruz County, were karyotyped. Methods Meristic characters studied were the numbers of lateral plates, gill rakers, and vertebrae. Dorsal, anal, and pectoral rays were not counted since Heuts Figure 1. Coastal drainages in central and southern California. The collecting sites are indicated by small circles. 1973 Systematics of GASTEROSTEUS ACULEATUS 5 (1949), Lindsey (1962), and Hagen (1967) have shown fin rays to be influ- enced greatly by water temperature. Dial calipers read to the nearest 0.1 mm were used in obtaining all morphometric data from preserved specimens. Magnification was used whenever necessary, generally with specimens under 25 mm SL and with larger G. a. williamsoni in determining spine length. Measuring and counting techniques followed Hubbs and Lagler (1958) with the following modifications. All lateral plates were counted, including those in the peduncular keel (when present), by using an air jet and a dissect- ing microscope. Small individuals were first stained with alizarin to aid plate enumeration. All gill rakers on the first gill arch were counted; determina- tion of rudimentary rakers was aided by staining the arch (in situ) with an aqueous solution of methylene blue (1:5,000). Vertebral counts (determined from radiographs) included the urostyle and the expanded hypural plate as one, and the first vertebra was regarded as the one to which the first neural spine attached. Orbit diameter was measured between the anterior and pos- terior bony margins. Body depth was measured anterior to the pelvic spines to avoid distortion in gravid females. Body width was measured anterior to the pectoral fins. Dorsal and pelvic spine lengths were taken along the anterior surfaces. Pubic-plate length (ventral shield. Nelson, 1971) was measured from a point between the anterior face of the insertions of the erected pelvic spines and the end of ossification. All morphometric data are presented as pro- portions with standard length as the numerator. Peduncular depth, first and second dorsal spine lengths, pelvic-spine length, pubic-plate length, and the number of lateral plates and gill rakers were combined in a character index using the following procedure. The ranges of the proportions formed from the three spine lengths and pubic-plate length were subdivided into 20 equal classes which were scored from 0 to 10 with increments of 0.5. A score of zero was reserved for situations in which the character was missing, but zero values were not used in determining the index which is the mean of the scored characters. Character states typical of G. a. williamsoni received the lowest scores, while character states typical of G. a. aculeatus received the highest scores. Lateral-plate number was not divided into 20 equal classes due to the extreme range. Counts of zero to seven, such as characterized freshwater populations of Gasterosteus, were accorded individual classes; thereafter, two or three plate counts made up a class. Ranges in plate counts in which no recorded individuals fell were lumped in groups of three. Scoring of classes was the same as described previously. Reciprocal crosses between G. a. microcephalus and G. a. williamsoni were incubated at 5, 10, 12, 15, 20, 25, and 30° C. Late in the study a female G. a. williamsoni was crossed with a male G. a. aculeatus at 20° C. Artificial fertilization was produced following the technique of Hagen (1967). The four masses of fertilized eggs resulting from the two reciprocal crosses were each washed onto a platform of fiberglass screen supported by a ring of surgical tubing. Each platform was placed in one of four 1.1 liter 6 Contributions in Science No. 243 jars which extended 6 cm above the water in a 76 liter tank. Water was air- lifted from the large tank to a reservoir 8 cm above the water and then returned through 1.1 liter jars to the main tank via siphons. Complete exchange of water in the 1.1 liter jars occurred every 12 minutes. The jars were removable so mat eggs and larvae could be counted by placing the jar over a grid. Eggs were kept in darkness prior to hatching and then received an irregular long- day photoperiod. Infusoria tablets were introduced three days after hatching, and after 10 to 12 days the larvae were fed washed nauplii of Artemia salina. Sticklebacks of 10 mm and larger were fed live immature and adult A. salina, adult frozen A. salina, and commercial dried food. Fish were kept in the con- trolled temperature systems for a minum of five months at which time the survivors were placed in 19 or 38 liter aquaria. After seven to nine months fish were fixed in 10% formalin for one to two weeks, washed and placed in 40% isopropanol. Only laboratory reared sticklebacks larger than 20 mm SL were utilized. Characters studied were plate number, first and second dorsal and pelvic spine lengths, and pubic-plate length. Gill rakers were omitted due to the large amount of overlap between natural populations of G. a. williamsoni and G. a. microcephalus, and the depth of the caudal peduncle was omitted since it was not useful in separating hybrids and controls. The term “hybrid,” as used in this paper, refers to crosses between different populations and does not imply an interspecific cross. Temporary chromosome preparations were made following McPhail and Jones (1966), and permanent slides were made with an air drying method modified from Ojima et al. (1964), using the same pretreatment. Gill arches were fixed in 3:1 ethanol-acetic acid (20 minutes), brushed across a micro- scope slide and dried over an alcohol flame. The dried preparation was flooded with 3% aceto-orcein for five minutes, rinsed in distilled water, dried, and mounted with diaphane. Chromosomes were counted from photographic enlargements. Results Morphological Studies Statistical comparisons of lateral-plate numbers for 18 stickleback pop- ulations using Tukey’s w' procedure (Steele and Torrie, 1960) show three distinct groups (Fig. 2). The large middle group shows considerable intra- group overlap, and the variation follows a north-south dine only approxi- mately. The three groups correspond with the subspecies G. a. aculeatus, G. a. microcephalus and G. a. williamsoni as defined by Miller and Hubbs (1969); however, the Arroyo Trabuco population will be shown to tend less toward G. a. microcephalus in certain other characters. Gill rakers differ significantly in number between the 18 sample popu- lations (p <0.005), and show an irregular decrease from north to south (Table 1). However, Tukey’s w' indicates a high degree of overlap between the means so that a statistical separation into taxonomically useful groups can- 1973 Systematics of GASTEROSTEUS ACULEATUS 7 SAN LORENZO RIVER BIG SUR RIVER TORO CREEK CARMEL RIVER MORRO CREEK ARROYO GRANDE CREEK PRICE CREEK SALSIPUEDES CREEK VENTURA RIVER NACIMIENTO RIVER SAN ANTONIO RIVER SAN LUIS OBISPO CREEK ARROYO TRABUCO ALAMO CREEK PIRU CREEK SANTA CLARA RIVER 2 MOHAVE RIVER SANTA CLARA RIVER I 14 30.5 15 8.3 12 7.1 30 6.5 30 6.5 30 5.4 29 5.2 25 5.2 30 4.9 10 4.9 25 4.7 84 4.6 30 3.5 27 1.0 7 1.0 25 0.4 30 0.4 69 0.2 Figure 2. Sample sizes and ranked means of lateral-plate numbers from 18 stickleback populations. The means connected by the same line are not significantly different according to Kramers’s (1956) modification of Tukey’s w' multiple range test (a = 0.05). 1. Santa Clara River, Soledad Canyon. 2. Santa Clara River, Ventura County. not be made. It should be noted that the use of multiple range tests modified for unequal sample sizes results in an increased probability of a significant difference occurring within a subset classified as homogeneous (Kramer, 1956). Vertebral numbers from 12 sample populations do not show a definite geographic trend nor a trend congruent to variation in other characters studied (range 29-33). Orbit diameter does not vary significantly among the populations sampled. However, the Santa Clara River, Salsipuedes, and Arroyo Grande creeks, and the San Antonio River samples display a tendency towards greater orbit size in some specimens. The variation in body depth also does not show statistic- ally significant differences. The ratio of standard length to body width decreases irregularly from north to south (Fig. 3a). Sample populations from the San Lorenzo, Carmel, and Big Sur rivers, and Toro, Morro, and Salispuedes creeks have more com- pressed bodies (x = 7.2; sd = 0.58). Plateless sticklebacks from the Santa Clara and Mohave rivers, and Alamo Creek have the roundest bodies (x = 6.61; sd = 0.59), but overlap is extreme with the southern partially plated sticklebacks. The ratio of standard length to peduncular depth generally decreases from north to south although Nacimiento and San Antonio river samples have deeper caudal peduncles in comparison with sticklebacks of adjacent latitudes 8 Contributions in Science No. 243 Counts of gill rakers Table 1 in different populations of Gasterosteus aculeatus from southern and central California Locality 12 13 14 15 16 17 18 19 20 21 22 23 24 25 N X San Lorenzo River 2 2 3 3 5 15 20.5 Carmel River 3 3 7 13 4 30 20.3 Big Sur River 4 4 5 1 1 15 20.1 San Antonio River 3 2 7 6 6 1 25 18.5 Nacimiento River 7 1 2 10 19.7 Toro Creek 1 2 4 5 12 20.1 Morro Creek 3 9 4 11 2 1 30 20.1 San Luis Obispo Cr. 1 4 8 25 21 14 4 4 1 1 83 19.9 Price Creek 6 11 5 6 1 29 19.5 Arroyo Grande Creek 1 3 10 12 4 30 19.4 Alamo Creek 1 3 7 6 7 3 27 17.9 Salsipuedes Creek 1 3 9 5 2 3 2 25 19.8 Ventura River 1 7 7 9 5 1 30 17.3 Piru Creek 2 1 1 4 17.0 Santa Clara River, Soledad Canyon 3 4 2 18 19 14 8 1 69 16.8 Santa Clara River, Ventura County 1 5 4 11 4 25 17.5 Mohave River 1 2 2 6 13 1 3 2 30 16.8 Arroyo Trabuco 3 3 7 11 3 3 30 18.6 (Fig. 3b). This is also true of the Arroyo Grande sample. The Salsipuedes Creek sticklebacks have more slender caudal peduncles, but overlap is exten- sive with the adjacent populations. The ratio of standard length to first dorsal spine and standard length to second dorsal spine show similar patterns of geographic variation (Fig. 3c, d). Samples from the San Lorenzo, Carmel, and Big Sur rivers, and Toro, Morro, San Luis Obispo, Arroyo Grande, Price, and Salsipuedes creeks, and the Ventura River have longer dorsal spines than other samples (I: x = 11.7; sd = 1.78; II: X = 10.1; sd = 1.43). Sticklebacks from Alamo Creek, the Santa Clara and Mohave rivers, and Arroyo Trabuco have the shortest dorsal spines (I: X = 19.1; sd = 3.82; II: x = 15.6; sd = 2.77). An intergroup separation with 91% joint nonoverlap is achieved with either spine . The San Antonio and Naci- miento river samples are intermediate between the two groups (I: x = 14.7; sd = 1.91; II: x = 12.9; sd = 1.62). The extremely short-spined Mohave River sample is statistically distinct from the Soledad Canyon sticklebacks 1973 Systematics of GASTEROSTEUS ACULEATUS 9 SAN LORENZO R. CARMEL R. BIG SUR R. NACIMIENTO R. SAN ANTONIO R. TORO CR. MORRO CR. SAN LUIS OBISPO CR. PRICE CR. ARROYO GRANDE CR. ALAMO CR. SALSIPUEDES CR. VENTURA R. PIRU CR.’ SANTA CLARA R. I SANTA CLARA R. 2 MOHAVE R. ARROYO TRABUCO A 5 6 7 8 9 10 S. LENGTH 12 SAN LORENZO R. CARMEL R. BIG SUR R. NACIMIENTO R. SAN ANTONIO R. TORO CR. MORRO CR. SAN LUIS OBISPO CR. PRICE CR. ARROYO GRANDE CR ALAMO CR. SALSIPUEDES CR. VENTURA R. PIRU CR. SANTA CLARA R. I SANTA CLARA R. 2 MOHAVE R. ARROYO TRABUCO B 13 15 17 19 21 23 25 S. LENGTH BODY WIDTH CAUDAL PEDUNCLE SAN LORENZO R. CARMEL R. BIG SUR R. NACIMIENTO R. SAN ANTONIO R. TORO CR. MORRO CR. SAN LUIS OBISPO CR. PRICE CR. ARROYO GRANDE CR. ALAMO CR. SALSIPUEDES CR. VENTURA R. PIRU CR. SANTA CLARA R. I SANTA CLARA R. 2 MOHAVE R. ARROYO TRABUCO c 8 12 16 20 24 28 32 36 S. LENGTH SAN LORENZO R. CARMEL R. BIG SUR R. NACIMIENTO R. SAN ANTONIO R. TORO CR. MORRO CR. SAN LUIS OBISPO CR. PRICE CR, ARROYO GRANDE CR. ALAMO CR. SALSIPUEDES CR. VENTURA R. PIRU CR. SANTA CLARA R. I SANTA CLARA R. 2 MOHAVE R. ARROYO TRABUCO _ 7 9 II 13 15 17 19 21 23 25 27 O a LENGTH FIRST DORSAL SPINE SECOND DORSAL SPINE Figure 3. Morphometric variation of populations of Gasterosteus aculeatus. The dia- grams indicate the mean (vertical lines), two standard errors on either side of the mean (closed rectangles), 1.5 standard deviations on either side of the mean (open rectangles), and the range (horizontal line). (p <0.001), although overlap is high. Variability for dorsal-spine lengths is extreme (CV=21-23). The ratio of standard length to pelvic-spine length also distinguishes the same two groups listed above, with 92% joint nonoverlap (Fig. 4a). The San Antonio and Nacimiento river samples again show intermediacy although the former is closer to the long-spined group than the latter. The Big Sur sample (x = 8.5) is also less extreme in spine length, possibly due to the small sample size (15), and to the low mean standard length of the series. In specimens smaller than 19 mm there is a decrease in pelvic-spine length relative to stan- dard length (Fig. 5a). The coefficients of variation are quite high for the Big 10 Contributions in Science No. 243 SAN LORENZO R. CARMEL R. BIG SUR R. NACIMIENTO R. SAN ANTONIO R. TORO OR. MORRO CR. SAN LUIS OBISPO CR. PRICE CR. ARROYO GRANDE CR. ALAMO CR. SALSIPUEDES CR. VENTURA R. PIRU CR. SANTA CLARA R. I SANTA CLARA R. 2 MOHAVE R. ARROYO TRABUCO L-tL r SAN LORENZO R. CARMEL R. BIG SUR R. NACIMIENTO R. SAN ANTONIO R. TORO CR. MORRO CR. SAN LUIS OBISPO CR. PRICE CR. ARROYO GRANDE CR. ALAMO CR. SALSIPUEDES CR. VENTURA R. PIRU CR. SANTA CLARA R. I SANTA CLARA R. 2 MOHAVE R. ARROYO TRABUCO Ltl 5 7 9 II 13 15 17 19 21 23 A S. LENGTH B 5 7 9 II 13 15 17 S. LENGTH PELVIC SPINE PUBIC PLATE SAN LORENZO R. CARMEL R. BIG SUR R. NACIMIENTO R. SAN ANTONIO R. TORO CR. MORRO CR. SAN LUIS OBISPO CR. PRICE CR. ARROYO GRANDE CR. ALAMO CR. SALSIPUEDES CR. VENTURA R. PIRU CR. SANTA CLARA R. I SANTA CLARA R. 2 MOHAVE R. ARROYO TRABUCO c xte I B I cBo 3 4 5 6 7 6 CHARACTER INDEX Figure 4. Morphometric variation and character index values of populations of Gaster- osteus aculeatus. See legend for figure 3 for explanation of diagrams, and text for ex- planation of character index. n lii 5'' / Sur sample (18.2), the samples from Alamo Creek (20.0), Soledad Canyon (19.0), and the Mohave River (22.5). Variation in the three latter populations ‘ is not a direct reflection of small size since almost all specimens were \ adults. A? Variation in the ratio standard length to pubic-plate length (Fig. 4b) is f quite similar to that shown by the dorsal and pelvic spines. The Big Sur | sample shows high variability (CV = 32.0), which is again a manifestation of the small specimens and sample size. A decrease in pubic-plate length rela- tive to standard length occurs between 20 and 22 mm (Fig. 5b). The San | Antonio and Nacimiento river samples are not clearly separable from the | 1973 Systematics of GASTEROSTEUS ACULEATUS 11 A z z % 12 u I 16_ »- ■ ■ ■ • • O 0 '* ','jr B “ 12_ 8_ ■ ■lO SUK R. • CARMIL R. O RRICf CR. ■ ■'AC •m ooa o* o • 10 ±J 14 LI- 18 22 1 1 ■ I 1 26 30 34 ,li-i 1 1 1 38 41 44 1 ‘ 1 1 1 ‘ 50 U-j 54 .L STANDARD LENGTH Figure 5. Pelvic-spine length (A), and pubic-plate length (B), vs. standard length. long-spined group by pubic-plate length. The short-spined group, composed of the Alamo and Piru creek, Santa Clara and Mohave river, and Arroyo Tra- buco samples, shows varying degrees of overlap with the northern group, with the latter sample being the most extreme. The character index delineates three principal groups, the most northern of which is composed only of the San Lorenzo River sample (Fig. 4c). The second group includes samples from Carmel, Big Sur, Nacimiento, and San Antonio rivers, and Toro, Morro, San Luis Obispo, Arroyo Grande, Price, and Salsipuedes creeks, and the Ventura River (range 5. 0-7. 6). The San Lorenzo sample shows 95% joint nonoverlap with the Carmel River sample, the north- ernmost location of the second group. There is a general trend towards a lower character index value in the more southerly populations. The Nacimiento and San Antonio river samples are most extreme in reduction of the index value but still are separable with 89% joint nonoverlap from Arroyo Trabuco. Stickle- backs from Alamo Creek and the Santa Clara and Mohave rivers comprise a third group which is distinct from the partially plated stickleback populations by well over 96% joint nonoverlap (CD = 1.97). The Arroyo Trabuco sticklebacks, when tested with Kramer’s (1956) modification of Tukey’s w' multiple range test using 18 sample means (Steele and Torrie, 1960), are significantly different from all other southern popula- tions of plateless and partially plated sticklebacks. However, figure 4c indi- cates a closer affinity with the G. a. williamsoni populations in comparison with G. a. microcephalus populations. Alamo Creek and the lower Santa Clara River sticklebacks are not significantly different when tested by the above 12 Contributions in Science No. 243 Table 2 Percent of fertilized eggs for reciprocal crosses between Gasterosteus aculeatus williamsoni and G. a. microcephalus MALES microcephalus williamsoni p(diag.) p(rows) 0.80 microcephalus X=91 sd=0.09 X=71 sd=0.34 0.05 FEMALES williamsoni X=67 sd=0.29 X=88 sd=:0.22 0.60 0.20 Combined control crosses X=90 sd=0.15 p<0.01 Combined test crosses X=69 sd=0.31 multiple range test, but they differ significantly from both the upper Santa Clara River and the Mohave River populations. Experimental Investigations The first postmating isolating mechanism, gamete incompatibility, was investigated by comparing the percent of eggs fertilized from each control and test cross. Control crosses showed higher mean fertilization values than the hybrid crosses (Table 2). Crosses using eggs from G. a. microcephalus females showed greater percent fertilization than the respective control and test crosses using females of G. a. williamsoni. Subsequent to fertilization, there were no statistically significant differences between control and hybrid crosses in embryo and larval mortality; t for paired comparisons .65 and .78 respectively (Table 3). Table 3 Summary of embryo- and larval mortality for reciprocal crosses between Gasterosteus aculeatus microcephalus and G. a. williamsoni. The results from 12° to 20° C incubation temperatures are included. The number of trials is shown in parentheses Fw. Mw. Fw. Mm. Em. Mm. Em. Mw. Total number hatched 136 131 362 304 Total number fertilized 158 (2) 191 (2) 491 (5) 405 (5) Percent mortality 13.9 31.4 26.3 24.9 Surv. to one week 125 118 315 242 Percent mortality 8.0 10.0 13.0 20.4 Frequency Frequency 1973 Systematics of GASTEROSTEUS ACULEA TVS 13 01 2345678 Plate Number Figure 6. A comparison of lateral plate numbers of wild populations of Gasterosteus aculeatus microcephalus and G. a. williamsoni with laboratory reared controls and hy- brids. The angle of hatching indicates the form of the female parent. G. a. williamsoni is shown by a positive slope, and G. a. microcephalus by a negative slope. 14 Contributions in Science No. 243 CHARACTER INDEX Figure 7. A comparison of character index values in wild populations of Gasterosteus aculeatus microcephalus and G. a. williamsoni with laboratory reared controls and hybrids. See legend for figure 6 for explanation of hatching. A cross between a female G. a. williamsoni and a male G. a. aculeatus from Scott Creek, Santa Cruz County, California, showed 88.3% fertilization for the control and 50% fertilization for the test cross. No decrease in embryo or larval viability occurred, and the test cross was, if anything, more vigorous than the control. All crosses had high embryo mortality at and below 10°C, and above 25° C, with the least mortality occurring between 15 and 20° C. Larval mor- tality of offspring from G. a. microcephalus females was lowest at 10° C and gradually increased with temperature from that point. Offspring from 1973 Systematics of GASTEROSTEUS ACULEATUS 15 G. a. williamsoni females remained fairly constant in percent mortality from 15 to 25° C. Both hybrid plate counts and character index values indicate partial dominance of G. a. microcephalus over G. a. williamsoni (Figs. 6 and 7). Hybrid individuals cannot all be distinguished morphologically from parental controls although the character index is slightly more reliable than plate number. Chromosome preparations were obtained from five G. a. williamsoni, six G. a. microcephalus, and two G. a. aculeatus. The diploid number in all three forms is 42 and morphology does not appear to differ from the karyotype of G. aculeatus published by Chen and Reisman (1970). Discussion Three principally allopatric forms of G. aculeatus exist in central and southern California. Both Jordan and Hubbs (1925) and Miller and Hubbs (1969) felt the differences warranted subspecific recognition. The character index described in this paper allows morphological separation of the two southernmost forms with 96% joint nonoverlap. This is well above the arbitrary 90% suggested by Mayr (1969) for distinguishing subspecies. How- ever, several populations show some degree of intermediacy. G. a. microcephalus from the Nacimiento and San Antonio rivers resemble the more southern G. a. microcephalus populations in most of the characters studied. The number of river miles from the Pacific Ocean is much greater for these streams compared to other locations sampled, thus a reduc- tion of gene flow with the coastal populations of G. a. microcephalus or with occasional stray aculeatus-microcephalus intergrades (“semi-armatus”) is likely. This isolation is now enhanced by the frequent lack of water in the Salinas River during the summer months. Natural selection must also play an important role to explain the diver- gence from the typical G. a. microcephalus phenotype of a similar latitude, assuming that the headwater populations are derived from sticklebacks on the coastal plain. If no selection were occurring, or if the characters studied were selectively neutral, one would not expect such a pronounced difference between the coastal and inland forms. Although the selective importance of predation has been demonstrated by several authors (e.g., Hoogland, Morris and Tinbergen, 1957; McPhail, 1969; Hagen and McPhail, 1970; and Hagen and Gilbertson, 1972), it is of questionable value in this instance. Ptycho- cheilus grandis, Salmo gairdneri, and Cottus asper were collected with gaster- osteus in the Salinas headwaters. All three species are potential predators on Gasterosteus. Cottus was observed feeding on G. a. williamsoni in the labor- atory. Cottus asper and C. aleuticus are also abundant in coastal streams from San Luis Obispo County northward. Consequently there is no apparent reduction in predation for the headwater populations of sticklebacks. Further south. Miller and Hubbs (1969) listed intergrades from the Ven- 16 Contributions in Science No. 243 tura River by citing Rutter (1896: 248), but commented (p. 62) that their counts from the Ventura River and a tributary, “pertain to G. a. microcepha- lus"' Rutter did not examine sticklebacks from the Ventura River. However, he did study 298 specimens from the Santa Clara River in Ventura County which have plate counts similar to those presented by Miller and Hubbs for the region near Fillmore. The Ventura River sticklebacks thus pertain clearly to G. a. microcephalus, although it is interesting that the gill raker counts are quite low compared to other microcephalus populations. G. a. microcephalus also occurs below the Los Angeles plain in southern Orange County, northern San Diego County and northwestern Baja California. Plate numbers gradually increase for populations in northern San Diego County and in northwest Baja California (Miller and Hubbs, 1969). Miller and Hubbs (1969) presented evidence for the introduce of G. a. microcephalus into Alamo Creek in 1940. Biological evidence of introduc- tion is the increased variation in lateral-plate numbers, and also a tendency towards longer pelvic spines and pubic plates. The Santa Clara River sample from the Los Angeles- Ventura County | line also tends towards G. a. microcephalus. Lateral-plate counts of specimens { collected near Fillmore (Miller and Hubbs, 1969) indicate an even greater shift towards G. a. microcephalus. The most obvious explanation is gene exchange with G. a. microcephalus. Both G. a. williamsoni and supposed G. a. microcephalus x williamsoni intergrades have been introduced into the Mohave River (Miller and Hubbs, 1969). The high morphological variability of the Mohave sample is likely due both to the combined gene pools and to the breakdown of normalizing selec- tion accompanying invasion of a new habitat. Sticklebacks were not pre- viously found in the Mohave River, an endorheic stream (Hubbs and Miller, 1948). Sticklebacks from Arroyo Trabuco, a tributary of San Juan Creek, also present taxonomic problems. Miller (1939) referred the San Juan Creek stickle- backs to G. a. williamsoni then later retracted his decision without assigning the form to G. a. microcephalus (Miller, 1960). Miller and Hubbs (1969) placed the form with G. a. microcephalus but suggested that a case could be made for calling it an intergrade. The character index suggests a closer affinity with G. a. williamsoni, but individual characters such as plate number, gill rakers, and pubic-plate length are intermediate. In the absence of a more complete study, it seems best to treat the Arroyo Trabuco-San Juan Creek sticklebacks as G. a. williamsoni x microcephalus, as suggested by Miller and Hubbs (1969). Miller and Hubbs interpreted the morphology of the San Juan Creek and Arroyo Trabuco sticklebacks, as well as other partially plated stickle- backs further south, to southward movement of G. a. microcephalus during a glaciation. However, many features of the G. a. microcephalus phenotype are absent or reduced in the Arroyo Trabuco population. Consequently, even if intergradation did occur, gene frequencies have been markedly altered by 1973 Systematics of GASTEROSTEUS ACULEATUS 17 selective forces and/or dilution by backcrossing with the native form. The great importance of selective pressure on an understanding of character differ- ences in Gasterosteus has been shown most recently by Hagen and Gilbert- son (1972). A potential selective force may be the presence of Fundulus par- vipinnis in San Juan and Arroyo Trabuco creeks (Miller, 1939), although direct predation, other than on eggs and larvae, seems unlikely. The extirpa- tion of G. a. williamsoni from most of its former range has been tentatively linked by Miller (1961) to the introduction of Gambusia afjinis. It is interesting that the southernmost distribution of Cottus asper (Hubbs, 1921; McPhail and Lindsey, 1970) corresponds with the southern- most continuous distribution of G. a. microcephalus in the Ventura River. Both C. asper and C. aleuticus are abundant in coastal streams from San Luis Obispo County northward and could act as potential predators on stickle- backs. C. asper was observed feeding on G. a. williamsoni in the laboratory. South of the Ventura River the streams are generally without piscivorous fishes, with the exception of exotic salmonids, recently introduced centrarch- ids, and Gambusia affinis. The only potentially piscivorous fish listed by Culver and Hubbs (1917) for the Santa Ana system is Salmo. However, fluvia- tile rainbow trout prefer swift currents (Lewis, 1969), while Gasterosteus shows a preference for slower water. Salmo is not found in Soledad Canyon of the Santa Clara River. South of the hiatus in the distribution of G. a. micro- cephalus are several fishes, listed by Follett (1960), which could act as preda- tors on sticklebacks. It is suggested that absence or reduction of predation may be a significant factor in the evolution of the williamsoni form. Clearly, the role of differential predation in the regulation of gene frequencies in stickleback populations of southern California needs to be investigated further. No potential postmating isolating mechanisms between G. a. microcepha- lus and G. a. williamsoni were substantiated in the laboratory investigations, with the exception of a difference in percentage of fertilization. Field studies have shown that the breeding seasons overlap almost completely. Assuming a lack of preferential mating, the percentage difference in fertilization between the control and test crosses would not prevent gene flow between the forms, but would decrease the rate of exchange. However, if contact between the two forms were broken following an initial invasion, the percent difference in fertilization could constitute a significant selective force, reducing the fre- quency of the invading form. Selection against Fi hybrids under natural con- ditions was not investigated, but laboratory produced hybrids developed normally. A high degree of ontogenetic compatibility is also indicated be- tween G. a. aculeatus and G. a. williamsoni, reported in this study, and between G. a. aculeatus and G. a. microcephalus (Hagen, 1967). Phenotypic dominance of G. a. microcephalus over G. a. williamsoni is shown by both meristic and morphometric characters. Consequently, Fi G. a. williamsoni x microcephalus hybrids cannot be separated from G. a. micro- cephalus on the basis of plate number. Hagen (1967) demonstrated that G. a. aculeatus is dominant over G. a. microcephalus and Miinzing (1963) showed 18 Contributions in Science No. 243 dominance of the fully plated European “trachurus” form over the partially plated “leiurus.” In conclusion, the evidence for postmating isolation between G. a. microcephalus and G. a. williamsoni is limited to a 21% differential in fertili- zation between intrasubspecific and intersubspecific crosses. Premating isolating mechanisms were not investigated. Taxonomically the forms are distinct so that subspecific recognition is useful. However, the trinomial is used in the sense of Mayr (1963) in that it does not necessarily infer an evolutionary unit. Acknowledgments This paper is derived from a thesis submitted to California State College, Fullerton, in partial fulfillment of the requirements for the MA degree. I gratefully acknowledge the advice, encouragement, and field assist- ance of D. W. Greenfield, my major professor, during the course of this study. Fellow graduate students, R. Hathaway Cornelius, David M. Wildrick, and James W. Wiley, also lent valuable assistance in collecting specimens. R. Hathaway Cornelius assisted in computer analysis of the data. I thank Dr. Lon McClanahan, Mr. Peter A. Bisson, and Dr. Bruce Cowell for reading the manuscript in its various stages and offering valuable suggestions. I am grateful to my wife Yvonne for her encouragement and for assistance in the preparation of this paper. Literature Cited Chen, T. R., and H. M. Reisman. 1970. A comparative chromosome study of the North American species of sticklebacks (Teleostei: Gasterosteidae). Cytogenetics 9: 321- 332. Culver, G. B., and C. L. Hubbs. 1917. The fishes of the Santa Ana system of streams in southern California. Lorquinia 1 (11): 82-83. Eigenmann, C. H. 1886. A review of the American Gasterosteidae. Proc. Acad. Nat. Sci. Phila. 1886: 233-252. Evermann, G. W., and H. W. Clark. 1931. A distributional list of the species of fresh- water fishes known to occur in California. Calif. Fish and Game, Fish. Bull. 35. , AND E. L. Goldsborough. 1906. The fishes of Alaska. Bull. Bur. Fish. 26: 219-360. Follett, W. I. 1960. The fresh-water fishes, their origins and affinities. Syst. Zool. 9 (3-4): 212-232. Gilbert, C. H. 1893. Report on the fishes of the Death Valley expedition, collected in southern California and Nevada in 1891, with descriptions of new species. N. Amer. Fauna 7: 229-234. Girard, C. 1854. Descriptions of new fishes collected by Dr. A. L. Heermann, natural- ist attached to the survey of the Pacific railroad route, under Lt. R. S. Williamson, U.S.A. Proc. Acad. Nat. Sci. Phila. 7, 1854-1855: 129-140. Hagen, D. W. 1967. Isolating mechanisms in threespine sticklebacks {Gasterosteus). J. Fish. Res. Bd. Canada 24 (8): 1637-1692. 1973 Systematics of GASTEROSTEUS ACULEATUS 19 and L.G. Gilbertson. 1972. Geographic variation and environmental selec- tion in Gasterosteus aculeatus L. in the Pacific Northwest, America. Evolution 26 (1); 32-51. , AND J. D. McPhail. 1970. The species problem within Gasterosteus acu- leatus on the Pacific Coast of North America. J. Fish. Res. Bd. Canada 27(1); 147- 155. Heuts, M. j. 1949. Racial divergence in fin ray variation patterns in Gasterosteus aculea- tus. J. Genet. 49 (3): 183-191. Hoogland, R., D. Morris, and N. Tinbergen. 1957. The spines of sticklebacks (Gasterosteus and Pygosteus) as means of defence against predators (Perea and Esox). Behaviour 10; 205-236. Hubbs, C. L. 1921. Notes on Cottus asper and Cottus aleuticus. Copeia 1921 (90); 7-8. 1929. The Atlantic American species of the fish genus Gasterosteus. Occ. Pap. Mus. Zook, Univ. Mich. No. 200; 1-9. , AND K. F. Fagler. 1958. Fishes of the Great Fakes region. Rev. ed. Cran- brook Inst. Sci. Bull. 26. , AND R. R. Miller. 1948. The Great Basin with emphasis on glacial and post- glacial times. IF The zoological evidence. Correlation between fish distribution and hydrographic history in the desert basins of western United States. Bull. Univ. Utah 38; 18-166. Jordan, D. S. 1894. Notes on the fresh-water species of San Fuis Obispo County, Cali- fornia. Bull. U.S. Fish. Comm. 1894, art. 14; 141-142. , AND H. E. Copeland. 1876. Check list of the fishes of the fresh-waters of North America. Bull. Buffalo Soc. Nat. Sci. 1876. , AND B. W. Evermann. 1896. The fishes of North and Middle America. U.S. Nat. Mus. Bull. 47 (11); 1-1240. , AND C. H. Gilbert. 1899. Fur seals and fur seal islands of the North Pacific Ocean. 3; 433-509. , AND C. F. Hubbs. 1925. Record of fishes obtained by D. S. Jordan in Japan, 1922. Mem. Carnegie Mus. 10 (2); 93-346. Kramer, C. Y. 1956. Extension of multiple range tests to group means with unequal numbers of replication. Biometrics 12; 307-310. Fewis, S. L. 1969. Physical factors influencing fish populations in pools of a trout stream. Trans. Am. Fish. Soc. 98 (1); 14-19. Findsey, C. C. 1962. Experimental study of meristic variation in a population of three- spine sticklebacks, Gasterosteus aculeatus. Can. J. Zook 40; 271-312. Mayr, E. 1963. Animal species and evolution. The Belknap Press, Harvard Univ. Press, Cambridge, Mass. 797 pp. 1969. Principles of systematic zoology. McGraw-Hill, New York. 428 pp. McPhail, J. D. 1969. Predation and the evolution of a stickleback (Gasterosteus). J. Fish. Res. Bd. Canada 26; 3183-3208. , AND R. F. Jones. 1966. A simple technique for obtaining chromosomes from teleost fishes. J. Fish. Res. Bd. Canada 23; 767-769. , AND C. C. Findsey. 1970. Freshwater fishes of northwestern Canada and Alaska. Fish. Res. Bd. Canada Bulk 173. 373 pp. Miller, R. R. 1939. Occurrence of the cyprinodont fish Fundulus parvipinnis in fresh- water in San Juan Creek, southern California. Copeia 1939 (3); 168. 1960. The type locality of Gasterosteus aculeatus williamsoni and its signif- icance in the taxonomy of Californian sticklebacks. Copeia 1960 (4); 348-350. 1961. Man and the changing fish fauna of the American Southwest. Pap. Mich. Acad. Sci., Arts, Fett. 1960, 46; 365-404. 1969. FU.C.N. Red Data Book, vok 4, Pisces. FF/222. , AND C. F. Hubbs. 1969. Systematics of Gasterosteus aculeatus, with par- ticular reference to intergradation and introgression along the Pacific Coast of North America; A commentary on a recent contribution. Copeia 1969 (1); 52-69. 20 Contributions in Science No. 243 Munzing, J. 1963. The evolution of variation and distributional patterns in European populations of the threespined stickleback, Gasterosteus aculeatus. Evolution 17; 320-332. Narver, D. W. 1969. Phenotypic variation in threespine sticklebacks (Gasterosteus aculeatus) of the Chignik River system, Alaska. J. Fish. Res. Bd. Canada 26: 405- 412. Nelson, J. S. 1971. Comparison of the pectoral and pelvic skeletons and of some other bones and their phylogenetic implications in the Aulorhynchidae and Gasterosteidae (Pisces). J. Fish. Res. Bd. Canada 28 (3): 427-442. Ojima, Y., S. Takayama, T. Minesita, and K. Yamaguchi. 1964. A modified air-drying method for a chromosome study of a solid tumor. Jap. J. Genet. 38 (5,6); 399-402. Regan, C. T. 1909. The species of three-spined sticklebacks. Ann. and Mag. Nat. Hist, ser. 8, 4; 435-437. Rutter, C. 1896. Notes on fresh water fishes of the Pacific slope of North America. Calif. Acad. Sci. ser. 2, 6: 245-267. Snyder, J.O. 1913. The fishes of the streams tributary to Monterey Bay, California. Bull. U.S. Bur. Fish. 22: 49-72. Steele, R.G.G. D. and J. H. Torrie. 1960. Principles and procedures in statistics. McGraw-Hill, New York. 481 pp. Accepted for publication March 7, 1973 Printed in Los Angeles, California by Anderson, Ritchie and Simon on Simpson Lee Recovei ^ 73 NUMBER 244 JUNE 28, 1973 A DISTRIBUTIONAL CHECKLIST OF THE TURTLES, CROCODILIANS, AND LIZARDS OF HONDURAS By John R. Meyer and Larry David Wilson CONTRIBUTIONS IN SCICNCE mrnsgrnmm NATURAL HISTORY MUSEUM LOS ANGELES COUNTY A DISTRIBUTIONAL CHECKLIST OF THE TURTLES, CROCODILIANS, AND LIZARDS OF HONDURAS' By John R. Meyer^ and Larry David Wilson^ Abstract: The turtles, crocodilians, and lizards of the Republic of Honduras are listed. For each of the 58 species present, brief synonymies, a statement of the entire range, a list of Honduran locality records, and a statement of the ecologic distribution in Honduras are given. Keys for identification are provided, and a list of species of questionable occurrence is included. This report represents the second in a series of papers designed to elucidate the composition and relationships of the herpetofauna of the Republic of Hon- duras. In an earlier paper (Meyer and Wilson, 1971a), we treated the 52 species of amphibians known from the country, and in preparation is a report on the snakes of Honduras. This paper is concerned with the 58 species of turtles, crocodilians, and lizards of the republic. Our earlier paper (Meyer and Wilson, 1971a) covered most of the same background material, and the reader is referred to it for information on pre- vious studies (Introduction, p. 1), extent of the various ecologic formations (Fig. 1, pp. 24-25), and localities (Gazetteer, p. 34). Synonymies given for each species include the following in the order indicated: 1) Author citation; 2) First citation of the present combination; 3) All references known to us in which Honduran specimens are mentioned. Additional localities, not included in the gazetteer of the previous paper, are given in the appendix. Acknowledgments This paper is a revised version of part of a doctoral thesis completed by Meyer at the University of Southern California. Thanks are extended to the chairman of his doctoral committee. Jay M. Savage, and to other members, Basil Nafpaktitis, Findlay Russell, John Wright, and Russel Zimmer. Curators of collections from which material was examined are Walter Auffenberg, Florida State Museum (UF); James E. Bdhlke, Academy of Natural Sciences, Philadelphia (ANSP); James R. Dixon, Texas A&M University (TCWC); William E. Duellman, Museum of Natural History, University of Kansas (KU); Robert F. Inger, Field Museum of Natural History (FMNH); Alan E. Leviton, California Academy of Sciences (CAS); C. J. McCoy, Carne- gie Museum (CM); the late Michael Ovchynnyk, Michigan State University 'Review Committee for this Contribution Clarence J. McCoy Roy W. McDiarmid John W. Wright ^Research Associate, Section of Herpetology, Natural History Museum of Los Angeles County, Los Angeles, California 90007 ^Division of Intercurricular Studies, Miami-Dade Junior College, South Campus, Miami, Florida 33156 1 1973 Turtles, Crocodilians, and Lizards of Honduras 2 (MSU); the late James A. Peters, National Museum of Natural History (USNM); Douglas A. Rossman, Museum of Zoology, Louisiana State Univer- sity (LSUMZ); Dorothy M. Smith, Museum of Natural History, University of Illinois (UIMNH); Charles F. Walker, Museum of Zoology, University of Michigan (UMMZ); Ernest E. Williams, Museum of Comparative Zoology, Harvard University (MCZ); John W. Wright, Natural History Museum of Los Angeles County (LACM); Richard G. Zweifel, American Museum of Natural History (AMNH). In addition, we would like to thank Ernest A. Liner, Houma, Louisiana for the donation of a portion of his private collection to the Louisiana State University Museum of Zoology at our request. Thanks are also extended to James DeWeese, University of Southern California, for his help in tracking down literature references. To the many people of Honduras whose hospitality and friendship we have enjoyed, we wish to express our deepest thanks. Particularly helpful were Rafael Becerra and family. La Ceiba; Oscar Crespo and family, Trujillo; faculty and students of the Escuela Nacional de Agricultura, Catacamas; per- sonnel of Standard Fruit Company, La Ceiba and Coyoles, and United Fruit Company, La Lima; H. E. Ostmark, La Lima; William Plowden, San Pedro Sula; faculty and students of the Escuela Americana, Tegucigalpa. Field work in 1967 was made possible by funds from the Biomedical Sciences Support Grant, FR 07012-01, from the National Institutes of Health (Meyer) and a Louisiana State University Field Research Fellowship (Wilson), and in 1968 by a Graduate School Travel Fellowship from the University of Southern California (Meyer) and a Sigma Xi Grant-in-Aid of Research (Meyer). Finally, we would like to thank our wives for their help and companion- ship during the field work. Order TESTUDINES KEY TO THE GENERA OF TURTLES IN HONDURAS 1. At least one scute of carapace in contact with pectoral scute 2 1. Scutes of carapace separated from pectoral scute of plastron either by a ligament or by one or more inframarginal scutes of bridge 3 2. Alveolar surface of upper jaw without a ridge; head and neck spotted or obscurely lined with dark on light ground color Rhinoclemmys 2. Alveolar surface of upper jaw with a ridge; head and neck with prominent bright yellow and black stripes on dark green or greenish black ground color Chrysemys 3. At least four inframarginal scutes Dermatemys 3. Fewer than four inframarginal scutes 4 4. Plastron of 1 1 scutes, with anterior and posterior portions hinged Kinosternon 3 Contributions In Science No. 244 4. Plastron of fewer than 11 scutes, not hinged 5 5. Plastron of 8 scutes; carapace with three strong keels Staurotypus j 5. Plastron of 10 scutes; carapace only weakly keeled, if at all Chelydra | Family Chelydridae Genus Chelydra Schweigger Chelydra serpentina (Linnaeus) Testudo serpentina Linnaeus, 1758: 199. Chelydra serpentina: Schweigger, 1812: 293. Chelydra rossignoni: Werner, 1896: 344; Schmidt, 1924: 86. Chelydra acutirostris: Campbell and Howell, 1965: 131. Range.-ho'w elevations of the Atlantic versant from southeastern Canada to Ecuador. Locality Recor ds. -COP 8 mi ENE Copan, LACM 48451. CORTES: Laguna Ticamaya, FMNK 5327; San Pedro Sula, FMNH 5322. EL PARAISO: Arenal, LACM 73821. OLANCHO: 4.5 km SE Catacamas, LACM 48449-50. Ecologic Distribtuion.-Sea level to 600 m in the Tropical Moist Forest, Tropical Dry Forest, and Subtropical Wet Forest formations. Individuals were found in a small, clear stream at night and in a drainage ditch in the daytime. This species is restricted to aquatic situations and is undoubtedly more wide- spread in the country than records indicate. Family Kinosternidae Genus Dermatemys Gray Dermatemys mawei Gray Dermatemys mawei Gray, 1847: 55; Boulenger, 1889: 28. Range.-how elevations of the Atlantic versant from Veracruz, Mexico to northwestern Honduras. Locality Records. -"Honduras'" (Boulenger, 1889). Ecologic Distribution.- Although there are no definite locality records for this species in Honduras, it is probable that it occurs in the lower reaches of at least the Rio Ulua-Rib Chamelecdn drainage. Genus Kinosternon Spix KEY TO THE SPECIES OF KINOSTERNON IN HONDURAS Two or three keels on carapace in adults, usually indicated only posteriorly; head mottled with dark and light markings (light markings reddish or orangish in life); no opposable patches of spines on thigh and calf in adult males scorpioides 1973 Turtles, Crocodilians, and Lizards of Honduras 4 Carapace never with more than a median keel; head uniformly light, with small light spots or blotches on a dark ground color, or with a distinct broad temporal light stripe (light areas yellowish in life) opposable patches of spines on thigh and calf in adult males leucostomum Kinosternon leucostomum Dumeril and Bibron Cinosternon leucostomum Dumeril and Bibron, 1851: 17; Werner, 1896: 345. Cinosternon brevigulare: Atkinson, 1907: 152. Cinosternon cobanum: Atkinson, 1907:152. Kinosternon leucostomum: Meyer, 1966: 174. Range.-Co'^ and moderate elevations of the Atlantic versant from Vera- cruz, Mexico to Panama'. Locality Records.- ATL/(l\: Lancetilla, AMNH 70422-29, 70431, MCZ 29830; La Ceiba, USNM 117607, 71721; Tela, MCZ 27601-05; Corozal, LSUMZ 21432; 8 km SE La Ceiba, KU 101406. CHOLUTECA: between San Lorenzo and Choluteca, AMNH 70404-06. COLON: 1-3 km W Trujillo, KU 101407; 1 km SSW Trujillo, KU 101408. COMAYAGUA: Las Limas, MCZ 38824; 1.8 km SW Comayagua, LSUMZ 24174, 24192-94; 3.7 km S Comayagua, LSUMZ 24196-97. COPAN: Copan, UMMZ 83046 (8), 83047 (3), FMNH 28523. CORTES: 1 mi W El Jaral, LACM 47288; 2 mi W San Pedro Sula, LACM 47289; Agua Azul, AMNH 70400-01; Amapa, AMNH 70503; San Pedro Sula, USNM 24374; Laguna Ticamaya, FMNH 5236-39, 5242-43; W of San Pedro Sula, FMNH 5249; Hacienda Santa Ana, FMNH 5246-48; Rio Santa Ana, FMNH 5250-52; Potrerillos, MCZ 29817-19; Santa Ana Mountains, MCZ 1973 Turtles, Crocodilians, and Lizards of Honduras 20 32208; E side Lago de Yojoa, KU 67155; 3 km W Cofradia, LSUMZ 24600. FRANCISCO MORAZXN: Zamorano, AMNH 70382-84, 70359-60, MCZ 48674, 49757, 49992, LACM 39769; Cantarranas, ANSP 26080-81; 1 mi W Talanga, TCWC 19189; Tegucigalpa, LACM 39770. GRACIAS A DIOS: Pa- tuca, USNM 20305; Rio Coco, USNM 24525-26; Tancin, LACM 47277-81. ISLAS DE LA BAHIA: Isla de Utila, 2 mi N Utila, TCWC 19190; Isla de Utila, Utila, UF 28403, 28440, LSUMZ 22273. LEMPIRA: Erandique (Hahn, 1971). OLANCHO: 1 km NW Catacamas, LACM 47276. SANTA BA'RBARA: Qui- mistan, USNM 128090. YORO: Subirana Valley, FMNH 21845 (2); Sulaco Mountains, UMMZ 77846; 0.5 km N Coyoles, LACM 47282-83; Coyoles, LACM 47284-87, LSUMZ 21421-31; 2 km S Coyoles, KU 101409. Ecologic Distribution.-SQ2i level to 1000 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical Moist Forest, and Subtropical Dry Forest formations. Generally encountered in open or edge situations, where it was found on the ground or on low vege- tation. Anolis sminthus Dunn and Emlen Anolis sminthus Dunn and Emlen, 1932: 26; Barbour, 1934: 150; Meyer and Wilson, 1971b: 108. -Intermediate elevations of the Pacific versant in El Salvador and Honduras. Locality Records.-FRANCISCO MORAZAN: San Juancito, ANSP 22946-49; 7 mi SW San Juancito, LACM 47676-78, LSUMZ 21415-16; Cerro La Tigra, LSUMZ 24200-01, 24212; Monte Crudo, AMNH 70392-93; Rancho Quemado, AMNH 70390-91 70394, 70396-97. INTIBUCA": 1.5 mi NE La Espe- ranza, LACM 47862. OCOTEPEQUE: 12.5 mi E Nueva Ocotepeque, LACM 47679-81. Ecologic Distribution.-¥vom 1700 to 2200 m in the Lower Montane Moist Forest formation. Found on the ground and on lower parts of vegetation in edge situations, clearings, and pine-oak forest. Anolis tropidonotus Peters Anolis tropidonotus Peters, 1863: 135; Werner, 1896: 346; Dunn and Emlen, 1932: 26; Barbour, 1934: 153; Smith, 1950: 55; Wilson and Meyer, 1969: 146; Hahn, 1971: 111. Anolis tropidonotus tropidonotus: Meyer, 1966: 175. Range.— ho'w, moderate, and intermediate elevations of the Atlantic versant from Veracruz, Me^xico to Nicaragua; also on the Pacific versant in Honduras. Locality i?ccorc/5.-ATLANTIDA: mountains above Corozal, LACM 47644- 47, LSUMZ 21390-96; 8 km SE La Ceiba, KU 101411-20; 13 km E La Ceiba, LACM 47649, LSUMZ 21397; Carmelina, USNM 62970; Lancetilla, AMNH 70432, 70434-36. COLON: 2 km E Trujillo, LACM 47650-54, LSUMZ 21400-09; 21 Contributions In Science No. 244 1 km SSE Trujillo, KU 101421-22; 1 km SSW Trujillo, KU 101423-29; 0.5 km SW Trujillo, LSUMZ 21410; mountains above Trujillo, LSUMZ 22464, 22490, 22449; Balfate, AMNH 58627, 58612-15. COMAYAGUA: Talaube, LSUMZ 24210. COPAN: 12 mi ENE Copan, LACM 47655; Copan, TCWC 19191, FMNH 28516, 28518-20, AMNH 70270. CORTES: 1 mi W El Jaral, LACM 45261-66, 47656-59; 1 mi SE El Jaral, LACM 47660-61, 47666; 2.5 mi ENE Villanueva, LACM 47663; 2 mi W San Pedro Sula, LACM 47665; 3 km WSW Cofradia, KU 67145; Lago de Yojoa, MSU 4644, AMNH 70289; Potrerillos, MCZ 29799-800; near Cofradia, MCZ 32202. EL PARAISO: Arenal, LACM 16850-54; Rio Guayambre, AMNH 70285; Valle de Jamastran, AMNH 70286; near Agua Fria, AMNH 70267-69; La Montanuela, AMNH 70265. FRANCISCO MORAZAN: La Montanita, AMNH 70266; Montana de Guaimaca, AMNH 70271; Zamorano, AMNH 70272-84; Cerro Uyuca, KU 103237-38, MCZ 49980-82, AMNH 70389, 70287-88, 70395, LSUMZ 24204; Rancho San Diego, AMNH 69087; Tegucigalpa, AMNH 69088; 9.9 km N Tegucigalpa, LSUMZ 24181-91; El Hatillo, LSUMZ 24205-06, 24211; 21 km NW Zamorano, LSUMZ 24601. LA PAZ: Marcala (Hahn, 1971). LEMPIRA: Gracias, FMNH 40869. OLANCHO: 1 km WNW Catacamas, LACM 47622-25, LSUMZ 21381-83, 2141 1; 1.5-3 km NW Catacamas, LACM 47626-32; 3 km N Catacamas, LACM 47633-34; 4.5 km SE Catacamas, LACM 47635, 45220-26, LSUMZ 21384-85; 4 km E San Jose del Rib Tinto, LACM 45144-45; Rib Guampusito, 7.5 mi E Dulce Nombre de Culmi', LACM 45153-54. SANTA BARBARA: 5.5 mi SW El Jaral, LACM 47662; El Sauce, KU 67148-50; mountains W of Lago de Yojoa, KU 67151-52; near Quimistan, USNM 128088-89. YORO: 2 km S Coyoles, KU 101430-33; 5 km E Coyoles, LACM 47636-38, LSUMZ 21386-87; Sopametepe, LSUMZ 21388, LACM 47639; Coyoles, LACM 47640-43, LSUMZ 21389; Subirana Valley, MCZ 38797, 38802-11; Mataderos Mountains, MCZ 38798-801. Ecologic Distribution. -Sea level to about 1900 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical Moist Forest, and Lower Montane Moist Forest formations. Gen- erally found in undisturbed forest, edge situations, or gallery forest along streams, where it was encountered on the ground and on the lower parts of vegetation. Genus Basiliscus Laurenti KEY TO THE SPECIES OF BASILISCUS IN HONDURAS Abdominal scales smooth plumifrons Abdominal scales keeled vittatus Basiliscus plumifrons Cope Basiliscus plumifrons Cope, 1876(1875): 125 Range.-Co\^ and moderate elevations of the Atlantic versant from eastern Honduras to Panama'. Locality Records.-GKACIAS A DIOS: Rib Coco, USNM 24523. 1973 Turtles, Crocodilians, and Lizards of Honduras 22 Ecologic Distribution. -Known only from below 100 m in the Tropical Moist Forest formation. Basiliscus vittatus Wiegmann Basiliscus vittatus Wiegmann, 1828: 373; Gunther, 1885: 55; Werner, 1896: 346; Dunn and Emlen, 1932: 28; Lynn, 1944: 190; Meyer, 1966: 175; Hahn, 1971: 111; Wilson and Hahn, in press. Cristasaura mitrella Gray, 1852: 439. Range.-Kow and moderate elevations from Veracruz, Me'xico on the Atlantic versant and Guerrero, Mexico on the Pacific to Panama. Locality T^^corJ^.-ATLANTlDA: La Ceiba, USNM 117604-06, 55243, 55246, LACM 48359; Carmelina, USNM 62969; Corozal, LACM 48361, LSUMZ 21651; 20 mi E Tela, LACM 48367; Lancetilla, AMNH 70438-44. CHOLUTECA: Choluteca, MSU 4648. COLON: 0.5-3 km W Trujillo, LSUMZ 21652-54, KU 101435; ca. 2 mi E Trujillo, LSUMZ 22492; mountains above Trujillo, LSUMZ 22461-63; Puerto Castilla, LSUMZ 22474; Balfate, AMNH 58610, 58628-29; Guaimoreto, LSUMZ 10287-91. COMAYAGUA: La Mision, MCZ 49976; Siguatepeque, FMNH 5212; 3 km W Comayagua, TCWC 23813. COPA'N: Rio Higuito, ANSP 22979-81, 26660; Copan, TCWC 23630; 8 mi ENE Copa'n, LACM 48362; 5 mi SE Santa Rosa de Copan, LACM 48363. CORTE'S: San Pedro Sula, USNM 24378; Lago de Yojoa, MSU 4645-47; 7 km SW La Lima, KU 67209; 5 km SW La Lima, KU 67210; 1 mi W El Jaral, LACM 48364; 4.5 mi ENE Villanueva, LACM 48365; La Lima, LACM 48366; 2 mi W San Pedro Sula, LACM 48368. EL PARAISO: near Ojo de Agua, AMNH 70350; Arenal, LACM 16838-43; 15.8 mi W Danlf, LACM 45069. FRANCISCO MORAZAN: Rancho San Diego, AMNH 69074-75, TCWC 19192-95; Tegu- cigalpa, AMNH 69085; Zamorano, AMNH 70347^9, 70352-53, LACM 39771; 5 mi ENE Zamorano, KU 103240-fl. GRACIAS A DIOS: Patuca, USNM 20297-303; Rio Coco, USNM 24512-22; Kiyras, LACM 16837; Tancin, LACM 48357-58, LSUMZ 21643. ISLAS DE LA BAHIA: Isla de Utila, USNM 28626; Isla de Utila, Utila, UF 28389-91, 28406-14, 28452-53, LSUMZ 22269-71; Isla de Roata'n, 0.5-1 km NNE Roatan, TCWC 21952-53; Isla de Roata'n, 0.5-1 km N Roatan, LSUMZ 21644-45, Isla de Roatan, Roata'n, LSUMZ 22318; Oak Ridge (Wilson and Hahn, in press); Isla de Guanaja, SE shore opposite Guan- aja, LSUMZ 21646, UF 28573; Isla de Guanaja, La Playa Hotel, LSUMZ 21647; Isla de Guanaja, KU 101434. LA PAZ: Marcala (Hahn, 1971). LEM- PIRA: Erandique (Hahn, 1971). OCOTEPEQUE: 3 km S Nueva Ocotepeque, TCWC 23732-34. OLANCHO: ca. 6 mi E Catacamas, LACM 45132; 0.5 km WNW Catacamas, LACM 48351-52; 12-14 mi W Catacamas, LACM 45166; 4.5 km SE Catacamas, LACM 48353-56; 5.9 km WSW Campamento, LSUMZ 22265. YORO: Subirana Valley, MCZ 38837-38; 0.5 km N Coyoles, LACM 48360; 5 km E Coyoles, LSUMZ 21650; Coyoles, LSUMZ 21648^9; 17 km NW Progreso, LSUMZ 24602. Ecologic Distribution.-S^a level to 1200 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical 23 Contributions In Science No. 244 Moist Forest, and Subtropical Dry Forest formations. Most frequently found near water; never encountered in closed forest, but in edge or open situations. Genus Corytophanes Boie Corytophanes cristatus (Merrem) Agama cristata Merrem, 1821: 50. {Corytophanes^ cristatus: Boie (by inference), 1827: 290. Corytophanes cristatus: Werner, 1896: 346; Dunn and Emlen, 1932: 28. Range.— ho'w and moderate elevations of the Atlantic versant from cen- tral Veracruz, Mexico to Colombia; also on the Pacific versant in Costa Rica. Locality i?ccor^/5.-ATLANTIDA: mountains above Corozal, LACM 47847-48; Guaymas District, UMMZ 58372, 58381; Tela, MCZ 21153; near Tela, USNM 101279; Lancetilla, MCZ 29393, 31483; La Ceiba, MCZ 32035. COLON: 15 mi E Trujillo, USNM 38379. CORTES: W of San Pedro Sula, MCZ 32036. GRACIAS A DIOS: between Rio Patuca and Rib Coco, MCZ 38840-41; Rib Coco, USNM 24524. YORO: Mataderos Mountains, FMNH 21794-95, MCZ 38842-44; Portillo Grande, FMNH 34754; Subirana Valley, MCZ 38839. Ecologic Distribution.—Sea. level to about 1300 m in the Tropical Moist Forest and Subtropical Wet Forest formations. This species was found on trees in undisturbed forest during the daytime. Genus Ctenosaura Wiegmann Ctenosaura similis (Gray) Iguana (Ctenosaura) Similis Gray, 1831: 38. Ctenosaura similis: Bailey, 1928: 32; Dunn and Emlen, 1932: 28; Meyer, 1966: 175; Wilson and Hahn, in press. Range.-Lo^ and moderate elevations of both versants from the Isthmus of Tehuantepec, Mexico to Panama. Locality Rccor^/^.-ATLANTIDA: Tela, MCZ 22669, UMMZ 69536. CHOLUTECA: Pespire, FMNH 5191-93; Choluteca, MSU 4649-53; Ola, TCWC 20813. COLON: 0.5-1. 5 km W Trujillo, LACM 48416-17, LSUMZ 21659-61; 1-3 mi E Trujillo, LSUMZ 22445-47, 22495; 2 km E Trujillo, LACM 48418, LSUMZ 21662; mountains above Trujillo, LSUMZ 22454; Puerto Cas- tilla, LSUMZ 22475-78. COMAYAGUA: 3 mi SSE Comayagua, LACM 48424. CORTES: San Pedro Sula, USNM 24376; near Cofradia, USNM 128086-87; La Lima, LACM 48419-22; 2-3 mi W San Pedro Sula, TCWC 19197, LACM 48423: Puerto Cortes, TCWC 19196; 1 mi W La Lima, TCWC 19198. FRAN- CISCO MORAZAN: Zamorano, MCZ 49927, AMNH 70342-45; 21 km SSW Sabana Grande, LSUMZ 24603. GRACIAS A DIOS: Patuca, USNM 20290-91; Tancin, LACM 48456. ISLAS DE LA BAHIA: Isla de Utila, near Utila, UF 28392-95, 28415-29, 28445-51, LSUMZ 22276-77, 22287-92, 22299-302; Isla de Guanaja, 5.5 mi W Sabana Bight, TCWC 21954; Isla de Guanaja, La Playa 1973 Turtles, Crocodilians, and Lizards of Honduras 24 Hotel, LACM 48412-14, 48458, LSUMZ 21655; Isla de Guanaja, KU 101438; Isla de Guanaja, SE shore opposite Guanaja, UF 28575-77, LSUMZ 22406-09; Isla de Guanaja, near Sabana Bight, UF 28581-89, LSUMZ 22410-11. LEMPIRA: Copan-Lempira border, FMNH 40866. SANTA BARBARA: near Quimistan, USNM 128094-97; 20.5 mi W Chamelecon, LACM 48458; 4 km SW Quimistan, KU 67222. VALLE: San Lorenzo, FMNH 5194 (8); Isla Zacate Grande, MSU 4654-59; 10 km WNW Nacaome, KU 67218-21. YORO: near Yoro, MCZ 29401-03; 0.5 km N Coyoles, LSUMZ 21656, LACM 48415; Rancho San Lorenzo, LSUMZ 21657-58. Ecologic Distribution— level to 800 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, and Subtropical Moist Forest forma- tions. Found in trees and on the ground in the daytime, and one individual was taken from a hollow stump at night. Genus Enyaliosaurus Gray • KEY TO THE SPECIES OF ENYALIOSAURUS IN HONDURAS 1. First two or more whorls of enlarged spinous caudal scales separated from each other by two rows of small, flat scales bakeri 1. First two or more whorls of enlarged spinous caudal scales separated from each other by one row of small, flat scales 2 2. Males and females with a prominent dewlap palearis 2. Males and females with a transverse gular fold, but no dewlap quinquecarinata Enyaliosaurus bakeri (Stejneger) Ctenosaura bakeri Stejneger, 1901: 467; Bailey, 1928: 38; Barbour, 1928: 56; Wilson and Hahn, in press. Enyaliosaurus bakeri: Cochmn, 1961: 105. (?) Ctenosaura completa: Gunther, 1885: 58 (In part). Range.— Known only from the Bay Islands of Honduras. Locality Records.-ISLAS DE LA BAHIA: Isla de Utila, USNM 26317, 25324; Isla de Utila, Utila, UF 28437, 28471, LSUMZ 22275, 22293; Isla de Roatan, French Harbor, FMNH 53831; Isla de Roatan, ca. 3 mi W Roatan, LSUMZ 22367-71, UF 28530-33; Isla de Roatan, near Roatan, LSUMZ 22399; Isla de Roatan, near French Harbor, UF 28553. Ecologic Distribution.— Known only from near sea level in the Tropical Moist Forest formation. This species was taken on rocks and in grass along beaches, and one individual was found asleep inland at night on the trunk of a thorn palm. Enyaliosaurus palearis (Stejneger) Ctenosaura palearis Stejneger, 1899: 381. Enyaliosaurus palearis: Smith and Taylor, 1950: 76; Echternacht, 1968: 151. 25 Contributions In Science No. 244 Range.-how^ elevations of the Motagua Valley in Guatemala and the Aguan Valley in Honduras. Locality Records.-\OKO\ 0.5 km N Coyoles, LACM 48425-27, LSUMZ | 21482-86; Rancho San Lorenzo, LACM 48428-30, LSUMZ 21487-88; 2 km S | Coyoles, KU 101439-41. | Ecologic Distribution.-Y^nov^n only from 150 to 250 m in the Tropical | Arid Forest formation. Adults were found in trees and juveniles on the ground | in the forest in the daytime. | Enyaliosaurus quinquecarinatus (Gray) Cyclura quinquecarinata Gray, 1842: 59. Enyaliosaurus quinquecarinata: Gray 1845: 192. Ctenosaura quinquecarinata: Gunther, 1885: 58; Bailey, 1928: 42. -Disjunct populations on the Pacific versant from the Isthmus of Tehuantepec, Mexico to Nicaragua. Locality -“Honduras” (Gunther, 1885). LA PAZ: La Paz, LACM 72088-89. Ecologic Distribution.-Y.no'wn only from about 750 m in the Subtropical Dry Forest formation. Genus Iguana Laurenti Iguana iguana (Linnaeus) Lacerta iguana Linnaeus, 1758: 206. Iguana [iguana^: Van Denburgh, 1898: 461. Iguana tuberculata: Gunther, 1885: 56. Iguana tuberculata var. rhinolopha: Werner, 1896: 346. Iguana iguana rhinolopha: Barbour, 1928: 56; Dunn and Emlen, 1932: 28; Dunn, 1934: 1; Smith, 1950: 55. Iguana iguana: Wilson and Hahn, in press. Range.-Lov^ and moderate elevations from Veracruz, Me'xico on the Atlantic versant and Sinaloa, Me^’xico on the Pacific to South America. Locality Records.-COLO^\ 2 km E Trujillo, LSUMZ 21507; ca. 8 mi NE Trujillo, LSUMZ 22502; Barranca, ANSP 24190. COPXN: Copan, TCWC 23634, UMMZ 83033 (2). CORTES: near Cofradia, USNM 128085. FRAN- CISCO MORAZAN: Zamorano, MCZ 49937; Cantarranas, ANSP 24191-93. GRACIAS A DIOS: Tancin, LSUMZ 21506; Patuca, USNM 20287-89. ISLAS DE LA BAHIA: Isla de Roatan, 3 km N Roatan, LACM 47849; Isla de Utila, Utila, LSUMZ 22294; Isla de Guanaja (Wilson and Hahn, in press). SANTA BA^RBARA: 12.3 km ENE Quimista'n, KU 67234-36. Ecologic Distribution.-Sea. level to about 800 m in the Tropical Moist Forest, Tropical Dry Forest, Subtropical Moist Forest, and Subtropical Dry Forest formations. Adults were found in trees in disturbed forest and edge situ- ations and juveniles were found on the ground and on low vegetation. In the drier formations this species always was encountered along streams. 1973 Turtles, Crocodilians, and Lizards of Honduras 26 Genus Laemanctus Wiegmann Laemanctus longipes Wiegmann Laemanctus longipes Wiegmann, 1834: 45. Laemanctus waiter si Schmidt, 1933: 20. Laemanctus longipes deborrei: McCoy, 1968: 668. Laemanctus longipes waiter si: McCoy, 1968: 670. Range.— Lo'w and moderate elevations of the Atlantic versant from cen- tral Veracruz, Mexico to Honduras. Locality /?ccorJ5.-ATLANTIDA: Tela, USNM 84026, 84550, MCZ 27907, 29334, 32034. COPXN: Copan, UMMZ 83032. CORTES: Laguna Ticamaya, FMNH 5213. Ecologic Distribution.-SQa level to about 600 m in the Tropical Moist Forest, Tropical Dry Forest, and Subtropical Moist Forest formations. Diurnal and arboreal. Genus Polychrus Cuvier Polychrus gutturosus Berthold Polychrus gutturosus Berthold, 1846: 5. Range.— and moderate elevations of the Atlantic versant from Hon- duras to Colombia and Ecuador. Locality /?ecorr/5.-ATLANTIDA: Guaymas District, UMMZ 58639. Ecologic Distribution.— Known only from about 20 m in the Tropical Moist Forest formation. The single specimen, according to field notes, was found on the trunk of a large tree immediately after it was felled. Genus Sceloporus Wiegmann KEY TO THE SPECIES OF SCELOPORUS IN HONDURAS 1. Postfemoral dermal pocket present variabilis 1. No postfemoral dermal pocket 2 2. Dorsal pattern gives the impression of a dorsolateral light stripe squamosus 2. Dorsal pattern not giving the impression of a dorsolateral light stripe malachiticus Sceloporus malachiticus Cope Sceloporus malachiticus Cope, 1864: 178; Dunn and Emlen, 1932: 28; Stuart, 1971: 238. Sceloporus schmidti Jones, 1927: 4. Sceloporus formosus malachiticus: Smith, 1939: 46. Sceloporus formosus smaragdinus: Smith, 1939: 41. Sceloporus malachiticus malachiticus: Smith, 1942: 356. 27 Contributions In Science No. 244 jR^z^gc.-Moderate and intermediate elevations from western Honduras on the Atlantic versant and El Salvador on the Pacific to Panama'. Locality Records. -CORTES: W San Pedro Sula, FMNH 5214. EL PAR- AISO: Monserrat, MCZ 49952. FRANCISCO MORAZA'N: 7.3 mi SW* San Juancito, LSUMZ 21505; La Montanita, MCZ 49934; Cerro Uyuca, MCZ 49939- 41, KU 103242-44; Tatumbla, MCZ 49984; 15 km N Sabana Grande, LSUMZ 24604-06; 5 km W Zambrano, KU 67270; Cerro La Tigra, LSUMZ 24202, 24215; El Hatillo, LACM 72090. LSUMZ 24207: 41 km NW Tegucigalpa, LSUMZ 24607. INTIBUCA: 1.5 mi NE La Esperanza, LACM 72103-16; 8 mi ESE La Esperanza, LACM 72116; vicinity of La Esperanza, LACM 45249. OCOTE- PEQUE: 12.5 mi E Nueva Ocotepeque, LACM 72096-98. SANTA BARBARA: Cerro Santa Barbara, E slope, LACM 72100-02. YORO: Subirana Ranch, MCZ 32283-92; Subirana Valley, MCZ 38846-50; Mataderos Mountains, MCZ 38845; Portillo Grande, MCZ 38851-54. Ecologic Distribution.-From about 900 to 2200 m in the Subtropical Wet Forest, Subtropical Moist Forest, Lower Montane Wet Forest, and Lower Mon- tane Moist Forest formations. Found on the sides of trees in clearings and edge situations in the daytime. Sceloporus squamosus Bocourt Sceloporus squamosus Bocourt, 1874: 212; Smith, 1939: 319; Hahn, 1971: 111. Range.-Lov^ and moderate elevations of the Pacific versant from Chi- apas, Mexico to Costa Rica; also on the Atlantic versant in Guatemala and Honduras. Locality Records.-COMAT AGE K: 3 mi SSL Comayagua, LACM 47294. COPA'N: Copan, UMMZ 84274; 17 km NE Cucuyagua, LSUMZ 24608. EL PARAISO: Monserrat, MCZ 49953; Ojo de Agua, AMNH 70387-88. FRAN- CISCO MORAZAN: Zamorano, KU 103245-48, LACM 39776; near Zam- orano, MCZ 49909, 49938, 49985, LACM 39775, AMNH 70385-86; 8 km ENE Zamorano KU 103249-50; 12 km SSW Sabana Grande, LSUMZ 24609; 21 km SSW Sabana Grande, LSUMZ 24610. LA PAZ: 1 km E La Paz (Hahn, 1971). LEMPIRA: Gracias, FMNH 28561, 40867; 7 mi NNW Gracias, LACM 47293. OCOTEPEQUE: 5 km N Nueva Ocotepeque, TCWC 23773. Ecologic Distribution.— From about 500 to 1000 m in the Tropical Arid Forest, Subtropical Moist Forest, and Subtropical Dry Forest formations. Found on fenceposts and rock walls in cleared areas in the daytime. Sceloporus variabilis Wiegmann Sc[eloporus] variabilis Wiegmann, 1834: 51. Sceloporus variabilis: Dunn and Emlen, 1932: 28; Wilson and Meyer, 1969: 146; Hahn, 1971: 111. Sceloporus variabilis olloporus: Smith, 1939: 282; Meyer, 1966: 175. Range.-Fov^ and moderate elevations from southern Texas on the Atlantic versant and Oaxaca, Mexico on the Pacific to Costa Rica. 1973 Turtles, Crocodilians, and Lizards of Honduras 28 Locality Records.-CWOLmECk: Respire, FMNH 5218. COMAYA- GUA: 13.7 km E Siguatepeque, KU 67286-89; Siguatepeque, FMNH 5219-25. COPAN; Copan, UMMZ 84275, AMNH 70319-21; 12 mi ENE Copan, LACM 47839^0; 6 km NE Copan, LSUMZ 24611; 13 km NE Cucuyagua, LSUMZ 24612. CORTES: 1 mi W El Jaral, LACM 47842, LSUMZ 24214; 1 mi SE El Jaral, LACM 47843; 3 mi W San Pedro Sula, LACM 47845-46; Agua Azul, MCZ 49965, AMNH 70326-27; 6 km N Agua Azul, LSUMZ 24613. EL PARAl- SO: Agua Fna, AMNH 70317; 6 km SE Los Limones, LSUMZ 24614; 15.8 mi W Danir, LACM 45066-68. FRANCISCO MORAZA'N: Rancho San Diego, TCWC 19199, AMNH 69076-79, 69069-73; 22 mi W Guaimaca, LACM 45095; Montana de Guaimaca, AMNH 70322; 18.7 km WSW Guaimaca, LSUMZ 22266; 18.4 km NE Talanga, LSUMZ 22267-68; Ventas, FMNH 5216-17; Za- morano, AMNH 70324-25, LACM 38774, MCZ 49764; near Zamorano, LACM 39772-73, MCZ 49910-19, AMNH 70323; 5 mi S Zamorano, AMNH 69641; 18.8 mi N Tegucigalpa, LACM 45242-43; 11-15 km N Sabana Grande, LSUMZ 24615-17; 12 km SSW Sabana Grande, LSUMZ 24618; 2-6 km SW Valle de Angeles, LSUMZ 24619-20; 16 km SW Valle de Angeles, LSUMZ 24621. GRACIAS A DIOS; Tancin, LSUMZ 21503-04. LA PAZ: Marcala (Hahn, 1971). LEMPIRA: Gracias (Hahn, 1971); 7 mi NNW Gracias, LACM 47841; Erandique (Hahn, 1971). OCOTEPEQUE: Nueva Ocotepeque, TCWC 23635- 36; 1 km W Nueva Ocotepeque, TCWC 23776. OLANCHO: Pataste, MSU 4660-78. SANTA BARBARA: 3 mi E Quimistan, LACM 47838; 1 mi NE El Sauce, LACM 47844. VALLE: 2 mi W Nacaome, LACM 47837. YORO: Por- tillo Grande, MCZ 32269, 32271-78, 38855; Subirana Ranch, MCZ 32279-82; Subirana Valley, MCZ 38857-90; Mataderos Mountains, MCZ 38856, FMNH 21869; 2 km S Coyoles, KU 101442-43. Ecologic Distribution. —Sea. level to about 1300 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical Moist Forest, and Subtropical Dry Forest formations. Found in the daytime on the ground, on lower parts of trees, and on rock walls and fences in cleared and disturbed areas. Family Scincidae Genus Eumeces Wiegmann Eumeces sumichrasti (Cope) Plistodon sumichrasti Cope, 1867; 321. Eumeces sumichrasti: Bocourt, 1879: 422; Taylor, 1946: 178; Marx, 1958: 459. Eumeces schmidti Dunn and Emlen, 1932: 30. Range.-Loy^ elevations of the Atlantic versant from central Veracruz, Mexico to Honduras. Locality 7?^cor<75.-ATLANTIDA: Lancetilla, ANSP 19877; Tela (Dunn and Emlen, 1932); mountains above Corozal, LACM 47295. Ecologic Distribution.-Y^no^n only from sea level to about 200 m in the Tropical Moist Forest formation. Found in the daytime on the ground and on fallen trees in undisturbed forest and edge situations. 29 Contributions In Science No. 244 Genus Mabuya Fitzinger Mabuya mabouya (Lacepede) Lacerta Mabouya Lacepede, 1788: 376. Mabuya [mabouya]: Fitzinger, 1826: 52. Mabuya agilis: Werner, 1896: 347; Dunn and Emlen, 1932: 31. Mabuya mabouya mabouya: Dunn, 1936: 544; Smith, 1950: 55. Mabuya mabouya: Meyer, 1966:176; Hahn, 1971: 111; Wilson and Hahn, in press. Range. -l.o'w and moderate elevations of both versants from central Mexico to Brazil; also in the Lesser Antilles. Locality 7?^cor^/5.-ATLANTIDA: Corozal, LACM 47753-54, 47756; 13 km E La Ceiba, LACM 47755. CHOLUTECA: 13.1 mi W Choluteca, UMMZ 123017. COLON: Barranca (Dunn and Emlen, 1932); Trujillo, LSUMZ 22428. COMAY AGUA: Siguatej^eque, FMNH 5063. COPAN: Copan, UMMZ 83029, AMNH 70339-40. CORTES: Agua Azul, TCWC 19211-12, AMNH 70337, MCZ 49966-67; Laguna Ticamaya, FMNH 5061; El Jaral, FMNH 5062; San Pedro Sula, FMNH 5060. EL PARAISO: Valle de Jamastran, AMNH 70380. FRAN- CISCO MORAZAN: Tegucigalpa, FMNH 5064-65; Zamorano, AMNH 70338. GRACIAS A DIOS: Patuca, USNM 20306-09; Kisalaya, LACM 16860; Kiwas Tara, LACM 16859; Tancin, LACM 47726-28. ISLAS DE LA BAHIA: Isla de Roatan, Jonesville, TCWC 21955; Isla de Utila, Utila, LSUMZ 22309; Isla de Guanaja, LSUMZ 21883. LEMPIRA: Gracias (Hahn, 1971). OLAN- CHO: 1 km WNW Catacamas, LACM 47720; 4.5 km SE Catacamas, LACM 45165, 47721-25; Rio Guampucito, 7.5 mi E Dulce Nombre de Culmf, LACM 45151. SANTA BARBARA: El Sauce, AMNH 70341. YORO: Subirana Valley, MCZ 32037-40, 38934-36, UMMZ 77848 (6); 5 km E Coyoles, LACM 47729; Coyoles, LACM 47730-52. Ecologic Distribution.-Sea. level to about 1100 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical Moist Forest, and Subtropical Dry Forest formations. Found on the forest floor in disturbed and cleared areas and under the bark of trees, logs, and fence posts. A large series was taken from bunches of bananas in a processing plant. Genus Scincella Mittleman Scincella cherriei (Cope) Mocoa cherriei Cope, 1893: 340. Scincella cherriei cherriei: Mittleman, 1950: 20. Leiolopisma assatum: Dunn and Emlen, 1932: 31. Lygosoma assatum cherriei: Stuart, 1940: 13. Lygosoma incertum Stuart, 1940: 10. (In part)^ ^The paratype of Lygosoma incertum (MCZ 38933), from Portillo Grande, Honduras, appears to us to represent Scincella cherriei. i! 1973 Turtles, Crocodilians, and Lizards of Honduras 30 i Leiolopisma cherriei cherriei: Smith, 1946: 111; Meyer, 1966: 175. I Scincella cherriei: Wilson and Meyer, 1969: 146. ' Range.— Lov^, moderate, and intermediate elevations from Veracruz, Mexico on the Atlantic versant and Chiapas, Mexico on the Pacific to Panama. Locality /?^cor<75.-ATLANTIDA: 9 mi E La Ceiba, LACM 47819; moun- ! tains above Corozal, LACM 47813-14, 47820, LSUMZ 21509-13; Lancetilla, MCZ 29397-400, 32214, TCWC 19210, AMNH 70445-46. COLON: mountains above Trujillo, LSUMZ 22453. COPAN: Copan, UMMZ 83030-31, TCWC 23637. CORTES: 1 mi W El Jaral, LACM 47815, LSUMZ 24213; 1 mi SE El Jaral, LACM 45299, 45349-51, 47816-18, 47882; 2 mi W San Pedro Sula, LACM 47821; Lago de Yojoa, MSU 4704-05; W of San Pedro Sula, FMNH 5054-59; Agua Azul, AMNH 70505-06. EL PARAISO: Arenal, LACM 20476. FRAN- CISCO MORAZAN: 9.9 km N. Tegucigalpa, LSUMZ 24180. OLANCHO: 4.5 km SE Catacamas, LACM 45156-57, 47810. YORO: Mataderos Mountains, UMMZ 77843, MCZ 38928; Subirana Valley, UMMZ 77850, MCZ 38927, 38932; 5 km E Coyoles, LACM 47811-12; Portillo Grande, MCZ 38933. Ecologic Distribution.-Sea level to 1600 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical Moist Forest, and Lower Montane Wet Forest formations. Found in the day- time on the forest floor and in leaf litter in undisturbed or slightly disturbed forest. Family Teiidae Genus Ameiva Meyer KEY TO THE SPECIES OF AMEIVA IN HONDURAS Enlarged gular scales larger than mental; generally a well-defined light medial stripe on the dorsum f estiva Enlarged gular scales much smaller than mental; never a light medial strip on the dorsum undulata Ameiva festiva (Lichtenstein) Cnemidophorus festivus Lichtenstein, 1856: 13. Ameiva festivus: Bocourt, 1874: 260. Ameiva festiva: Barbour and Noble, 1916: 473; Dunn and Emlen, 1932: 30. Ameiva festiva edwardsi: Echternacht, 1968: 152. Range. -Lov/ and moderate elevations of the Atlantic versant from the Isthmus of Tehuantepec, Mexico to Panama. Locality 7?ccort/5.-ATLANTIDA: Lancetilla, AMNH 70456; Colorado District, UMMZ 58376-77; Guaymas District, UMMZ 58378, 58382; mountains above Corozal, LACM 48072-76, 48083-85, LSUMZ 21622-27; 12 km SSE La Ceiba, KU 10195-96; Carmelina, USNM 62972. COLON: mountains above Trujillo, LSUMZ 22465; ca. 2 mi E Trujillo, LSUMZ 22499. COPAN: 12 mi ENE Copan, LACM 72084-85. CORTES: Agua Azul, AMNH 70336; Lago de 31 Contributions In Science No. 244 Yojoa, MSU 4695-703; W of San Pedro Sula, FMNH 5067-71, 5074; El Jaral, FMNH 5073; Laguna Ticamaya, FMNH 5072. GRACIAS A DIOS: Rio Coco, USNM 24527-28. OLANCHO: ca. 40 km E Catacamas, TCWC 23638. SANTA BARBARA: 5.5 mi SW El Jaral, LACM 48077-82. YORO: Mataderos Moun- tains, MCZ 38924, FMNH 21781; Subirana Valley, MCZ 38926; Portillo Grande, MCZ 38925. Ecologic Distribution.-Sea level to about 1400 m in the Tropical Moist Forest, Tropical Dry Forest, and Subtropical Wet Forest formations. Found in the daytime on the forest floor in undisturbed and edge situations. Ameiva undulata (Wiegmann) Cn.[emidophorus] undulatus Wiegmann, 1834: 27. Ameiva undulatus: Gray, 1845: 20. Ameiva undulata: Dunn and Emlen, 1932: 30; Wilson and Meyer, 1969: 146; Hahn, 1971: 111. Ameiva undulata pulchra: Stuart, 1942: 146; Smith, 1950: 55. Ameiva undulata hartwegi: Echternacht, 1968: 152. Range.-Lov/ and moderate elevations from Veracruz, Me'xico to Nica- ragua on the Atlantic versant and Nayarit, Mexico to Costa Rica on the Pacific. Locality /^ccorc/^.-ATLANTIDA: Corozal, LACM 48106-07, 48125, 72086; mountains above Corozal, LACM 48104-05, LSUMZ 21713; 2 km SE La Ceiba, KU 101228-38; 8 km SE La Ceiba, KU 101239-42; 12 km SSE La Ceiba, KU 101252-53. COLON: 0.5-3 km W Trujillo, LSUMZ 21714, KU 101243-48; 2 km E Trujillo, LACM 48108, LSUMZ 21715-17; 0.5 km SW Trujillo, LACM 72087; 1 km SSW Trujillo, KU 101249-51; Trujillo, LSUMZ 22427; ca. 2 mi. E Tru- jillo, LSUMZ 22497-98; Balfate, AMNH 58607-09. COMAYAGUA: La Liber- tad, MCZ 38923; 8.7 km S Comayagua, LSUMZ 24199. COPAN: Copan, UMMZ 83035 (16), FMNH 28525-30; Rio Higuito, ANSP 22195-98; 2 km NE Copan, LSUMZ 24622. CORTES: 1 mi W El Jaral, LACM 48113-15, 48117- 18; 1 mi SE El Jaral, LACM 48116, 48119, LSUMZ 22514; 4.5 mi ENE Villanueva, LACM 48120; 4 mi NE La Lima, LACM 48121-24; 2-3 mi W San Pedro Sula, LACM 48126-28, MCZ 29387-88; Amapa, AMNH 70458; 3 km WSW Cofradia, KU 67330-31; E side Lago de Yojoa, KU 67332; 7 km SW La Lima, KU 67333; Agua Azul, MCZ 49964. EL PARAISO: 15 km E Guadalupe, LACM 39777; Valle de Jamastran, AMNH 70333-35. FRANCISCO MORA- ZAN: Tegucigalpa, MSU 4690; Cantarranas, ANSP 22199-212; Zamorano, MCZ 49766-67, 100001-03, KU 101260-67, AMNH 70466, 70470-73; near Zamorano, AMNH 70369-79; above Agua Amarilla, AMNH 70368. GRACIAS A DIOS: Tancin, LACM 48093-94. LEMPIRA: 7 mi NNW Gracias, LACM 48109-12; Gracias, FMNH 40865; Erandique (Hahn, 1971). OLANCHO: 0.5-1 km WNW Catacamas, LACM 48086, LSUMZ 21696-99; 1-3 km NW Catacamas, LACM 48087-89, LSUMZ 21700-03; 4.5 km SE Catacamas, LACM 48090-92, LSUMZ 21704-06; 6.5 km SE Catacamas, LSUMZ 21707-08; Pataste, MSU 4691-94; 7.6 mi SW Juticalpa, LACM 45235-37. SANTA BARBARA: 1973 Turtles, Crocodilians, and Lizards of Honduras 32 4 km SW Quimistan, KU 67326-29; near Quimistan, USNM 128091-92. YORO: 2 km S Coyoles, KU 101216-26, 101254-59, 109973, 107911-14; 0.5 km N Coyoles, LACM 48095-97, LSUMZ 21709; 5 km E Coyoles, LACM 48098-103, LSUMZ 21710-11; Rancho San Lorenzo, LSUMZ 21712; Progreso, UMMZ 58375. Ecologic Distribution.-Sea. level to about 1200 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical Moist Forest, and Subtropical Dry Forest formations. Found in disturbed situations in the wetter formations, and in the forest and along streams in the drier formations during the day. Genus Cnemidophorus Wagler KEY TO THE SPECIES OF CNEMIDOPHORUS IN HONDURAS 1. With one interparietal and four parietals lemniscatus 1. With one interparietal and two parietals 2 2. Fewer than nine rows of longitudinal ventral plates; tail pinkish to orange in life motaguae 2. More than nine rows of longitudinal ventral plates; tail bluish in life deppei Cnemidophorus deppei Wiegmann Cnemidophorus Deppii Wiegmann, 1830: 28. Cnemidophorus deppii deppii: Burt, 1931: 56. ? Cnemidophorus lemniscatus lemniscatus: Dunn and Emlen, 1932: 30. Cnemidophorus deppei deppei: Meyer, 1966: 176; Echternacht, 1968: 152. Range.— and moderate elevations from Veracruz, Mexico on the Atlantic versant and Michoacan, Mexico on the Pacific to Costa Rica. Locality Records. -CnOWTECN. Respire, MCZ 49986, FMNH 5101; Ola, TCWC 19200; Choluteca, MSU 4679-81. COLON: Puerto Castilla, LSUMZ 22482. COMAYAGUA: 4 mi S Comayagua, LACM 48037-40; 11 mi NW Comayagua, LACM 48041-43. CORTES: San Pedro Sula, USNM 24367- 70; 3 km WSW Cofradia, KU 67360-71; 2 mi W San Pedro Sula, LACM 48046- 47; Lago de Yojoa, MCZ 29811; Hacienda Santa Ana, FMNH 5147, 5102-13, 5115-39, 5141-46. EL PARAISO: near Ojo de Agua, AMNH 70329, 70331-32. FRANCISCO MORAZXN: Zamorano, MCZ 49769-72, 49931; 21 km SSW Sabana Grande, LSUMZ 24623. GRACIAS A DIOS: Tancin, LACM 48023-25, LSUMZ 21628-31. INTIBUCA': 5 mi S Jesus de Otoro, LACM 48048-50. OLANCHO: between Juticalpa and Catacamas, AMNH 70330. SANTA BA'RBARA: 4 km SW Quimistan, KU 67359; 3 mi E Quimistan, LACM 48044- 45; 4 km S Cofradia, LSUMZ 24624; ca. 10 km N Santa Bairbara, LACM 45324- 25; ca. 12 km N Santa Barbara, LACM 45326-32. VALLE: San Lorenzo, FMNH 5098-100. YORO: 2 km S Coyoles, KU 101312-16, 101318-26; 6 km N Coy- oles, KU 101317; 0.5 km N Coyoles, LACM 48026-32, LSUMZ 21632-38; 5 km 33 Contributions In Science No. 244 E Coyoles, LACM 48033, LSUMZ 21639; Rancho San Lorenzo, LACM 48034- 36, LSUMZ 21640-42. Ecologic Distribution.-SQa. level to 850 m in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, Subtropical Moist Forest, and Subtropical Dry Forest formations. A diurnal species found in cleared and edge situations in the wetter formations, and in the open and in the forest in the drier formations. Cnemidophorus lemniscatus (Linnaeus) Lacerta Lemniscata Linnaeus, 1758: 209. Cnemidophorus lemniscatus: DumeTil and Bibron, 1839: 123; Echternacht, 1968: 152; Wilson and Hahn, in press. Cnemidophorus lemniscatus ruatanus Barbour, 1928: 60; Rand, 1954: 260. Cnemidophorus lemniscatus lemniscatus: Burt, 1931: 30; Lynn, 1944: 190; Dunn and Saxe, 1950: 157; Rand, 1954: 260; Meyer, 1966: 176. Range.-Co' 1961 (R. O. Schuster, #96; UCD); 2 mi N Riverton, El Dorado Co., 12 Dec. 1970 (F. G. Andrews; LACM), Berlese ex pine duff; 1.6 mi W Quintetle, El Dorado Co., 4 Apr. 1971 (F. G. Andrews; LACM); 5.6 mi SE Dorrington, Cal- averas Co., 16 June 1959 (L. M. Smith & R. O. Schuster, #142; UCD); Forest- hill, Placer Co., 6 June 1959 (F. C. Raney, #139; UCD); Tahoe City, Placer Co., 3 July 1961 (H. L. McKenzie, #112; UCD); Michigan Bluff, 3500', Placer Co., 24 June and 5 Nov. 1962 (R. R. Snelling; LACM); Bieber, Lassen i , Co., 17 Apr. 1958 (R. W. Gerhardt, #101; UCD); Post Pile Camp, 5800', | Tehama Co., 30 Aug. 1960 (R. O. Schuster & A. A. Grigarick; UCD); Chico, ji Butte Co., 4 Sept. 1958 (L. M. Smith & R. O. Schuster, #192; UCD); 2 mi w|. Springville, Tulare Co., 13 May 1959 (L. M. Smith #86; UCD); 5 mi S Gorman, | Los Angeles Co., 25 Mar. 1959 (H. L. McKenzie, #34; UCD). | DISCUSSION. When adequate samples, including entire nest populations, | become available it may prove that what is here treated as a single, highly j variable species is actually two species. The type series consists of a uniformly ^ similar assemblage in which the integument is moderately to strongly shiny on i 1973 Studies on California nts 25 the thorax and petiolar nodes. The thoracic rugulae are coarse and well de- fined. The first gastric sternite has a number of widely spaced, rather coarse longitudinal striae and the corresponding tergite has numerous coarse striae basally, these about 0.2 mm long. The nodes of the petiole and postpetiole are conspicuously longitudinally rugulose. In addition to the types, specimens from Michigan Bluff (24 June 1962), Strawberry and Post Pile Camp fall into this type of sculpturation. Most of the remaining specimens are, on the average, smaller, with thor- acic rugulae quite fine and the interspaces closely punctulate, hence only slightly shiny. The first gastric sternite is very finely and closely longitudinally striate with a few punctulae between striae, especially toward the sides. The base of the first tergite is closely and finely striate, the striae 0.13 mm long, or less. These closely punctulate specimens also usually have short, triangular propodeal spines, but the series from the Tehachapi Mountains has the spines about as long as in the types. A few specimens (Quintetle, Foresthill) appear to be intermediate. The Quintetle series have short propodeal spines, the interrugular spaces are mod- erately shiny and the first sternite, while mostly finely striate, has a few coarse striae. The Foresthill specimens are as shiny and coarsely rugulose as the types, the propodeal spines are long, but the sternal striae are fine and punctu- late between striae at the sides. Among the species with sculptured gastric segments S. dyscheres most closely resembles S. punctatoventre and S. californicum; S. punctatoventre may be immediately separated by the shape of the median clypeal lobe, which is produced and appears truncate in frontal view. In S. californicum the rugulae on the sides of the thorax are much reduced and very obscure because of the fine, dense punctulation; this is especially evident on the mesopleura which usually is without rugulae and on the propodeal sides which may have a few rugulae below the spiracle but none above it. In this species, too, the basal one-fourth or more of the first tergite is finely punctulate and without evident striations except the usual ones at the basal articulation. Finally, the sculpture of the first sternite consists of fine, dense punctulation. Those specimens of S. dyscheres which possess dense punctulae on the sides of the thorax retain the coarse, conspicuous rugulae on the mesopleura and propodeal sides and the base of the first tergite is conspicuously striate, the articular striae inter- spersed with longer, finer supplementary striae. The sternite is striate, the striae coarse or fine and punctulae may be present laterally between striae. Stenamma occidentale M. Smith Figures 30, 31, 36, 37, 60 Stenamma neoarcticum Mayr, 1886. Zool.-Bot. Gesell. Wien. 36:454.$^ (not ?). ? Stenamma (Stenamma) westwoodi neoarcticum, Emery, 1895. Zool. Jahrb. Abt. f. Syst. 8:299-300. 9 9 . 26 Contributions in Science No. 245 Stenamma (Stenamma) neoarcticum, Ford, 1901. Soc. Ent. Belg. Ann. 45:347. $ $ . Stenamma neoarcticum, Wheeler, 1903. Psyche 10:165-166. 9 . Creigh- ton, 1950. Bull. Mus. Comp. Zool. 104:, 135, 138. 9 9 5. Stenamma occidentale M. R. Smith, 1957. Amer. Midi. Nat. 57:142, 146- 150. 9 95 . Most of the confusion between this species and S. diecki has been discussed under that species and need not be repeated. Material which, in my opinion, belongs to S. occidentale has been examined from the following States: Arizona, New Mexico, Colorado, Utah, Idaho, California, South Dakota, Oregon and Figures 28, 29: S. dyscheres, worker, lateral view and frontal view of head; Figures 30, 31: 5’. occidentale, worker, same; Figures 32, 33: S. sequoiarum, worker, same. Figures by Ruth Ann DeNicola. 1973 Studies on California Ants 27 Figures 34, 35; 5'. dyscheres, female, lateral view and frontal view of head; Figures 36, 37; S. occidentale, female, same (cotype of S. neoarcticum Mayr); Figures 38, 39; S. sequoiarum, female, same. Figures by Ruth Ann DeNicola. 28 Contributions in Science No. 245 Washington. Although there are no records from Nevada, the ant is likely to be j found there. ' The following California specimens have been examined: 1?, \$, “Calif. | Oct. 31.85” (Cotypes of S. neoarcticum Mayr; USNM): Auburn, Placer Co., f 12 Mar. 1958 (L. M. Smith & R. O. Schuster; UCD); Davis, Yolo Co., 2 May L 1932 (T. W. Cook; LACM) (Note: These are the specimens from Davis re- | corded by Cook (1953) as S. brevicorne heathi). | I Stenamma sequoiarum Wheeler j Figures 32, 33, 38, 39, 59 Stenamma brevicorne sequoiarum Wheeler, 1917. Proc. Amer. Acad. | Arts. Sci. 52:520. ? $ . ji Stenamma diecki sequoiarum, Creighton, 1950. Bull. Mus. Comp. Zool. | 104: 136, 137. 9 . | Stenamma sequoiarum, M. Smith, 1957. Amer. Midi. Nat. 57: 142, 156- J 158. 9 9. I The most important characteristics are adequately covered in the above | key and in the redescription of Smith (1957), who studied the types from Mt. Tamalpias and specimens from Sam P. Taylor State Park, both in Marin County, California. I have seen specimens from the following additional Cali- fornia localities: Steven’s Creek, Santa Clara Co.; 28 and 31 m W of Paso Robles, San Luis Obispo Co.; Pine Cyn., Monterey Co.; Donomore Meadow, i 5600', Siskiyou Co. The Davis, Yolo Co., record cited by Cook (1953) cannot be verified as there are no such specimens under this name in his collection. Group III Stenamma exasperatum, new species j Figures 40, 41, 46, 47, 63 DIAGNOSIS. Eyes of worker with three facets in greatest diameter; first tergite densely and finely punctulate, with basal longitudinal striae and scat- tered coarse piligerous punctures; first sternite densely, finely punctulate; scapes and tibiae with abundant fully erect hairs. WORKER. Measurements. HL 0.87-0.93 (0.93); HW 0.70-0.77 (0.77); SL 0.60-0.66 (0.66); WL 1.00-1.13 (1.13); PW 0.47-0.53 (0.53). Head. A little longer than broad. Cl 81-83 (82), longer than scape, a little narrowed toward occiput. Eyes small, with three facets in greatest diameter, OMD 2.16-2.33 (2.33) x EL; OI 10-1 1 (10). Scape stout, a little shorter than HW, SI 86-89 (87), distinctly thickened near apex. Mandible with longitudinal rugulae over most of length, shiny and sparsely punctate, cutting margin with six or seven teeth. Median lobe of clypeus not exceeding clypeal margin, longitudin- ally depressed and laterally carinulate, without transverse carinula. Clypeus and depressed frontal area shiny; frontal lobes roughened, slightly shiny; 1973 Studies on California Ants 29 remainder of cephalic dorsum slightly shiny, finely punctulate, evenly and coarsely reticulo-rugose, with numerous coarse piligerous punctures. Thorax. Pronotal neck slightly shiny, densely punctulate and with a few coarse punctures; laterally reticulo-rugose, interspaces punctate and slightly shiny; promesonotum regularly reticulo-rugose, interspaces shiny, with scat- tered obscure punctures. Mesopleura and sides of propodeum coarsely longi- tudinally rugose, interspaces closely punctate and slightly shiny. Metanotal depression broad and deep. Propodeum basally with a poorly defined trans- verse welt; basal face closely punctulate and with irregular rugulae tending to form reticulae; spines short, sharp, about two-thirds as long as distance between them; declivity closely punctulate and slightly shiny on upper half, lower half shiny and impunctate. Petiole. Dorsal face of anterior peduncle about as long as anterior face of node; summit of node narrowly rounded, anterior and posterior faces about equal; peduncle without ventral tooth anteriorly. Postpetiole, from above, about as broad as long, sides slightly convex. Sides and venter of petiole and postpetiole dull, densely punctulate; nodes sharply reticulo-rugose and dull, with closely punctulate interspaces. Gaster. First tergite densely punctulate and dull on basal half, or more; entire segment with numerous coarse, piligerous punctures; punctulate area with numerous fine longitudinal rugulae, forming reticulae toward base. First sternite similar, but striatopunctulate on basal half, rugulae less conspicuous. Remainder of gaster as usual in genus. Pilosity. Scapes, femora and tibiae with abundant fully erect whitish hairs of even length; cephalic and thoracic dorsa with numerous evenly spaced fully erect hairs of uniform length. Gastric hairs a little denser than in S. heathi. Color. Yellowish ferruginous, median flagellar segments a little darker; mandibular margins darker; antennal club and tibiae yellowish. FEMALE. See Discussion below. MALE. Unknown. TYPE MATERIAL. Holotype and three paratype workers; Calaveras Big Trees, Calaveras Co., CALIF., June 1952 (R. R. Snelling), under stone in Sequoia grove, all specimens in LACM. ETYMOLOGY. Exasperatum, L., so named to express my feeling upon discovering that these specimens, thought to be S. heathi, represent still another species. ADDITIONAL MATERIAL. Three workers, Mariposa Camp, 5600', Yosem- ite National Park, 3 August 1952 (E. O. Wilson; USNM) “under rock, pine forest”. DISCUSSION. This species is most closely related to S. heathi with which it shares details of clypeal structure and gastric sculpture. The sculpturation of the head and thorax are similar but in S. exasperatum both are more clearly reticulate. The reticulae become less defined on the sides of the head and thor- ax of S. heathi, usually replaced by a series of irregular longitudinal rugulae. Although the hairs of the scapes and tibiae are fully erect in S. exasper- 30 Contributions in Science No. 245 atum studied, there may be some variation not now evident. In S. heathi these appendages usually have the hairs strongly decumbent, but in some speci- mens the hairs may be subdecumbent and with a few which are suberect. The hairs of the hind tibiae apparently are consistently decumbent in nH Simon on Simncon T Printed in I.os Aneeles Gnlifornin hv Andercon Ritehii NUMBER 246 JUNE 30,1973 C. xLSi^3 LUMINESCENT SILHOUETTING IN STOMIATOID FISHES By William T. O’Day CONTRIBUTIONS IN SCICNCE NATURAL HISTORY MUSEUM • LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- ^ | tists on an exchange basis. Copies may also be purchased at a nominal price. f Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los !: Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. | INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life ! or Earth Sciences. Acceptance of papers will be determined by the amount and char- ! acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent i specialists outside the Museum. | Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- i j cate that the primary type has become the property of a scientific institution of their j choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition I (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. ^ (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not \ I be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- ? | able footnotes must be typed on separate sheets. Several of one kind may be placed on ■ I a sheet. (6) An abstract must be included for all papers. This will be published at the i head of each paper. (7) A Spanish summary is required for all manuscripts dealing ' I with Latin American subjects. Summaries in other languages are not required but are | strongly recommended. Summaries will be published at the end of the paper. (8) A A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of , | manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic I prints of good contrast. Original illustrations and art work will be returned after the | manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor LUMINESCENT SILHOUETTING IN STOMIATOID FISHESi By William T. O’Day^ Abstract: The luminescence which silhouettes three sto- miatoid fishes — Stomias atriventer, Idiacanthus antrostomus, and Chauliodus macouni — has been seen repeatedly in adrenalin-stim- ulated specimens. It has been photographed in S. atriventer. The silhouetting luminescence derives from minute, unpigmented or- gans, found along the dorsal and ventral margins of these fishes and on all the fins. These luminous organs are composed of in- nervated aggregations of secretory cells. Stomiatoid fishes inhabit the depths of the oceanic midwaters. Among their remarkable adaptations is their multiplicity of luminescent organs (Brauer, 1908; Gibbs, 1964, 1969; Morrow, 1964a, 1964b, 1964c; Morrow and Gibbs, 1964; Goodyear and Gibbs, 1969). I report new observations of biolumines- cence in three species of stomiatoid fishes— atriventer Garman, 1889, Idiacanthus antrostomus Gilbert, 1890, and Chauliodus macouni Bean, 1890 (Families Stomiatidae, Idiacanthidae, and Chauliodontidae, respectively, all belonging to the Suborder Stomiatoidei, Order Salmoniformes). The classifica- tion referred to is the one proposed by Greenwood et al. ( 1966). These fishes can be outlined by bioluminescence from numerous organs of the simple, un- pigmented type. The organs are composed of innervated masses of secretory cells which emit light after stimulation with adrenalin. Materials and Methods Fishes were collected in deep waters off southern California by the R/V Velero IV of the University of Southern California. Some were also obtained during one cruise of the R/V Alpha Helix of the Scripps Institution of Ocean- ography. Observations of luminescence were made in the darkroom on board ship. Tactile stimulation was occasionally effective in eliciting luminescence from these three species but my usual method of stimulation was to add a small quantity of adrenalin solution (roughly 10 ml of 0.01% to 0.1% adren- alin in seawater) to 1-4 liters of cooled seawater in a tank containing the speci- men. Sometimes, however, I applied a few drops of adrenalin solution directly on the skin of the fish. I also injected Stomias intraperitoneally and intramus- 1 Review Committee for this Contribution Jules M. Crane, Jr. Robert H. Gibbs, Jr. Robert J. Lavenberg ^Dept. Biol. Sciences, University of Southern California, Los Angeles, Calif. 90007; and Research Associate in Ichthyology, Natural History Museum of Los Angeles County, Los Angeles, Calif. 90007 1 2 Contributions in Science No. 246 cularly with adrenalin (about 14 to Vi ml of 0.1% solution of adrenalin in phosphate buffer, 0.15 M, pH 7.5), while subsequently injecting controls with an equivalent volume of buffer alone. Photographs of bioluminescence were made by placing the glowing fish directly on a large sheet of Kodak Plus-X Pan film, which had been sealed inside a plastic bag, for about 30 to 60 seconds. Specimens I Stomias atriventer Three specimens, 149 to 228 mm standard length (SL), were used for most of the histology. Bioluminescence was observed in 23 specimens, 137 to 250 mm SL. | Idiacanthus antrostomus One specimen, 95 mm SL, was used for histology. Bioluminescence was | observed in eight specimens, 258 to 335 mm SL. k Chauliodus macouni Three specimens, 71 to 230 mm SL, were stimulated with adrenalin with- j out eliciting a luminous response and subsequently used for histology. These I specimens, however, were nearly dead. Another specimen, 115 mm SL, did luminesce when stimulated with adrenalin. Appearance of the Luminous Organs I! Each of these fishes has several kinds of luminescent organs. Large body photophores form ventral and ventrolateral rows. Prominent light organs lie near the eye and on close inspection very small photophores appear in various areas all over the body. In Stomias and Idiacanthus a luminous barbel trails ( from the lower jaw. All of these luminescent organs are easily visible in pre- | served specimens. However, an additional kind of luminescent organ is in- I conspicuous in preserved specimens of Stomias and Chauliodus, although it is ! quite evident in undamaged living specimens. Brauer (1908) referred to it as ' the simple, unpigmented type since no reflector or pigment sheath is present. | In both Stomias and Chauliodus luminous organs of this type are embedded in a gelatinous sheath that envelops the fish. Spherical to ovoid bodies of simple, unpigmented luminous tissue form an irregular row along the dorsal and ven- tral margins of Stomias (Fig. la) and they extend along the rays of all the fins (Fig. lb), in Chauliodus spherical bodies of luminous tissue are dis- tributed along the dorsal and ventral margins of the fish and on all the fins. By contrast, Idiacanthus lacks the gelatinous sheath and instead has a row of chevron-shaped patches of luminous tissue along its dorsal margin and rows of small patches of luminous tissue along its ventral surface, with similar tissue along the rays of all the fins. In all three species this luminous tissue appears bright pink in life. 1973 Luminescent Silhouetting in Stomiatoid Fishes 3 Figure la. Minute balls of naked luminous tissue along the ventral margin of the body in a specimen of Stomias atriventer 149 mm SL. Photographed under a dis- secting microscope. The scale equals one mm. Figure lb. Luminous tissue along two branches of a ray from the anal fin of the same specimen. Photographed under a compound microscope with transmitted light. The scale equals 0.5 mm. Observations of Luminescence There have been no published observations of luminescence of this tissue in either Stomias or Idiacanthus. However, P. J. Herring of the National Insti- tute of Oceanography in England has made some related observations (per- sonal communication). There is a single report of luminescence from the spherical bodies of tissue on Chauliodus sloani by Skowron (1928), who saw them emitting a bluish light. He also saw blue luminescence from the large body photophores of this fish. It remained alive in a tank for 45 minutes, dur- ing which a steady pale bluish luminescence issued from the ventral region when it remained undisturbed. Touched with a glass rod, the fish emitted pulses of luminescence which lit up the contours of its body. The unpigmented luminous bodies found on Stomias, Idiacanthus, and Chauliodus emit light when the fishes are placed in a weak solution of adren- alin in seawater. Injection or topical application of adrenalin solution was also effective in stimulating luminescence. Injection of adrenalin elicited lumines- cence from the ball-shaped bodies of Stomias without noticeable delay near the point of injection; then the luminescence spread in the course of several seconds to the rest of the ball-shaped bodies on the dorsal and ventral margins of the fish and on the fins. Tactile stimulation following injection of adrenalin 4 Contributions in Science No. 246 spread the glow more rapidly. Topical application of adrenalin elicited lumi- nescence rapidly from the ball-shaped bodies around the site of stimulation. The glow spread gradually to other parts of the body. The Stomias I have ob- served usually luminesced for several seconds to about a minute. Repeated stimulation with adrenalin sometimes evoked weaker subsequent responses. I found on one occasion that a few of the luminescing ball-shaped bodies of Stomias continued to glow for a few seconds after being removed from the fish. The luminescence I have seen in Idiacanthus lasted only a few seconds. In rare cases, merely touching Stomias or Idiacanthus caused them to light up brightly. Adrenalin stimulates not only the small unpigmented luminous organs but also the large body photophores and the postorbital photophores. No emission spectra have been recorded for the luminescence of these three stomiatoid fishes. To my dark-adapted eyes, however, the color of the luminescence of Stomias was usually yellowish, although one especially bright- glowing specimen looked distinctly bluish. In Idiacanthus, too, the lumines- cence has sometimes appeared yellowish, at other times blue-green. Figure 2a. Section of one of the minute ball-shaped bodies of Stomias atriventer where a nerve (arrow) approaches it. Figure 2b. Section of one of the minute ball-shaped bodies of Chauliodus macouni. Specimens fixed with glutaraldehyde and osmium, embedded in Epon, and stained with toluidine blue. The scale for both figures equals 20 microns. 1973 Luminescent Silhouetting in Stomiatoid Fishes 5 Histological Structure The suggestion that the ball-shaped bodies and masses of tissue around the fins are luminescent was put forward many years ago by several classical histologists, especially Brauer (1908), who noted their similarity to tissue in the photophores, barbels, and subocular light organs of these fishes. Micro- scopically the luminous bodies of Stomias and Idiacanthus appear similar: they are solid balls of secretory cells in which the secretory product is usually intracellular. A thin layer of a second kind of cell— the “bindegewebige Hiille” of Brauer— encloses the mass of secretory cells. The outermost layer of each ball consists of a well-developed basal lamina. Nerves terminate at the surfaces of these bodies but do not penetrate them (Fig. 2a). The spherical luminous bodies of Chauliodus elaborate their secretory product in a peripheral layer of cells, after which it accumulates in a central lumen (Fig. 2b). When fresh undamaged specimens of Stomias or Chauliodus are viewed with a dissecting microscope, the nerves appear as white threads running through the gelatinous sheath, their branches going to the luminous bodies. Figure 3 shows a cross section of such a nerve fiber from the gelatinous sheath of Chauliodus which contains several myelinated axons of varying diameter. The gelatinous sheath of Chauliodus and Stomias is really a highly hy- Figure 3. Cross section of a nerve fiber in the mucous sheath of Chauliodus macouni, a specimen 230 mm SL. The myelin sheaths of the axons appear black. Fixed with glutaraldehyde and osmium, embedded in Epon, and stained with toluidine blue. The scale equals 20 microns. 6 Contributions in Science No. 246 drated part of the dermis, as Brauer ( 1908) realized. In addition to the spheri- cal or ovoid bodies, it contains nerves, blood vessels, a sparse network of fibro- blasts and collagen, and mucoid substance. In life the mucous sheath is per- fectly transparent. It is well developed along the dorsal and ventral margins of the body. However, it is very thin or absent on the lateral surfaces of the body. A thin epithelium lies over the mucous sheath and the other exposed parts of the body. The secretory nature of these luminescent bodies and their pink appear- ance in life are properties that they share with the large, serially arranged photophores on the ventral part of the body. The serial ventral photophores, however, lack innervation according to Brauer’s (1908) study of Idiacanthus fasciola, Chauliodus sloani and C. pammelas, and Stomias valdiviae (=^ S. affinis). Bassot (1966) found no innervation in his study of the serial ventral photophores of a species of Chauliodus with the light and electron microscope. The serial ventral photophores are also considerably more elaborate in struc- ture (Fig. 4), being composed of pigment sheath, reflector, and at least two kinds of secretory cells (Bassot, 1966). The position of the photophores on the Figures 4a and 4b. Transverse section of one of the large body photophores of Chauliodus macouni, viewed with normal brightfield illumination and between crossed polarizers, respectively. The crystals of the reflector are strongly birefringent. Fixed with glutaraldehyde, embedded in glycol methacrylate, stained with acid fuchsin and toluidine blue. The scale equals 100 microns. 1973 Luminescent Silhouetting in Stomiatoid Fishes 7 body and their association with pigment sheath and reflector ensure that their luminescence is ventrally directed. The ventral position of the large serial photophores and the ventral direc- tion of their luminescence have been explained in terms of the hypothesis that ventral luminescence in midwater animals serves to camouflage them, when they are viewed from below by potential predators against the dimly lit upper layers of water (Fraser, 1962; Jerzmanska, 1960; Clarke, 1963; McAllister, 1967; Hastings, 1971). Camouflage does not seem a likely function, however, for the naked luminous tissue on the fins and on the dorsal and ventral margins of these fishes. This kind of luminescent organ seems to do just the opposite: to render the outline of the fishes visible (Fig. 5). Luminescent silhouetting may aid them in mating, spacing themselves out as they hunt, maintaining conspecific aggregations, warning potential predators of their own formidable size, or perhaps allowing them to escape from predators by temporarily blind- ing them. These functions, however, remain speculative. Acknowledgments I especially thank Mr. Jules Crane and Dr. Basil Nafpaktitis. I also thank Dr. Robert Bils, Dr. Hector Fernandez, Dr. Jay Savage, Dr. L. Daniel Schaffer, Dr. Bernard Strehler, and Dr. Russel Zimmer. I acknowledge partial sup- Figure 5. Drawing of the deep-sea fish Stomias atriv enter Carman, and photograph of a specimen taken by the light from its own bioluminescence following stimula- tion with adrenalin. The specimen was 192 mm SL. 8 Contributions in Science No. 246 port from the following sources: NIH Predoctoral Fellowship No. 5 FOl GM45 106-02; Biomedical Sciences Support Grant RR-07012-04 from the General Research Support Branch, Division of Research Resources, Bureau of Health Professions Education and Manpower Training, National Institutes of Health; and the Society of the Sigma Xi. The vessels R/ V Velero IV of the University of Southern California and R/ V Alpha Helix of the Scripps Institu- tion of Oceanography were supported by funds from the National Science Foundation. Literature Cited Bassot, J. M. 1966. On the comparative morphology of some luminous organs, p. 557-610. In F. H. lohnson and Y. Haneda (Eds.) Bioluminescence in progress. Princeton Univ. Press, Princeton. Brauer, a. 1908. Die Tiefsee-Fische. 2. Anatomischer Teil. Wiss. Erg. Deut. Tief- see-Exped. “Valdivia” 1898-1899, 15 (2): 1-266. Clarke, W. D. 1963. Function of bioluminescence in mesopelagic organisms. Na- ture 198 (4887): 1244-1246. Fraser, J. H. 1962. Nature adrift. G. T. Foulis, London. Gibbs, R. H., Ir. 1964. Family Idiacanthidae. Fishes of the Western North Atlantic. Mem. Sears Found. Mar. Res. 1 (4) : 512-522. . 1969. Taxonomy, sexual dimorphism, vertical distribution, and evolution- ary zoogeography of the bathypelagic fish genus Stomias (Stomiatidae). Smith- sonian Contrib. Zool. No. 31. Goodyear, R. H., and R. H. Gibbs, Jr. 1969. Ergebnisse der Forschungsreisen des FFS Walter Herwig nach Siidamerika. X. Systematics and zoogeography of stomiatoid fishes of the Astronesthes cyaneus species group (family Astro- nesthidae), with descriptions of three new species. Arch. Fischereiwiss. 20 (2/3): 107-131. Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bull. Amer. Mus. Nat. Hist. 131 (4) : 339-456. Hastings, J. W. 1971. Light to hide by: Ventral luminescence to camouflage the silhouette. Science 173: 1016-1017. Jerzmanska, a. 1960. The structure and biological significance of light organs in the Teleostei. Przeglad Zoologiczny 4: 112-118. In Polish. Not seen. McAllister, D. E. 1967. The significance of ventral bioluminescence in fishes, J. Fish. Res. Bd. Canada 24 (3) : 537-554. Morrow, J. E., Jr. 1964a. Family Chauliodontidae. Fishes of the Western North Atlantic. Mem. Sears Found. Mar. Res. 1 (4) : 274-289. . 1964b. Family Stomiatidae. Ibid. 1 (4) : 290-310. . 1964c. Family Malacosteidae. Ibid. 1 (4): 523-549. , AND R. H. Gibbs, Jr. 1964. Family Melanostomiatidae. Ibid. 1 (4): 351- 511. Skowron, S. 1928. Uber das Leuchten des Tiefseefisches, Chauliodus sloani. Biologisches Zentralblatt 48 : 680-685. I Accepted for publication May 17, 1973 I ''i' I '.lii..' Printed in Los Angeles, California by Anderson, Ritchie and Simon on Simpson Lee Recoye^^ NUMBER 247 JUNE 30 1973 C -a PRAEKOGIA CEDROSENSIS, A NEW GENUS AND SPECIES OF FOSSIL PYGMY SPERM WHALE FROM ISLA CEDROS, BAJA CALIFORNIA, MEXICO By Lawrence G. Barnes CONTRIBUTIONS IN SCICNCE NATURAL HISTORY MUSEUM • LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. 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All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor PRAEKOGIA CEDROSENSIS, A NEW GENUS AND SPECIES OF FOSSIL PYGMY SPERM WHALE FROM ISLA CEDROS, BAJA CALIFORNIA, MEXICO^- ^ By Lawrence G. Barnes^ Abstract: The new genus and species, Praekogia cedrosen- sis, is an extinct fossil pygmy sperm whale. Its known osteology is based upon a cranium without the rostrum, recovered from marine sedimentary rocks identified as the Almejas Formation on Isla Cedros, Baja California, Mexico. The geologic age of Prae- kogia cedrosensis is concluded to be early Pliocene in the broad sense. The base of the Almejas Formation, stratigraphically be- low the stratum that produced Praekogia, rests unconformably upon Temblor-age Miocene rocks, and mollusk-bearing beds of middle to early late Pliocene age overlie the site. Praekogia is an ideal antecedent for the living pygmy sperm whale, Kogia Gray with a less telescoped braincase and fewer cranial speciali- zations. Introduction During the summers of 1964 and 1965, members of the Department of Geological Sciences at the University of California at Riverside were engaged in extensive collecting in late Tertiary marine sedimentary deposits on Isla Cedros which lies olf the western coast of Baja California, Mexico. The existence of vertebrate fossils was brought to the attention of vertebrate pale- ontologists by Frank H. Kilmer of Humboldt State University, Areata, Cali- fornia, who has been studying the geology and invertebrate paleontology of Isla Cedros. In 1964 Kilmer, with David P. Whistler and George Jefferson, of the University of California at Riverside, travelled to Isla Cedros and col- lected the fossil skull of the pygmy sperm whale that forms the basis of this paper. The following year (1965) Kilmer was joined by Whistler and Richard H. Tedford from the University of California at Riverside and Juan Felix from the University of Baja California at Ensenada. They made a large collec- tion of fossil marine vertebrates, including one of the most diverse fossil ceta- cean assemblages known from the Pacific coast of North America (Barnes, 1972: 114-264). lA contribution from the University of California Museum of Paleontology and the Natural History Museum of Los Angeles County. Research supported in part by the University of California Annie M. Alexander Fund. ^Review Committee for this Contribution Robert L. Brownell Albert C. Myrick, Jr. David P. Whistler Frank C. Whitmore, Jr. ^Associate Curator of Vertebrate Paleontology, Natural History Museum of Los Angeles County, Los Angeles, California 90007 1 2 Contributions in Science No. 247 Marine sediments of Miocene and Pliocene age are exposed at the south- ^ east end of Isla Cedros, Baja California, and near the middle of the east side of the island (Hanna, 1927). The younger parts of these sediments have been I|! called the Salada Formation (Hertlein, 1925) and the Almejas Formation |j (Durham and Allison, 1960), both of which are generally regarded as Plio- 5' cene in age and have their type sections on the mainland of Baja California. | The name Salada Formation was considered not applicable for the younger I sediments on the island by Jordan and Hertlein ( 1926) and the name Almejas I Formation is used in this paper. The fossil vertebrates collected in 1964 and 1965, including the pygmy j| sperm whale studied herein, were quarried from the exposures in deep canyons | and on ridges near the base of the escarpment that forms the northwest wall ! of Valle Blanca at the southeast end of Isla Cedros. Here, about 800 feet of | lightly colored sands and gravels, of Pliocene age, unconformably overlie | Miocene strata (Luisian age) of the Tortuga Formation (Hanna, 1927). The ! geology and invertebrate paleontology of the island have been studied by sev- j eral writers (Hertlein, 1925; Jordan and Hertlein, 1926; Hanna, 1927; Beal, 1949) and the Pliocene sediments were correlated with mainland exposures of the Almejas Formation at Bahia de las Tortugas (Mina, 1957; Durham and , Allison, 1960) and with the San Diego Formation, which has an age of mid- j die Pliocene to early late Pliocene (Jordan and Hertlein, 1926). i From their studies, Jordan and Hertlein concluded that the Almejas in- vertebrate fauna on Isla Cedros contained a mixture of some cool water species and mostly warm water species, with an ocean water temperature like that prevalent today around Isla Cedros. The marine vertebrates from the Almejas Formation, however, are not associated with the invertebrates. All the invertebrates occur in algal con- glomerate beds stratigraphically above the basal 100 to 120 feet of strata bearing the vertebrates. Kilmer has suggested (in litt. to Charles A. Repen- ning) that an unconformity may lie above the yertebrate-bearing part of the formation. Therefore the cetaceans must be considered older than the age determination of middle to early late Pliocene based on invertebrates in over- lying strata, and younger than the Luisian age placed on the disconformably underlying Miocene beds. Howard (1971) has recently presented a study of the fossil birds that are part of the marine vertebrate assemblage from the Almejas Formation. She concluded that the birds appear to be early Pliocene in age, and are different from those comprising late Miocene and late Pliocene avifaunas. The present paper is the first to describe a mammal from the formation. The other ceta- ceans will be described in other studies in preparation. The pygmy sperm whale described here is an extinct genus and species, represented by a braincase, and is the first fossil pygmy sperm whale cranial material reported in the literature. Within the descriptive parts of this report I have made extensive com- 1973 New Genus and Species of Fossil Pygmy Whale 3 parisons with both living species of pygmy sperm whales, Kogia breviceps Blainville, 1838 and Kogia simus Owen, 1866 (see Handley, 1966). For the purposes of comparison with Praekogia the two living species of Kogia Gray, 1846 are almost identical. This does not imply the invalidity of either living species, but instead, indicates that the fossil skull differs from those of both living species in the same characters. Morphological comparisons will there- fore be made at the generic level. I have referred extensively to the published articles dealing with the anatomy of modern Kogia presented by Schulte (1917), Kernan and Schulte (1918), Raven and Gregory (1933), and Handley (1966), although I have not in all cases used the anatomical terminology used by any one of these writers. Abbreviations The following abbreviations are used in text: LACM — Natural History Museum of Los Angeles County, Vertebrate Pale- ontology Section UCMP — University of California, Museum of Paleontology. UCR — University of California at Riverside, Department of Geological Sciences mm — millimeter cm — centimeter The following abbreviations are used illustrations: A1 — alisphenoid Bo — basioccipital Fr — frontal Ju — jugal Mx — maxilla Oc — occipital Pmx — premaxilla Pt — pterygoid Sq — squamosal Vo — vomer for bones and anatomical structures in aon — antorbital notch cc — carotid canal earn — external auditory meatus fhy — hypoglossal foramen fio — infraorbital foramen fop — orbital fissure fp — falcate process of basioccipital fmx — maxillary foramen g — glenoid fossa ji — jugular incisure n — narial passage occ — occipital condyle pp — paroccipital process Acknowledgments For their assistance in recording and interpreting the geologic and geo- graphic source of the specimen I thank Richard H. Tedford, Frank H. Kilmer, and David P. Whistler. The cooperation and generosity of the former director of the Instituto de Geologia of the Universidad Nacional Autonoma de Mexico Ing. Guillermo P. Salas, the former acting director Luis Blasquez L., and the Mexican government are hereby acknowledged and recognized as having made possible the expeditions from the University of California at Riverside. Addi- 4 Contributions in Science No. 247 tionally, the assistance of personnel from the University of Baja California at Ensenada is gratefully acknowledged. I thank Richard H. Tedford and Michael O. Woodburne for the privilege of studying the cetacean fossils from Isla Cedros that are housed at the Uni- versity of California, Riverside. For their special assistance in obtaining mate- rials I thank George Jefferson, Jacquelin Schoenwald, Robert T. Orr, and Rob- ert E. Jones. The line drawings were prepared by J. Patricia Lufkin (Figs. 1, 3, 5), staff artist in UCMP and by myself (Figs. 2, 4, 6). The photographic plates (Figs. 7-10) were prepared by Lawrence Reynolds of LACM. The Spanish summary was prepared by Alvaro Mones of the Museo Nacional de Historia Natural in Montevideo, Uruguay. Systematics Family Physeteridae Gray, 1821 Subfamily Kogiinae Gill, 1871 Praekogia, new genus Diagnosis of ge/iM^.-Kogiinae with small skull, slightly more than one- half the size of that of adult Kogia breviceps Blainville, 1838, cranium not fore- shortened (telescoped) as in Kogia Gray, 1846, lateral maxillary crests high and with sharp margins, not rounded and inflated, forming nearly straight lateral margins of the supracranial basin, left premaxilla wraps around poste- rior side of left naris and joins in formation of sagittal facial crest in supra- cranial basin, left margin of this crest less developed than in Kogia, right mar- gin of right premaxilla not forming a crest as in Kogia with deep fossa in its position, temporal fossa large with anteroposterior dimension greater than vertical dimension, lateral surface exposure of jugal between maxilla and supraorbital process of frontal very slender, bone texture of skull not inflated and porous as in Kogia. Type species. -Praekogia cedrosensis X Praekogia cedrosensis, new species Figures 1-10 Diagnosis of species. Sdccae as that for the genus until further species of Praekogia are described. Holotype.-\JCK 15229, an incomplete cranium lacking the rostrum, oc- cipital crest, pterygoids, and with the left zygomatic process of the squamosal separated from the braincase. > Type locality. -\JCK RV-7315, on the ridge west of “Arroyo Delphin,” the first prominent drainage system to reach the shoreline north of the point where Valle Blanca reaches the shoreline, south of the cannery village, Isla Cedros, Baja California, Mexico. ■ New Genus and Species of Fossil Pygmy Whale 5 !' 1973 Formation and age.-Almejas Formation, probably early Pliocene in the I sense of Weaver et al., (1943). The holotype of Praekogia cedrosensis was collected from an ochre-yellow sand designated as unit O in Richard H. Ted- i ford’s field notes of 1965. The horizon is about 6 feet stratigraphically below the first and lowest prominent pebble conglomerate in the Almejas Formation I as exposed at locality UCR RV-73 15. This pebble conglomerate forms a prom- inent break in slope above the arroyo below. The site is also located approxi- i mately 120 feet stratigraphically above the base of the Almejas Formation I (Tedford’s field notes, 1965). Below the base of the Almejas Formation unconformably lies the Mio- cene Tortuga Formation of Luisian age (Hanna, 1927). Molluscan fossils have been collected higher stratigraphically in the Almejas Formation than the type locality of Praekogia cedrosensis. They have been favorably compared with Mollusca from the San Diego Formation which is traditionally considered to be middle Pliocene to early late Pliocene in age (Jordan and Hertlein, 1926). Kilmer (in litt. to C.A. Repenning) has suggested that an unconformity may exist between the vertebrate-bearing strata and the overlying mollusk-bearing conglomeratic beds. A time gap may be indicated by this. The age of Praekogia cedrosensis must lie somewhere between Luisian age and middle Pliocene age, and may provisionally be placed at early Pliocene. Rocks traditionally cited as being of early Pliocene age on the Pacific coast of North America (Weaver et al., 1943) may currently best be called late Miocene in age in light of recent interpretations of the Miocene-Pliocene boundary (Berggren, 1969: Table 1). Collector s.-T>SL\id P. Whistler and George Jefferson, 15 to 16 July 1964. Osteology Skull.-Thc holotype consists of a cranium lacking the rostrum anterior to the nares. It is typically kogiine in being brachycephalic, and having a steeply ascending supracranial basin with a sagittal crest posterior to the nares, small supraorbital processes of the frontals, and broad basioccipital with large de- scending falcate processes. Unlike modern Kogia, the skull bones are not por- ous and inflated. Only the posterior walls of the narial passages remain intact, and the left passage is much larger and located more posteriorly than is the right (Fig. 2). The right narial opening appears larger, relative to the overall skull size, than in Kogia breviceps. As in Kogia and members of the sperm whale subfamily Physeterinae, the right premaxilla extends posteriorly into the supracranial basin, bends around and partly encompasses the right naris, expands laterally in the center of the basin behind both nares, and reaches the occipital crest. Praekogia resembles the Miocene sperm whale Aulophyseter Kellogg, 1927 in that the left premaxilla extends around the left naris, forms its poste- rior wall, then meets the right premaxilla to which it is closely appressed in Figure 1. Praekogia cedrosensis, new genus and species, holotype, skull, UCR 5229, locality UCR RV-7315, dorsal view. the anterior part of the sagittal facial crest posterior to the left naris. In Kogia and Physeter the left premaxilla does not extend as far posteriorly and usually only extends as far posteriorly as the middle of the lateral wall of the left naris. In those genera, the left maxilla then forms the bony surface of the posterior wall of the left naris (Schulte, 1917:371). On its medial (left) margin, the right premaxilla meets the left premaxilla and the left maxilla to produce an elevated sagittal facial crest with a steep left side, but there is no great upward flare and overhang dorsal to the left maxil- lary fossa as is developed in the homologous structure of Recent Kogia. The right premaxillary posterior apex is situated left of the midline, and continues New Genus and Species of Fossil Pygmy Whale 7 1973 Figure 2. Praekogia cedrosensis, new genus and species, restoration of skull based on holotype, UCR 15229, and Recent Kogia, dorsal view. toward the occipital crest in a low posterior extension of the sagittal facial crest between the right and left maxillae. This part of the crest consists only of the right premaxilla and is not contributed to by the maxillae, nor formed into a tuberosity as inKogia (Schulte, 1917:372). Slightly to the right of the center of the supracranial basin, behind the right naris, is a large premaxillary fossa (Fig. 1), with its deepest portion lo- cated near the center of the skull in the right side of the right premaxilla. The same fossa also extends shallowly onto the medial portion of the adjacent right 8 Contributions in Science No. 247 Figure 3. Praekogia cedrosensis, new genus and species, holotype, skull, UCR 15229, locality UCR RV-7315, right lateral view. Table I. Measurements (in cm) of the holotype skull of Praekogia cedrosensis, UCR 15229. Parentheses indicate estimated measurements. Rostral width (12. ) Interorbital width (20. ) Zygomatic width (24. ) Exoccipital width (23. ) Condyle width 7.2 Length, antorbital notch to occipital condyle 1 4.9 Height of temporal fossa 6.1 Length of temporal fossa 7.7 maxilla and there is no distinct right maxillary fossa as is present in Kogia. The left margin of the premaxillary fossa of Praekogia is formed by a steeply over- hanging ridge in the center of the right premaxilla which runs obliquely poste- riorly and to the left from the external margin of the right naris (Figs. 1-2). A suture between the right premaxilla and right maxilla runs generally through the right part of this premaxillary fossa, then curves medially to meet the apex of the premaxilla. In its center, the fossa is floored by a smooth bone sur- face, but toward its right side the bone surface is rugose, pitted, and a faint, curving, medially concave ridge in the middle of the right maxilla marks the right margin of the fossa. 1973 New Genus and Species of Fossil Pygmy Whale 9 This central premaxillary fossa in Praekogia does not appear to be homol- ogous with the large, shallow fossa in the elevated right premaxilla of Kogia, but instead is probably homologous with a small depression in Kogia present in some animals (especially in adults) around the medial side of the right pos- terior maxillary foramen. The above-mentioned steeply overhanging ridge in the center of the right premaxilla of Praekogia seems homologous with a simi- larly positioned ridge in Recent Kogia that extends posteriorly and to the left from the lateral side of the right naris to the anterior part of the premaxillary basin. The more shallow, lateral part of the fossa on the right maxilla of Praekogia therefore probably corresponds with the large fossa in the center of the facial part of the right maxilla of Kogia. In modern Kogia, the posterior part of the right premaxilla not involved in the premaxillary fossa of Praekogia, is elevated, with upturned margins, in- clined dorsally to the left side, and shallowly excavated into a depression. No such depression is present in Praekogia, in which the premaxillary margins are not upturned, and the premaxillary surface while inclined dorsally to the left side is neither elevated nor excavated to the extent seen in Kogia. On each side of the supracranial basin, the lateral maxillary crests are up- turned into sharp-topped ridges with slight medial overhang and flat external surfaces. These crests are much more inflated, rounded, and curved in Kogia. 10 Contributions in Science No. 247 Figure 5. Praekogia cedrosensis, new genus and species, holotype, skull, UCR 15229, locality UCR RV-7315, ventral view. Typically in Kogia, the thickened external margin of the maxilla has a wide exposure over the supraorbital process of the frontal, and is much thicker and more elevated on the left side than on the right. In Praekogia, however, the maxilla is not widely exposed on the external surface of the lateral maxillary crest (Figs. 3-4), has a straight ventral sutural margin, does not thicken over the supraorbital process of the frontal, and the right and left sides are equally developed. The lateral maxillary crests are straighter, and the supracranial basin is correspondingly more anteroposteriorly elongate and not as rounded as in Kogia. 1973 New Genus and Species of Fossil Pygmy Whale 11 Figure 6. Praekogia cedrosensis, new genus and species, restoration of skull based on holotype, UCR 15229, and Recent Kogia, ventral view. In the relative positions of the posterior maxillary foramina only the left one of Praekogia resembles that of Kogia. The right maxillary foramen of Praekogia is placed farther laterally than in Kogia. A vestige of the broken left posterior maxillary foramen lies to the left of the posterior margin of the left naris. On the right maxilla two connecting apertures of the posterior maxillary foramen lie lateral to the posterior margin of the right naris. The anterior one of these is not located as close to the premaxilla as is common in Kogia. The 12 Contributions in Science No. 247 antorbital notch appears not to be as deeply incised into the dorsal surface of the maxilla as in Kogia. In Kogia, the jugal is large and a considerable portion of the bone is visi- ble laterally as a wedge extending posteriorly between the maxilla and the antorbital process of the frontal (Schulte, 1917:pl. 35). In Praekogia, a very slender section of the jugal extends as a wedge (Fig. 3) farther posteriorly be- tween the maxilla and frontal than in Kogia. The supraorbital processes of the frontals of Praekogia and Kogia are almost identical in shape but the ventro- lateral margin dorsal to the orbit is turned laterally and is thinner in Praekogia than in Kogia, and the lateral bone surface is concave and rugose rather than inflated and smooth. The temporal fossa of Praekogia is relatively and abso- lutely larger, more rounded, and not as roofed over by the frontal as in Kogia. The difference between the temporal fossae of Praekogia and Kogia is associated with a relatively more posterior position of the orbit in the latter. Compared with Praekogia, the bony orbit of Kogia is located more over the anterior end of the zygomatic process of the squamosal and encroaches on the anterior part of the temporal fossa. The fossa, which in Praekogia is entirely open in lateral view (Fig. 3), is partly hidden in a lateral view of Kogia. Con- commitantly, the posterior margin of the temporal fossa extends more poste- riorly into the occipital shield in Kogia than in Praekogia. The fossa is higher than wide in Kogia and more anteroposteriorly expanded in Praekogia, fur- ther emphasizing the different positions of the orbit. Praekogia resembles Kogia when viewed posteriorly (Fig. 10b) in pos- sessing large, ventrolaterally flaring exoccipitals, large falcate processes of the basioccipital, and occipital condyles which are not distinctly set off from the occipital shield. In Praekogia, though, the zygomatic processes of the squa- mosals flare farther laterally, the paroccipital processes of the exoccipitals ex- tend farther posteriorly, and the occipitals bear rugose fossae just lateral to the occipital condyles. Praekogia differs from Kogia by having the occipital condyles more set off from the occipital shield and the foramen magnum and condyles nearly circular, not transversely compressed as in Kogia (Schulte, 1917: plate 37; Handley, 1966: Fig. 1). Much of the ventral surface of the cranium is badly damaged (Fig. 5) and most of the pterygoids are missing. The proportions and relationships of the preserved parts resemble those of Kogia. Subtle differences are that the pterygoid-basioccipital descending processes do not diverge as widely poste- riorly and the orbits are positioned more anteriorly so that the tract of the optic nerve enters the braincase directed posteromedially and not so nearly perpen- dicular to the long axis as in Kogia. The ventral part of the internarial septum, formed by the presphenoid and vomer, is relatively thick compared with Kogia and appears to have been entirely sheathed ventrally by the posterior part of the vomer which extends across, and farther posterior than, the open basisphenoid-presphenoid fissure. 1973 New Genus and Species of Fossil Pygmy Whale 13 In the holotype of Praekogia, the right side of the internarial part of the presphenoid is covered by vomer at least to the midline, and appears to be in- completely covered to the left side only because parts of the vomer are broken away. In Kogia, the vomer is applied to either side of the internarial septum, but the ventral keel is exposed ( Schulte, 1917:386). The transverse dimension of the lateral border of the orbit, the tract of the optic nerve, and the orbital fissure are all relatively larger than those in Kogia, indicating that the eye of Praekogia was larger. Schulte (1917:373) has noted that the lateral process of the pterygoid underlies the medial part of the frontal and forms a shelf in the wall of the orbit. This is similar to the struc- ture in Praekogia. Ventral to this shelf and directly anterior to the orbital fissure in Praekogia, there is a small (10 mm diameter) recess or sinus in the lateral surface of the pterygoid that does not exist in Kogia. In Kogia there exists a fossa marking the position of a sinus in the medial part of the alisphenoid and the pterygoid, medial to the foramen ovale. There is an indication of a smaller sinus in the same position in Praekogia, but the pterygoid is broken away and impairs discernment of a fossa. A small canal for the carotid artery lies posterior to the foramen ovale and is larger and closer to the latter than in Kogia. The aperture of the poste- rior lacerate foramen of Praekogia is situated more medially and is more re- cessed dorsal to the falcate process of the basioccipital than in Kogia. With this, the aperture is farther from the lateral margin of the recess for the mas- toid process and there is more space in the roof of the auditory region formed by the squamosal than in Kogia. The posterior process, or what Schulte (1917: 394) called the tympano-mastoid, of the auditory bulla must have been rela- tively much larger than in Kogia for the excavation in the lateral part of the squamosal and exoccipital for its attachment is much wider than in Kogia. The posterior wall of this excavation, which is formed by the paroccipital process of the exoccipital is much thinner than, and not inflated as in Kogia. At the ventral extremity of a descending transverse strut in the squamosal is a vestigial groove marking the passage of the external auditory meatus. The groove is slightly larger (2.5 mm across) than in most specimens of Kogia, and is directed anterolaterally posterior to the postglenoid process, but is broken and incomplete. Unlike the condition in Kogia, there is a bony fossa marking the position of a middle ear air sinus developed in the squamosal on the posteromedial part of the postglenoid process. The sinus is well marked because the posteroventral part of the articular surface of the glenoid fossa is set off from the squamosal and the sinus runs around it posteriorly and medially. Relationships Praekogia cedrosensis is a kogiine physeterid or pygmy sperm whale that is more generalized than either of the two living species, Kogia breviceps and K. simus, and may be ancestral to the genus Kogia. Praekogia exhibits nearly 14 Contributions in Science No. 247 Figure 7. Praekogia cedrosensis, new genus and species, holotype, skull, UCR 15229, locality UCR RV-7315, dorsal view. all of the structural peculiarities in the skull of Recent Kogia except that the bone is not porous and inflated, and the skull is less telescoped or foreshortened (see Miller, 1923). Concomitant with this lesser degree of telescoping, and in contrast with Kogia, the supracranial basin is longer with straighter sides, the supraorbital process of the frontal and the optic nerve tract are directed more anterolaterally, the orbit is more anteriorly located, the squamosal is relatively longer, the temporal fossa is larger, the sagittal facial crest is lower, and the narial openings are not located as far posteriorly. In the retention of the more primitive proportions and structures the skull of Praekogia resembles certain extinct Miocene sperm wales (for example Orycterocetus Leidy, 1853; Aulo- physeter Kellogg, 1927). Two previous publications allude to fossil Kogiinae. Kellogg (1929) based a new genus and species, Kogiopsis floridanus, upon the symphyseal por- tion of a fossil mandible from the Bone Valley Gravels of Polk County, Flor- 1973 New Genus and Species of Fossil Pygmy Whale 15 Figure 8. Praekogia cedrosensis, new genus and species, holotype, skull, UCR 15229, locality UCR RV-7315, ventral view. ida. In his description Kellogg stressed the similarity between Kogiopsis and fossil physeterines and hoplocetines with which it compares favorably also in its large size. The large size of Kogiopsis alone precludes synonymy, and pos- sibly close relationship, between Kogiopsis floridanus and Praekogia cedro- sensis. Kogia prisca is a name applied by Matsumoto (1936) to isolated teeth, apparently of Miocene age, from Japan. These teeth are large (50 to 90 mm long) and bear enamel. The teeth of K. prisca are too large to belong to Prae- kogia cedrosensis, and like Recent Kogia, to which it is obviously closely re- lated, Praekogia probably had no enamel on the teeth. In light of present meager knowledge of North Pacific Miocene cetaceans, I would withhold any 16 Contributions in Science No. 247 Figure 9. Praekogia cedrosensis, new genus and species, holotype, skull, UCR 15229, locality UCR RV-7315; a, left lateral view; b, right lateral view. 1973 New Genus and Species of Fossil Pygmy Whale 17 Figure 10. Praekogia cedrosensis, new genus and species, holotype, skull, UCR 15229, locality UCR RV-7315, a, anterior view; b, posterior view. 18 Contributions in Science No. 247 positive statement regarding the identity of the teeth figured by Matsumoto, and the name is here declared a nomen vanum. Praekogia, therefore, is the first described fossil to show positive close phylogenetic relationships with living pygmy sperm whales of the genus Kogia. All the structural peculiarities of the skull are present in both genera, and Praekogia differs by being less highly modified from a primitive condition. The major structural modifications required in the evolution of a skull like that of Kogia would be accomplished by further telescoping of the skull of Prae- kogia and by the inflation and increase in the porosity of the bone. Miller (1923) has convincingly postulated, using the fossil record, the manner in which members of various cetacean groups appear to have evolved highly modified skulls through telescoping. The process of telescoping involves the extension of bones over and under one another to reach positions on the skull where they do not exist in more generalized and less highly evolved animals. Telescoping also involves the posterior movement on the skull of the dorsal narial apertures and of the orbit with a resulting shortened braincase. The skull of Praekogia had telescoped to the point where the cranial bones were roughly in the same position and had the same sutural relationships as in Recent Kogia, but further foreshortening of the skull was still possible to bring the eye and narial apertures farther posterior upon the braincase. This is well displayed by the fact that the orbit of Kogia is in a position dorsal to the anterior part of the zygomatic process of the squamosal. This is an unusual situation, because in most fossil and Recent cetaceans (including Praekogia) , the anterior tip of the zygomatic process articulates with or is very close to the ventral extremity of the postorbital process of the frontal. If telescoping of the skull of Kogia progresses further, it is possible that the orbit may proceed even farther pos- teriorly into the temporal fossa. Other cranial features possessed by Praekogia, that differ from Kogia, serve to point out the relatively more primitive nature of Praekogia. These in- clude the less elevated sagittal facial crest, more prominent occipital con- dyles, relatively larger zygomatic process of the squamosal, more expansive vomer on the ventral surface of the internarial septum, and uninflated nature of the bone surface. I believe that the pygmy sperm whales are best included as a subfamily Kogiinae within the Physeteridae as was done by Simpson (1945: 102) and Handley (1966: 64). This contrasts with some authors (Miller, 1923; Kellogg, 1928, 1929; Miller and Kellogg, 1955) who recognized a family Kogiidae. Conclusions 1. Praekogia cedrosensis, a new genus and species of extinct pygmy sperm whale, is based on the first cranial remains of a pygmy sperm whale doc- umented in the fossil record. Kogiopsis floridanus Kellogg, 1929, described from mandibular sections found in Florida’s Bone Valley Gravel, is an animal 1973 New Genus and Species of Fossil Pygmy Whale 19 ; much larger than, and probably distantly related to, Praekogia. Kogia prisca I Matsumoto, 1936, known only by isolated teeth from probable Miocene rocks in Japan is herein designated a nomen vanum. 2. Praekogia cedrosensis is known only from one site in the Pliocene Almejas Formation on Isla Cedros, Baja California, Mexico. The strata bear- I ing the holotype are unconformably underlain by Luisian age (Miocene) rocks of the Tortuga Formation, and overlain (possibly unconformably) by mollusk- bearing beds correlated with the middle Pliocene to early late Pliocene-age San I Diego Formation of California. The geologic age of Praekogia cedrosensis is therefore estimated to be early Pliocene. 3. The braincase of Praekogia indicates possible morphological ancestry to both of the living species of pygmy sperm whale Kogia breviceps and K. simus. The osteological changes that would be necessary for the skull of Prae- kogia to come to resemble one of a Recent Kogia are an overall telescoping (foreshortening) of the braincase, enlargement of the maxillary crests around the supracranial basin, elevation of the sagittal facial crest formed by the maxillae and premaxillae posterior to the nares, and inflation and general in- crease in porosity of the bone tissue of the cranium. 4. Both Kogia and Praekogia are included in the Subfamily Kogiinae of the Family Physeteridae. Cranial morphology of the Kogiinae is sufficiently similar to the true sperm whales (Physeterinae, Hoplocetinae) to argue against the use of a family solely for the pygmy sperm whales. Resumen El nuevo genero y especie Praekogia cedrosensis es un cachalote pigmeo extincto, conocido por una porcidn de craneo sin el rostro, descubierto en depositos marinos sedimentarios identificados como Formacion Almejas, en la isla Cedros, Baja California, Mexico. La antigUedad geologica de Praekogia cedrosensis es considerada como Plioceno Inferior en sentido amplio. La base de la Formacion Almejas, estratigraficamente 36 metros por debajo de hori- zonte que proporciono Praekogia, descansa discordantemente sobre rocas Miocenas de edad Temblor. Por encima del mencionado horizonte existen depositos con moluscos marinos de edad Pliocena Media-Superior. Praekogia es un antecesor ideal para el cachalote pigmeo actual, Kogia Gray, con una caja craneana menos telescopica y una menor especializacion craneana. Literature Cited Barnes, L. G. 1972. Late Tertiary Cetacea of the northeast Pacific Ocean. Unpub- lished Ph.D. dissertation, University of California, Berkeley, California. 494 pp. Beal, C. H. 1948. Reconnaisance of the geology and oil possibilities of Baja Cali- fornia, Mexico. Geol. Soc. Amer., Mem. 31: 1-138. Berggren, W. a. 1969. Cenozoic chronostratigraphy, planktonic foraminiferal zona- tion and the radiometric time scale. Nature 224: 1072-1075. 20 Contributions in Science I ■ i: No. 247 1; Durham, J. W., and E. C. Allison. 1960. Symposium: The biogeography of Baja i California and adjacent seas. Part I. Geologic history. The geologic history of j Baja California and its marine faunas. Systematic Zool. 9(2) : 47-91. l Handley, C. O., Jr. 1966. A synopsis of the genus Kogia (pygmy sperm whales). ’ pp. 62-69. In K. S. Norris, (Ed.), Whales, Dolphins, and Porpoises. Univ. Calif, j Press, Berkeley and Los Angeles, California, i-xv, + 789 pp. Hanna, G. D. 1927. Geology of the west Mexican islands. Pan-Am. Geol. 48: 1-24. Hertlein, L. G. 1925. Pectens from the Tertiary of lower California. Proc. Calif. Acad. Sci. Ser. 4, 14( 1 ) : 1-35. Howard, H. 1971. Pliocene avian remains from Baja California. Los Angeles Co. Mus., Contrib. Sci. 217: 1-17. Jordan, E. K., and L. G. Hertlein. 1926. Contribution to the geology and paleon- tology of the Tertiary of Cedros Island and adjacent parts of Lower California. Proc. Calif. Acad. Sci. Ser. 4, 15(14) : 409-464. Kellogg, A. R. 1927. Study of the skull of a fossil sperm-whale from the Temblor , Miocene of Southern California. Contrib. to Paleontol., Carnegie Inst. Wash- i ington 346: 1-23. | 1928. The history of whales, their adaptation to life in the water. Quar- terly Rev. Biol. 3( 1 ): 29-76, 3(2) : 174-208. 1929. A new fossil toothed whale from Florida. Amer. Mus. Novit. 389: 1-10. Kernan, j. D., Jr., and H. von W. Schulte. 1918. Memoranda upon the anatomy of the respiratory tract, foregut, and thoracic viscera of a foetal Kogia breviceps. , Bull. Amer. Mus. Nat. Hist. 28(8) : 231-267. , j Matsumoto, H. 1936. On some fossil cetaceans of Japan. Sci. Repts. Tohoku Imp. J Univ., Sendai, Japan, 2nd Ser. (Geology) 10( 1 ) : 17-27. Miller, G. S. 1923. The telescoping of the cetacean skull. Smithsonian Misc. Coll. ^ ' 76(5) 1-71. ^ ? , AND A. R. Kellogg. 1955. List of North American Recent mammals. « Bull. U.S. Nat. Mus. 205 : i-xii, 1-954. Mina, U. F. 1957. Bosquejo geologica del Territorio de la Baja California. Assoc. Mexicana Geol. Petrol., Boll. 9: 141-269. { Raven, H. C., and W. K. Gregory. 1933. The spermaceti organ and nasal passages | of the sperm whale (Physeter catodon) and other odontocetes. Amer. Mus. | Novit. 677: 1-18. ^ Schulte, H. von W. 1917. The skull of Kogia breviceps Blainv. Bull. Amer. Mus. Nat. Hist. 37(17): 361-404. Simpson, G. G. 1945. The principles of classification and a classification of mam- | mals. Bull. Amer. Mus. Nat. Hist. 85: 1-350. Weaver, C. E. et al. 1944. Correlation of the marine Cenozoic formations of west- ern North America. Bull. Geol. Soc. Amer. 55 : 569-598. Yamada, M. 1954. Some remarks on the pygmy sperm whale Kogia. Sci. Repts., i Whales Res. Inst. 9: 37-58. I li Accepted for publication June 4, 1973 Printed in Los Angeles. California hv Anderson Ritrhip Kimnr, r,r> Q.mr.con T NUMBER 248 JUNE 30, 1973 NEW SPECIES OF SALAMANDERS, GENUS BOLITOGLOSSA, FROM PANAMA By David B. Wake, Arden H. Brame, AND William E. Duellman CONTRIBUTIONS IN SCI6NCE NATURAL HISTORY MUSEUM • LOS ANGELES COUNTY NEW SPECIES OF SALAMANDERS, GENUS BOLITOGLOSSA, FROM PANAMA^ By David B. Wake,^ Arden H. Brame, Jr.,^ and William E. Duellman'^ Abstract: Three species of salamanders, Bolitoglossa cuna, B. minutula, and B. compacta, are described from Panamanian populations. Bolitoglossa cuna, from the Territorio de San Bias, is a lowland species with fully webbed hands and feet. It is a member of the sima group. Bolitoglossa minutula is the smallest member of the genus. It occurs at elevations between 1800 and 2000 m near the Costa Rican border, and has some similarity to the larger Costa Rican species, B. epimela. Bolitoglossa compacta is a large species with slightly webbed hands and feet which is sympatric with B. minutula. It is a relative of B. cerroensis of Costa Rica. Gradually the notion that salamanders are rare and lack diversity in the tropics is being dispelled. The increased field work of recent years has dis- closed that salamanders are widely distributed in tropical America, but densi- ties are often low. As a result, most species are represented by rather small series in collections. As field work continues, the diversity of the group is be- coming apparent. Often in a given restricted area several species live in broad sympatry and have varied structure and ecology. In the present paper we describe specimens of three new species collected principally by Charles W. Myers, William E. Duellman, and Linda Trueb since 1963. These species en- compass much of the range of morphological diversity in Panamanian species of Bolitoglossa and add considerably to our knowledge of the Panamanian salamander fauna. The first species occurs only in eastern Panama, along the Caribbean low- lands. It is named for the humans indigenous to the region. ^Review Committee for this Contribution Richard Highton George B. Rabb John W. Wright ‘-Museum of Vertebrate Zoology, Univ. Calif., Berkeley, Calif. 94720 (address for reprint requests); and Research Associate, Section of Herpetology, Natural History Museum of Los Angeles County, Los Angeles, Calif. 90007 3Eaton Canyon Nature Center, 1750 North Altadena Dr., Pasadena, Calif. 91107; and Research Associate, Section of Herpetology, Natural History Museum of Los Angeles County, Los Angeles, Calif. 90007 ■^Curator, Division of Herpetology, Museum of Natural History, Univ. Kansas, Lawrence, Kansas 66044 1 1973 New Salamanders from Panama 2 Bolitoglossa cuna, new species Figures 1 and 2 Holotype: KU 116519, an adult female from Camp Sasardi, 12 m (39 ft) elevation, Territorio de San Bias, Panama, obtained by Charles W. Myers on 6 February 1967. Paratypes: USNM 150035-36, adult females from Armila, near sea level, Territorio de San Bias, Panama, collected by C. O. Handley, Jr. and F. M. Greenwell on 26 February, 1963. Diagnosis: A moderate-sized species (3 adult females: 46.6-55.7, mean 50.3 SL^) with high numbers of maxillary (mean 72) and vomerine (mean 35) teeth; distinguished from B. silverstonei by its more numerous teeth and nar- rower head, hands and feet; from B. biseriata by its more numerous teeth and narrower head; from B. sima by its shorter legs, more numerous teeth and narrower head. B. cuna is distinguished from all other members of the genus Figure 1. Dorsal and ventral views of the holotype of Bolitoglossa cuna, new spe- cies, KU 116519. ^Standard length, the distance in mm from the snout to the posterior end of the vent (all measurements are in millimeters). 3 Contributions in Science No. 248 by the combination of its nearly completely webbed hands and feet, its size and dentitional features, and its coloration (light golden tan mottled with some darker color dorsally; lighter venter with a peppering of small, dark spots). Description of Holotype: Adult female, snout of moderate length. Nostril moderately small; labial protuberances of nasolabial grooves small and poorly developed. Canthus rostralis of moderate length, gently arched. Standard length 7.2 times head width; standard length 5.0 times snout-gular fold length. Deep groove below eye extends almost full length of opening, following curva- ture of eye, but does not communicate with lip. Eye moderately small, not protuberant. Well defined postorbital groove extends posteriorly from eye as shallow depression for 1.6 mm; proceeds sharply ventrad at level of posterior end of mandible and extends across gular area as nuchal groove parallel to and 3.3 mm anterior to sharply defined gular fold. Vomerine teeth 33, patched, extending to distal borders of the internal nares; patches extend from distal end in a straight line to near the center of the palate, where they are separated by 0.8 mm from the right and left parasphenoid tooth patches. Maxillary teeth 77, extend posteriorly to a point about three-fourths through eye. Five mod- erately small premaxillary teeth protrude through the lip. Tail relatively long, 0.9 times standard length; postiliac gland clearly evident. Limbs moderately short, limb interval 3.5; standard length 4.8 times right forelimb; 4.7 times right hind limb; 11.1 times right foot width; webbing of hands and feet so ex- tensive as to be nearly complete, with only the longer digits protruding from webbing (Fig. 2). No subterminal pads. The fingers are in order of decreasing length; 3, 2, 4, 1; toes in order of decreasing length: 3, 4, 2, 5, 1. Measurements (in mm): Head width 1.1 \ snout to gular fold (head length) 11.1; head depth at posterior angle of jaw 3.7; eyelid length 2.9; eyelid width 1.4; anterior rim of orbit to snout 3.8; horizontal orbital diameter 2.1; inter- orbital distance 3.8; distance between vomerine teeth and parasphenoid tooth patch 0.8; snout to forelimb 14.3; distance separating internal nares 2.3; dis- tance separating external nares 3.1; snout projection beyond mandible 1.1; snout to posterior angle of vent (standard length) 55.7; snout to anterior angle of vent 52.5; axilla to groin 32.5; tail length 52.2; tail width at base 4.1; tail depth at base 4.1; forelimb length 11.5; hind limb length 11.9; width of right hand 3.8; width of right foot 5.0. Coloration in Alcohol: The holotype is light reddish brown dorsally with dark brown lateral bands extending from the neck along the body and well onto the tail. The venter is cream with many small brown spots which leave some cream areas unmarked. There is a peppered effect on both dorsal and ventral surfaces consisting of tiny scattered dark brown melanophores. The head is light reddish brown dorsally with the same general coloring as the dorsum of the trunk, although the lateral regions between the eyes and snout are distinctly paler. The ventral coloration of the gular area is like the trunk venter, mottled cream and dark brown with many tiny dark brown melanophores clearly discernible. The small eyes appear to have a gold ring 1973 New Salamanders from Panama 4 around the horizontally elliptical pupil. The dorsal surfaces of the limbs are mottled with dark brown, and are darker than the trunk dorsally. Ventrally the limbs are similar in color to the trunk venter. The ventral surfaces of the hands and feet are rather light grey brown. Variation: The poorly preserved paratypes have more dorsal mottling than the holotype, with large blotches of light color which contrast with the dark brown ground color. There are no dark brown lateral bands. The dorsal colora- tion gradually becomes lighter along the lower lateral surfaces, finally fading to a uniform light grey ventrally. USNM 150036 has larger feet than the holo- type or other paratype and USNM 150035 has a somewhat broader head (see Table I) but otherwise the external morphology of the three specimens is similar. Osteology: All information has been derived from radiographs, and many details of the skull cannot be discerned. The skull is well developed with well articulated bones. The premaxilla is relatively slender and has processes of moderate length which are slightly dilated at their distal tips. The nasals are large and well developed, but are only slightly protuberant. They are closely articulated to the facial part of the maxillae. No prefrontals can be seen. The vomers are well separated for their entire length. The preorbital processes are long and extend well beyond the lateral margins of the internal nares. Teeth extend along these processes to about the level of the lateral margins of the nares. The moderately long maxillae extend to the posterior margin of the eyes. The operculum has no stilus. All but the last trunk vertebrae bear well devel- oped ribs. There are one cervical, fourteen trunk, one sacral, two caudosacral and thirty-two caudal vertebrae in the single specimen (USNM 150036) which has a complete tail.® The holotype has a tail which is regenerated distal to the fifteenth caudal vertebra. The regenerated portion has twenty-one vertebrae. USNM 150035 lacks a tail. The long, stout transverse processes on the first caudosacral vertebra are nearly perpendicular to the long axis of the body. The shorter and stouter processes of the second caudosacral vertebra are angled anteriorly, with blunt tips. The processes of the first caudal vertebra are very long and slant sharply in an anterior direction. They are much longer in the holotype than in the paratype, and extend nearly to the anterior end of the second caudosacral vertebra. The processes of adjacent vertebrae do not overlap. Processes on suc- ceeding vertebrae are progressively shorter. They are present to about the twelfth vertebra in the holotype and to the nineteenth in the paratype. All lie at the anterior end of the vertebra and slant anteriorly. In the holotype the second caudosacral and first caudal vertebrae are shorter than any trunk vertebrae and all of the unregenerated tail vertebrae. The mid-trunk vertebrae are the longest in the column, and all but the first trunk vertebrae are longer than any post-trunk vertebrae. No tibial spurs are present, but a distinct ridge is visible in one paratype. 6See Wake (1966) for definitions of osteological characters. 5 Contributions in Science No. 248 Table I. Measurements and data for specimens of new species of Bolitoglossa OJU C M o p- £ X c (L> c o X) C 4> hJ 13 > k. JS X o> ^ [— ( •d 5 o o bu B. cuna KU 116519* $ 55.7 32.5 7.7 11.9 11.5 3.5 11.1 52.2 77 33 5.0 USNM 150035 $ 48.7 28.4 7.4 10.7 10.2 3.5 11.6 — 69 33 4.4 USNM 150036 B. minutula $ 46.6 — 6.8 9.8 9.8 3.? 10.4 45.0* 66 38 4.6 KU 116564 $ 36.5 20.0 5.4 8.2 7.8 3 7.8 33.0 36 15 3.3 KU 116575 9 36.3 19.8 5.6 7.7 6.8 3 8.1 35.9 48 16 3.2 KU 116576 9 36.0 20.0 5.3 7.8 7.8 3 8.2 36.1 43 18 3.2 KU 116577 9 34.8 19.7 5.6 8.7 7.8 2.5 8.1 31.8 51 16 3.1 KU 116608 9 34.2 18.9 5.2 7.8 7.2 4 7.5 * ❖ 33 15 3.0 KU 116556 9 34.1 18.2 5.6 8.1 7.7 2 8.1 34.2 42 16 3.3 KU 116595 9 33.9 18.6 5.3 8.0 7.0 3 7.8 32.7 43 16 3.1 KU 116554* 9 33.6 18.5 5.2 8.0 7.8 3 8.0 32.2 46 19 3.1 KU 116574 9 33.2 18.7 5.3 7.8 7.2 3 7.2 34.3 47 19 3.2 KU 116607 9 33.2 18.3 5.1 8.0 7.0 3 7.8 29.9 37 16 2.9 KU 116582 9 31.9 17.6 4.9 7.3 6.3 3 7.2 27.8 34 12 2.9 KU 116611 9 31.9 16.9 5.2 7.9 6.9 3 7.3 29.4 42 17 3.2 LACM 78731 9 31.4 17.5 5.0 7.3 6.3 2.5 7.2 29.2 45 17 2.8 KU 116610 9 30.1 16.7 4.9 6.8 6.3 3 7.2 25.8 37 23 3.0 KU 116555 $ 36.0 19.7 5.3 9.6 8.9 1 8.4 40.2 47 — 3.7 KU 116552 $ 35.7 19.3 5.7 9.8 8.9 1.5 8.0 38.2 46 22 3.7 KU 116579 $ 34.8 19.2 5.4 9.0 7.8 2 7.8 35.2 42 26 3.4 KU 116609 $ 34.7 19.2 5.6 9.2 9.0 2 8.2 ** 46 15 3.7 KU 116557 $ 34.5 19.2 5.6 8.9 8.6 1.5 8.0 ** 51 17 3.6 KU 116605 $ 34.1 18.2 5.1 8.3 7.8 2 7.9 33.9 48 22 3.1 KU 116578 $ 34.1 18.0 5.2 9.3 8.2 1 8.0 35.0 41 18 3.3 KU 116580 $ 33.9 17.4 5.2 8.5 7.8 1.5 7.8 37.0 45 15 3.2 KU 116606 $ 33.3 17.1 5.2 8.1 7.1 1.5 8.0 36.2 47 23 3.1 KU 116601 $ 33.1 17.1 5.4 8.1 6.9 1 7.8 33.6 46 20 3.0 KU 116604 $ 33.0 17.1 5.1 8.7 7.8 1.5 8.0 32.0 36 17 3.0 KU 116592 $ 32.7 17.1 5.4 8.4 8.0 1.5 8.2 32.2 55 40 3.0 KU 116553 $ 32.5 17.4 5.5 9.0 8.3 0.5 7.9 13.3** 43 16 3.2 LACM 78732 $ 32.5 16.2 5.0 8.3 7.8 1 8.1 35.1 39 17 2.9 KU 116597 $ 32.1 16.5 5.2 8.2 8.0 1 7.8 33.2 40 16 2.8 LACM 78729 B. compacta $ 28.4 14.1 4.8 7.4 7.2 1 7.4 26.7 34 19 2.9 KU 116662* 9 74.2 40.0 10.3 17.7 16.2 2 14.9 68.8 36 20 7.4 GML 9 73.1 39.4 10.0 16.2 15.0 2 13.9 65.0 44 22 7.4 KU 116659 9 71.3 37.6 10.2 16.8 15.7 2.5 15.5 59.0 40 26 7.1 KU 116663 9 70.6 39.6 10.4 16.8 15.8 2.5 15.1 70.1 50 19 7.2 KU 116664 9 68.5 35.9 10.1 17.8 16.9 1.5 15.2 63.2 48 33 7.2 LACM 78728 5 53.4 27.4 8.3 13.3 12.3 1.5 12.7 46.8 11 19 5.8 KU 116660 $ 44.9 24.5 7.3 12.4 11.8 1.5 10.9 36.0 20 22 4.9 * = holotype; * * = regenerated tail stubs. 1973 New Salamanders from Panama 6 Phalangeal formulae are 1, 2, 3, 2 and 1, 2, 3, 3, 2. This species has extensive webbing, and the hands and feet are very flattened. Accordingly, digits are in- distinct, except at their tips. Metapodial elements are broad and flat (Fig. 2) and the phalanges become smaller in a markedly progressive way toward the tips. The basic dumbell shape of the elements is obscured by lateral bony webs. Terminal phalanges are small and irregular in shape, but tend to be pointed (Fig. 2). Figure 2. Hands and feet of new species of Bolitoglossa, drawn from cleared and stained, and x-rayed specimens, with aid of microprojector. Cartilage stippled, (a) Left hand of specimen of B. minutula. (b) Left foot of specimen of B. minutula. (c) Left foot of specimen of B. cornpacta. (d) Left hand of specimen of B. cuna (Holotype). (e) Left foot of specimen of B. cuna (Holotype). 7 Contributions in Science No. 248 Habitat: The holotype was on a leaf of an herb less than 1 m above ground at night in lowland rainforest composed of tall, buttressed trees with a lower story of palms and saplings. The herbaceous layer was poorly devel- oped, and the ground had a deep layer of litter. At Camp Sasardi B. biseriata also was found in the same habitat. Range: Known only from the narrow Caribbean lowland region of east- ern Panama. A large series of a particularly small species of Bolitoglossa was collected on the slopes of Cerro Pando. As this is the smallest known species of the genus, we name it; Bolitoglossa minutulor new species Figures 2, 3, and 4 Holotype: KU 116554, an adult female, from the north slope of Cerro Pando, 1920 m (6298 ft) elevation, Provincia de Bocas del Toro, western Panama, obtained by Charles W. Myers on 3 May !1 ( Figure 3. Dorsal and ventral views of the holotype of Bolitoglossa minutula, new species, KU 116554. 1973 New Salamanders from Panama 8 Paratypes: KU 104330 (cleared and stained), KU 116555-61, KU 116564-85, KU 116607-11, LACM 78729-35, all from type locality; KU 116562-63, 2000 m (6560 ft); KU 116593-605, 1920-1970 m (6298-6562 ft) ; KU 116606, 1810m (5937 ft) ; all from north slopes of Cerro Pando, Provin- cia de Bocas del Toro, western Panama collected by William E. Duellman, Charles W. Myers, and Linda Trueb on 3-10 May 1966. Diagnosis: The smallest known species of Bolitoglossa (16 adult males : I 28.4—36.0, mean 33.5 SL; 14 adult females: 30.1—36.5, mean 33.7 SL) with j moderate numbers of maxillary (means; males, 44.1; females, 41.7) and moderately low numbers of vomerine (means; males, 20.2; females, 16.9) I teeth; distinguished from B. epimela by its smaller size, relatively broader I head, and its smaller, more extensively webbed hands and feet. B. minutula is distinguished from all other species of the genus by the combination of its small size, extensively webbed hands and feet which retain digital integrity, and dentitional features. Description of Holotype: Adult female, snout moderately short. Nostril relatively small; labial protuberances of nasolabial groove moderately devel- oped. Canthus rostralis moderately short, slightly arched. Standard length 6.5 times head width; standard length 4.2 times snout-gular fold length. Deep groove below eye extends almost full length of opening, following curvature of eye, but does not communicate with lip. Eye moderately large, originally slightly protuberant. Strongly defined postorbital groove extends posteriorly from eye as shallow depression for 1.7 mm, proceeds sharply ventrad at level of posterior end of mandible and extends across gular area as a moderately defined nuchal groove, parallel to and 2.4 mm anterior to sharply defined gular fold. Vomerine teeth 19, arranged in single row on each side, extend from distal edge of internal choanae in strongly arched pattern toward midline then posteriad towards parasphenoid tooth patch, from which separated by 0.8 mm. Maxillary teeth 46, extend posteriorly to a point about four-fifths through eye. Three premaxillary teeth protrude from behind the lip in the same general plan as maxillary teeth; same size as maxillary teeth. Tail moderately long, 0.96 times standard length, strong lateral compression, strongly constricted at base. Postiliac gland not evident. Limbs of medium length, limb interval 3; standard length 4.3 times right forelimb, 4.2 times right hind limb, and 10.8 times right foot width; webbing of hands and feet extensive, and nearly complete but tips of all digits protrude slightly from webbing (Fig. 2). Pattern one of zygodac- tyly and webbing combined. Subterminal pads weakly developed but present. The fingers in order of decreasing length: 3, 2, 4, 1 ; toes in order of decreasing length: 3, 2, 4, 5, 1. Measurements (in mm): Head width 5.2; snout to gular fold (head length) 8.0; head depth at posterior angle of jaw 2.8; eyelid length 2.6; eyelid width 1.1; anterior rim of orbit to snout 2.2; horizontal orbital diameter 1.9; interorbital distance 1.8; distance between vomerine teeth and parasphenoid tooth patch 0.8; snout to forelimb 9.3; distance separating internal nares 1.2; 9 Contributions in Science No. 248 distance separating external nares 1.8; snout projection beyond mandible 0.4; | snout to posterior angle of vent (standard length) 33.6; snout to anterior | angle of vent 31.1; axilla to groin 18.5; tail length 32.3; tail width at base 2.6; I tail depth at base 3.2; forelimb length 7.8; hind limb length 8.0; width of right | hand 1.9; width of right foot 3.1. I Coloration in Alcohol: The dorsum of the head is black, except for the I whitish nasolabial protuberances. The dorsal surfaces of the neck, trunk and I tail have a pale yellow-white ground color. This area appears essentially un- ! pigmented, with the exception of a fine punctation of black melanophores. j Dorsal surfaces of the limbs are black, but the corresponding surfaces of the | hands and feet are mottled with black and yellow-white. Lateral surfaces of the j head and neck are black, but the same portions of the trunk and tail are colored like the dorsal surfaces. The gular region and anterior part of the trunk venter are black. The posterior part of the trunk venter is basically yellowish white, with a few large areas of black pigment located midventrally. Ventral surfaces of the tail base region are mottled black and pale yellow-white. The anterior half of the tail venter is a nearly immaculate yellow-white, but the posterior j half, as well as the ventral surfaces of the limbs, are mottled black and white. Ventral surfaces of the hands and feet are a dirty gray-yellow. Coloration in Life: Body and tail pale dull orange with dark gray flecks; head dark brown above and below; chest and belly heavily suffused with dark brown. Limbs brown, becoming orange on digits, minutely flecked with silver. I Each cirrus paler brown than rest of head. Several silver flecks on upper lip below eye. Iris pale brown. j Variation: This species is sexually dimorphic, particularly in limb and j foot differences. Males have longer limbs (19 males SL 3.5— 4.1, mean 3.8, j times hind limb lengths; 15 females SL 4.0— 4.7, mean 4.3, times hind limb length). Males have 0.5—2 (mean 1.3) costal folds uncovered by appressed i limbs (limb interval) versus 2—4 (mean 2.9) for females. Males also have larger feet (SL 9.4— 1 1.5, mean 10.3, times right hind foot width compared to | 10.0—12.5, mean 11.0, for females). Heads of males are somewhat broader | (SL 5.7— 6.8, mean 6.2, times head width, versus 6. 1—6.8, mean 6.4, for females). The species is highly variable in coloration, the holotype being at one ex- treme, mostly devoid of black pigment save head, limbs and front half of venter. Nine paratypes approach the holotype color but have more melanin present in varying degrees, but always to a greater extent than the holotype. Twenty paratypes are intermediate between the light (largely pigmentless) group (including the holotype) and the nearly all black group. This intermedi- ate sample is characterized by being light to medium brown dorsally with in- distinct, narrow stripes of darker black melanin midventrally and dorsolater- ally. The venters are variously mottled dirty yellow and black; tail venters usually have more yellow than trunk venters. The final group of 29 paratypes is mostly very dark, but some (12) have much yellow mottling on ventral or , 1973 New Salamanders from Panama 10 ventrolateral surfaces. Dorsally they are nearly totally uniform brown-black (see Fig. 3 and Table I for additional information) . Osteology: Information concerning osteology has been derived from two cleared and stained specimens (an adult male and female) and from radio- graphs of a number of additional specimens. The skull is compact and well ossified. The premaxillary is small, with a narrow dental portion and virtually no palatal process. The bone is narrower and more anteriorly placed in the male than in the female. Frontal processes are very narrow and slender, but they remain separated for their entire length. They terminate at a point just a bit beyond the extreme anterior margin of the orbit, but fall short of the posterior edge of the nasals. Tips of the processes are undilated. There is only a very narrow internasal fontanelle, for the proc- esses diverge only slightly. Contact with the frontals is very restricted. Nasals are large and protuberant. There is marked sexual dimorphism, with the male having inflated nasal capsules and enlarged nasal bones. The bones conform to the shape of the capsule to some extent, and are extended ventrally along the medial border of the capsule. Posteriorly the nasals extend to the orbit. They occupy most of the facial portion of the skull. On their posterolateral aspect the nasals are deeply evacuated by the foramen of the nasolacrimal duct. The evacuation is bordered posterolaterally by the small, triangular, facial process of the maxilla. Prefrontals are absent, and the nasal, and a small process of the frontal, occupy the space usually filled by the prefrontal. The maxillaries are slender bones with small processes. They fall short of the posterior margin of the eyeball. The vomers are small bones which are widely separated from each other for their entire lengths. The intervomerine fontanelle is broad and has a spin- dlelike shape. Preorbital processes extend beyond the lateral margins of the vomerine bodies, and these processes bear small teeth which extend in series to about the mid-point of the internal nares. Frontals are well developed, but have rather small facial portions. The two bones are in close proximity for their entire lengths, but direct contacts are rather restricted. A slight lateral lobe overlaps the parietal. Parietals are well developed and have the lateral spurs that are characteristic of the genus. They are in close proximity medially, and are rather well articulated to one another. The otic capsules are relatively large. They bear no crests or projec- tions. The large parasphenoid is narrow anteriorly, but the orbitosphenoids are well separated from each other. The tip of the parasphenoid is blunt. Posterior vomerine teeth are in large patches on the parasphenoid. These patches ap- proach each other closely, but do not touch. In the two cleared and stained individuals the patches bear 72 (left) and 70 (right), and 60 and 61 bicuspid, ankylosed teeth in the male and female, respectively. The operculum has no stilus. Quadrates are small, and are joined to the skull by the weak, narrow squamosals, and the cartilaginous suspensorium. The hyobranchial apparatus is typical of other members of the genus 11 Contributions in Science No. 248 (Wake, 1966), and there are no features worthy of special note in the lower jaw. Vertebral structure is similar to that of other species of the genus. There are one cervical, fourteen trunk, one sacral, two caudosacral and from 25 to 30 (mean of 10 adults is 27.2, mode 28) caudal vertebrae. In small individuals, ^ such as those characteristic of this species, differentiation according to centrum | length is difficult to discern in the types of preparation available. However, the first few caudal vertebrae nearly equal in length the longest trunk vertebrae. Ribs are present on all trunk vertebrae except the last one (8 specimens) or two (4 specimens). Transverse processes are present on all but the last four or five caudal vertebrae, which are very poorly developed. Transverse proc- esses of the first caudosacral vertebra are long and directed almost perpendicu- j larly to the body axis, or slightly in a posterior direction. Those of the second \ are much shorter and slenderer, and are directed strongly in an anterior direc- tion. The long, slender and non-bifurcated processes of the first caudal verte- bra arise at its anterior end. They are even more sharply oriented in an ante- j rior direction than those of the preceding vertebra. They do not cross the im- | mediately anterior pair of processes, but do extend to the origin of those proc- esses at the midvertebral level of the second caudosacral vertebra. More pos- i terior processes have a similar origin and orientation, but are much smaller and become increasing short as one moves down the column. The distinctive hands and feet have rather well formed digits which are comprised of well developed phalangeal elements. The amount of skeletal ma- terial relative to webbing is high. The usual phalangeal formula in Bolitoglossa \ is 1, 2, 3, 2 for the hand and 1, 2, 3, 3, 2 for the foot. This pattern is seen in hands of all individuals studied (12) but one, which has a formula of 1,2, 2, 2. The typical foot pattern is encountered in six of the twelve animals studied. In | four of the specimens the formula is 1, 2, 3, 2, 2. One individual is assymetrical for the two formulas. A final individual has a formula of 1,2, 2, 2, 2. Thus there is variation in a total of three digits, the longest (number 3) in the hand, and the longest (number 3) and next to longest (number 4) in the foot. Re- ductions do not occur in the terminal phalanges, but apparently in the penulti- mate phalanges in all instances. In no case is there a reduction of more than one phalanx per digit. The trend toward reduction is apparent even in those individuals which have the typical formula (Fig. 1). It is particularly evident in the very small amounts of bone in the penultimate phalanges of the third digits in both hands and feet. The terminal phalanges, in contrast, are not only well developed, but also display specialization. Most are distally expanded, with small lateral processes of bone. They probably are functionally significant in these climbing animals. The other phalanges are round in cross-section and unspecialized. The metapodial elements are only a little flattened, and are gen- erally unspecialized. There are seven carpals and eight tarsals in the typical j pattern of generalized Bolitoglossa. There is no tibial spur, and only a minute : crest. 1973 New Salamanders from Panama 12 Figure 4. Dorsal views of male (left, KU 116555) and female (right, KU 116575) paratypes of Bolitoglossa mimitula, new species. Habitat: With the exception of one individual found in a bromeliad on a log by day, all of these salamanders were obtained at night. One was on a boulder, and one was on a bare tree trunk; all others were on leaves of herba- ceous vegetation. None was more than 1 m above the ground. The habitat is described further in the description of the following species. Range: Known only from the vicinity of the type locality from 1810-2100 m (5937-6888 ft) elevation, north slope of Cerro Pando, Provincia de Bocas del Toro, Panama. A small series of rather stout-bodied, generalized species was obtained from the slopes of Cerro Pando. In allusion to its body form it is named: Bolitoglossa compacfa, new species Figures 2 and 5 Holotype: KU 116662, an adult female from the north slope of Cerro 13 Contributions in Science No. 248 Pando, 1920-1970 m (6298-6462 ft) elevation, Provincia de Bocas del Toro, western Panama, obtained by Charles W. Myers on 9 May 1966. Paratypes: KU 116663, same data as holotype; KU 116659-60, KU 104334 (cleared and stained), LACM 78728, 1920 m (6298 ft); KU 116664, 1810m (6937 ft) ; all from north slope of Cerro Pando, Provincia de Bocas del Toro, Panama, collected by William E. Duellman, Charles W. Myers, and Linda Trueb on 3-9 May 1966. Referred Material: Gorgas Memorial Laboratory (no number), trail to Changena, 1829-2134 m (6000-7000 ft), presumably on the north slope of Cerro Pando, Provincia de Bocas del Toro, Panama, obtained on 10 August, 1962, by Ratibor Hartmann. Diagnosis: A moderately large, stout species ( 1 adult male: 53.4 standard length; 5 adult females 68.5—74.2, mean 71.5 standard length) with moder- ately low numbers of maxillary (male, 11; mean for females, 43.6) and mod- erate numbers of vomerine (mean 23) teeth; distinguished from B. cerroensis by its stouter body form, somewhat shorter legs, and much darker general coloration (very dark black, with some light orange and yellow blotches dor- solaterally which may fuse to form an indistinct pair of stripes or a poorly defined dorsal band) ; from B. marmorea and B. sooyorum by its smaller hands and feet and less numerous teeth; from B. subpalmata by its large size and less numerous maxillary teeth. B. compacta is distinguished from all other mem- bers of the genus by the combination of slightly webbed hands and feet, its coloration, its size and dentitional features, and its relatively simple vertebral pattern in the tail base (see Wake and Brame, 1969). Description of Holotype: Adult female, snout moderately short. Nostril relatively small; labial protuberances of nasolabial grooves small and poorly developed. Canthus rostralis moderately short in length, gently arched. Stand- ard length 7.2 times head width; standard length 5.0 times snout-gular fold length. Deep unpigmented groove below eye extends almost full length of open- Figure 5. Dorsal view of holotype of Bolitoglossa compacta, new species, KU 116662. 1973 New Salamanders from Panama 14 ing, following curvature of eye, but does not communicate with lip. Eye mod- erately large, originally protuberant. Well defined postorbital groove extends posteriorly from eye as shallow depression for 2.2 mm, proceeds sharply ventrad at level of posterior end of mandible and extends across gular area as nuchal groove parallel to and 5.6 mm anterior to sharply defined gular fold. Vomerine teeth 20, arranged in a single row on each side, extending from distal edge of internal nares and curving gently toward the midline. Near the midline both rows angle sharply in the direction of the parasphenoid tooth patches, from which they are separated by 0.8 mm. Maxillary teeth 36, extend posteriorly to a point about half way through eye. Six moderately small pre- maxillary teeth protrude from well behind the lip. Tail strongly laterally com- pressed, of moderate length, 0.93 times standard length; postiliac gland clearly evident, pale and crescent-shaped. Limbs moderately long, limb interval 2; standard length 4.6 times right forelimb; 4.2 times right hind limb, 10.0 times right foot width. Webbing of hands and feet slight to moderate. Digits broad and truncate at tips. Subterminal pads present on all digits, very small on digit 1. The fingers in order of decreasing length: 3, 2, 4, 1; toes in order of decreas- ing length: 3, 4, 2, 5, 1. Measurements (in mm): Head width 10.3; snout to gular fold (head length) 14.9; head depth at posterior angle of jaw 5.3; eyelid length 4.2; eyelid width 2.3; anterior rim of orbit to snout 4.2; horizontal orbital diameter 2.5; inter- orbital distance 4.0; distance between vomerine teeth and parasphenoid tooth patch 0.8; snout to forelimb 19.9; distance separating internal nares 2.7; dis- tance separating external nares 3.6; snout projection beyond mandible 1.0; snout to posterior angle of vent (standard length) 74.2; snout to anterior angle of vent 67.2; axilla to groin 40.0; tail length 68.8; tail width at base 4.5; tail depth at base 5.3; forelimb length 16.2; hind limb length 17.7; width of right hand 5.4; width of right foot 7.4. Coloration in Alcohol: The ground color of the holotype is deep black with irregular orange-yellow spots and blotches arranged in dorsolateral rows so as to form imperfect stripes. A few large orange patches occur about the base of the tail and there are some small spots on the top of the head behind the eyes and on the snout. Eyelids are mottled orange and black. A few small scattered yellow-orange spots are present middorsally on the trunk. Ventro- lateral and ventral surfaces are uniformly black, almost as dark as the dorsum. No spots of lighter color are present ventrally. Limbs are uniformly black, but for one small light spot on the left hind limb just above the ankle. Palms of hands and soles of feet are greyish. Coloration in Life: The dorsum varies from dark dull brown with con- spicuous reddish brown blotches to deep brownish red. The venter is dull brown, and the iris is pale brown. Variation: KU 116664 and KU 116660 are colored much as the holotype. Three other paratypes have more extensive orange patching. KU 116659 has extensive yellow-orange mottling on the dorsum of the head and numerous 15 Contributions in Science No. 248 yellow-orange spots middorsally on the trunk. KU 116663 is almost banded yellow-orange, the head being nearly completely tan-orange dorsally and the trunk dorsum mottled yellow-orange and black with a large bright orange patch at the base of the tail, and smaller streaks and spots on the rest of the tail dorsally. KU 1 16661 has a broad yellow-orange stripe (almost banded) but the head is mostly black except for orange-black mottled eyelids and snout. The first third of the tail is blotched with orange-yellow. Ventrolateral and ventral surfaces are uniformly black in all. The two male paratypes are much smaller than the five females (including holotype). In addition, the male para- types have considerably broader heads (SL 6.2— 6.4, mean 6.3, times head width; females 6.8— 7.3, mean 7.0). Males have larger feet (SL 9.2 times right foot width as compared to 9.5—10.0, mean 9.8 for females) and males have longer hind limbs (SL 3.6— 4.0, mean 3.8 times right hind limb length in con- trast to 3.8— 4.5, mean 4.2, for females). Appressed limbs leave 1.5 costal folds uncovered in males, but 1.5— 2.5 (mean 2.1) in females. The males have con- siderably fewer maxillary teeth but they are also much smaller specimens than the females. See table I for additional data and measurement differences. Osteology: Information concerning osteology has been derived from one cleared and stained specimen and from radiographs of KU 1 1 6664. The skull is generalized for the genus Bolitoglossa. The premaxillary is the normal rather small structure with a moderately narrow dental process and a small palatal shelf. Frontal processes are slender and divergent. They are weak and have virtually no distal expansion. Contact with the expanded facial portions of the frontals is limited. The processes extend to about the level of the extreme anterior margin of the orbit, but do not reach the posterior margins of the nasal bones. The internasal fontanelle is very large and has a dis- tinctive shape. The fontanelle is in the form of an inverted triangle, with two long sides extending anteriorly. The posterior border is straight, and is formed by the strongly divergent facial portions of the frontal bones. The nasals are moderately large but do not protrude from the general outline of the skull. ThuSj the nasal capsules are small in a relative sense. Posteriorly the pointed margins of the nasals extend beyond the anterior-most extension of the orbit, but they do not reach the orbit, from which they are separated by the pre- frontals. The prefrontal bones are well developed and regular in outline. They overlap the frontals and are in contact with the nasals. The prefrontals are about one-fourth to one-fifth the area of the nasals, and a little less than one- half the area of the facial processes of the maxillaries. The nasolacrimal duct enters the nasal capsule through a space bounded by the posterolateral margin of the nasal, the anterolateral margin of the prefrontal, and the dorsoanterior margin of the facial process of the maxilla. The maxillaries are moderately developed, and have a large facial process. Dentition is weak, and only a part of the bone is toothed. The maxillary extends about three-fourths of the dis- tance through the eyeball. The vomers are moderate-sized bones which are separated from each 1973 New Salamanders from Panama 16 other for their entire lengths. The intervomerine fontanelle is of moderate breadth, and is only a little broader at its midpoint than near either end. Pre- orbital processes are stout and bear a rather strong dentition which is patched in the cleared and stained individual. The processes extend well beyond the lateral margins of the vomerine bodies. Teeth extend to about the midpoint of the internal nares. Frontal bones are well developed. Anteriorly the facial processes are re- stricted to lateral positions by the internasal fontanelle. The processes are long and narrow. The frontals are in close contact medially, but with rather slight articulation. The bones extend farther in a posterior direction laterally than medially. Parietals are separated by a slight gap from each other medially. They are extensively overlapped by the frontals. The parietal spurs typical of the genus are present. Otic capsules are of moderate size for the genus. There are no crests or projections. The blunt tip of the large parasphenoid is rela- tively broad, and the orbitosphenoids are well separated from each other medially. Posterior vomerine teeth are in large patches on the parasphenoid. These patches do not touch. In the single cleared and stained individual the patches contain 75 (left) and 76 (right) ankylosed, bicuspid teeth. These teeth are about the same size as the marginal teeth on the jaws. The operculum has no stilus. Quadrates are of moderate size and are joined to the skull by un- specialized, rectangular squamosals, and the cartilaginous suspensorium. The jaws, both upper and lower, are quite weak and poorly developed for an animal of this size, relative to other species of this genus. The hyobranchial apparatus is typical of members of the genus. Vertebrae have no special features. There are one cervical, fourteen trunk, one sacral, two caudosacral and 24 or 34 (two individuals) caudal vertebrae. The second trunk vertebra is the longest in the body, but caudal vertebrae 2 through 8 are as long as the mid trunk vertebrae. The first caudal vertebra shorter than the shortest trunk vertebra (the first) is the tenth. This does not include the first caudal vertebra which, together with the second caudosacral vertebra is much shortened in the constricted area at the base of the tail. Trans- verse processes are present on all but the last few vertebrae. The processes are long and nearly perpendicular to the body axis on the first caudosacral verte- bra. On the second the somewhat shorter processes are directed first in an anterior and then in a lateral direction. The processes of the first caudal verte- bra arise at the anterior end of the vertebra and proceed anteriorly and later- ally, reaching a level somewhat short of the midpoint of the second caudosacral vertebra. They are not long relative to other members of the genus. They do not cross those of the more anterior vertebra. More posterior processes have a similar origin at the anterior end of the vertebra, and they become progres- sively shorter. The hands and feet have a quite generalized structure. The phalangeal formulae are 1, 2, 3, 2 for the hands and 1, 2, 3, 3, 2 for the feet. The digits are discrete and comprised of well ossified, cylindrical phalangeal elements. 17 Contributions in Science No. 248 The terminal phalanges are well developed, with a generalized, moderately j large expansion (Fig. 2). There are seven carpals and eight tarsals. There is no tibial spur. Habitat: All specimens were found in undisturbed montane cloud forest (see Myers, 1969, for detailed description). The area, around 1920 m on the north slope of Cerro Pando, supports broad-leaved evergreen forest with a canopy height of about 20 m and an understory of palms and tree ferns. The area was extremely wet in May 1966; probably moisture is abundant con- tinuously, as evidenced by thick growths of mosses on tree trunks and logs. Four individuals were found on leaves of low herbs (< 1 m) at night; one was on a palm stem at night. At an elevation of 1810m one individaul was found by day beneath the decomposing thatch of a former shelter. j Three species of Bolitoglossa (compacta, marmorea, and minutula) and J Oedipina grandis occur in sympatry on the high northern slopes of Cerro Pando. All are active at night. The Oedipina is terrestrial, and Bolitoglossa marmorea is arboreal; individuals move about in the moss covering trunks and limbs of trees. Bolitoglossa compacta and minutula were found on leaves of low herbaceous plants; no ecological differences between the two species were noted. ! On the lower Caribbean slopes (830-910 m) of the cordillera the four highland species are replaced by an assemblage of species having broad dis- tributions in lower Central America: Bolitoglossa biseriata, colonnea, robusta, j; schizodactyla, and Oedipina collaris. I Relationships The three species described in this paper are only remotely related, de- spite the fact that they are all members of the same genus. The genus is large and diverse, and new species are being discovered regularly. Any detailed con- sideration of relationships is premature, but at least some general comments ^ can be presented. The relationships of Bolitoglossa cuna are with members of the sima group (Brame and Wake, 1972), which occur mainly west of the continental divide in Panama, Colombia and northern Ecuador. The other members of the group include B. sima, B. chica, B. biseriata, and B. silverstonei. All are similar in size, extent of hand and foot webbing, general structure of the hands and feet, head proportions and general aspects of coloration. B. cuna has more teeth than any other members of this group and is larger than B. chica. It also has a narrower head than species of similar size. In general this group is so poorly known that further comments are inappropriate. Only skeletons of B. sima and B. chica have been available for study. There is no basis for postu- lation of relationship with any other species group. Bolitoglossa minutula is a very distinctive species that does not fit in any currently recognized group. It is most similar to Bolitoglossa epimela, a species known from the Atlantic slope of Costa Rica at a considerably lower (915 m) 1973 New Salamanders from Panama 18 elevation (Wake and Brame, 1963). However, B. epimela has very large, somewhat less extensively webbed hands and feet and is a larger species, with a relatively narrower head. B. epimela has relatively few teeth for its size. The only other species that seems to be a possible close relative of B. minutula is a sympatric associate which appears to be undescribed. It is a somewhat larger species with less fully webbed hands and feet, but with similar numbers of teeth, and limbs of similar length. This species is somewhat intermediate between B. epimela and B. subpalmata in structural features (see Wake and Brame, 1963), having more teeth, a broader head and less hand and foot webbing than B. epimela, but being smaller and much slenderer than B. sub- palmata. Thus, in a structural sense only, there is something of a morphologi- cal transition from B. subpalmata to B. minutula. In contrast, we do not see any indications of relationship of B. minutula to any of the fully webbed species of the neighboring lowlands (for example, B. colonnea or B. biseriata). Bolitoglossa compacta falls in the B. cerroensis-B. marmorea-B. sooyo- rum section of the B. subpalmata group. There is a close similarity among these species in gross morphology, but structural distinctions can be recog- nized. Further, on both Cerro de la Muerte in Costa Rica (B. sooyorum, B. cerroensis) and on Cerro Pando (B. marmorea, B. compacta) two members of the section occur in sympatry. B. sooyorum and B. marmorea are apparent close relatives, as are B. cerroensis and B. compacta. Distinctions between members of the first pair are subtle, and they should probably be considered races. B. cerroensis and B. compacta are more distinct. B. compacta has shorter legs and a broader head, but the two species are similar in number of maxillary teeth, vomerine teeth and foot shape. The low numbers of maxillary teeth in this pair set them apart from other members of the subpalmata group. B. compacta is a darker salamander, but elements of the coloration, especially pattern, are similar. Light coloration tends to be yellow-orange to reddish brown in B. compacta, but lavender in B. cerroensis. Finally there are some osteological differences between the two species. It must be remembered that these are based on detailed study of only one B. compacta and two B. cerroen- sis, in addition to radiographs. However, B. compacta has prefrontal bones, which are fused with the nasals in B. cerroensis. In addition, the internasal fontanelle is much larger in B. compacta, and the premaxillary bones are less well developed. Structure of the phalangeal elements is nearly identical in the two species. Acknowledgments Duellman gratefully acknowledges the field companionship of Drs. Charles W. Myers and Linda Trueb, who worked with him in Panama. The field studies were supported by the National Institutes of Health (GM-12020) in cooperation with the Gorgas Memorial Laboratory in Panama City. Labora- tory work on Panamanian collections was supported by the National Science Foundation (NSF-GB 8139 to Duellman and NSF-GB 17112 to Wake). Fig- ure 2 was prepared by Gene M. Christman. 19 Contributions in Science No. 248 Resumen I Tres especies de salamandras, Bolitoglossa cuna, B. minutula, y B. com- I pacta, son descritas para poblaciones de Panama. I Bolitoglossa cuna es una especie con manos y pies totalmente palmeados I que habita a escasa altitud en el Territorio de San Bias. Es una especie miem- k bro del grupo sima. | Bolitoglossa minutula es la especie mas pequena del genero. Se le ubica : 5 a altitudes entre 1800 y 2000 m cerca de la frontera con Costa Rica. Tiene | algunas similitudes con la especie costaricense de tamano mas grande, B. I epimela. Bolitoglossa com pacta es una especie de gran tamano con manos y pies escasamente palmeados y simpatrica con B. minutula. Esta emparentada con j B. cerroensis de Costa Rica. ■ Literature Cited i Brame, a. H., Jr., and W. E. Duellman. 1970. A new salamander (Genus Oedi- pina) of the uniformis group from western Panama. Los Angeles Co Mus 1 Contrib. Sci. 201: 1-8. ’ Brame, A. H., Jr., and D. B. Wake. 1972. New species of salamanders (Genus Bolitoglossa) from Colombia, Ecuador and Panama. Contrib. Sci. 219: 1-34. Myers, C. W. 1969. The ecological geography of cloud forest in Panama. Amer. A Mus. Novit. 2396: 1-52. i Wake, D. B. 1966. Comparative osteology and evolution of the lungless salaman- ders, family Plethodontidae. So. Calif. Acad. Sci., Mem. 4: 1-111. Wake, D. B., and A. H. Brame, Jr., 1963. A new species of Costa Rican salamander, | genus Bolitoglossa. Rev. Biol. Trop. 1 1 : 63-73. P Wake, D. B., and A. H. Brame, Jr. 1969. Systematics and evolution of neotropical j. ,| salamanders of the Bolitoglossa helmrichi group. Los Angeles Co. Mus. Con- ^ trib. Sci. 175: 1-40. ’ p Accepted for publication June 8, 1973 1' I' I Printed in Los Angeles. California by Anderson. Ritchie and Simon on Simmnn T Rprnvpj^i i i>0 f.-fS I NUMBER 249 JUNE 30, 1973 NEW SPECIES OF MEMBRANOBALANVS HOEK AND HEXACREUSIA ZULLO (CIRRIPEDIA, BALANIDAE) FROM THE GALAPAGOS ARCHIPELAGO By Victor A. Zullo and Dea B. Beach ^ CONTRIBUTIONS IN SCIGNCE CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- able footnotes must be typed on separate sheets. Several of one kind may be placed on a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing with Latin American subjects. Summaries in other languages are not required but are strongly recommended. Summaries will be published at the end of the paper. (8) A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF. — Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to the author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor NEW SPECIES OF MEMBRANOBALANUS HOEK AND HEXACREUSIA ZULLO (CIRRIPEDIA, BALANIDAE) FROM THE GALAPAGOS ARCHIPELAGO^ By Victor A. Zullo^ and Dea B. Beach^ Abstract: A second eastern Pacific membranobalanid occurs at several localities in the Galapagos Archipelago off Ecuador. It is found in association with a small clionid sponge that burrows in nonliving parts of such hermatypic corals as Porites and Pavona. Balanus (Membranobalanus) nebrias, new species, is morpholog- ically similar to B. (M.) orcutti Pilsbry, known from southern California and the Gulf of California, and to B. (M.) declivis Darwin of the tropical western Atlantic. The new species is nota- ble for the presence of conspicuous chitinous laminae in the apices of the opercular valves. A second species of the coral-inhabiting barnacle Hexa- creusia Zullo is described on the basis of three specimens living on balanophyllid corals that were dredged from 55 to 90 m. Hexacreusia straeleni, new species, differs from the type species, H. durhami (Zullo) in its association with ahermatypic rather than hermatypic corals and in its conic rather than depressed shape. The two new species described herein are part of a broader study of the cirriped fauna of the Galapagos Archipelago and of the adjacent areas of the tropical eastern Pacific. The overall study is based on collections made by Zullo during the 1964 Galapagos International Scientific Project, on previously unstudied collections in various museums, and on collections recently made by various individuals. This preliminary report is presented to draw attention to two cryptic barnacles in the hope that knowledge of their existence will bring to light additional information on their distribution. The subgenus Membranobalanus Hoek (of the genus Balanus Da Costa) is comprised of a small number of species inhabiting sponges in tropical and warm temperate waters. Heretofore, only a single species, Balanus orcutti Pilsbry, was known from the eastern Pacific. Its known range includes south- ern California, Cape San Lucas, Baja California, and the Gulf of California, ^Review Committee for this Contribution James H. McLean William A. Newman Arnold Ross ^Research Associate in Invertebrate Paleontology, Natural History Museum of Los Angeles County, Los Angeles, California 90007; and Univ. of North Carolina, Wilmington, North Carolina 28401 3Dept. of Psychology, Duke Univ., Durham, North Carolina 27708 1 2 Contributions in Science No. 249 |( i| but B. orcutti probably will be found to occur throughout the tropical Pana- l| mic province. Balanus orcutti is related to B. declivis Darwin, which is found | at scattered localities throughout the tropical western Atlantic, and is the only I other membranobalanid known previously from the New World. These two I species, together with the new Galapagan species, form a close knit group | that is morphologically distinct from membranobalanids in other parts of the ^ world. Unlike B. orcutti and B. declivis which are found in a variety of sponges, the new species appears to be restricted to a single species of boring clionid > sponge. The sponge usually occurs in nonliving, generally basal parts of such hermatypic corals as Porites and Pavona, but in one instance it was discov- ered burrowed in weathered vesicular basaltic cobbles. Corals from several f other tropical eastern Pacific localities have been examined, but to date this I new species has been found only in the Galapagos Archipelago, where it is , widespread and abundant. I The six plated, coral-inhabiting barnacle Hexacreusia Zullo, heretofore | known only by its type species, H. durhami (Zullo), is a common symbiont of Porites californica Verrill in the Gulf of California and the Tres Marias Islands, and of P. californica and possibly P. lobata (Dana) on the Pacific coast of Panama (Zullo, 1961; Zullo, Beach and Carlton, 1972). Corals from the Galapagos, Cocos Island off Costa Rica, and Clipperton Island off Mexico have been examined with some care, yet none has yielded symbiotic barnacles. | The new Galapagan species was obtained under unexpected circum- ! stances. The three known specimens were found attached to two ahermatypic I balanophyllid corals dredged from 55 to 90 m off Beagle Island. Although this habitat is similar to that of the northeastern Atlantic coral barnacle Pyrgomina anglica (Sowerby), the possibility of this type of association had been overlooked by us in searching for additional occurrences of Hexacreu- sia. It is, thus, likely that other eastern Pacific localities will yield Hexacreusia \ upon re-examination. . Systematic Descriptions Suborder BALANOMORPHA Pilsbry, 1916 Family BALANIDAE Leach, 1817 Genus BALANUS Da Costa, 1778 Subgenus MEMBRANOBALANUS Hoek, 1913 Balanus (Membranobalanus) nebrias, new species Figures 1-17 Diagnosis. Conspicuous chitinous laminae intercalated with shelly laminae in apices of opercular valves; adductor ridge of scutum absent; scutal articular ridge short, but prominent; tergal spur short and broad; labrum narrow, 1973 New Species from the Galapagos 3 pointed, triangular in shape; mandible with five distinct teeth on cutting edge; armature of Cirrus IV simple; five pairs of spines on medial articles of Cirrus VI; reddish purple ciri marked with large white spots. Description. Shell (Fig. 5) high conic or cylindric, white, with toothed orifice; compartmental plates solid, easily disarticulated; parietes ornamented externally with fine, closely spaced, irregular growth lines crossed by finer, closely spaced longitudinal striae; rostrum elongate, half again as long as other plates, roughly diamond-shaped with lower half broader and basal mar- gin either broadly rounded or pointed; apices of rostrum and carina in un- broken specimens with deeply incised V-shaped notches (probably caused by cirral rasping); sheath of rostrum one-half to two-thirds length of plate; sheath on other plates two-thirds to three-fourths length of plate; basal edge of sheath not free in most specimens, but merging into parietes below; inte- rior of parietes smooth, basal edge not denticulate; radii absent, or represented by narrow, irregular ledges; alae wide with oblique summits, and without den- ticulation on sutural edges; basis membranous; a single specimen possesses an incomplete ‘basal plate’ formed by the inward projection of secondary basal ledges from each compartmental plate (Fig. 2); the width of the basal ledge varies, with that of the carina being broadest, that of the rostrum narrowest, and those of the other plates intermediate in width; the ledge apparently is laid down secondarily after formation of the compartmental wall, and con- sists only of the continuation of the inner lamina (ae) of the compartmental plate in a different growth plane, and, therefore, does not represent a true balanid basis. Scutum (Figs. 1, 3, 6, 8) thick, convex, white, with yellow adherent epidermis on exterior; basal margin shorter than or equal to tergal margin; tergal margin reflexed; exterior ornamented with sharp, closely spaced growth ridges, every other ridge carrying over onto occludent margin as an occludent tooth; growth ridges incised by fine, closely spaced radial striae, especially evident in median part of valve; articular ridge prominent, triangular, reflexed, extending far beyond border of scutal margin, and restricted to upper half or two-thirds of scutal margin; articular furrow narrow, deep; adductor ridge absent or represented by slightly raised border of large, well delimited, lenticu- lar adductor muscle pit; depressor muscle pit large, deep, triangular, extending well up under inner lamina of scutum; apex of scutum often divided by two or three radiating furrows into longitudinal ridges; apical shell laminae of scutum above top of adductor muscle pit intercalated with prominent chitinous lami- nae imparting yellow-brown color to apical part of scutum; upper margin of articular ridge often chitinous also. Tergum (Figs. 4, 7, 9) heavy, broad, convex, white, with basal margin shorter than scutal margin; exterior ornamented by fine, closely spaced growth ridges, often with an adherent yellow-brown epidermis bearing golden hairs; spur fasciole broad, open, well delimited by an abrupt change in elevation on either side; carinal side of exterior often crossed by one or two incised longi- 4 Contributions in Science No. 249 li Figures 1-9. Shell and opercular valves of Balanus (Membranobalanus) nebrias, n. j sp. (1) scutum interior, paratype no. 4; (2) carina interior, paratype no. 5, showing i inflected basal margin; (3) scutum interior, holotype; (4) tergum exterior, holotype; j (5) shell profile, paratype no. 6; (6) scutum exterior, holotype; (7) tergum interior, ' holotype; (8) scutum exterior, paratype no. 4; (9) tergum interior, paratype no. 4. ; I Scale A: Figs. 1, 3, 4, 6-9; Scale B: Fig. 2; Scale C: Fig. 5. | i 1973 New Species from the Galapagos 5 j tudinal furrows near and parallel to fasciole border; tergal spur short, trun- cate, not parallel to basal margin, close to but distinct from basiscutal angle; ’ depressor muscle crests absent; articular ridge short, straight, restricted to center of apical third of valve, with chitinous laminae intercalated with cal- i careous laminae as in scutum; chitinous laminae especially conspicuous as I lining of articular furrow. Labrum (Figs. 14, 15) sharply triangular, divided at apex by deep notch; two or three teeth usually on either side of notch; sides of notch lined with fine hairs; Paratype no. 2 with two teeth at either edge of notch and a third along * outer slope on either side. Palps (Fig. 10) kidney-shaped; superior margin concave, lined by numer- ous equal sized setae; inner and inferior margins bearing fewer, but longer I setae. i Mandible (Fig. 12) with five teeth on cutting edge; first three teeth of I approximately equal size, though varying in individuals; second tooth bifid, in center of cutting edge; third tooth in center of lower half of cutting edge, often bifid with blunt accessory denticles; fourth and fifth teeth smaller, of about equal size, molariform to varying degrees; inferior angle and cutting edge immediately above often provided with sharp, multi-pronged spines; inferior margin lined with row of long setae; superior margin with a few long setae. Maxilla I (Fig. 13) with straight edge; uppermost pair of spines large, with upper spine largest of pair; notch below upper pair reduced to slit, some- times occupied by small spine; middle section of cutting edge with five to eight smaller, right-left alternating spines; middle section followed by a pair of spines in lower third equal in size to apical pair, with upper spine largest of pair; inferior angle with five to six short spinules; inferior and superior margins clothed with long setae. Maxilla II (Fig. 11) bilobed; outer lobe tipped with reddish purple, long, narrow, with parallel sides, superior and posterior margins densely setose; inner lobe protrudent, circular in outline, densely setose. Cirri and trophi appear to exhibit some asymmetry, which in cirri may be result of nipping by predators; Cirrus I with grossly unequal rami; anterior ramus about three times length of posterior ramus (in Paratype no. 3, anterior more than four times length of posterior); anterior ramus reversed, antenni- form; posterior ramus more setose than anterior, with somewhat protuberant articles; Cirrus II with outer ramus one-third again as long as inner ramus; articles of both rami slightly protuberant, covered with setae somewhat longer than those of Cirrus I; Cirrus III (Fig. 7) with outer ramus longer than inner; similar in size and structure to Cirrus II; proximal articles fused or partially fused; Cirrus IV (Fig. 16) with inner ramus half again as long as outer; pedicel nearly as long as outer ramus, imparting whiplike appearance to cirrus; outer ramus bearing large, recurved teeth, usually three in number, on anterior sur- faces of proximal and medial articles; distal articles and pedicel bearing numer- ous, small, upright teeth along distal margins of anterior surfaces; inner ramus 6 Contributions in Science No. 249 Figures 10-17. Trophi and cirral detail of Balanus (Membranobalaniis) nebrias, n. sp. (10) palp, paratype no. 4; (11) second maxilla, holotype; (12) mandible, para- type no. 4; (13) first maxilla, paratype no. 4; (14) labrum, holotype; (15) labrum, paratype no. 4; (16) detail of armature of Cirrus IV, paratype no. 1; (17) detail of armature of Cirrus III, paratype no. 4. 1973 New Species from the Galapagos 7 I also with small upright teeth at distal anterior margins of articles; Cirrus V I with subequal to equal, long, slender rami; outer ramus bearing small erect teeth on distal margins^ of medial articles; Cirrus VI with long, slender, equal I rami, medial articles bearing four large pairs and a fifth minute pair of spines on the anterior surfaces; cirri reddish purple in color, and marked by large, I white, circular spots; penis without basidorsal point, bearing four longitudinal rows of setae. The number of articles on individual cirri are tabulated for the following specimens (those in parentheses indicate broken rami) : Holotype : Cirrus I II III IV V VI outer (ant.) ramus 28 - 14 13 20 22 inner (post.) ramus 8-9 - 12 17 18 22 Paratype no. 1 : outer (ant.) ramus 24 11 14 13 19 21 inner (post.) ramus 8 9 11 18 20 22 Paratype no. 4: outer (ant.) ramus 27 11 13-15 (10-17) 16 16 inner (post.) ramus 9 9-10 11-13 (10-11) 16 17 Dimensions. Paratype no. 2: height of rostrum, 8.9 mm; carinorostral diameter of base, 6.4 mm; lateral diameter of base, 4.9 mm; greatest diameter of orifice, 1.8 mm. Disposition of Type Material. The holotype and all paratypes are de- posited in the Natural History Museum of Los Angeles County. Referred specimens have been deposited in the California Academy of Sciences, San Francisco, the University of California Museum of Paleontology, Berkeley, and the United States National Museum, Washington, D. C. Holotype: South Channel (channel between Baltra and Santa Cruz Islands), Baltra Island, Gala- pagos; in red clionid sponge burrowed in basaltic rock at 2 m depth; Feb. 12, 1964, V. Zullo collector. Paratype no. 1: Sullivan Bay, James Island, Gala- pagos; in red clionid sponge burrowed in Leptoseris at 2 m depth; Feb. 16, 1964, V. Zullo collector. Venedig, north of Conway Bay, northwestern Santa Cruz Island, Galapagos; in red clionid sponge burrowed in Porites at 2-3 m depth; Feb. 19, 1964, V. Zullo collector. Paratype no. 3: Academy Bay, Santa Cruz Island, Galapagos; in red clionid burrowed in Pavona; Feb. 25, 1964, J. W. Durham collector. Paratype nos. 4 and 5 : locality same as for holotype. Discussion. The following species are presently included in the subgenus Membranobalanus : Balanus basicupula Suhaimi (1966: 65), Singapore. Balanus cuneiformis Hiro (1936: 627), Port Darwin, Australia; Seto, Japan (Hiro, 1939). 8 Contributions in Science No. 249 Balanus declivis Darwin (1854: 275), West Indies, Jamaica; Cape Sable, Florida, Bermudas (Pilsbry, 1916: 230); includes “var. cuspidatus” of Verrill (1902: 22), Bermudas. Balanus longirostrum Hoek (1913: 205), Celebes: west coast of Sumatra (Nilsson-Cantell, 1921: 340) ; Sunda Strait (Broch, 1931: 85) Gulf of Manaar, Pamban (Nilsson-Cantell, 1938: 54); includes “var. krusa- daiensis” of Daniel (1956: 26) , Bay of Bengal. Balanus orcutti Pilsbry (1907: 361; 1916: 233), Gulf of California. Barnard (1924: 74) recorded this species from Algoa Bay, South Africa, but comparison of the cirri and trophi of B. orcutti from south- ern California (see below) with Barnard’s description suggests that the South African specimens represent a species similar to B. longirostrum. Balanus nebrias most closely resembles the New World species B. declivis and B. orcutti, but differs from both in the presence of large white spots on the reddish purple cirri and of chitinous lamellae in the apices of the opercular j valves. The concentration of chitinous material in the apex of the opercular pyramid is apparently a device to inhibit the decalcification of the exposed parts of the opercular valves by the boring sponge inhabited by the barnacle. The erosive effects of this sponge on the calcareous lamellae is quite evident I (Figs. 4, 6), especially on the exterior surfaces of the valves. It also appears | that the chitinous lamellae are not secondary deposits, but are laid down alter- nately with the calcareous lamellae and retain the form and structure of the j valve. The shell wall of the barnacle does not require protection against dissolu- tion by the sponge, as those sponge cells involved in boring are concentrated on the sponge’s exterior. The opercular valves would require protection pri- marily against damage by newly settled, actively boring sponges. Damage that might occur at the apices of the compartmental plates would be compen- sated by continued growth at their basal margins. However, such damage to the opercular pyramid would be irreparable. Specifically, B. nebrias differs from B. declivis in the shorter articular ridge of the scutum, the more nearly equilateral tergum with a shorter, broad- er spur, a more pointed, triangular labrum, a less complex armature on the anterior surface of the outer ramus of Cirrus IV, and five rather than four pairs of spines on the median articles of Cirrus VI. The new species differs from B. orcutti in the much shorter rostrum which is not ribbed below the sheath, the absence of an adductor ridge on the scu- tum, and the shape of the articular ridge of the tergum. The trophi and cirri of B. orcutti have not been described. Based on specimens in the Allan Han- cock Foundation collection from southern California (described below), B. nebrias is seen to have a more angular and narrower labrum, a less angular palp, a mandible with five rather than four distinct teeth, a simpler armament of Cirrus IV, and five rather than four pairs of spines on the median articles of Cirrus VI. 1973 New Species from the Galapagos 9 Figures 18-23. Trophi and Cirrus IV of Balamis (Membranobalanus) orcutti Pilsbry. (18) mandible; (19) first maxilla; (20) labrum; (21) second maxilla; (22) palp; (23) detail of armature of Cirrus IV. (Figs. 18, 19, 21, hypotype no. 1, AHF loc. 1153; Figs. 20, 22, 23, hypotype no. 2, AHF loc. 1 170; southern California). Balanus nebrias differs markedly from B. basicupula, B. longirostrum, and B. cuneiformis in the presence of prominent recurved teeth on Cirrus IV, and in the form of the tergal articular ridge. In addition, B. longirostrum lacks a scutal depressor muscle pit, and the second and third cirri bear small, erect teeth along the distal margin of the articles. The shell of B. basicupula dif- fers in being essentially membranous and flexible. Supplementary Description: mouth parts and cirri of Balanus orcutti Pils- bry (Figs. 18-23). Labrum (Fig. 20) with deep central notch lined with fine setae; labral crest rounded, with sloping sides, but not acutely triangular; two teeth on one side of notch only. Palps rectangular; superior margin concave, lined by short setae; inner and inferior margins bearing slightly longer setae. Mandible with four teeth and molariform area above inferior angle; first three teeth approximately equal in size, with second tooth in center of cutting edge and third tooth in center of lower half of cutting edge; second and third teeth bifid; fourth tooth small, often merging with molariform area 10 Contributions in Science No. 249 below; small spinules present at inferior angle in Hypotype no. 1; superior and inferior margins lined by short, straight spines. Maxilla I with single pair of large spines at top of cutting edge, followed by well-developed notch bearing one or two small spines; central area of cut- ting edge with six to ten right-left alternating spines; lowermost pair of spines in Hypotype no. 1 not distinguishable from those of central part of cutting edge; cutting edge of maxilla straight. Maxilla II bilobed; outer lobe ellipsoid, inner lobe circular; both lobes densely setose. Cirrus I with grossly unequal rami; anterior ramus three times longer than posterior, antenniform, reversed; posterior ramus densely setose, with protuberant articles; Cirrus II with outer ramus longer than inner, densely setose; Cirrus III subequal to equal, similar in structure to Cirrus II; Cirrus IV with outer ramus shorter than inner ramus, bearing large recurved teeth on anterior surfaces of proximal and medial articles, and large erect teeth on distal anterior surfaces of proximal and medial articles and pedicel; pedicel long, over half the length of outer ramus; inner ramus with small erect spines on distal anterior surfaces of medial articles; Cirrus VI with equal rami; an- terior surfaces of medial articles with four pairs of spines; penis bearing four longitudinal rows of setae, and with prominent basidorsal point. The number of articles for the individual cirri are as follows (those in parentheses indicate broken rami) : Hypotype AHF no. 1 : Cirrus I II III IV V VI outer (ant.) ramus 30 15 14 20-21 (28)-31 (24-30) inner (post.) ramus 10 12 15 30-24 35-31 37-34 Hypotype AHF no. 2 : outer (ant.) ramus 27-28 11-14 13-14 22-19 (23)-32 (30)-36 inner (post.) ramus 9 10-11 14 23-25 (30-31) (33-32) Material Examined. Two hypotypes are deposited in the Allan Hancock Foundation (AHF) collection. Hypotype no. 1: AHF locality 1153-40, 35-46 fathoms on mud bottom, east of Long Point, Santa Catalina Id., Los Angeles County, California. (33° 24' 10" N., 118° 21' 25" W.). July 5, 1940. Hypo- type no. 2: AHF locality 1 170-40, 80-100 fathoms on sand and gravel. Isthmus Cove, Santa Catalina Id., Los Angeles County, California. (33° 26' 24" N., 118° 27' 35" W.). August 19, 1940. On the basis of the specimens of B. orcutti and B. nebrias described here- in, it would appear that certain characters that have been judged significant in the differentiation of species of Membranobalanus are quite variable. The specific characters implied are the form and length of the rostrum and the width and development of the tergal spur. In B. orcutti the rostrum is usually twice the height of the other compartmental plates and tongue- or fingernail- 1973 New Species from the Galapagos 11 shaped, but a number of specimens examined have a short rostrum which may be either broadly rounded or narrow and pointed. The terga of speci- mens from the Allan Hancock Foundation localities have spurs much more like that figured for B. declivis (Pilsbry, 1916, pi. 55, figs. Ic, d) than that of the type of B. orcutti (Pilsbry, 1916, pi. 55, figs. 2a, c). The width of the spur in relation to the total width of the basal margin is also misleading, and de- pends particularly on the point of measurement: whether at the root of the spur or at its extremity. In the former case the tergum of B. orcutti figured by Pilsbry equals two-thirds of the width of the basal margin, but that of B. declivis is larger, occupying nearly three-fourths of the basal margin. When measured at the extremity, the same tergal spur is 55 percent of the margin in B. orcutti and 53 percent, or smaller in B. declivis. It is probable that the form both of the rostrum and the opercular valves is governed by age and individual habitat of the specimen. The identification of B. orcutti from Algoa Bay, South Africa has always appeared doubtful because of the known distribution of membranobalanid species. Although Barnard (1924) did not illustrate his material, comparison of his description of the mandible and second maxilla of the South African specimens with those described herein from southern California indicate significant differences. Barnard’s specimens appear to be more closely related to B. longirostrum with which they also have closer geographic affinity. Etymology. The specific name “nebrias” is from the Greek for “spotted like a fawn,” referring to the color pattern of the cirri. Genus HEXACREUSIA Zullo, 1961 Hexacreusia straeleni, new species Figures 24-34 Diagnosis. Solid, high conic to globular shell with large, untoothed ori- fice and regularly ribbed shell wall; basis solid, cup-shaped; scutum thick, nar- row, with well-developed internal adductor shelf; tergum thin, narrow, with truncate tergal spur continuous with basiscutal angle. Description. Shell (Fig. 25) small, light pink to light purple, high conic to globular; orifice large, subtrigonal to diamond-shaped, not toothed; parietes solid, exteriors regularly ribbed; radii broad, with horizontal summits; sheath long, extending nearly to base of shell wall; basis cup-shaped. Scutum (Figs. 24, 28) thick, narrow, with reflexed tergal margin; ex- terior ornamented by growth increments only; basal margin short, about three- fourths the length of tergal margin; occludent margin straight, toothed; articu- lar ridge low, long; articular furrow narrow, shallow; a low, thin, arched ridge extends vertically across lower third of valve on tergal side of center; an “adductor shelf” continuous with occludent margin and occludent side of articular ridge projects out over the inner surface of the scutum, leaving a deep pocket beneath it; shelf bears deep, narrow, V-shaped notch centrally; 12 Contributions in Science No. 249 Figures 24-28. Shell and opercular valves of Hexacreusia straeleni, n. sp., paratype no. 1. (24) interior of scutum; (25) shell (holotype in background) attached to balanophyllid coral; (26) interior of tergum; (27) exterior of tergum; (28) exterior of scutum. 1973 New Species from the Galapagos 13 small, deep, oval adductor muscle pit on lower half of “adductor shelf”; de- pressor muscle pit small, shallow, situated on basal margin at basitergal angle. Tergum (Figs. 26, 27) thin, narrow; carinal side of exterior radially ribbed; articular ridge low, thin; articular furrow broad, shallow; depressor muscle crests numerous, well developed; scutal and carinal margins straight; tergal spur broad, nearly one-half width of basal margin, situated at and not differentiated from basiscutal angle; spur furrow open, differentiated only by change in direction of growth lines. Labrum (Fig. 30) deeply notched, with three teeth on the left and two teeth on the right side of notch. Palps rectangular, with short, stout, curved, coarsely pectinate setae on the superior margin, and long, slender, nonpectinate setae on the inner mar- gin. Mandibles (Fig. 29) with five distinct teeth, not including inferior angle; first three teeth of equal size; bifid second tooth at middle of cutting edge; third tooth trifid; fourth tooth with accessory denticles; inferior and superior margins of mandible setose. Maxilla I (Fig. 31) with straight cutting edge and with small notch be- low uppermost two large spines; center of cutting edge bears four smaller spines followed by a large pair of spines; base of cutting edge bears tuft of spinules; inferior and superior margins setose. Maxilla II (Fig. 32) bilobed, anterior lobe small, rounded, posterior lobe large, attenuated; both lobes densely clothed with pectinate setae. Outer ramus of Cirrus I antenniform, twice as long as stout inner ramus; rami densely setose, with pectinate distal setae; rami of Cirrus II short, stout; outer ramus one-fourth again as long as inner; rami densely setose with pec- tinate distal setae; rami of Cirrus III (Fig. 33) short, stout; inner ramus one- third again as long as outer; both rami bearing prominent sharp teeth along anterior margins of articles; Cirri IV-VI unarmed, except for occasional tufts of minute spinules near distal edges of articles; long, slender, subequal; a maximum of one small proximal pair and three large distal pairs of setae per article on Cirrus VI (Fig. 34). The number of articles per cirrus of the holotype is as follows : Cirrus I II III IV V VI outer ramus 13-14 8-9 12 18 21 24 inner ramus 7 7-8 9-10 21 21 24 Occurrence and Type Disposition. Holotype and paratype nos. 1 and 2 are deposited in the Natural History Museum of Los Angeles County. The specimens were dredged by Fritz and Carmen Angermeyer off Beagle Island, Galapagos at depths between 55 and 90 m (date of collection unknown). Discussion. Hexacreusia straeleni can be distinguished from H. durhami 14 Contributions in Science No. 249 Figures 29-34. Trophi and cirri of Hexacreiisia straeleni, n. sp., paratype no. 1. (29) mandible; (30) labrum; (31) first maxilla; (32) second maxilla; (33) detail of armature of Cirrus III; (34) setation of median articles of Cirrus VI. 1973 New Species from the Galapagos 15 by its choice of coral host, by shell shape, and by opercular valve morphology. Shell shape can be attributed in part to partial embedment in the balanophyllid coral. However, the high conic to globular shell wall of H. straeleni would ap- pear to be independent of habitat and differs, therefore, from the patelliform wall of H. durhami. The opercular valves of H. straeleni are both thicker and narrower than those 'of H. durhami. The scutum of H. straeleni has a nar- rower “adductor shelf,” a shorter central internal ridge that is restricted to the basal part of the valve, and smaller adductor and depressor muscle pits. The tergum of H. straeleni is notable for its truncate tergal spur that is con- fluent with the basiscutal angle. Etymology. This species is named for the late Dr. Victor van Straelen, first president of the Charles Darwin Foundation for the Galapagos Islands. Acknowledgments We thank Mr. and Mrs. Fritz Angermeyer of Academy Bay, Galapagos for providing the specimens of Hexacreusia straeleni, and the organizers of the 1964 Galapagos International Scientific Project for making possible the col- lection of specimens of Membranobalanus nebrias. Drawings were executed by Miss Susan P. Heller and Mrs. Ruth von Arx of the Marine Biological Laboratory, Woods Hole, Massachusetts. Support for this study was pro- vided by the National Science Foundation (grant nos. GB-4507, GE-2370), the California Academy of Sciences, the Marine Biological Laboratory, Woods Hole, and the University of North Carolina at Wilmington. Resumen Dos especies nuevas de Cirripedia Thoracica de las Isles Galapagos son descrites. Balanus (Membranobalanus) nebrias, especie nueva, se halla en las esponjas amadrigadas en las bases de corales. La especie nueva se parece a B. declivis Darwin y B. orcutti Pilsbry de la America tropical. Hexacreusia straeleni, especie nueva, fue encontrada a los fondos de 55 m a 90 m en los corales balanophyllides, y es la segunda especie conocida de Hexacreusia. Literature Cited Barnard, K. H. 1924. Contribution to the crustacean fauna of South Africa, no. 7. Cirripedia. Ann.S. Afr. Mus. 20( 1 ) : 1-103, pi. 1. Broch, H. 1931. Papers from Dr. Mortensen’s Pacific Expedition 1914-16, LVI. Indomalayan Cirripedia. Vidensk. Meddel. fra Dansk naturhist. Foren. Koben- havn 91 : 1-146, text figs. 1-41. Daniel, A. 1956. The Cirripedia of the Madras coast. Bull. Madras Gov’t. Mus. (n.ser.) (Nat. Hist.) 6(2): 1-40. pis. 1-10. Darwin, C. 1854. A monograph on the sub-class Cirripedia, Balanidae, Verrucidae. Ray Soc. London. 684 pp., 30 pis., 1 1 text figs. Hiro, F. 1936. Report on the Cirripedia collected in Malayan waters by the ship “Zuiho-maru.” Japan. J. Zool. 6(4) :621-636, text figs. 1-9. 16 Contributions in Science No. 249 . 1939. Studies on the cirripedian fauna of Japan, III. Supplementary notes on the cirripeds found in the vicinity of Seto. Mem. Col. Sci., Kyoto Imp. Univ. (ser. B) 15(2) :237-244, text figs. 1-2. Hoek, P. P. C. 1913. The Cirripedia of the Siboga Expedition, B. Cirripedia sessilia. 5//70g^2-Expeditie, 31b:i-xxv, 129-275, pis. 11-27, text figs. 1-2. Nilsson-Cantell, C. A. 1921. Cirripeden-Studien. Zool. Bidrag Uppsala 7:75-390, pis. 1-3, text figs. 1-89. . 1938. Cirripedes from the Indian Ocean in the collection of the Indian Museum, Calcutta. Mem. Indian Mus. 13(1): 1-81, pis. 1-3, text figs. 1-28. PiLSBRY, H. A. 1907. Notes on some Pacific cirripedes. Proc. Acad. Nat. Sci. Phila- delphia 59:360-362, pi. 29. . 1916. The sessile barnacles (Cirripedia) contained in the collections of the U. S. National Museum; including a monograph of the American species. Bull. U. S. Nat. Mus. 93:i-xi, 1-366, pis. 1-76, text figs. 1-99. SuHAiMi, A. 1966. A new species of Balanus (Crustacea: Cirripedia) from Singa- pore. Bull. Nat. Mus. Singapore 33(9) :65-68, text figs. 1-2. Verrill, a. E. 1902. Additions to the fauna of the Bermudas from the Yale Expedi- tion of 1901, with notes on other species. Trans. Connecticut Acad. Arts and Sci. 11(1): 15-62, pis. 1-9, text figs. 1-6. ZuLLO, V. A. 1961. A new subgenus and species of coral-inhabiting barnacle. Veli- ger 4(2) :71-75, pi. 17, text figs. 1-2. , D. B. Beach, and J. Carlton. 1972. New barnacle records (Cirripedia, Thoracica). Proc. Calif. Acad. Sci. (4)39(6) :65-74, figs. 1-22. Accepted for publication May 15, 1973 Printed in Los Angeles, California by Anderson. Ritchie and Simon on Simpson Lee 5c^7‘73 Gal NUMBER 250 JULY 5, 1973 ADDITIONAL TYPE SPECIMENS OF FOSSIL INVERTEBRATA IN THE COLLECTIONS OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY By Gale G. Sphon W- CONTRIBUTIONS IN SCICNCE NATURAL HISTORY MUSEUM • LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and char- acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- um, other technical papers will be considered. All manuscripts must be recommend- ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM. — (1) The 1972 CBE Style Manual, third edition (AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- able footnotes must be typed on separate sheets. Several of one kind may be placed on a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing with Latin American subjects. Summaries in other languages are not required but are strongly recommended. Summaries will be published at the end of the paper. (8) A diagnosis must accompany any newly proposed taxon. (9) Submit two copies of manuscript. ILLUSTRATIONS. — All illustrations, including maps and photographs, will be referred to as figures. All illustrations should be of sufficient clarity and in the proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE Style Manual, third edition (AIBS) in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photographic prints of good contrast. Original illustrations and art work will be returned after the manuscript has been published. PROOF.— Authors will be sent galley proof which should be corrected and returned promptly. Any changes or alterations, other than typographical corrections, will be billed to Ae author. Unless otherwise requested, page proof also will be sent to the author. One hundred copies of each paper will be given free to each author or divided equally among multiple authors. Orders for additional copies must be sent to the Editor at the time corrected galley proof is returned. Appropriate order forms will be included with the galley proof. Virginia D. Miller Editor ADDITIONAL TYPE SPECIMENS OF FOSSIL INVERTEBRATA IN THE COLLECTIONS OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY^ By Gale G. Sphon^ Abstract; This catalogue lists additional type and figured specimens of invertebrate fossils (including paleoentomology) in the collections of the Natural History Museum of Los Angeles County. Each entry includes name, author and date of original publication, bibliographic reference, type category and museum type number, age, formation (if known), and locality; some entries are annotated. The list is arranged phylogenetically from phyla to orders and alphabetically below that level. Species have been listed in their original genera and have been placed in what is currently considered to be the correct family. An index to species is provided. iReview Committee for this Contribution Charles L. Hogue Edward C. Wilson ^Curatorial Assistant, Invertebrate Zoology, Natural History Museum of Los Angeles County, Los Angeles, California 90007 INTRODUCTION This is the* second list of type and figured specimens of invertebrate fossils in the Natural History Museum of Los Angeles County and is published in compliance with article 7 2D of the International Code of Zoological Nomenclature. The first list (Wilson and Bing, 1970) excluded paleoentomological fossils while the present list includes both insects and subsequent additions from other phyla. These specimens are primarily from the collections of: W. Dwight Pierce; Georg Statz; and the California Institute of Technology, Division of Geological and Planetary Sciences. The first two collections are the property of the museum and the third has been deposited on permanent loan. The 673 entries in this list are spread among eight phyla and range in age from Devonian through Pleistocene. There are 339 holotypes, 560 syntypes, 2 lectotypes, 529 paratypes, 31 paralectotypes, 2 allotypes (an entomological term for a paratype of the opposite sex of the holotype), 353 figured specimens not in the above categories, and 41 “plastotypes” (casts or copies of type specimens housed in other institutions). Fourteen holo- types, 26 syntypes, 23 paratypes, and 42 figured specimens have not been located and are listed as MISSING. Information about the whereabouts of missing specimens should be communicated to the museum. 1 2 Contributions in Science No. 250: The list is arranged phylogenetically by phylum, class, and order and alphabetically by family, genus, and species group names. Each entry in- ■ eludes the genus and species group names of the original author and date of ■ publication, bibliographic reference, type category and The Natural History i Museum of Los' Angeles County type number, age, formation (if known), I and locality. Each specimen is numbered separately with the rare exception of an indeterminable number being present on a rock face, in which case a single number is given the aggregate. Species included in new genera are marked by an asterisk ( * ) . Acknowledgements I wish to extend my thanks to Dr. Edward C. Wilson for the opportunity to work with this material and my appreciation for the help given me with j the project to the museum entomologists Julian Donahue, Roy Snelling, and in particular. Dr. Charles Hogue who helped bring the insect classification up to date. I especially wish to thank Mrs. Gerda MacGregor for her help I with the Statz translations and Mrs. Betty Ponti for having had the foresight to have compiled Dr. Pierce’s bibliography and a list of his new taxa before his death in 1967. Phylum PROTOZOA Class SARCODINA Order FORAMINIFERIDA ActinocycUna aster Family DISCOCYCLINIDAE WOODRING, 1930:152, pi. 14, figs. 3-6; pi. 16, figs. 2 and 4; pi. 17, fig. 1. ' Holotype 4024 (pi. 14, figs. 3 and 5); 4 paratypes 4025 (pi. 14, fig. ,6); i 4026 (pi. 16, fig. 2); 4027 (pi. 16, fig. 4); all Eocene; W side of Canada j de los Sauces, Santa Barbara County, California; 4028 (pi. 17, fig. 1); Eocene; S slope of Jalama Creek, Santa Barbara County, California. Dioscocylina psila WOODRING, 1930:148, pi. 14, figs. 2, 4-6; pi. 15, figs. 1-4; pi. 17, fig. 1. i 15-Hsyntypes 4028 (pi. 17, fig. 1); 4029-4033 (pi. 14, fig. 2); 4034 and 4311 (uncounted number of syntypes; single type numbers assigned to the two lots; pi. 14, figs. 5-6); 4035 (pi. 15, fig. 1); 4036 (pi. 15, fig. 2); 4037 (pi. 15, Eg. 3); 4038 (pi. 15, fig. 4); 4039-4042 (unfigured); Eocene; Canada de los Sauces, Santa Barbara County, California. Family NUMMULITIDAE Operculina cf. ocalana Cushman, 1921 WOODRING, 1930 as pi. 14, fig. 1. Figured specimen 4043; Eocene; Canada de los Sauces, Santa Barbara County, California. 1973 Additional Type Specimens 3 Phylum COLENTERATA Class ANTHOZOA Order SCLERACTINIA Family DENDROPHYLLIIDAE Dendrophyllia hannibali NOMLAND, 1916:67. 3 paratypes 4252-4254 (all unfigured); Oligocene, Lincoln Formation; lyz mi above Porter, Grays Harbor County, Washington. Phylum PLATYHELMINTHES Class TURBELLARIA ^Gibronea miocenae PIERCE, 1964:82, fig. 1-4. Holotype 2494; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. Phylum ASCHELMINTHES Class ROTIFERA ^Pararotifera enigmatica PIERCE, 1964:85, figs. 6-7. Holotype 2496; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. Class NEMATODA Family APHELENCHIDAE ^Mioaphelenchus gamagei PIERCE, 1964:84, fig. 5. Holotype 2495; Miocene, Barstow Formation; southern Calico Mountains San Bernardino County, California. Phylum BRACHIPODA Class ARTICULATA Order TEREBRATULIDA Family CRANAENIDAE '^Maclarenella maculosa STEHLI, 1955:869, pi. 87, fig. 1-2, 4-5, 7. Paratype 4044; Devonian, Waterways Formation; 4-6 mi N of McMurray, Athabaska River, Alberta, Canada. Family DIELASMATIDAE Dielasma sp. STEHLI, 1955 as pi. 87, figs. 14-19. 4 figured specimens a255 (fig. 14-15 and 18); 4256 (fig. 16); 4257 (fig. 17); 4258 (fig. 19); Permian; Getaway Member, Cherry Canyon Formation; Guadalupe Mountains, Texas. Dielasma elongatum (Schlotheim, 1816) STEHLI, 1956 as pi. 40, fig. B4. Figured specimen 4045; Permian; Zechstein, near Possneck, Thuringa, Germany. 4 Contributions in Science No. 250 Fletcherina amygdala (Dana, 1847) STEHLI, 1961a as pi. 61, fig. 10. Figured specimen 4046; Permian, Upper Marine Series; Gerringong, New South Wales, Australia. Fletcherina inversa (de Koninck, 1877) STEHLI, 1961a as pi. 61, figs. 8-9, 12-13, 16 and 26. 2 figured specimens 4047 (fig. 13); 4312 (figs., 8-9, 12, 16 and 26); Permian, Upper Marine Series; Wyro, near Ulladulla, New South Wales, Australia. *Lowenstamia texana STEHLI, 1961b:461, pi. 62, group d, figs. 1-10. 9 syntypes 4048 (figs. 4-5, 8-10); 4049 (figs. 1-3, 6-7); 4050-4056 (un- figured); Permian, Putnam Formation; San Angelo junction, 4 mi SE of Coleman, Coleman County, Texas. Family HETERELASMINIDAE Beecheria sp. STEHLI, 1956, pi. 40, fig. A6. Figured specimen 4057; Pennsylvanian; Grapevine Canyon, Otero County, New Mexico. Gilledia cymbaeformis (Morris, 1845) STEHLI, 1961a as pi. 61, figs. 1-2, 5-6. Figured specimen 4058; Permian, Upper Marine Series; Wyro, near Ulladulla, New South Wales, Australia. Family NOTOTHYRIDIDAE Timorina broili STEHLI, 1961b:465, pi. 62, group E, figs. 1-11. Holotype 4060 (fig. 10); 5 paratypes 4061 (figs. 4-5); 4062 (fig. 1); 4063 (fig. 2); 4064 (figs. 3 and 6); 4065 (figs. 7-8); Permian; Basleo, Timor, Dutch East Indies. NOTE: I am unable to match figs. 9 and 11 with the specimens. Phylum MOLLUSCA Class PELECYPODA Order CONOCARDIDOIDA Family CONOCARDIIDAE Conocardium langenheimi WILSON, 1970:1, fig. 8. Figured paratype 2443; Lower Permian, McCloud Limestone; Shasta County, California. Order HIPPURITOIDA Family CAPRINIDAE '*Coralliochama orcutti WHITE, 1885:10. 7 plastosyntypes 4302-4308; Cretaceous, Rosario Formation; north side of Punta Banda, “shore of Todos Santos Bay,” Baja California, Mexico. 1973 ADDITIONAL Type Specimens 5 Order NUCULOIDA Family NUCULIDAE Acila (Truncacila) demessq Finlay, 1927 SCHENCK, 1943 as pi. 9, figs. 1, 3 and 7. 3 figured specimens 4066 (fig. 1); 4067 (fig. 3); 4068 (fig. 7); Cretaceous, Santa Ana Mountains, Orange County, California. Order ARCOIDA Family ARCIDAE Anadara (Anadara) carrizoensis REINHART, 1943:56, pi. 5, figs. 1, 4 and 10. Holotype 4069 (pi. 5, fig. 10); paratype 4070 (pi. 5, figs. 1 and 4); Pliocene, Imperial Formation; south slope of Carrizo Mountain, Imperial County, California. Anadara aff. A. (Anadara) montereyana (Osmont, 1905) STANTON, 1966 as pi. 5, fig. 1. Figured specimen 4071; Miocene, Castaic Formation; bottom of small can- yon tributary to Haskell Canyon, Los Angeles County, California. Area (Area) santamariensis REINHART, 1937:183, pi. 28, figs. 4-5, 7-8 and 11. REINHART, 1943 as pi. 3, figs. 7-11. Holotype 4072; Pliocene; Fugler Point, IV2 mi SE of Santa Maria, Santa Barbara County, California. Area (Area) sisquoeensis REINHART, 1937: 182, pi. 28, figs. 1-3. REINHART, 1943 as pi. 2, fig. 8. Holotype 4073; Pliocene; Fugler Point, 7V^ mi SE of Santa Maria, Santa Barbara County, California. A rca vancouverensis MEEK, 1864:40. Plastoparatype 4074; Cretaceous; Nanaimo, Vancouver Island, British Colum- bia, Canada. Barbatia (Fugleria) pseudoillota REINHART, 1937:184, pi. 28, figs. 6, 9 and 10. REINHART, 1943 as pi. 3, figs. 4-6. Holotype 4075; paratype 4076 (unfigured); Pliocene; Fugler Point, IV2 mi SE of Santa Maria, Santa Barbara County, California. Family GLYCYMERIDIDAE Glycymeris cf. G. gigantea (Reeve, 1843) STANTON, 1966 as pi. 5, figs. 2-3. 2 figured specimens 4077 (fig. 2); 4259 (fig. 3); Miocene, Castaic Formation; tributary to Elderberry Canyon, Los Angeles County, California. Glycymeris meganosensis Clark and Woodford, 1927 SMITH, 1945 as fig. 6. Figured specimen 4078; Eocene, Meganos Formation; Mount Diablo Quad- rangle, Contra Costa County, California. 6 Contributions in Science No. 250 Glycymeris veatchii (Gabb, 1864) SMITH, 1945 as figs. 1 and 9. 2 figured specimens 4079 (fig. 1); Cretaceous; Tuscan Springs, Tehama County, California; 4080 (fig. 9); Cretaceous; Santa Monica Mountains, Los Angeles County, California. Glycymeris veatchii anae SMITH, 1945:42, figs. 3 and 7. 2 syntypes 4081 (fig. 3); 4082 (fig. 7); Cretaceous; Santa Ana Mountains, Orange County, California. Glycymeris veatchii major (Stanton, 1896) SMITH, 1945 as figs. 5 and 8. 2 figured specimens 4083 (fig. 5); Eocene, “Martinez” Formation; Simi Valley, Ventura County, California; 4084 (fig. 8); Cretaceous, Garzas Formation; Garzas Creek, Merced County, California. Glycymeris veatchii reddingensis SMITH, 1945:42, figs. 2 and 4. 2 syntypes 4085 (fig. 2); Cretaceous; near Redding, Shasta County, California. Family LIMOPSIDAE T rinacria cor POPENOE, 1937:380. Plastoholotype 4087; Plastoparatype 4088; Cretaceous, Williams Formation; Santa Ana Mountains, Orange County, California. Order MYTILOIDA Family MYTILIDAE Brachidontes bifurcatus POPENOE, 1937:383. Plastoholotype 4089; Cretaceous, Williams Formation; Santa Ana Mountains, Orange County, California. Inoperna bellarugosa POPENOE, 1937:382. Plastoholotype 4090; Plastoparatype 4091; Cretaceous, Ladd Formation; Santa Ana Mountains, Orange County, California. Order PTERIOIDA Family ANOMIIDAE Pododesmus cf. P. macroschisma (Deshayes, 1839) STANTON, 1966 as pi. 5 fig. 7. Figured specimen 4092; Miocene, Castaic Formation; Small valley E of Dry Canyon Dam, Los Angeles County, California. Family LIMIDAE Lima (Acesta) beta POPENOE, 1937:382. Plastoholotype 4093; Cretaceous, Ladd Formation; Santa Ana Mountains, Orange County, California. 1973 Additional Type Specimens 7 Family PECTINIDAE Chlamys hodgei (Hertlein, 1925) STANTON, 1966 as pi. 5, figs. 4-5. Figured specimen 4094; Miocene, Castaic Formation; Humphreys Quadran- gle, Los Angeles County, California. Hinnites midtirugosiis \2lv. crassiplicatus (Gale, 1928) STANTON, 1966 as pi. 5, fig. 6. Figured specimen 4095; Miocene, Castaic Formation; Humphreys Quadran- gle, Los Angeles County, California. Pecten (Amusium) condoni HERTLEIN, 1925b:41. Paratype 4260 (unfigured); Miocene, Montesano Formation; at dam #35, W Wishkah River, Grays Harbor County, Washington. Pecten (Propeamusium) cowpen WARING, 1917:63. Paratype 4096 (unfigured); Cretaceous, Chico Formation; Bells Canyon, Ventura County, California. Pecten (Plagioctenium) cristobalensis HERTLEIN, 1925a: 19. Paratype 4097 (unfigured); Pliocene, Salada Formation; 3 mi SE of Turtle Bay, Baja California, Mexico. Pecten (Plagioctenium) hakei HERTLEIN, 1925a: 18. Paratype 4098 (unfigured); Pliocene, Salada Formation; 5 mi N of Abreojos Point, Ballenas Bay Quadrangle, Baja California, Mexico. Pecten (Camptonectes) harfordus DAVIS, 1913:456. Paratype 4099 (unfigured); “Jurassic”; 6 mi N of Port Hartford, San Luis Obispo County, California. Pecten (Aeqiiipecten) pe reams HERTLEIN, 1925a: 13. Paratype 4100 (unfigured); Pliocene, Salada Formation; mouth of large arroyo NE of Elephant Mesa, Scammon’s Lagoon Quadrangle, Baja Cali- fornia, Mexico. Pecten stearnsii Dali, 1878 WOODRING, BRAMLETTE, and KEW, 1946 as pi. 30, figs. 9-10. Figured specimen 4101; Pleistocene, Lomita Marl; Hilltop Quarry, San Pedro, Los Angeles County, California. Family SPONDYLIDAE Spondylus sp. STANTON, 1966 as pi. 5, fig. 8. Figured specimen 4102; Miocene, Castaic Formation; E of junction of Castaic and Fish Creeks, Los Angeles County, California. Order VENEROIDA Family ASTARTIDAE Astarte concentrica Conrad, 1834 POPENOE and FINDLAY, 1933 as pi. 19, figs. 5-7. 3 figured specimens 4103 (fig. 5); 4104 (fig. 6); 4105 (fig. 7); Miocene, Yorktown Formation; Grove Wharf, James City County, Virginia. 8 Contributions in Science No. 250 Family CARDITIDAE V enericardia parva Lea, 1833 POPENOE and FINDLAY, 1933 as pi. 19, figs. 1-4 and 8-9. 6 figured specimens 4106 (fig. 1); 4107 (fig. 2); 4108 (fig. 3); 4109 (fig. 4); 4110 (fig. 8); 4111 (fig. 9); Eocene; Clairborne Bluff, Monroe County, Alabama. “V enericardia” ventricosa (Gould, 1850) POPENOE and FINDLAY, 1933 as pi. 19, fig. 17. Figured specimen 4112; Pleistocene; San Pedro, Los Angeles County, Cali- fornia. Family CRASS ATELLIDAE Crassatella gamma POPENOE, 1937:388. Plastoholotype 4113; 2 plastoparatypes 4114 and 4115; Cretaceous, Ladd Formation; Santa Ana Mountains, Orange County, California. Crassatella grand (Wiedey, 1928) WOODRING, 1942 as pi. 13, fig. 1. Figured specimen 4116; Miocene, Vaqueros Formation; Cajon Pass, San Bernardino County, California. Crassinella nuculiformis BERRY, 1940:149. Paratype 4117; Pleistocene; W side of Gaffey Street in cut just below General St., San Pedro, Los Angeles County, California. Eucrassatella subgibbosa (Hanna, 1926) STANTON, 1966 as pi. 5, figs. 9-12. 4 figured specimens 4118 (fig. 9); 4119 (fig. 10); 4120 (fig. 11); 4121 (fig. 12); Miocene, Castaic Formation; Humphreys Quadrangle, Los Angeles County, California. Family ISOCARDIIDAE Clisocolus corrugatus POPENOE, 1937:390: Plastoholotype 4122; Cretaceous, Ladd Formation; Santa Ana Mountains, Orange County, California. Isocardia delta POPENOE, 1937:389. Plastoholotype 4123; 2 Plastoparatypes 4124-4125; Cretaceous, Ladd Forma- tion; Santa Ana Mountains, Orange County, California. Family PSAMMOBIIDAE Sanguinolaria cf. S. nuttalli Conrad, 1837 STANTON, 1966 as pi. 6, figs. 3-4. Figured specimens 4126; Miocene, Castaic Formation; NE side of Castaic Canyon, Los Angeles County, California. Family TELLINIDAE Macoma ? sp. STANTON, 1966 as pi. 6, figs. 6-7. 2 figured specimens 4127 (fig. 6); 4128 (fig. 7); Miocene, Castaic Formation; W bank at junction of Castaic and Elizabeth Lake Canyons, Los Angeles County, California. 'i ] 1973 Additional Type Specimens 9 Psammotreta (Florimetis) biangulata (Carpenter, 1855) STANTON, 1966 as pi. 6, fig. 5. Figured specimen 4129; Miocene, Castaic Formation; NE sid*e of Castaic Canyon, Los Angeles County, California. Family VENERIDAE *Calva regina POPENOE, 1937;395. 2 plastosyntypes 4130-4131; Cretaceous, Ladd Formation; Santa Ana Moun- tains, Orange County, California. Cyprimeria moorei POPENOE, 1937:391. 4 plastosyntypes 4132-4135; Cretaceous, Ladd Formation; Santa Ana Moun- tains, Orange County, California. Dosinia sp. STANTON, 1966 as pi. 5, figs. 13-14. 2 figured specimens 4136 (fig. 13); 4137 (fig. 14); Miocene, Castaic Forma- tion; NE side of Castaic Canyon, Los Angeles County, California. Flaventia zeta POPENOE, 1937:393. Plastoholotype 4138; 2 platsoparatypes 4139-4140; Cretaceous, Ladd Forma- tion; Santa Ana Mountains, Orange County, California. Macrocallista stantoni WARING, 1971:77. Paratype 4141 (unfigured); Eocene, “Martinez” Formation; Simi Hills, Ventura County, California. Mercenaria perlaminosa (Conrad, 1855) WOODRING, BRAMLETTE and KEW, 1946 as pi. 31, figs. 16-17. Figured specimen 4142; Pleistocene, Lomita Marl; San Pedro, Los Angeles County, California. Pitar (LamelHconcha) sp. STANTON, 1966 as pi. 6, figs. 1-2. Figured specimen 4143; Miocene, Castaic Formation; Humphreys Quadran- gle, Los Angeles County, California. Tivela gastonensis CLARK, 1925:93. Paratype 4144 (unfigured); Oligocene; Scroggins Canon, Gaston, Washington County, Oregon. Tivela scarificata BERRY, 1940:5. 2 paratypes 4145-4146 (unfigured); Pleistocene; NW corner of Beacon and 2nd Sts., San Pedro, Los Angeles County, California. Order MYOIDA Family HIATELLIDAE Panomya beringianiis Dali, 1916 WOODRING, BRAMLETTE and KEW, 1946 as pi. 33, fig. 13. Figured specimen 4147; Pleistocene, Timms Point Silt; Timms Point, San Pedro, Los Angeles County, California. 10 Contributions in Science No. 250 Order PHOLADOMYOIDA Family PANDORIDAE Pandora grandis Dali, 1877 WOODRING, BRAMLETTE and KEW, 1946 as pi. 33, fig. 18. Figured specimen 4148; Pleistocene, Timms Point Silt; Timms Point, San Pedro, Los Angeles County, California. Family PERiPLOMATlDAE Periploma cf. P. discus Stearns, 1890 STANTON, 1966 as pi. 6, figs. 8-9. Figured specimen 4149; Miocene, Castaic Formation; Bottom of small can- yon tributary to Haskell Canyon, Los Angeles County, California. Family THRACIIDAE Thracia trapezoides Conrad, 1849 WOODRING, BRAMLETTE and KEW, 1946 as pi. 33, fig. 15. Figured specimen 4150; Pleistocene, Timms Point Silt; Timms Point, San Pedro, Los Angeles County, California. Class GASTROPODA Order ARCHAEOGASTROPODA Family ACMAEIDAE Acmaea lepisma BERRY, 1940:155. 4 paratypes 4151-4154 (all unfigured); Pleistocene; Hilltop Quarry, San Pedro, Los Angeles County, California. Family HALIOTIDAE Haliotis lomaensis ANDERSON, 1902:75. Plastoholotype 4155; Cretaceous; Point Loma, San Diego, San Diego County, California. Haliotis palaea WOODRING, 1931:38, pi. 6, figs. 1-3. Holotype 4156 (fig. 1); paratype 4157 (fig. 2); Miocene; Brown Canyon, Santa Monica Mountains, Los Angeles County, California. Family LIOTIIDAE Liotia cnrmn/fl Carpenter, 1857 STANTON, 1966 as pi. 6, figs. 15-16. Figured specimen 4158; Miocene, Castaic Formation; Humphreys Quadran- gle, Los Angeles County, California. Family NERITIDAE Nerita sp. STANTON, 1966 as pi. 6, figs. 13-14. Figured specimen 4159; Miocene, Castaic Formation; NE bank of Castaic Canyon, Los Angeles County, California. 1973 Additional Type Specimens 11 Family TROCHIDAE Lirularia magna T. S. OLDROYD, 1925:36. 9 paratypes 2460-2468 (unfigured); Pleistocene; San Pedro, Los Angeles County, California. Turcica imperialis brevis Stewart in Woodring, Stewart and Richards, 1941. WOODRING and BRAMLETTE, 1950 as pi. 12, figs. 12 and 14. Figured specimen 4160; Pliocene, Careaga sandstone; Fugler Point, Santa Barbara County, California. Tiircicula santacruzana ARNOLD, 1908:373. Paratype 4161 (unfigured); Oligocene, San Lorenzo Formation; San Lorenzo River, 3 mi above Boulder Creek, Santa Cruz County, California. Family TURBINIDAE Astraea (Pomaulax) cf. A. (Pomaulax) gradata Grant and Gale, 1931 STANTON, 1966 as pi. 6, figs. 10-12. 2 figured specimens; 4162 (figs. 10-11) Miocene, Castaic Formation; Crest of ridge SE of Elderberry Canyon, Los Angeles County, California; 4163 (fig. 12) Miocene, Castaic Formation; Humphreys Quadrangle, Los Angeles County, California. Astraea (Pomaulax) petrothanma BERRY, 1940:156. 2 paratypes 4164-4165 (unfigured); Pleistocene; Hilltop Quarry, San Pedro, Los Angeles County, California. Homalopoma cf. H. paucicostata (Dali, 1871) WOODRING and BRAMLETTE, 1950 as pi. 12, fig. 4. Figured specimen 4166; Pliocene, Careaga sandstone; Fugler Point, Santa Barbara County, California. Leptothyra siibobsoleta WILLETT, 1937:63, pi. 25. 9 paratypes 2471-2479; Pleistocene, Timms Point Silt; Timms Point, San Pedro, Los Angeles County, California. NOTE: The holotype, but no paratypes, was reported in Wilson and Bing (1970:7). Order MESOGASTROPODA Family CALYPTRAEIDAE Crepidula princeps Comzid., 1856 WOODRING, BRAMLETTE and KEW, 1946 as pi. 32, figs. 5-6. Figured specimen 4167; Pleistocene, Timms Point Silt; Timms Point, San Pedro, Los Angeles County, California. Trochita cf. T. trochiformis (Born, 1778) STANTON, 1966 as pi. 7, fig. 8. Figured specimen 4168; Miocene, Castaic Formation; Humphreys Quadrangle, Los Angeles County, California. Family CERITHIIDAE IBittium arnoldi Bartsch, 1911 STANTON, 1966 as pi. 6, fig. 17. Figured specimen 4169; Miocene, Castaic Formation; W side of northerly trending tributary to Elizabeth Lake Canyon, Los Angeles County, California. 12 Contributions in Science No. 250 Family FICIDAE Ficus (Trophosycon) ocoyana (Conrad, 1855) STANTON, 1966 as pi. 7, figs. 9-10. 2 figured specimens; 4170 (fig. 9); Miocene, Castaic Formation; W side of northerly trending tributary to Elizabeth Lake Canyon, Los Angeles County, California; 4171 (fig. 10); Miocene, Castaic Formation, Humphreys Quadran- gle, Los Angeles County, California. Family NATICIDAE Ampullina packardi POPENOE, 1937:399. Plastoholotype 4172; Cretaceous, Ladd Formation; Santa Ana Mountains, Orange County, California. Family TURRITELLIDAE Tunitella andersoni mulleri MERRIAM, 1941:80. 3 paratypes 4173-4175 (all unfigured); Eocene; Mount Pinos Quadrangle, Ventura County, California. Turritella cooperi Carpenter, 1864 STANTON, 1966 as pi. 7, figs. 1-5. 5 figured specimens 4176 (fig. 1); 4177 (fig. 2); 4178 (fig. 3); 4179 (fig. 4); 4180 (fig. 5); Miocene, Castaic Formation; On' ridge W of junction of Reynier and Sand Canyons, Los Angeles County, California. Turritella aff. T. freya Nomland, 1917 STANTON, 1966 as pi. 7, figs. 6-7. 2 figured specimens 4181 (fig. 6); 4182 (fig. 7); Miocene, Castaic Formation; on ridge W of junction of Reynier and Sand Canyon, Los Angeles County, California. Turritella inezana Conrad, 1857 WOODRING, 1942 as pi. 13, figs. 3-4 and pi. 14, fig. 3. 3 figured specimens 4183 (pL 13, fig. 3); 4184 (pi. 13, fig. 4); 4185 (pi. 14, fig. 3); Miocene; Cajon Pass, San Bernardino County, California. Turritella iota POPENOE, 1937:401. Plastoholotype 4186; Cretaceous, Ladd Formation; Santa Ana Mountains, Orange County, California. Turritella ocoyana var. topangaensis MERRIAM, 1941:115. Plastoparatype 4187; Miocene, Topanga Formation; La Panza Mountains, San Luis Obispo County, California. Turritella ossa POPENOE, 1937:401. Plastoholotype 4188; 2 plastoparatypes 4189-4190; Cretaceous, Ladd Forma- tion; Santa Ana Mountains, Orange County, California. Turritella uvasana subsp. inf era MERRIAM, 1941:90. Plastoparatype 4191; Eocene, Santa Susana Formation; Simi Valley, Ventura County, California. 1973 Additional Type Specimens 13 Turritella uvasana subsp. stewarti MERRIAM, 1941:95. Plastoparatype 4192; Eocene, Cowlitz Formation; Coal Ct^eek, Cowlitz County, Washington. Family VIVIPARIDAE Viviparus washingtoniamis ARNOLD and HANNIBAL in HANNIBAL, 1912:194. Paratype 4193 (unfigured); Eocene; Olequa Creek, 2 mi N of Little Falls, Washington. Order NEOGASTROPODA Family BUCCINIDAE Calicantharus fortis var. cf. angulata (Arnold, 1907) WOODRING and BRAMLETTE, 1950 as pi. 14, fig. 10. Figured specimen 4194; Pliocene, Careaga sandstone; Fugler Point, Santa Barbara County, California. Gyrineiim lewisii CARSON, 1926:54. Paratype 4195 (unfigured); Lower Pliocene; Fugler’s Point, Santa Maria District, Santa Barbara County, California. Family FASCIOLARIDAE Lirofusus ash ley i ARNOLD, 1908:372. Paratype 4196 (unfigured); Oligocene, San Lorenzo Formation; 3 mi above Boulder Creek, Santa Cruz County, California. Family MARGINELLIDAE Marginella cf. M. albuminosa Dali, 1919 STANTON, 1966 as pi. 7, figs. 14-15. Figured specimen 4197; Miocene, Castaic Formation; Humphreys Quadrangle, Los Angeles County, California. Family MURICIDAE Jaton cf. i. carpenteri (Dali, 1899) WOODRING and BRAMLETTE, 1950 as pi. 15, figs. 9 and 11. Figured specimen 4198; Pliocene, Careaga sandstone; Fugler Point, Santa Barbara County, California. Family NEPTUNEIDAE Kelletia kanakoffi HERTLEIN, 1970:3, figs. 1-3. Holotype 2456 (figs. 1-2); paratype 2457 (fig. 3); Pleistocene, Lomita Marl; San Pedro, Los Angeles County, California. Family OLIVIDAE Oliva spicata (Roding, 1798) STANTON, 1966 as pi. 7, fig. 13. Figured specimen 4199; Miocene, Castaic Formation; W of Haskell Canyon, Los Angeles County, California. 14 Contributions in Science No. 250 Family THAIDIDAE Thais (Nucella) shumanensis CARSON, 1926:56. Paratype 4200 (unfigured); Pliocene, Fernando Formation; 1/2 mi N of Schuman, Santa Barbara County, California. Family TURRIDAE Anomalosipho sp. STANTON, 1966 as pi. 7, figs. 11-12. 2 figured specimens 4201 (fig. 11); 4202 (fig. 12); Miocene, Castaic Forma- tion; Bottom of small canyon tributary to Haskell Canyon, Los Angeles County, California. Antiplanes sp. STANTON, 1966 as pi. 7, fig. 16. Figured specimen 4203; Miocene, Castaic Formation; On ridge W of junction of Reynier and Sand Canyons, Los Angeles County, California. Clatlwrella (Glyphostoma) tridesmia BERRY, 1941:8. 3 paratypes 4204-4206 (all unfigured); Pleistocene; Hilltop Quarry, San Pedro, Los Angeles County, California. Mitromorpha barbarensis woodfordi BERRY, 1941:10. 2 paratypes 4207-4208 (both iinfigured); Pleistocene; Hilltop Quarry, San Pedro, Los Angeles County, California. Mitromorpha galeana BERRY, 1941:12. 2 paratypes 4209-4210 (both unfigured); Pleistocene; Hilltop Quarry, San Pedro, Los Angeles County, California. Moniliopsis chacei BERRY, 1941:6. Paratype 4211 (unfigured); Pleistocene; Hilltop Quarry, San Pedro, Los Angeles County, California. Family VOLUTIDAE Lyria andersoni WARING, 1917:97. Paratype 4212 (unfigured); Eocene, Tejon Formation; McCray Wells, Ventura County, California. Psephaea oregonensis (Dali, 1907) WOODRING and BRAMLETTE, 1950 as pi. 14, fig. 4-5. Figured specimen 4213; Pliocene, Careaga Sandstone; Fugler Point, Santa Maria, Santa Barbara County, California. Order ENTOMOTAENIATA Family PYRAMIDELLIDAE Iselica fenestrata (Carpenter, 1864) WOODRING and BRAMLETTE, 1950 as pi. 13, figs. 17-18. Figured specimen 4214; Pliocene, Careaga sandstone; Fugler Point, S side of Santa Maria River, Santa Barbara County, California. 1973 Additional Type Specimens 15 Order STYLOMMATOPHORA Family FRUTICICOLIDAE , Helminthoglypta ? stocki HANNA, 1934:539, figs. l-3a. Holotype 4261 (fig. 1); 2 paratypes 4262 (figs. 2, 3, 3a); 4263 (unfigured); Eocene; N of Simi Valley, Ventura County, California. Order CEPHALASPIDIA Family ACTEONIDAE Acteon cf. A. boulderdna Etherington, 1931 STANTON, 1966 as pi. 7, fig. 17. Figured specimen 4215; Miocene, Castaic Formation; NE side of Castaic Canyon, Los Angeles County, California. Class SCAPHOPODA Order SOLENOCONCHA Family DENTALIIDAE Dentalium sp. STANTON, 1966 as pi. 7, fig. 21. Figured specimen 4216; Miocene, Castaic Formation; N side of small W trending valley E of Canyon Dam, Los Angeles County, California. Class CEPHALOPODA Order AMMONOIDEA Family NOSTCERATIDAE '^'Ainoceras kamuy MATSUMOTOand KANIE, 1967:351. Plastoparatype 4309; Cretaceous; Hattori Ranch, Urakawa, Hidaka Province, Hokkaido, Japan. Family PACHYDISCIDAE Pachydiscus seppenradensis LANDOIS, 1895:99. Plastosyntype 4310; Cretaceous; Seppenrade, Westphalia, Germany. Phylum ECHINODERMATA Class STELLEROIDEA Order PAXILLOSIDA Family ASTROPECTINIDAE Astropecten matilijaensis DURHAM and ROBERTS, 1948:435, pi. 65, figs. 1-5; pi. 66, figs. 2, 4-5. DURHAM, 1948: 28. Holotype 4217 (Durham and Roberts 1948, pi. 65, figs. 3-4); 29 paratypes 4218 (Durham and Roberts 1948, pi. 65, fig. 1; pi. 66, fig. 4); 4219 (Durham and Roberts 1948, pi. 65, fig. 2; Durham 1948:28); 4220 (Durham and Roberts 1948, pi. 65, fig. 5); 4221 (Durham and Roberts 1948, pi. 66, fig. 2); 4222 (Durham and Roberts 1948, pi. 66, fig. 5); 4223 (Durham and Roberts 1948, pi. 65, fig. 3, upper); 4224 (Durham and Roberts 1948, pi. 65, fig. 3, lower); 4225-4247 (unfigured); Cretaceous, Chico Formation [Eocene, Cozy Dell Formation}; North fork of Matilija Creek, Ventura County, California. 16 Contributions in Science No. 250 Order FORCIPULATIDA Family ECHINASTERIDAE Henricia (?) venturana DURHAM and ROBERTS, 1948:437, pi. 66, figs. 1 and 3. Holotype 4248; Cretaceous, Chico Formation {Eocene, Cozy Dell Formation}; North fork of Matilija Creek, Ventura County, California. Class ECHINOIDEA Order CIDAROIDA Family CIDARIDAE Cidarid spines STANTON, 1966 as pi. 7, figs. 18-20. 2 figured specimens 4249 (pi. 7, fig. 18; Miocene, Castaic Formation; Hum- phreys Quadrangle, Los Angeles County, California) 4250 (pi. 7, figs. 19-20); Miocene, Castaic Formation; Castaic Creek, N of Cordova Ranch, Los Angeles County, California, Order CLYPEASTEROIDA Family ECHINARACHNIIDAE Astrodapsis fernandoensis Pack, 1909 STANTON, 1966 as pi. 7, fig. 22. Figured specimen 4251; Miocene, Castaic Formation; Bottom of small canyon tributary to Haskell Canyon, Los Angeles County, California. Phylum ARTHROPODA Class ARACHNIDA Order PEDIPALPIDA Family SCHIZOMIDAE *Caloschizonius latisternum PIERCE, 1951a:41, pi. 14, fig. 4; pi. 15, fig. 7. PETRUNKEVITCH, 1955, fig. 90, la. Holotype 2505; Tertiary; Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. '•‘Onychothelyphonus bonneri PIERCE, 1950d:3, pi. 34, fig. 1. PETRUNKEVITCH, 1955, fig. 90, lb. Holotype 2509; Tertiary; Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. Thelyphonus hadleyi PIERCE, 1945c:7, pi. 5. Holotype 2504; Middle Miocene; Cabrillo Beach, San Pedro, Los Angeles County, California. Order ARANEIDA Family ARGYRONETIDAE Argyroneta antiqua Heyden, 1859 FURST, 1959:12, p. 12, pi. and fig. unnumbered. FURST, 1970:23, p. 23, bottom fig., unnumbered. Figured specimen 3086; Oligocene; Rott, Rheinland, Germany. Family SALIICIDAE Salticus seymoiiri PIERCE, 1950a: 3 (unfigured). 5 syntypes 2499-2503; Pleistocene; Seymour River, 8 mi above Vancouver Channel, British Columbia, Canada. 1973 Additional Type Specimens 17 Class CRUSTACEA Order CLADOCERA Family DAPHNIDAE Daphinia {sic] fossilis Heyden STATZ, 1938b: 56, fig. 13. Figured specimen 3087; Oligocene; Rott, Rheinland, Germany. Order DECAPODA Family CANCRIDAE Cancer branneri Rathbun, 1926 MENZIES, 1951:165, figs. 2a-2b. 2 figured specimens 2448 (fig. 2a); 2449 (fig. 2b); Pleistocene, Timm’s Pojnt Formation; San Pedro, Los Angeles County, California. Cancer jordani Rathbun, 1900 MENZIES, 1951:165, figs. 5a-5d. 2 figured specimens 2446 (figs. 5a-5b); 2447 (figs. 5c-5d); Pleistocene; San Pedro, Los Angeles County. California. Cancer productus Randall, 1839 MENZIES, 1951:165, figs. 4a-4d. 2 figured specimens 2444 (figs. 4a-4b) (MISSING); 2445 (fip. 4c-4d) (MIS- SING); Pleistocene; San Pedro, Los Angeles County, California. Family PALAEMONIDAE Micropsalis papyracea Meyer, 1859 STATZ, 1938a:29, fig. 12. Figured specimen 3088; Oligocene; Rott, Rheinland, Germany. Class DIPLOPODA Order JULIDA Family PARAJULIDAE Parajulus lasti PIERCE, 1945a: 1, pi. 1. Holotype 2497; Upper Miocene; Oxnard, Ventura County, California. Parajulus onychis PIERCE, 1951b:41, pi. 15, figs. 8-9. Holotype 2498; Tertiary; Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. Class INSECTA Order COLLEMBOLA Family PALAEOSMINTHURIDAE ^Palaeosminthurus juliae PIERCE and GIBRON, 1962:147, fig. 4. Holotype 2512; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. Order DIPLURA Family CAMPODEIDAE * Onychocampodea onychis PIERCE, 1951c:48, pi. 17, fig. 11. Holotype 2506; Tertiary; Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. 18 Contributions in Science No. 250 Family JAPYGIDAE ^Onychojapyx schmidti PIERCE, 1950e: 104, pi. 34, fig. 2. Holotype 2508; Tertiary; Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. Family PROJAPYGIDAE *Plioprojapyx primitivus PIERCE, 1951c:48, pi. 18, fig. 16. Holotype 2511; Tertiary; Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. Order THYSANURA Family LEPISMATIDAE ^Onycholepisma arizonae PIERCE, 1951c;45, pi. 17, fig. 14. Holotype 2510; Tertiary; Bonner Quarry, Kaibab National Forest, Yavpai County, Arizona. Family MACHILIDAE *Onychomachilis fisheri PIERCE, 1951c:45, pi. 17, figs. 12 and 15. Holotype 2507 (MISSING); Tertiary, Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. Order EPHEMEROPTERA Family APHELOPHLEBODIDAE *A phelophlebodes stocki PIERCE, 1945b: 4, pi. 2. Holotype 2513; Upper Miocene; Puente Hills, SE of Puente, Los Angeles County, California. Order ODONATA Family AGRIONIDAE Lestes sp. STATZ, 1938b:58, fig. 16. Figured specimen 3089; Oligocene; Rott, Rheinland, Germany. Lestes grand is STATZ, 1930:11, fig. 1. Holotype 3090; Oligocene; Rott, Rheinland, Germany. NOTE: The figured plate is in the Kastenholz collection; the counterplate is in the Natural History Museum of Los Angeles County. Family COENAGRIONIDAE *Sobobapteron kirkbyae PIERCE, 1965:160, fig. 2. Holotype 2514 (MISSING); Upper Miocene; Woodcrest Drive, Sherman Oaks, Los Angeles County, California. NOTE: Carpenter (1968) moved this species from the order Protorthoptera where it was originally placed to the present position, order Odonata, family Coenagrionidae. 1973 Additional Type Specimens 19 Family LIBELLULIDAE ?Libellulidae (nymph) MOUNT, 1971:3, fig. 1. Figured specimen 3085; Miocene, Altamira shale; Float between Cabrillo Beach and Point Fermin, San Pedro, Los Angeles County, California. Celithemis ceUulosa (Hagen, 1863) STATZ, 1938b:57, fig. 15. Figured specimen 3091; Oligocene; Rott, Rheinland, Germany. Libellula ceres Hagen, 1863 STATZ, 1938b:56, fig. 14. Figured specimen 3092 (MISSING); Oligocene; Rott, Rheinland, Germany. Order PLECOPTERA Family LEUCTRIDAE Leuctra antiqua STATZ, 1938b:58, fig. 17. Figured specimen 3093; Oligocene; Rott, Rheinland, Germany. Family LIPOSCELIDAE ^Miotroctes rousei PIERCE, 1960:46, pi. 16, fig. 7. Holotype 2526 (MISSING); Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. Order ORTHOPTERA Family BLATTIDAE Blaberites rhenana STATZ, 1939: 10, pi. 4, figs. 14-15. 2 syntypes 3094 (fig. 14) and 3095 (fig. 15); Oligocene; Rott, Rheinland, Germany. Ectobius glabellas STATZ, 1939:12, pi. 5, figs. 18-22. 3 syntypes 3096 (figs. 18 and 19), 3097 (figs. 20-21), 3098 (fig. 22); Oligocene; Rott, Rheinland, Germany. Nyctibora ? elongata STATZ, 1939:11, pi. 4, figs. 16-17. Holotype 3099; Oligocene: Rott, Rheinland, Germany. Family EXAERETOPTERIDAE *Exaeretoptera fosteri pierce: 1944c: 15, pis. 5-6. Holotype 2516; Upper Miocene; Fort MacArthur, San Pedro, Los Angeles County, California. Family GRYLLIDAE Liogryllus rottensis STATZ, 1939:8, pi. 2, figs. 8-9; pi. 3, figs. 12-13. 3 syntypes 3100 (pi. 2, figs. 8-9); 3102 (pi. 3, figs. 12-13); 3103 (unfigured); Oligocene; Rott, Rheinland, Germany. 20 Contributions in Science No. 250 Famliy PHASMIDAE ^Calcibacunculus tenuis PIERCE, 1951c:48, pi. 17, fig. 13; pi. 18, fig. 17. Holotype 2526; Tertiary; Bonner Quarry, Kaibab National Forest, Yavapai County, Arizona. Family TETTIGONIIDAE ^'Protosegestes Iloydi PIERCE, 1944c:13, pl.4. Holotype 2517; Upper Miocene; Fort MacArthur, San Pedro, Los Angeles County, California. Tettigoniidarum spec, indet. Nr. 1 STATZ, 1939, pi. 1, fig. 1. Figured specimen 3103; Oligocene: Rott, Rheinland, Germany. Tettigoniidarum spec, indet. Nr. 2 STATZ, 1939, pi. l,fig. 2. Figured specimen 3104; Oligocene; Rott, Rheinland, Germany. Order ISOPTERA Family HODOTERMITIDAE Calotermes rottensis STATZ, 1930:12. STATZ, 1939 as pi. 7, figs. 32-33. STATZ, 1940a as figs. 3-4. EMERSON, 1968 as figs. 2a-2c and 2f. Lectotype 3l'05 (unfigured; Emerson, 1968: 16); 27 paralectotypes, 3106 (Statz, 1939, pi. 7, fig 32-1); 3107 (Statz, 1939., pi. 7, fig. 32-2); 3108 (Statz, 1939, pi. 7, fig. 32-3); 3109 (Statz, 1939, pi. 8, fig. 33-1); 3110 (Statz, 1939, pi. 8, fig. 33-2); 3111 (Statz, 1939, pi. 8, fig. 33-3); 3112 (Statz, 1940a, fig. 4); 3113 (Emerson, 1968, fig. 2C); 3114 (Emerson, 1968, fig. 2F); 3115-3130 (unfigured); 4021-4022 (unfigured); Oligocene; Rott, Rheinland, Gerrnany. NOTE 1: In 1939 Statz changed the generic name from Calotermes to Ulmeriella and credited the species name to Meunier. Emerson (1968:16) when selecting the lectotype could find no reason for attributing the species to Meunier. NOTE 2: Statz stated that the original figured specimen for this species was in the Rastenholz collection. Emerson (1968) has selected the counterplate of this specimen as the lectotype. Ulmeriella bauckhorni Meunier, 1920 STATZ, 1939, as pi. 6, figs. 28-30 and fig. 35. STATZ, 1940a as fig. 1-2. EMERSON, 1968 as figs. 2A-2B. 5 figured specimens 3131 (Statz, 1939, pi. 6, fig. 28); 3132 (Statz, 1939, pi. 6, fig. 29); 3133 (Statz, 1939, pi. 6, fig. 26; Statz, 1940a, fig. 1); 3134 (Statz, 1940a, fig. 2); 3135 (Emerson, 1968, figs. 2A-2B); Oligocene; Rott, Rheinland, Germany. Ulmeriella rottensis (See Calotermes rottensis) 1973 Additional Type Specimens 21 Family KALOTERMITIDAE Calotermes rhenanus Hagen, 1863 STATZ, 1939 as pi. 8, figs. 34-35. STATZ, 1940a as fig. 6. STATZ, 1941c as fig. 5. 3 figured specimens 3136 (Statz, 1939, pi. 8, fig. 34); 3137 (Statz, 1939, pi. 8, fig. 35); 3138 (Statz, 1940a, fig. 6; 1941c, fig. 5); Oligocene; Rott, Rheinland, Germany. Cryptotermes ryshkoffi PIERCE, 1958:19, pi. 5, fig. 1; pi. 6, fig. 1. Holotype 2418; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. ^'Eotermes grandaeva STATZ, 1939: 13, pi. 6, figs. 23-25. EMERSON, 1969 as figs. 1-2. Lectotype 3139 (Emerson, 1969, fig. 2); 4 paralectotypes 3140 (Statz, 1939, fig. 23); 3141 (Statz, 1939, fig. 24); 3142 (Emerson, 1969, fig. la); 3143 (Emerson, 1969, figs. Ib-lc); Oligocene; Rott, Rheinland, Germany. Family RHINOTERMITIDAE Parastylotermes calico PIERCE, 1958:19, pi. 5, fig. 2; pi. 6, fig. 7. EMERSON, 1971 as fig. 15. Holotype 2520; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. Recticulitermes laurae PIERCE, 1958:20, pi. 5, fig. 3; pi. 6, fig. 8; pi. 10, fig. 10. EMERSON, 1971 as figs. 8-9. Holotype 2521 (Pierce, 1958, pi. 5, fig. 3; pi. 7, fig. 10; Emerson, 1971, fig. 8); 4 paratypes 2522 (Pierce, 1958, pi. 6, fig. 8; Emerson, 1971, fig. 9); 2523-2525 (unfigured; 2524 MISSING); Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. “Termes” adustus STATZ, 1939:20, pi. 10, figs. 47-48. Holotype 3144; Oligocene; Rott, Rheinland, Germany. “Termes" aethiops STATZ, 1939:19, pi. 9, fig. 39. STATZ, 1940a as fig. 10. Holotype 3145; Oligocene; Rott, Rheinland, Germany. “Termes” atomus STATZ, 1939:21, pi. 10, figs. 49-50. Holotype 3146; Oligocene; Rott, Rheinland, Germany. “Termes” blandus STATZ, 1939:19, pi. 9, figs. 41-42. Holotype 3147; paratype 3148 (unfigured); Oligocene; Rott, Rheinland, Germany. “Termes” concinnus STATZ, 1939:20, pi. 10, fig. 43. STATZ, 1940a as fig. 11. Holotype 3149; Oligocene; Rott, Rheinland, Germany. No. 250 22 Contributions in Science “Termes” contractulus STATZ, 1939:20, pi. 10, figs. 45-46. Holotype 3150; Oligocene; Rott, Rheinland, Germany. “Termes” holmgreni STATZ, 1939:18, pi. 8, fig. 36. STATZ, 1940a as fig. 9. EMERSON, 1971 as fig. 7b. Holotype 3151; Oligocene; Rott, Rheinland, Germany. Family TERMITIDAE Gnathamitermes magnoculus rousei PIERCE, 1958:21, pi. 5, fig. 9; pi. 6, fig. 9. Holotype 2519; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. Order HETEROPTERA Family ANTHOCORIDAE Temnostethus blandus STATZ in STATZ and WAGNER, 1950: 103, pi. 24, fig. 17. Holotype 3152; Oligocene; Rott, Rheinland, Germany. Family COREIDAE Coreide sp. A. [j/c] Coreidae In STATZ and WAGNER, 1950 as pi. 27, fig. 47. Figured specimen 3153; Oligocene; Rott, Rheinland, Germany. Coreide sp. B. [5ic] Coreidae In STATZ and WAGNER, 1950 as pi. 27, fig. 48. Figured specimen 3154; Oligocene; Rott, Rheinland, Germany. Syromastes punctiventris STATZ in STATZ and WAGNER, 1950:1 19, pi. 27, fig. 46. Holotype 3155; Oligocene; Rott, Rheinland, Germany. Family CORIXIDAE Corixa cf Nr. 1 STATZ, 1950b as pi. 20, figs. 75-76. 2 figured specimens 3156 (pi. 20, fig. 75); 3157 (pi. 20, fig. 76) (MISSING); Oligocene; Rott, Rheinland, Germany. Corixa d Nr. 2 STATZ, 1950b as pi. 20, fig. 78. Figured specimen 3158; Oligocene; Rott, Rheinland, Germany. Corixa cf Nr. 3 STATZ, 1950b as pi. 20, fig. 80. Figured specimen 3159; Oligocene; Rott, Rheinland, Germany. Corixa cf Nr. 4 STATZ, 1950b as pi. 20, fig. 81. 3 figured specimens 3160 (fig. 81; MISSING); 3161 (fig. 82); 3162 (fig. 83); Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 23 Corixa rhenana STATZ, 1950b:71, pi. 11, fig. 16; pi. 20, figs. 77 and 79. 2 syntypes 3163 (pi. 11, fig. 16; pi. 20, fig. 79); 3164 (pi. 30, fig. 77); Oligocene; Rott, Rheinland, Germany. NOTE: Syntype 3163 was figured by Statz (1938b, fig. 27) as Corixa elegans von Schlechtendal, 1898. Corixa vidua STATZ, 1950b:72 (no figs.) 7 syntypes 3165-3171 (MISSING); Oligocene; Rott, Rheinland, Germany. Family CYDNIDAE Crocistethus ? madens STATZ in STATZ and WAGNER, 1950:126, pi. 22, fig. 9; pi. 28, fig. 56. Holotype 3172; Oligocene; Rott, Rheinland, Germany. Cydnus ? picatus STATZ in STATZ and WAGNER, 1950:124, no figs. Holotype 3173 (MISSING); Oligocene; Rott, Rheinland, Germany. Sehirus lygaeus STATZ in STATZ and WAGNER, 1950:125, pi. 28, figs. 58-59. Holotype 3174; Oligocene; Rott, Rheinland, Germany. Sehirus paludosus STATZ in STATZ and WAGNER, 1950: 125, pi. 28, fig. 57. Holotype 3175; Oligocene; Rott, Rheinland, Gerrpany. Sehirus spinitibialis STATZ in STATZ and WAGNER, 1950:126, pi. 22, fig. 11; pi. 28, fig. 55. Holotype 3176; Oligocene; Rott, Rheinland, Germany. Family VELIDAE Velia bouatia MEUNIER, 1914:191, fig. 3; pi. 4, fig. 6. 10 syntypes 4314 (fig. 3; pi. 4, fig. 6); 4315 (fig. 3); 4316 (fig. 3); 4317 (fig. 3); 4318 (fig. 3); 4319 (fig. 3); 4320-4323 (unfigured); Oligocene; Aix, Provence, France. Family GERRIDAE Gerris sp. (Nymph) STATZ, 1950b as pi. 20, fig. 84. Figured specimen 3177 (MISSING); Oligocene; Rott, Rheinland, Germany. Family LYGAEIDAE Lygaeide sp. A In STATZ and WAGNER, 1950 as pi. 26, fig. 36. Figured specimen 3178; Oligocene; Rott, Rheinland, Germany. Lygaeide sp. B In STATZ and WAGNER, 1950 as pi. 36, fig. 37. Figured specimen 3179; Oligocene; Rott, Rheinland, Germany. Lygaeide sp. C In STATZ and WAGNER, 1950 as pi. 26, fig. 38. Figured specimen 3180; Oligocene; Rott, Rheinland, Germany. Lygaeide sp. D In STATZ and WAGNER, 1950 as pi. 26, fig. 39. Figured specimen 3181; Oligocene; Rott, Rheinland, Germany. 24 Contributions in Science No. 250 JLygaeide sp. E In STATZ and WAGNER, 1950 as pi. 26, fig. 40. Figured specimen 3182; Oligocene; Rott, Rheinland, Germany. Lygaeide sp. I Lygaeide sp. H In STATZ and WAGNER, 1950 as pi. 26, fig. 42. Figured specimen 3183; Oligocene; Rott, Rheinland, Germany. In STATZ and WAGNER, 1950 as pi. 26, fig. 43. Figured specimen 3184 (MISSING); Oligocene; Rott, Rheinland, Germany. Heterogasterine sp. In STATZ and WAGNER, 1950 as pi. 25, figs. 27-28. Figured specimen 3185; Oligocene; Rott, Rheinland, Germany. Chi lads ? univestris STATZ in STATZ and WAGNER, 1950: 108, pi. 24, fig. 22. Holotype 3186; Oligocene; Rott, Rheinland, Germany. Drymus punctatus STATZ in STATZ and WAGNER, 1950: 113, pi. 26, fig. 35. Holotype 3187; Oligocene; Rott, Rheinland, Germany. Ischnodemus rottensis STATZ in STATZ and WAGNER, 1950: 108, pi. 24, figs. 24-26. 4 syntypes; 3188 (fig. 24); 3189-3191 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: Statz refers to an unspecified number of specimens (syntypes), four in his collection and the remainder in the Kastenholz collection. Only one of those figured is in the LACM collection. The other two figured specimens are presumed to be in the Kastenholz collection. Lygaeosoma wagneri STATZ in STATZ and WAGNER, 1950:107, pi. 21, fig. 2; pi. 24, fig. 23. Holotype 3192; Oligocene; Rott, Rheinland, Germany. Pionosomus ? pusiUus STATZ in STATZ and WAGNER, 1950:110, pi. 25, fig. 31. Holotype 3193; Oligocene; Rott, Rheinland, Germany. Raglius austerus STATZ in STATZ and WAGNER, 1950: 1 1 1, pi. 25, fig. 32. Holotype 3194; Oligocene; Rott, Rheinland, Germany. Raglius ? decoratus STATZ in STATZ and WAGNER, 1950:1 12, pi. 25, figs. 33-34. Holotype 3195; Oligocene; Rott, Rheinland, Germany. Raglius pulchellus STATZ in STATZ and WAGNER, 1950:1 10, pi. 25, fig. 30. Holotype 3196; Oligocene; Rott, Rheinland, Germany. Scolopostethus statzi WAGNER in STATZ and WAGNER, 1950:116, pi. 26, fig. 41. Holotype 3197; Oligocene; Rott, Rheinland, Germany. Family MIRIDAE Calocoris antennatus STATZ and WAGNER, 1950 as pi. 24, fig. 21. 24, fig. 18. Holotype 3198; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 25 Heterocordylus ? sp. STATZ and WAGNER, 1950 as pL 24, fig. 21. Figured specimen 3199; Oiigocene; Rott, Rheinland, Germany. Lygus ? oligocaenica STATZ in STATZ and WAGNER, 1950: 105, pi. 24, fig. 19. Holotype 3200 (MISSING); Oiigocene; Rott, Rheinland, Germany. Trapezonotus ? riguus STATZ in STATZ and WAGNER, 1950: 112, pi. 25, fig. 29. Holotype 3201; Oiigocene; Rott, Rheinland, Germany. Trapezonotus ? striatus WAGNER in STATZ and WAGNER, 1950:105, pi. 24, fig. 20. Holotype 3202; Oiigocene; Rott, Rheinland, Germany. Family NAUCORIDAE Naucoris rottensis Schlechtendal, 1899 STATZ, 1938b, fig. 28. STATZ, 1950b, pi. 12, figs. 24-31; pi. 13, figs. 32-34; pi. 14, figs. 37-40; pi. 15, figs. 41-42, 44-47. FURST, 1959:17, p. 17, pi. and fig. unnumbered. 19 figured specimens 3203 (Statz, 1938b, fig. 28; Statz, 1950b, pi. 13, fig. 34; Furst, 1959:17); 3204 (Statz, 1950b, pi. 12, fig. 24); 3205 (Statz, 1950b, pi. 12, fig. 25); 3206 (Statz, 1950b, pi. 12, fig. 26); 3207 (Statz, 1950b, pi. 12, fig. 27); 3208 (Statz, 1950b,‘ pi. 12, fig. 28); 3209 (Statz, 1950b, pi. 12, fig. 29); 3210 (Statz, 1950b, pi. 12, fig. 30); 3211 (Statz, 1950b, pi. 12, fig. 31); 3212 (Statz, 1950b, pi. 13, figs. 32 and 33); 3213 (Statz, 1950b, pi. 14, fig. 37); 3214 (Statz, 1950b, pi. 14, fig. 38); 3215 (Statz, 1950b, pi. 14, fig. 39); 3216 (Statz, 1950b, pi. 14, fig. 40); 3217 (Statz, 1950b, pi. 15, fig. 41); 3218 (Statz, 1950b, pi. 15, fig. 42); 3219 (Statz, 1950b, pi. 15, fig. 44); 3220 (Statz, 1950b, pi. 15, figs. 45 and 46); 3221 (Statz, 1950b, pi. 15, fig. 47); Oiigocene; Rott, Rheinland, Germany. Family NEPIDAE Ranatra asphalti PIERCE, 1948b:29,pl. 6, figs. 13-15. Holotype 2564; Pleistocene; asphalt deposit at McKittrick, Kern County, California. Renatra bessomi PIERCE, 1948b:27, pi. 6, figs. 11-12; pi. 7, figs. 16-18. Holotype 2565, 6 paratypes 2566-2571; Pleistocene; asphalt deposit at McKittrick, Kern County, California. NOTE: I am unable to match the specimens with the drawings which appear to be composite. Family NOTONECTIDAE Notonecta (Paranecta) badia PIERCE, 1948b:24, pi. 5, figs. 4-9. Holotype 2527; allotype 2528; 36 paratypes 2529-2562; Pleistocene; asphalt deposit at McKittrick, Kern County, California. NOTE: I am unable to match the specimens with the drawings. Notonecta jubata Schlechtendal, 1894 STATZ, 1938b as fig. 29. Figured specimen 3222, Oiigocene; Rott, Rheinland, Germany. 26 Contributions in Science No. 250 Notonecta primaeva Heyden, 1859 STATZ, 1950b as pi. 16, figs. 49-53; pi. 17, figs. 58-59. 6 figured specimens 3223 (pi. 16, fig. 49); 3224 (pi. 16, fig. 50); 3225 (pi. 16, fig. 51); 3226 (pi. 16, fig. 52, MISSING); 3227 (pi. 16, fig. 53); 3228) pi. 17, figs. 58 and 59); Oligocene; Rott, Rheinland, Germany. Soevenia hey deni (Deichmuller, 1881) STATZ, 1950b as pi. 13, figs. 35-36; pi. 17, figs. 54-57; pi. 18, figs. 60-67; pi. 19, figs. 68-74. 21 figured specimens 3229 (pi. 13, fig. 35); 3230 (pi. 13, fig. 36); 3231 (pi. 17, fig. 54); 3232 (pi. 17, fig. 55); 3233 (pi. 17, fig. 56); 3234 (pi. 17, fig. 57); 3235 (pi. 18, fig. 60); 3236 (pi. 18, fig. 61); 3237 (pi. 18, fig. 62); 3238 (pi. 18, fig. 63); 3239 (pi. 18, fig. 64); 3240 (pi. 18, fig. 65); 3241 (pi. 18, fig. 66); 3242 (pi. 18, fig. 67); 3243 (pi. 19, fig. 68); 3244 (pi. 19, fig. 69); 3245 (pi. 19, fig. 70, MISSING); 3246 (pi. 19, fig. 71); 3247 (pi. 19, fig. 72); 3248 (pi. 19, fig. 73); 3249 (pi. 19, fig. 74); Oligocene; Rott, Rheinland, Germany. Family PENTATOMIDAE Dinidorine sp. A STATZ and WAGNER, 1950 as pL 28, fig. 51. Figured specimen 3250 (MISSING); Oligocene; Rott, Rheinland, Germany. Pentatomine sp. A STATZ and WAGNER, 1950 as pi. 28, fig. 52. Figured specimen 3251 (MISSING); Oligocene, Rott, Rheinland, Germany. Pentatomine sp. B STATZ and WAGNER, 1950 as pi. 28, fig. 53. Figured specimen 3252; Oligocene; Rott, Rheinland, Germany. Pentatomine sp. C STATZ and WAGNER, 1950 as pi. 28, fig. 54. Figured specimen 3253; Oligocene; Rott, Rheinland, Germany. Family PYRRHOCORIDAE Pyrrhocoris ? tibialis STATZ in STATZ and WAGNER, 1950:118, pi. 27, fig. 44. Holotype 3257; Oligocene; Rott, Rheinland, Germany. Family REDUVIIDAE Rhinocoris michalki STATZ in STATZ and WAGNER, 1950:102, pi. 23, fig. 16. Syntype 3258 (MISSING); Oligocene; Rott, Rheinland, Germany. NOTE: There is another figured syntype (pi. 21, fig. 3; pi. 23, fig. 15) in the Kastenholz collection. Family SALDIDAE *Oligosaldina aquatilis STATZ in STATZ and WAGNER, 1950: 101, pi. 23, fig. 14. Holotype 3259 (MISSING); Oligocene; Rot(, Rheinland, Germany. *Oligasaldina rhenana STATZ in STATZ and WAGNER, 1950: 100, pi. 23, fig. 13. Holotype 3260 (MISSING); Oligocene; Rott, Rheinland, Germany. ^Oligosaldina rottensis STATZ in STATZ and WAGNER, 1950:99, pi. 23, fig. 12. Holotype 3261 (MISSING); Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 27 Family SCUTELLERIDAE Scutellerine sp. A STATZ and WAGNER, 1950 as pi. 27, fig. 45. Figured specimen 3254 (MISSING); Oligocene; Rott, Rheinland, Gerdiany. Scutellerine sp. B STATZ and WAGNER, 1950 as pi. 27, fig. 49. Figured specimen 3255; Oligocene; Rott, Rheinland, Germany. Poecilocoris ? rottensis STATZ in STATZ and WAGNER, 1950: 121, pi. 27, fig. 50. Holotype 3256; Oligocene; Rott, Rheinland, Germany. Order HOMOPTERA Family AETHIALIONIDAE Aethalion ? oligocaenica STATZ, 1950a: 6, pi. 3, figs. 32-33. Holotype 3262; Oligocene; Rott, Rheinland, Germany. Family CERCOPIDAE Cercopis ? sp. STATZ, 1950a as pi. 1, fig. 3; pi. 3, fig. 34. Figured specimen 3263; Oligocene; Rott, Rheinland, Germany. Family CHERMIDAE ^Draycotia cordovae PIERCE, 1948a:43,pl. 9. Holotype 2572; Pleistocene; Cordova Bay, Vancouver Island, British Colum- bia, Canada. Family CICADELLIDAE *Miochlorotettix gibroni PIERCE, 1963:73, figs. 1-2, 15-17. Holotype 2573 (figs. 1 and 15); allotype 2574 (figs. 16-17-); figured paratype 2576 (fig. 2, MISSING); 9 unfigured paratypes 2575 and 2577-2584 (2577- 2584 MISSING); Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. ^Miochlorotettix kirkbyi PIERCE, 1963:77, figs. 5 and 10-12. Holotype 2585 (MISSING); Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. ^Miomesamia juliae PIERCE, 1963:81, figs. 8-9. Holotype 2586; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. Phlepsius weissmanae PIERCE, 1963:80, figs. 13-14. Holotype 2563; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. ^Protochlorotettix calico PIERCE, 1963:78, figs. 3-4 and 18. Holotype 2587; Miocene, Barstow Formation; southern Calico Mountains, San Bernardino County, California. 28 Contributions in Science No. 250 Family CIXIIDAE Cixius ? sp. STATZ, 1950a as pi. 1, fig. 1; pi. 3, fig. 28. Figured specimen 3264; Oligocene; Rott, Rheinland, Germany. “Cixius” sp. STATZ, 1950a as pi. 1, fig. 2; pi. 3, fig. 29. Figured specimen 3265; Oligocene; Rott, Rheinland, Germany. Hyalesthes ? rottensis STATZ, 1950a:4, pi. 3, fig. 30. Holotype 3266 (MISSING); Oligocene; Rott, Rheinland, Germany. Family DELPHACIDAE “Delphax” rhemma STATZ, 1950a:5, pi. 3, fig. 31. Holotype 3267; Oligocene; Rott, Rheinland, Germany. Family JASSIDAE Batrachomorphiis ? femoralis STATZ, 1950a: 9, pi. 4, fig. 44. Holotype 3268; Oligocene: Rott. Rheinland, Germany. Bythoscopus lumitus STATZ, 1950a: 13, pi. 5, fig. 54. Holotype 3269; Oligocene; Rott, Rheinland, Germany. Bythoscopus robustus STATZ, 1950a: 14, pi. 5, fig. 55; pi. 6, fig. 56. 2 syntypes 3270 (pi. 5, fig. 55 MISSING); 3271 (pi. 6, fig. 56); Oligocene; Rott, Rheinland, Germany. Bythoscopus scpultus (Meunier, 1917) STATZ, 1950a as pi. 4, fig. 45; pi. 5, fig. 46-51. 7 figured specimens 3272 (pi. 4, fig. 45, MISSING); 3273 (pi. 5, fig. 46, MISSING).; 3274 (pi. 5, fig. 47); 3275 (pi. 5, fig. 48); 3276 (pi. 5, fig. 49); 3277 (pi. 5, fig. 50, MISSING); 3278 (pi. 5, fig. 51); Oligocene; Rott, Rhein- land, Germany. Bythoscopus scpultus austerus STATZ, 1950a; 13, pi. 5, figs. 52-53. 11 syntypes 3279 (pi. 5, fig. 52); 3280 (pi. 5, fig. 53); 3281-3289 (unfigured); Oligocene; Rott, Rheinland, Germany. Deltocephalus aberrans STATZ, 1950a: 18, pi. 2, fig. 19; pi. 6, figs. 63 and 65. Holoytpe 3290 (MISSING); Oligocene; Rott, Rheinland, Germany. Deltocephalus ? maculipennis STATZ, 1950a:20, pi. 7, figs. 68-69. 2 syntypes 3291 (fig. 68); 3292 (fig. 69); Oligocene; Rott, Rheinland, Germany. Deltocephalus ? macidosus STATZ, 1950a:22, pi. 8, fig. 72. Holotype 3293; Oligocene; Rott, Rheinland, Germany. STATZ, 1950a;20, pi. 7,fig. 67. Deltocephalus nervosus Holotype 3294; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 29 Deltocephalus ? pennatus STATZ, 1950a:22, pi. 7, fig. 71. Holotype 3295; Oligocene; Rott, Rheinland, Germany. Deltocephalus robustiis STATZ, 1950a: 19, pi. 2, fig. 22; pi. 7, fig. 66. Holotype 3296; Oligocene; Rott, Rheinland, Germany. Deltocephalus rottensis STATZ, 1950a: 18, pi. 2, figs. 20-21; pi. 6, fig. 64. Holotype 3297; Oligocene; Rott, Rheinland, Germany. Deltocephalus ? spinitibialis STATZ, 1950a:21, pi. 7, fig. 70. Holotype 3298; Oligocene; Rott, Rheinland, Germany. Euscelis plateosus STATZ, 1950a:23,pl. 8, fig. 73. 4 syntypes 3299 (pi. 8, fig. 73); 3300-3302 (unfigured); 3302 (MISSING); Oligocene; Rott, Rheinland, Germany. "^■'Gyponites abdominalis STATZ, 1950a: 11, pi. 4, fig. 43. Holotype 33.03; Oligocene; Rott, Rheinland, Germany. ■•'Gyponites pronota STATZ, 1950a: 10, pi. 4, fig. 42. Holotype 3304; Oligocene’; Rott, Rheinland, Germany. Idiocerus ? sp. STATZ, 1950a, as pi. 6, fig. 59. Figured specimen 3305; Oligocene; Rott, Rheinland, Germany. Idiocerus goeckei STATZ, 1950a: 17, pi. 6, figs. 60-62. 2 syntypes 3306 (figs. 60-61); 3307 (fig. 62); Oligocene; Rott, Rheinland, Germany. Macropsis pectoral is STATZ, 1950a: 15, pi. 6, fig. 57. Holotype 3308; Oligocene; Rott, Rheinland, Germany. '^'Oligogypona haupti STATZ, 1950a:8, pi. 1, figs. 8-9; pi. 4, fig. 39. Holotype 3309; Oligocene; Rott, Rheinland, Germany. "^•'Oligoidiocerus pronotumalis STATZ, 1950a: 16, pi. 2, fig. 17; pi. 6, fig. 58. Holotype 3310; Oligocene; Rott, Rheinland, Germany. '^Oligopenthimia ovalis STATZ, 1950a: 9, pi. 1, fig. 6; pi. 4, figs. 40-41. Holotype 3311; Oligocene; Rott, Rheinland, Germany. Paropia ? fortivenosa STATZ, 1950a: 6, pi. 1, fig. 4; pi. 3, figs. 35-36. Holotype 3312; Oligocene; Rott, Rheinland, Germany. Tettigoniella ? capita ta STATZ, 1950a:7, pi. 1, fig. 5; pi. 4, figs. 37-38. Holotype 3313; Oligocene; Rott, Rheinland, Germany. Thamnotettix sp. STATZ, 1950a as pi. 8, fig. 79. Figured specimen 3314; Oligocene; Rott, Rheinland, Germany. 30 Contributions in Science No. 250 Thamnotettix ampliventris STATZ, 1950a:24, pi. 8, fig. 76. 3 syntypes 3315 (pi. 8, fig. 76); 3316-3318 (unfigured; 3318 MISSING); Oligocene; Rott, Rheinland, Germany. Thamnotettix angustipennis STATZ, 1950a:24, pi. 2, fig. 25; pi. 8, fig. 75. Holotype 3319; Oligocene; Rott, Rheinland, Germany. Thamnotettix ? anthr acinus STATZ, 1950a:26, pi. 8, fig. 78. Holotype 3320; Oligocene; Rott, Rheinland, Germany. Thamnotettix clypealis STATZ, 1950a: 26, pi. 8, fig. 80. 2 syntypes 3321 (pi. 8, fig. 80); 3322 (unfigured); Oligocene; Rott, Rheinland, Germany. Thamnotettix gracilentus STATZ, 1950a:25, pi, 8, fig. 77. 12 syntypes 3323 (pi. 8, fig. 77); 3324-3334 (unfigured; 3334 MISSING); Oligocene; Rott, Rheinland, Germany. Thamnotettix ustus STATZ, 1950a:23, pi. 2, fig. 24; pi. 8, fig. 74. Holotype 3335; Oligocene; Rott, Rheinland, Germany. Order THYSANOPTERA Family THRIPIDAE Thrips sp. STATZ, 1941b as figs. 1-5. 5 figured specimens 3336 (fig. 1); 3337 (fig. 2); 3338 (fig. 3); 3339 (fig. 4); 3340 (fig. 5); Oligocene; Rott, Rheinland, Germany. Order NEUROPTERA Family CHRYSOPIDAE Notochrysa [sic Nothochrysa] praeclara STATZ, 1936a:215, figs. 3-4. Holotype 3341; Oligocene; Rott, Rheinland, Germany. Family MYRMELEONTIDAE Dendroleon septemmontanus STATZ, 1936a:212, figs. 1-2. Holotype 3342; Oligocene; Rott, Rheinland, Germany. Order MECOPTERA Family PANORPIDAE Panorpa gracilis STATZ, 1936a:216, figs. 5-7. Holotype 3343; Oligocene; Rott, Rheinland, Germany. Panorpa maculatuosa STATZ, 1936a:220, figs. 10-11. Holotype 3344; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 31 Panorpa rottensis STATZ, 1936a:218. Holotype 3345; Oligocene; Rott, Rheinland, Germany. NOTE: The figured half of the holotype (figs. 8-9) is in the Kastenholz collection. Order TRICHOPTERA Family PHRYGANEIDAE Neuronia excellenta STATZ, 1936a:221. Holotype 3346; Oligocene; Rott, Rheinland, Germany. NOTE: The figured half of the holotype (figs. 12-13) is in the Kastenholz collection. Phryganea pupa Nr, 1 STATZ, 1936a as fig. 14. Figured specimen 3347 (MISSING); Oligocene; Rott, Rheinland, Germany. Phryganea pupa Nr. 2 STATZ, 1936a as fig. 16. Figured specimen 3348; Oligocene; Rott, Rheinland, Germany. Phryganea pupa Nr. 3 STATZ, 1936a as fig. 19. Figured specimen 3349; Oligocene; Rott, Rheinland, Germany. Phryganea elegantula Meunier, 1920 STATZ, 1936a as figs. 21-22. Figured specimen 3350; Oligocene; Rott, Rheinland, Germany. Phryganea Uthophila STATZ, 1936a:233, figs. 23-24. 2 syntypes 3351 (unfigured); 3352 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: According to the label with the specimen (LACM 3351) it is the counterplate to the figured specimen. The figured portion is missing. Statz made no mention of more than one specimen in the type lot, but I consider a second specimen labeled as this species in the collection (LACM 3352) to be an additional syntype. Phryganea meiiniere Statz, 1936a:236. Holotype 3353; Oligocene; Rott, Rheinland, Germany. NOTE: The figured plate (figs. 27-28) is in the Kastenholz collection. Our specimen is the counterplate. Phryganea submersa STATZ, 1936a:235, figs. 25-26. Holotype 3354; Oligocene; Rott, Rheinland, Germany. Order LEPIDOPTERA Family HEPIALIDAE *Protohepialus comstocki PIERCE, 1945b:5, pi. 3-4. Holotype 3072; Upper Miocene; Puente Hills, SE of Puente, Los Angeles County, California. 32 Contributions in Science No. 250 Order DIPTERA Family ANTHOMYZIDAE Anthomyza ? rhenana STATZ, 1940b: 150, pi. 26, fig. 85. 2 syntypes 3355 (pi. 22, fig. 85); 3356 (unfigured); Oligocene; Rott, Rhein- land, Germany. Eamily BIBIONIDAE Bibio d Nr. 2 STATZ, 1943 as pi. 4, fig. 36 and pi. 12, fig. 95. Figured specimen 3357; Oligocene; Rott, Rheinland, Germany. Bibio c? Nr. 3 STATZ, 1943 as pi. 4, fig. 37, and pi. 12, fig. 94. Figured specimen 3358; Oligocene; Rott, Rheinland, Germany. Bibio cT Nr. 4 STATZ, 1943 as pi. 4, fig. 38 and pi. 12, fig. 96. Figured specimen 3359; Oligocene; Rott, Rheinland, Germany. Bibio d Nr. 5 STATZ, 1943 as pi. 4, fig. 39 and pi. 12, fig. 97. Figured specimen 3360; Oligocene; Rott, Rheinland, Germany. Bibio d Nr. 6 STATZ, 1943 as pi. 4, fig. 40 and pi. 12, fig. 98. Figured specimen 3361 (MISSING); Oligocene; Rott, Rheinland, Germany. Bibio aerosa STATZ, 1943:50, pi. 4, fig. 32; pi. 11, fig. 91. Holotype 3362; Oligocene; Rott, Rheinland, Germany. Bibio auripes STATZ, 1943:51, pi. 4, fig. 34; pi. 11, fig. 92. Holotype 3363; Oligocene; Rott, Rheinland, Germany. Bibio comosella STATZ, 1943:47, pi. 11, fig. 87. 2 syntypes 3364 (pi. 11, fig. 87); 3365 (unfigured); Oligocene; Rott, Rhein- land, Germany. Bibio compta STATZ, 1943:52, pi. 4, fig. 35; pi. 12, fig. 93. Holotype 3366; Oligocene; Rott, Rheinland, Germany. Bibio rohli STATZ, 1943:46, pi. 11, fig. 86. Holotype 3367; Oligocene; Rott, Rheinland, Germany. NOTE: The figured plate is in the Kastenholz collection. Our specimen is the counterplate. Bibio rottensis STATZ, 1943:48, pi. 11, fig. 88. 6 syntypes 3368 (pi. 11, fig. 88); 3369-3373 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: There are 4 additional syntypes in the Kastenholz collection and 1 in the Siegburger collection. Bibio spadicea STATZ, 1943:51, pi. 4, fig. 33; pi. 11, fig. 90. Holotype 3374; Oligocene; Rott, Rheinland, Germany. Bibio vulpina STATZ, 1943:49, pi. 4, fig. 31; pi. 11, fig. 89. Holotype 3375; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 33 Penthetria cT Nr. 1 STATZ, 1943 as pi. 10, fig. 81. Figured specimen 3376; Oligocene; Rott, Rheinland, Germany. Penthetria cf Nr. 2 STATZ, 1943 as pi. 10, fig. 85. Figured specimen 3377; Oligocene; Rott, Rheinland, Germany. Penthetria sp. STATZ, 1943 as pi. 3, fig. 25. Figured specimen 3378; Oligocene; Rott, Rheinland, Germany. Penthetria aestimata STATZ, 1943:39, pi. 10, figs. 79-80. 3 syntypes 3379 (pi. 10, fig. 79); 3380 (pi. 10, fig. 80); 3381 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: The counterplate to LACM 3379 is in the Kastenholz collection. Penthetria nigra STATZ, 1943:38, pi. 3, fig. 20. Holotype 3382; Oligocene; Rott, Rheinland, Germany. Penthetria rottensis STATZ, 1943:40, pi. 3, fig. 27; pi. 10, fig. 83. Holotype 3383; Oligocene; Rott, Rheinland, Germany. Penthetria scita STATZ, 1943:40, pi. 3, fig. 23; pi. 10, fig. 84. Holotype 3384; Oligocene; Rott, Rheinland, Germany. Penthetria tropica STATZ, 1943:39, pi. 3, fig. 22; pi. 10, fig. 82. Holotype 3385; Oligocene; Rott, Rheinland, Germany. Plecia STATZ, 1942a as fig. 5. Figured specimen 3386 (MISSING); Oligocene; Rott, Rheinland, Germany. Plecia c? Nr. 1 STATZ, 1943 as pi. 8, fig. 67. Figured specimen 3387; Oligocene; Rott, Rheinland, Germany. Plecia d' Nr. 2 STATZ, 1943 as pi. 1, fig. 10; pi. 8, fig. 70. Figured specimen 3388; Oligocene; Rott, Rheinland, Germany. Plecia d Nr. 3 STATZ, 1943 as pi. 8, fig. 68. Figured specimen 3389; Oligocene; Rott, Rheinland, Germany. Plecia cT Nr. 4 STATZ, 1943 as pi. 1, fig. 11; pi. 8, fig. 69. Figured specimen 3390; Oligocene; Rott, Rheinland, Germany. Plecia d Nr. 5 STATZ, 1943 as pi. 1, fig. 12; pi. 8, fig. 72. Figured specimen 3391; Oligocene; Rott, Rheinland, Germany. Plecia d Nr. 6 STATZ, 1943 as pi. 1, fig. 13; pi. 8, fig. 71. Figured specimen 3392; Oligocene; Rott, Rheinland, Germany. Plecia cf Nr. 7 STATZ, 1943 as pi. 2, fig. 14; pi. 9, fig. 73. Figured specimen 3393; Oligocene; Rott, Rheinland, Germany. Plecia d Nr. 10 STATZ, 1943 as pi. 2, fig. 15; pi. 9, fig. 74. Figured specimen 3394; Oligocene; Rott, Rheinland, Germany. 34 Contributions in Science No, 250 Plecia d Nr. 12 STATZ, 1943 as pL 2, fig. 18; pi. 9, fig. 75, Figured specimen 3395; Oligocene; Rott, Rheinland, Germany. Plecia Nr. 13 STATZ, 1943 as pi. 2, fig. 19; pi. 9, fig. 78. Figured specimen 3396; Oligocene; Rott, Rheinland, Germany. Plecia d Nr. 14 STATZ, 1943 as pi. 2, fig. 17; pi. 9, fig. 77. Figured specimen 3397; Oligocene; Rott, Rheinland, Germany. Plecia cf Nr. 15 STATZ, 1943 as pi-. 2, fig. 16; pi. 9, fig. 76. Figured specimen 3398; Oligocene; Rott, Rheinland, Germany. Plecia sp. STATZ, 1942a as fig. 4. STATZ, 1943 as pi. 7, fig 61. 2 figured specimens 3399 (Statz, 1942a, fig. 4; MISSING); 3400 (Statz, 1943, pi. 7, fig 61); Oligocene; Rott, Rheinland, Germany. Plecia exigiia STATZ, 1943:26, pi. 7, fig. 63. 2 syntypes 3401 (pi. 7, fig. 63); 3402 (unfigured); Oligocene; Rott, Rheinland, Germany. Plecia expositia (Heyden, 1865) STATZ, 1943 as pi. 7, fig. 59. Figured specimen 3404; Oligocene; Rott, Rheinland, Germany. Plecia grossa (Heyden, 1865) STATZ, 1943 as pi. 7, fig. 58. Figured specimen 3404; Oligocene; Rott, Rheinland, Germany. Plecia grandaeva (Heyden, 1865) STATZ, 1943 as pi. 7, fig. 65. Figured specimen 3405; Oligocene; Rott, Rheinland, Germany. Plecia hypogaea (Heyden, 1865) STATZ, 1943 as pi. 6, fig. 51. Figured specimen 3406; Oligocene; Rott, Rheinland, Germany. Plecia imperialis STATZ, 1943:13, pi. 1, fig. 3; pi. 5, fig. 48. Holotype 3407; Oligocene; Rott, Rheinland, Germany. Plecia lapidaria Heyden, 1865 STATZ, 1943 as pi. 7, fig. 60. Figured specimen 3408; Oligocene; Rott, Rheinland, Germany. Plecia lapidaria nigra STATZ, 1943:25, pi. 7, fig. 64. 4 syntypes 3409 (pi. 7, fig. 64); 3410-3412 (unfigured); Oligocene; Rott, Rheinland, Germany. Plecia luctuosa (Heyden, 1865) STATZ, 1943 as pi. 6, fig. 53. Figured specimen 3413; Oligocene; Rott, Rheinland, Germany. Plecia pennata STATZ, 1943:19, pi. 1, fig. 4; pi. 6, fig. 57. Holotype 3414; Oligocene; Rott, Rheinland, Germany. Plecia philippi STATZ, 1943:10, pi. l,fig. 1; pi. 5, fig. 50. Holotype 3415; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 35 Plecia pinguis (Heyden, 1865) STATZ, 1943 as pi. 6, fig. 56. Figured specimen 3416; Oligocene; Rott, Rheinland, Germany. Plecia proserpina (Heyden, 1865) STATZ, 1943 as pi. 6, fig. 55. Figured specimen 3417; Oligocene; Rott, Rheinland, Germany. Plecia rhenana Heyden, 1865 STATZ, 1943 as pi. 5, figs. 45-46. FURST, 1959, p. 16, top two figures, unnumbered. Figured specimen 3418; Oligocene; Rott, Rheinland, Germany. Plecia satyrus STATZ, 1943:12, pi. 5, fig. 47. 6 syntypes 3419 (pi. 5, fig. 47); 3420-3424 (unfigured, 3423 and 3424 MISSING); Oligocene; Rott, Rheinland, Germany. Plecia star mi STATZ, 1943:20, pi. 1, fig. 5; pi. 6, fig. 54. Holotype 3425; Oligocene; Rott, Rheinland, Germany. Plecia stygia (Heyden, 1865) STATZ, 1942a as fig. 6. STATZ, 1943 as pi. 7, fig. 62. Figured specimen 3426; Oligocene; Rott, Rheinland, Germany. Plecia veterana (Heyden, 1865) STATZ, 1943 as pi. 5, fig. 49. Figured specimen 3427; Oligocene; Rott, Rheinland, Germany. Plecia vulpina STATZ, 1943:27, pi. 7, fig. 66. 2 syntypes 3428 (pi. 7, fig. 66); 3429 (unfigured); Oligocene; Rott, Rhein- land, Germany. Plecia winnertzi (Heyden, 1865) STATZ, 1943 as pi. 6, fig. 52. Figured specimen 3430; Oligocene; Rott, Rheinland, Germany. Family BOMBYIIDAE Spongostylum sp. STATZ, 1940b as pi. 19, fig. 11; pi. 23, fig. 62. Figured specimen 3431; Oligocene; Rott, Rheinland, Germany. Usia atra STATZ, 1940b:131,pl. 19, fig. 10; pi. 23, fig. 61. Holotype 3432; Oligocene; Rott, Rheinland, Germany. Family CALLIPHORIDAE '^Protoclirysomyia howardae PIERCE, 1945d:8, pi. 6. ? 11 syntypes 3073-3083; Pleistocene; Pit 3, Rancho La Brea, Los Angeles, Los Angeles County, California. NOTE: There may be more fragments or whole specimens imbedded in the bone matrix. Family CERATOPOGONIDAE A trichopogon brunnescens STATZ, 1944c: 141, pi. 8, fig. 57. 9 syntypes 3433 (pi. 8, fig. 57); 3434-3441 (unfigured; 3441 MISSING); Oligocene; Rott, Rheinland, Germany. 36 Contributions in Science No. 250 Bezzia ? longipennis STATZ, 1944c: 151, pi. 7, fig. 55; pi. 10, fig. 80. Holotype 3442; Oligocene; Rott, Rheinland, Germany. “Ceratopogon*' sp. STATZ, 1941b as fig. 11. STATZ, 1944c as pi. 10, figs. 82-83. 3 figured specimens 3443 (Statz 1941b, fig. 11, MISSING); 3444 (Statz, 1944c, pi. 10, fig. 82); 3445 (Statz 1944c, pi. 10, fig. 83); Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 1 STATZ, 1944c as pi. 7, fig. 54; pi. 10, fig. 84, figs. 86-87. 4 figured specimens 3446 (pi. 7, fig. 54); 3447 (pi. 10, fig. 85); 3448 (pi. 10, fig. 86); 3449 (pi. 10, fig. 87); Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 2 STATZ, 1944c as pi. 10, fig. 88. Figured specimen 3450; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 3 STATZ, 1944c as pi. 10, fig. 89. Figured specimen 3451; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 4 STATZ, 1944c as pi. 1 1, fig. 90. Figured specimen 3452; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 5 STATZ, 1944c as pi. 11, fig. 91. Figured specimen 3453; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 6 STATZ, 1944c as pi. 11, fig. 92. Figured specimen 3454; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 7 STATZ, 1944c as pi. 11, figs. 93-95. 3 figured specimens 3455 (fig. 93); 3456 (fig. 94); 3457 (fig. 95); Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 8 STATZ, 1944c as pi. 11, fig. 96. Figured specimen 3458; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 9 STATZ, 1944c as pi. 11, fig. 97. Figured specimen 3459; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 10 STATZ, 1944c as pi. 11, fig. 98. Figured specimen 3460; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupa Nr. 1 1 STATZ, 1944c as pi. 11, fig. 99. Figured specimen 3461; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” 'Hr. 12 STATZ, 1944c as pi. 11, fig. 100. Figured specimen 3462; Oligocene; Rott, Rheinland, Germany. “Ceratopogon” pupdi'Hr. 13 STATZ, 1944c as pi. 11, fig. 101. Figured specimen 3463; Oligocene; Rott, Rheinland, Germany. Culicoides sp. STATZ, 1941b as fig. 6. Figured specimen 3464 (MISSING); Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 37 Culicoides abhreviatipennis STATZ, 1944c: 142, pi. 8, fig. 59. ^ 7 syntypes 3465 (pi. 8, fig. 59); 3466-3471 (unfigurecl; 3471 MISSING); Oligocene; Rott, Rheinland, Germany. Culicoides ? abdominalis STATZ, 1944c: 145, pi. 8, fig. 66. 3 syntypes 3472 (pi. 8, fig. 67); 3473-3474 (unfigured); Oligocene; Rott, Rheinland, Germany. Culicoides ? atratus STATZ, 1944c: 147, pi. 9, fig. 69. Holotype 3475; Oligocene; Rott, Rheinland, Germany. Culicoides austerus STATZ, 1944c: 144, pi. 7, figs. 48-49; pi. 8, figs. 63-64. 9 syntypes 3476 (pi. 7, fig. 38; pi. 8, fig. 63); 3477 (pi. 7, fig. 49; pi. 8, fig. 64); 3478-3484 (unfigured); Oligocene; Rott, Rheinland, Germany. Culicoides bicolor STATZ, 1944c: 146, pi. 8, fig. 68; pi. 9, fig. 70. 8 syntypes 3485 (pi. 8, fig. 68); 3486 (pi. 9, fig. 70); 3487-3492 (unfigured); 3489-3492 (MISSING); Oligocene; Rott, Rheinland, Germany. Culicoides elongatulus STATZ, 1944c: 143, pi. 8, figs. 60 and 62. 11 syntypes 3493 (fig. 60); 3494 (fig. 62); 3495-3503 (unfigured); Oligocene; Rott, Rheinland, Germany. Culicoides ? gracilior STATZ, 1944c: 143, pi. 8, fig. 61 3 syntypes 3504 (pi. 8, fig. 61); 3505 (unfigured); 3506 (unfigured; MISSING); Oligocene; Rott, Rheinland, Germany. Culicoides jucundus STATZ, 1941b as fig. 8 (nomen nudum). STATZ, 1944c: 147, pi. 9, figs. 72-74. 4 syntypes 3507 (Statz 1941b, fig. 8; Statz 1944c, pi. 9, figs. 72-73); 3508 (Statz, 1944c, pi. 9, fig. 74); 3509 (unfigured); 3f 10 (unfigured; MISSING); Oligocene; Rott, Rheinland, Germany. NOTE: In 1941 Statz figured a specimen with the name Culicoides jucundus with no accompanying description invalidating his later use of the name in 1944. Culicoides liliputanus STATZ, 1944c: 142, pi. 8, fig. 58. 3 syntypes 3511 (pi. 8, fig. 58); 3512 and 3513 (unfigured); Oligocene; Rott, Rheinland, Germany. Culicoides obesus STATZ, 1944c: 145, pi. 8, fig. 67. Holotype 3514; Oligocene; Rott, Rheinland, Germany. Culicoides obscuratus STATZ, 1944c: 144, pi. 8, fig. 65. 9 syntypes 3515 (pi. 8, fig. 65); 3516-3523 (unfigured; 3522 and 3523 MISSING); Oligocene; Rott, Rheinland, Germany. Culicoides tenuipennis STATZ, 1944c: 147, pi. 7, fig. 50; pi. 9, fig. 71. Holotype 3524; Oligocene; Rott, Rheinland, Germany. Serromyia sp. STATZ, 1944c as pi. 9, fig 75. Figured specimen 3525; Oligocene; Rott, Rheinland, Germany. 38 Contributions in Science No. 250 Serromyia austera STATZ, 1941b as fig. 10 (nomen nudum). Holotype 3526; Oligocene; Rott, Rheiland, Germany. NOTE: In 1941 Statz figured a specimen with the name Serromyia austera with no accompanying description invalidating his later use of the narre in 1944. Serromyia colorata STATZ, 1944c: 150, pi. 9, fig. 78. Holotype 3527; Oligocene; Rott, Rheinland, Germany. Serromyia spinofemorata STATZ, 1941b as fig. 9 (nomen nudum). STATZ, 1944c: 151, pi. 10, fig. 81. Holotype 3528; Oligocene; Rott, Rheinland, Germany. NOTE: In 1941 Statz figured a specimen with the name Serromyia spino- femorata with no accompanying description invalidating his later use of the name in 1944. Stilobezzia goetghebueri STATZ, 1944c: 148, pi. 7, fig. 52; pi. 9, fig. 77. Holotype 3529; Oligocene; Rott, Rheinland, Germany. Stilobezzia veterana (Meunier, 1920) STATZ, 1941b as fig. 7. STATZ, 1944c as pi. 9, fig. 76. Eigured specimen 3530 (MISSING); Oligocene; Rott, Rheinland, Germany. Family CHIRONOMIDAE “Orthocladius” cf Nr. 1 STATZ, 1944c as pi. 5, figs. 35-36. 2 figured specimens 3531 (pi. 5, fig. 35); 3532 (pi. 5, fig. 36; MISSING); Oligocene; Rott, Rheinland, Germany. “Orthocladius” d" Nr. 2 STATZ, 1944c as pi. 5, fig. 37. Figured specimen 3533; Oligocene; Rott, Rheinland, Germany. “Orthocladius” cf Nr. 3 STATZ, 1944c as pi. 5, fig. 38. Figured specimen 3534; Oligocene; Rott, Rheinland, Germany. “Orthocladius” cf Nr. 4 STATZ, 1944c as pi. 6, fig. 39. Figured specimen 3535; Oligocene; Rott, Rheinland, Germany. Pelopia ? sp. STATZ, 1944c as pi. 5, fig. 32. Figured specimen 3536; Oligocene; Rott, Rheinland, Germany. Pelopia dorminans (Heyden, 1870) STATZ, 1944c as pi. 3, figs. 16-21. 6 figured specimens 3537 (fig. 16); 3538 (fig. 17); 3539 (fig. 18); 3540 (fig. 19); 3541 (fig. 20); 3542 (fig. 21); Oligocene; Rott, Rheinland, Germany. Pelopia ? obscura STATZ, 1944c: 129, pi. 4, figs. 28-29. 10 syntypes 3543 (fig. 28); 3544 (fig. 29); 3545-3552 (unfigured); Oligocene; Rott, Rheinland, Germany. Pelopia pagasti STATZ, 1944c: 130, pi. 5 figs. 30-31. 4 syntypes 3553 (fig. 30); 3554 (fig. 31); 3555-3556 (unfigured); Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 39 Pelopia ? palaemon (Heyden, 1870) STATZ, 1944c as pi. 5, figs. 33-34. Figured specimen 3557; Oligocene; Rott, Rheinland, Germany. Pelopia thienemanni STATZ, 1944c: 128, pi. 4, figs. 22-24 and 27. 37 syntypes 3558 (fig. 22); 3559 (fig. 23); 3560 (fig. 24); 3561 (fig. 27); 3562-3594 (unfigured); Oligocene; Rott, Rheinland, Germany. Pelopiina (pupa) STATZ, 1944c as pi. 6, figs. 40-45. 6 figured specimens 3595 (fig. 40); 3596 (fig. 41); 3597 (fig. 42); 3598 (fig. 43); 3599 (fig. 44); 3600 (fig. 45, MISSING); Oligocene; Rott, Rhein- land, Germany. Pelopiina (exuvia) STATZ, 1944c as pi. 6, fig. 46. Figured specimen 3601; Oligocene; Rott, Rheinland, Germany. Procladius ? cf Nr. 1 STATZ, 1944c as pi. 2, figs. 7-13. 7 figured specimens 3602 (fig. 7); 3603 (fig. 8, MISSING); 3604 (fig. 9, MISSING); 3605 (fig. 10); 3606 (fig. 11); 3607 (fig. 12, MISSING); 3608 (fig. 13); Oligocene; Rott, Rheinland, Germany. Procladius d" Nr. 2 STATZ, 1944c as pi. 3, figs. 14-15. 2 figured specimens 3609 (fig. 14); 3610 (fig. 15); Oligocene; Rott, Rhein- land, Germany. Family CHLOROPIDAE “Chlorops” anthracinus STATZ, 1940b: 153, pi. 22, fig. 48; pi. 27, fig. 92. Holotype 3611; Oligocene; Rott, Rheinland, Germany. “Oscinella” dormescens STATZ, 1940b: 153, pi. 22, fig. 49. 2 syntypes 3612 (pi. 22, fig. 49); 3613 (unfigured); Oligocene; Rott, Rhein- land, Germany. Family CLUSIIDAE Clusiodes ? petreficata STATZ, 1940a: 150, pi. 22, fig. 42. Holotype 3614; Oligocene; Rott, Rheinland, Germany. Family CULICIDAE Anopheles ? rottensis STATZ, 1944b:109, pi. 2, fig. 19; pi. 5, fig. 38. Holotype 3615; Oligocene; Rott, Rheinland, Germany. Mansonia martinii STATZ, 1941b as fig. 13 (nomen nudum). STATZ, 1944b: 1 10, pi. 2, fig. 17; pi. 5, fig. 37. Holotype 3616; Oligocene; Rott, Rheinland, Germany. NOTE: In 1941 Statz figured a specimen with the name Mansonia martinii with no accompanying description invalidating his later use of the name in 1944. 40 Contributions in Science No. 250 Mansonia varivestita STATZ, 1941b as fig. 14 (nomen nudum). STATZ, 1944b: 111. 1 syntype 3617 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE 1: In 1941 Statz figured a specimen with the name Mansonia vari- vestita with no accompanying description invalidating his later use of the name in 1944. NOTE 2: There is a figured syntype (Statz, 1941b, fig. 14; Statz, 1944b, pi. 2, fig. 18; pi. 5, fig. 36) in the Kastenholz collection. Eamily DOLICHOPODIDAE “Dolichopus” minutus STATZ, 1940b: 136, pi. 20, fig. 17; pi. 25, fig. 70. Holotype 3618; Oligocene; Rott, Rheinland, Germany. NOTE: The counterplate is in the Kastenholz collection. “Dolichopus” scitus STATZ, 1940b: 136, pi. 20, fig. 18; pi. 24, fig. 68. Holotype 3619; Oligocene; Rott, Rheinland, Germany. Dolichopus spinosus STATZ, 1940b: 135, pi. 20, fig. 16. Holotype 3620; Oligocene; Rott, Rheinland, Germany. “Oncopygius” gracilior STATZ, 1940b: 137, pi. 20, fig. 20; pi. 25, fig. 71. Holotype 3622; Oligocene; Rott, Rheinland, Germany. “Oncopygius” oligocaenicus STATZ, 1940b: 137, pi. 24, fig. 69. 2 syntypes 3623 (pi. 24, fig. 69); 3624 (unfigured); Oligocene; Rott, Rhein- land, (Germany. “Oncopygius” venustus STATZ, 1940b: 138, pi. 20, fig. 21; pi. 25, fig. 72. Holotype 3621; Oligocene; Rott, Rheinland, Germany. Sciapus ? amabilis STATZ, 1940b: 139, pi. 20, fig. 23. Holotype 3625; Oligocene; Rott, Rheinland, Germany. Sciapus rottensis STATZ, 1940b: 139, pi. 21, fig. 24; pi. 25, fig. 74. Holotype 3626; Oligocene; Rott, Rheinland, Germany. Xanthochlorus ? tenellus STATZ, 1940b: 138, pi. 20, fig. 22; pi. 25, fig. 73. Holotype 3627; Oligocene; Rott, Rheinland, Germany. NOTE: The unfigured counterplate is in the Kastenholz collection. Family DROSOPHILIDAE “Drosophila” blanda STATZ, 1940b: 152, pi. 22, fig. 47; pi. 27, fig. 90. Holotype 3628; Oligocene; Rott, Rheinland, Germany. “Drosophila” elegans STATZ, 1940b: 152, pi. 22, fig. 46; pi. 27, fig. 91. Holotype 3629; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 41 Family DRYOMYZIDAE Dryomyza pelidua STATZ, 1940b: 146, pi. 22, fig. 35; pi. 26, fig. 81. Holotype 3630; Oligocene; Rott, Rheinland, Germany. Family EMPIDIDAE Empis spinifera STATZ, 1940b: 132, pi. 20, fig. 12; pi. 24, fig. 65. FURST, 1959:16, bottom figure, unnumbered. Holotype 3631; Oligocene; Rott, Rheinland, Germany. Empis umbonata STATZ, 1940b: 133, pi. 24, fig. 66. 2 syntypes 3632 (pi. 24, fig. 66); 3633 (unfigured); Oligocene; Rott, Rhein- land, (Germany. Hilara heeri MEUNIER, 1915b: 12, pi. 4, figs. 12-12a. 2 syntypes 4324 (fig. 12); 4325 (fig. 12a); Oligocene; Aix, Provence, France. Rhamphomyia fulvescens STATZ, 1940b: 133, pi. 20, fig. 14. Holotype 3640; Oligocene; Rott, Rheinland, Germany. Rhamphomyia latipennata STATZ, 1940b: 134, text fig. on p. 134. Holotype 3641; Oligocene; Rott, Rheinland, Germany. Rhamphomyia rottensis STATZ, 1940b: 135, pi. 24, fig. 67. 6 syntypes 3634 (pi. 24, fig. 67); 3635-3639 (unfigured); Oligocene; Rott, Rheinland, Germany. Family HELOMYZIDAE Leria bauckhorni (Meunier, 1915a) STATZ, 1940b as pi. 22, figs. 40-41; pi. 27, fig. 87. 2 figured specimens 3642 (pi. 22, fig. 41); 3643 (pi. 27, fig. 87); Oligocene; Rott, Rheinland, Germany. Family HIPPOBOSCIDAE Ornithoponus rottensis STATZ, 1940b: 154, pi. 22, fig. 50; pi. 27, fig. 93. STATZ, 1941b as figs. 16-17. MAA, 1966 as figs. 1-2. Holotype 3644; Oligocene; Rott, Rheinland, Germany. Family MYCETOPHILIDAE Asindulum ? pygmaea STATZ, 1944a:70, pi. 1, fig. 3; pi. 3, fig. 30. Holotype 3645; Oligocene; Rott, Rheinland, Germany. Boletina angustipennis STATZ, 1944a:78, pi 2, fig. 18; pi. 5, fig. 43. Holotype 3646; Oligocene; Rott, Rheinland, Germany. Boletina brunnescens STATZ, 1944a:78, pi. 2, fig. 17; pi. 5, fig. 42. Holotype 3647; Oligocene; Rott, Rheinland, Germany. 42 Contributions in Science No. 250 Boletina giaciosa STATZ, 1944a:79, pi. 2, fig 19; pi. 5, figs. 44-45. Holotype 3648; Oligocene; Rott, Rheinland, Germany. Boletina longicornis STATZ, 1944a:77, pi. 2, fig. 16; pi. 5, fig. 41. Holotype 3649; Oligocene; Rott, Rheinland, Germany. Ditomyia pilosella STATZ, 1944a:69, pi. 1, fig. 1; pi. 3, fig. 28. Holotype 3650; Oligocene; Rott, Rheinland, Germany. Docosia pilosa STATZ, 1944a:82, pi. 2, fig. 23; pi. 6, fig. 50. 1 syntype 3651; Oligocene; Rott, Rheinland, Germany. NOTE: The syntype figured as pi. 6, fig. 51 is in the Kastenholz collection. Dziedzickia ? lepida STATZ, 1944a:77, pi. 1, fig. 13; pi. 5, fig. 40. Holotype 3652; Oligocene; Rott, Rheinland, Germany. Ectrepesthoneura rottensis STATZ, 1944a:81, pj. 2, fig. 22; pi. 6, fig. 48. Holotype 3653 (MISSING); Oligocene; Rott, Rheinland, Germany. NOTE: ? second specimen is in the Kastenholz collection. Fungivorites problematicum STATZ, 1944a: 85, pi. 7, fig. 56. Holotype 3654; Oligocene; Rott, Rheinland, Germany. Leia aberrans STATZ, 1944a:79, pi. 2, fig. 20; pi. 6, fig. 46. Holotype 3655; Oligocene; Rott, Rheinland, Germany. Leia exhumata STATZ, 1944a:80, pi. 2, fig. 21; pi. 6, fig. 47. Holotype 3656; Oligocene; Rott, Rheinland, Germany. Macrocera umbonata STATZ, 1944a:70, pi. 1, fig. 2; pi. 3, fig. 29. Holotype 3657; Oligocene; Rott, Rheinland, Germany. Manota concolor STATZ, 1944a:71, pi. 1, fig. 4; pi. 3, fig. 32. Holotype 3658; Oligocene; Rott, Rheinland, Germany. Mycomyia sp. STATZ, 1944a as pi. 1, fig. 6; pi. 7, fig. 57. Figured specimen 3659; Oligocene; Rott, Rheinland, Germany. Mycomyia fossilis STATZ, 1944a:73, pi. 1, fig. 7; pi. 3, fig. 33. Holotype 3660; Oligocene; Rott, Rheinland, Germany. Mycomyia kuhni STATZ, 1944a:73, pi. 1, fig. 8; pi. 4, fig. 34. Holotype 3661; Oligocene; Rott, Rheinland, Germany. Mycomyia reisingeri STATZ, 1944a:75, pi. 1, fig. 11; pi. 4, fig. 37. Holotype 3662; Oligocene; Rott, Rheinland, Germany. Mycomyia umbonata STATZ, 1944a:74, pi. 1, fig. 9; pi. 4, fig. 36. Holotype 3663; Oligocene; Rott, Rheinland, Germany. Mycomyia unicolor STATZ, 1944a:71, pi. 1, fig. 5; pi. 3, fig. 31. Holotype 3664; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 43 Phthinia longipoda STATZ, 1944a:76, pi. 2, fig. 15; pi. 4, fig. 38. Holotype 3665; Oligocene; Rott, Rheinland, Germany. Polyxena hastata STATZ, 1944a:84, pi. 2, fig. 26; pi. 7, fig. 55. Holotype 3666; Oligocene; Rott, Rheinland, Germany. Polyxena (Cordyla) rhenana (Meunier, 1923) STATZ, 1944a as pi. 7, fig. 54. Figured specimen 3667; Oligocene; Rott, Rheinland, Germany. Family OPOMYZIDAE “Opomyza” pelidua STATZ, 1940b: 151, pi. 22, fig. 44; pi. 27, fig. 89. Holotype 3668; Oligocene; Rott, Rheinland, Germany. “Opomyza” venusta STATZ, 1940b: 151, pi. 27, fig. 88. 2 syntypes 3669 (pi. 27, fig. 88); 3670 (unfigured); Oligocene; Rott, Rhein- land, Germany. Family PHORIDAE Phalacrotophora tertiaria STATZ, 1940b: 140, pi. 21, fig. 25; pi. 25, fig. 75. Holotype 3671; Oligocene; Rott, Rheinland, Germany. Family PLATYPEZIDAE Opetia atra STATZ, 1940b: 141, pi. 21, fig. 26. Holotype 3672; Oligocene; Rott, Rheinland, Germany. Family RHAGIONIDAE Chrysopilus stigmaticus STATZ, 1940b: 129, pi. 19, figs. 6-7. Holotype 3673; Oligocene; Rott, Rheinland, Germany. Family SCATOPHAGIDAE Norellisoma setaria STATZ, 1940b: 145, pi. 21, fig. 33. Holotype 3674; Oligocene; Rott, Rheinland, Germany. Scopeuma punctata STATZ, 1940b: 146, pi. 22, fig. 34; pi. 26, fig. 80. Holotype 3675; Oligocene; Rott, Rheinland, Germany. Family SCIARIDAE Lycoria cf Nr. 3 STATZ, 1944c as pi. 13, fig. 127; pi. 17, fig. 153. Figured specimen 3676; Oligocene; Rott, Rheinland, Germany. Lycoria amicula STATZ, 1944c: 176, pi. 17, fig. 152. 4 syntypes 3677 (pi. 17, fig. 152); 3678-3679 (unfigured); 3680 (unfigured; MISSING); Oligocene; Rott, Rheinland, Germany. 44 Contributions in Science No. 250 Lycoria antennaria STATZ, 1944c: 169, pi. 15, fig. 134. 2 syntypes 3681 (pi. 15, fig. 134); 3682 (unfigured; MISSING); Oligocene; Rott, Rheinland, Germany. Lycoria aulica STATZ, 1944c: 165, pi. 14, fig. 128. 12 syntypes 3683 (pi. 14, fig. 128); 3684-3694 (unfigured); 3693 and 3694 (MISSING); Oligocene; Rott, Rheinland, Germany. Lycoria austera STATZ, 1944c: 166, pi. 14, fig. 131. 2 syntypes 3695 fpl. 14, fig. 131); 3696 (unfigured); Oligocene; Rott, Rhein- land, Germany. Lycoria bella STATZ, 1944c: 177, pi. 17, fig. 149. 5 syntypes 3697 (pi. 17, fig. 149); 3698-3701 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: One additional syntype is in the Kastenholz collection. Lycoria bicolora STATZ, 1944c: 173, pi. 16, fig. 144. 3 syntypes 3702 (pi. 16, fig. 144); 3703-3704 (unfigured); Oligocene; Rott, Rheinland, Germany. Lycoria compta STATZ, 1944c: 174, pi. 16, fig. 143. 2 syntypes 3705 (pi. 16, fig. 143); 3706 (unfigured); Oligocene; Rott, Rhein- land, Germany. Lycoria cryptocephala STATZ, 1944c: 168, pi. 12, fig. 107; pi. 14, fig. 133. Holotype 3707; Oligocene; Rott, Rheinland, Germany. Lycoria exilis STATZ, 1944c: 178, pi. 13, fig. 126; pi. 17, fig. 150. Holotype 3708; Oligocene; Rott, Rheinland, Germany. Lycoria femuralis STATZ, 1944c: 175, pi. 13, fig. 120; pi. 16, fig. 146. Holotype 3709; Oligocene; Rott, Rheinland, Germany. Lycoria gracilenta STATZ, 1944c: 172, pi. 13, fig. 116; pi. 16, fig. 142. Holotype 3710; (pi. 13, fig. 116; pi. 16, fig. 142); 3 paratypes 3711-3713 (unfigured); Oligocene; Rott, Rheinland, Germany. Lycoria gracilior STATZ, 1944c: 176, pi. 13, fig. 123; pi. 17, fig. 147. Holotype 3714; Oligocene; Rott, Rheinland, Germany. Lycoria jucunda STATZ, 1944c:170, pi. 15, fig. 140. 3 syntypes 3715 (pi. 15, fig. 140); 3716-3717 (unfigured); Oligocene; Rott, Rheinland, Germany. Lycoria laticornis STATZ, 1944c: 169, pi. 12, fig. 110; pi. 15, fig. 137. Holotype 3718; Oligocene; Rott, Rheinland, Germany. Lycoria lengersdorfi STATZ, 1944c: 172, pi. 16, fig. 141. 3 syntypes 3719 (pi. 16, fig. 141); 3720-3721 (unfigured); Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 45 Lycoria longa STATZ, 1944c: 168. 2 paratypes 3722-3723 (unfigured); Oligocene; Rott, Rheinland,^ Germany. NOTE: The holotype (pi. 15, fig. 135) is in the Kastenholz collection. Lycoria lugens STATZ, 1944c: 173, pi. 13, fig. 117; pi. 16, fig. 145. Holotype 3724; Oligocene; Rott, Rheinland, Germany. Lycoria melanopa STATZ, 1944c: 171, pi. 13, fig. 114; pi. 15, fig. 138. Holotype 3725; Oligocene; Rott, Rheinland, Germany. Lycoria minuscula STATZ, 1944c: 177, pi. 13, fig. 125; pi. 17, fig. 151. Holotype 3726; Oligocene; Rott, Rheinland, Germany. Lycoria ovalipennata STATZ, 1944c: 171, pi. 12, fig. 113; pi. 15, fig. 139. Holotype 3727; Oligocene; Rott, Rheinland, Germany. Lycoria pelidua STATZ, 1944c: 167, pi. 14, fig. 130. 9 syntypes 3728 (pi. 14, fig. 130); 3729-3736 (unfigured); Oligocene; Rott, Rheinland, Germany. Lycoria pennata STATZ, 1944c: 167, pi. 14, fig. 132. 2 syntypes 3737 (pi. 14, fig. 132); 3738 (unfigured); Oligocene; Rott, Rhein- land, Germany. Lycoria thoracica STATZ, 1944c: 175, pi. 13, fig. 121; pi. 17, fig. 148. Holotype 3739; Oligocene; Rott, Rheinland, Germany. Lycoria weylandi STATZ, 1944c: 165, pi. 14, fig. 129. 1 syntype 3740 (pi. 12, fig. 129); Oligocene; Rott, Rheinland, Germany. NOTE: An additional syntype is in the Heimatmuseum der Stadt Siegburg. Family SEPSIDAE Sepsis ? atra STATZ, 1940b: 147, pi. 22, fig. 37; pi. 26, fig. 83. Holotype 3741; Oligocene; Rott, Rheinland, Germany. Sepsis ? bicolorata STATZ, 1940b: 147, pi. 22, fig. 36; pi. 26, fig. 82. Holotype 3742; Oligocene; Rott, Rheinland, Germany. Sepsis ? ignobilis STATZ, 1940b: 148, pi. 26, fig. 84. 4 syntypes 3743 (pi. 26, fig. 84); 3744-3746 (unfigured); Oligocene; Rott, Rheinland, Germany. Family STRATIOMYIIDAE Eulalia sp. Nr. 1 STATZ, 1940b as pi. 19, fig. 3; pi. 23, figs. 52-54. 4 figured specimens 3747 (pi. 19, fig. 3); 3748 (pi. 23, fig. 52); 3749 (pi. 23, fig. 53); 3750 (pi. 23, fig. 54); Oligocene; Rott, Rheinland, Germany. 46 Contributions in Science No. 250 Eulalia sp. Nr. 2 STATZ, 1940b as pi. 19, fig. 4; p. 23, figs. 55-56. 3 figured specimens 3751 (pi. 19, fig. 4); 3752 (pi. 23, fig. 55); 3753 (pi. 23, fig. 56); Oligocene; Rott, Rheinland, Germany. Eulalia sp. Nr. 3 STATZ, 1940b as pi. 23, fig. 57. Figured specimen 3754; Oligocene; Rott, Rheinland, Germany. Hermione sp. STATZ, 1940b as pi. 23, fig. 58. Figured specimen 3755; Oligocene; Rott, Rheinland, Germany. Stratiomya sp. STATZ, 1940b as pi. 19, fig. 2; pi. 23, fig. 51. 2 figured specimens 3756 (pi. 19, fig. 2); 3757 (pi. 23, fig. 51); Oligocene; Rott, Rheinland, Germany. “Stratiomyia” pentadiscalia STATZ, 1940b: 124, pi. 19, fig. 1; pi. 23, fig. 59. Holotype 3758; Oligocene; Rott, Rheinland, Germany. Family SYRPHIDAE Chilosia picata STATZ, 1940b: 141. Holotype 3759; Oligocene; Rott, Rheinland, Germany. NOTE: The figured counterplate (pi. 21, fig. 27) is in the Kastenholz col- lection. Epistrophe kastenholzi STATZ, 1940b: 142, pi. 21, fig. 28. Holotype 3760; Oligocene; Rott, Rheinland, Germany. “Eristalis” sp. STATZ, 1940b as pi. 25, fig. 76. Figured specimen 3761; Oligocene; Rott, Rheinland, Germany. Myiolepta elisabethae STATZ, 1940b: 144, pi. 21, fig. 31; pi. 26, fig. 79. Holotype 3762; Oligocene; Rott, Rheinland, Germany. Spilomyia sp. STATZ, 1940b as pi. 21, fig. 30. Figured specimen 3763; Oligocene; Rott, Rheinland, Germany. Syrphus aeneus STATZ, 1940b: 142, pi. 21, fig. 29. Holotype 3764; Oligocene; Rott, Rheinland, Germany. NOTE: The counterplate is in the Kastenholz collection. Temnostoma sacki STATZ, 1940b: 145, pi. 21, fig. 32; pi. 25, fig. 77. Holotype 3765; Oligocene; Rott, Rheinland, Germany. Tropidia sp. STATZ, 1940b as pi. 26, fig. 78. Figured specimen 3766; Oligocene; Rott, Rheinland, Germany. Family TABANIDAE Tahanus fulvescens STATZ, 1940b: 130, pi. 19, fig. 8; pi. 24, fig. 63. STATZ, 1941b as fig. 15. Holotype 3767; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 47 Family THEREVIDAE Paraclia ? tarsalis STATZ, 1940b: 130, pi. 19, fig. 9; pi. 24, fig. 64. Holotype 3768; Oligocene; Rott, Rheinland, Germany. Family TIPULIDAE Dactylolabis tenuis STATZ, 1944b:99, pi. 1, fig. 6; pi. 4, fig. 27. Holotype 3769; Oligocene; Rott, Rheinland, Germany. Dicranoptycha rottensis STATZ, 1944b:95,pl. 1, fig. 2; pi. 3, fig. 22. Holotype 3770; Oligocene; Rott, Rheinland, Germany. Elephantomyia weigandi STATZ, 1934b:97, figs. 4-5. Holotype 3771; Oligocene; Rott, Rheinland, Germany. Erioptera ? pagasti STATZ, 1944b:97, pi. 1, fig. 5; pi. 3, fig. 25. Holotype 3772; Oligocene; Rott, Rheinland, Germany. Gonomyia munda STATZ, 1944b;96, pi. 1, fig. 3; pi. 3, fig. 21. Holotype 3773; Oligocene; Rott, Rheinland, Germany. Helius tenera STATZ, 1944b:95, pi. 1, fig. 1; pi. 3, fig. 20. Holotype 3774; Oligocene; Rott, Rheinland, Germany. Limnobia antiqua STATZ, 1934b:98, fig. 7. Holotype 3775; Oligocene; Rott, Rheinland, Germany. Limnobia horioni STATZ, 1934b:98,fig.6. Holotype 3776; Oligocene;. Rott, Rheinland, Germany. Limnophila sp. Nr. 1 STATZ, 1944b as pi. 1, fig. 9; pi. 3, fig. 23. Figured specimen 3777 (MISSING); Oligocene; Rott, Rheinland, Germany. Limnophila sp. Nr. 2 STATZ, 1944b as pi. 1, fig. 10; pi. 4, fig. 29. Figured specimen 3778; Oligocene; Rott, Rheinland, Germany. Limnophila sp. Nr. 3 STATZ, 1944b as pi. 2, fig. 14; pi. 4, fig. 30. Figured specimen 3779; Oligocene; Rott, Rheinland, Germany. Limnophila sp. Nr. 4 STATZ, 1944b as pi. 2, fig. 11; pi. 4, fig. 32. Figured specimens 3780 (MISSING); Oligocene; Rott, Rheinland, Germany. Limnophila ? pterotrichia STATZ, 1934b: 103, fig. 13. Holotype 3781; Oligocene; Rott, Rheinland, Germany. Limnophila robusta STATZ, 1934b: 102, fig. 12. Holotype 3782; Oligocene; Rott, Rheinland, Germany. Limnophila umbonata STATZ, 1944b:99, pi. 1, fig. 7; pi. 4, fig. 28. Holotype 3783; Oligocene; Rott, Rheinland, Germany. 48 Contributions in Science No. 250 Limnophila veterana STATZ, 1934b; 102, fig. 11. Holotype 3784; Oligocene; Rott, Rheinland, Germany. Rhabdomastix oligocaenica STATZ, 1944b:98, pi. 1, fig. 8; pi. 4, fig. 26. Holoytpe 3785; Oligocene; Rott, Rheinland, Germany. Tipiila sp. Nr. 1 STATZ, 1944b as pi. 2, fig. 16; pi. 5, fig. 34. Figured specimen 3786; Oligocene; Rott, Rheinland, Germany. Tipiila sp. Nr. 2 STATZ, 1944b as pi. 2, fig. 12; pi. 5, fig. 33. Figured specimen 3787 (MISSING); Oligocene; Rott, Rheinland, Germany. Tipiila sp. Nr. 3 STATZ, 1944b as pi. 2, fig. 13; pi. 5, fig. 35. Figured specimen 3788 (MISSING); Oligocene; Rott, Rheinland, Germany. Tipula bouata MEONIER, 1915b: 14; pi. 4, fig. 15. Holotype 4326; Oligocene; Aix, Provence, Erance. Tipula maxima STATZ, 1934b:95, figs. 1-3. Holotype 3789; Oligocene; Rott, Rheinland, Germany. Tipula peusi STATZ, 1944b: 101, pi. 2, fig. 15; pi. 4, fig. 31. Holotype 3790; Oligocene; Rott, Rheinland, Germany. Via rottensis STATZ, 1934b: 100, fig. 9. Holotype 3791; Oligocene: Rott, Rheinland, Germany. Eamily TRICHOCERIDAE T richocera miocaenica STATZ, 1934b: 100, fig. 10. Holotype 3792; Oligocene; Rott, Rheinland, Germany. Eamily TYLIDAE Trepidaria ? rottensis STATZ, 1940b: 148, pi. 22, fig. 39; pi. 27, fig. 86. Holotype 3793 (pi. 22, fig. 39; pi. 27, fig. 86); Paratype 3794 (unfigured); Oligocene; Rott, Rheinland, Germany. Order COLEOPTERA Eamily CARABIDAE Bembidion (Oclitliedromus) davidae PIERCE, 1944a:7, pi. 1, figs. a-c. Holotype 2590; Lower Miocene; W bank of the Cuyama River near the junction with Husana Creek, San Luis Obispo County, California. Bembidion ev ere s t a e PIERCE, 1944b: 11, pi. 2; pi. 3, figs. a-b. Holotype 2591; Lower Miocene; near Rosedale, 6 mi W of Bakersfield, Kern County, California. 1973 Additional Type Specimens 49 Elaphrus clairvillei lynni PIERCE, 1948a:52, pi. 12. Holotype 2948; Pleistocene; Lynn Creek, north of Vancouver, British Columbia, Canada. Elaphrus ruscarius foveatus PIERCE, 1948c: 54, pi. 13, figs. 1-3. Holotype 2949; Pleistocene; asphalt deposit at McKittrick, Kern County, California. Noiiophilus vancouveri PIERCE, 1950b: 5, pi. 2, figs. 1-3. Holotype 3056; Pleistocene; Lynn Creek, N of Vancouver, British Columbia, Canada. Family CERAMBYCIDAE Anaesthelis ? wapiti ficus STATZ, 1938c- 176, figs. 5-6. Holotype 3795; Oligocene; Rott, Rheinland, Germany. Haplocnemia sophia STATZ, 1938c: 175, fig. 2. Holotype 3796; Oligocene; Rott, Rheinland, Germany. Leptura loupipcunis STATZ, 1938c; 175, fig. 1. Holotype 3797; Oligocene; Rott, Rheinland, Germany. Sapcrda ? pclrorum STATZ, 1938c: 177, fig. 3. Holotype 3798; Oligocene; Rott, Rheinland, Germany. NOTE: The unfigured counterplate is in the Kastenholz collection. Te traps rotteusis STATZ, 1938c: 178, fig. 4. Holotype 3799; Oligocene; Rott, Rheinland, Germany. Family CH R YSOMELIDAE Donacia (Donacia) cordovae PIERCE, 1950c:8 (unfigured). Holotype 2887; 3 paratypes 2888-2890; Pleistocene; Cordova Bay, Vancouver Island, British Columbia, Canada. Donacia (Donacia) draxcoti PIERCE, 1950c:7(unfigured). Holotype 2891; 54 paratypes 2892-2945; Pleistocene; Lynn Creek, N of Vancouver, British Columbia, Canada. Donacia (Donacia) lynni PIERCE, 1950c;8 (unfigured). Holotype 2946; paratype 2947; Oligocene; Lynn Creek, N of Vancouver, British Columbia, Canada. Family COCCINELLIDAE Coccinclla sp. FURST, 1959:14, p. 14, top figure, unnumbered. Figured specimen 3800 (MISSING); Oligocene; Rott, Rheinland, Germany. 50 Contributions in Science No. 250 FamUy CURCULIONIDAE Rhyncolus kathrynae SLEEPER, 1968:196, figs. 1-2. Holotype 3084; Pliocene; Sam Swarz Ranch, Fernley, Lyon County, Nevada. Rhyssematus miocenae PIERCE, 1965:157, fig. 1. Holotype 3070; Upper Miocene; Woodcrest Drive, Sherman Oaks, Los Angeles County, California. Family DRYOPIDAE Helmis decoratus STATZ, 1938b, fig. 26 (both plate and counterplate figured); (nomen nudum). STATZ, 1939:68, pi. 20, figs. 92-94. Holotype 3801; Oligocene; Rott, Rheinland, Germany. NOTE: In 1938 Statz figured a specimen with the name Helmis decoratus with no accompanying description invalidating his later use of the name in 1944. Family DYTISCIDAE Dytiscites sp. (larva) STATZ, 1939 as pi. 16, fig. 73. Figured specimen 3802; Oligocene; Rott, Rheinland, Germany. Agabus hoc hi STATZ, 1938b as fig. 21; (nomen nudum). STATZ, 1939:56, pi. 13, figs. 61-62. Holotype 3803; Oligocene; Rott, Rheinland, Germany. NOTE: In 1938 Statz figured a specimen with the name Agabus hochi with no accompanying description invalidating his later use of the name in 1944. Hydroporus ? maculatiis STATZ, 1939:55, pi. 12, fig. 58. Holotype 3804; Oligocene; Rott, Rheinland, Germany. Hydroporus multipiinctatus STATZ, 1939:55, pi. 13, figs. 59-60. Holotype 3805; Oligocene; Rott, Rheinland, Germany. Oreodites cryptolineatus STATZ, 1938b as fig. 20 (2 specimens figured); (nomen nudum). STATZ, 1939:53, pi. 11, figs. 51-54; pi. 12, figs. 55-57. 76 syntypes 3806 (Statz, 1938b, fig. 20, ventral view; Statz, 1939, pi. 12, fig. 56); 3807 (Statz, 1938b, fig. 20, dorsal view); 3808 (Statz, 1939, pi. 1 1, fig. 51); 3809 (pi. 11, fig. 52); 3810 (pf. 11, fig. 53); 3811 (pi. 11, fig. 54); 3812 (pi. 12, fig. 55); 3813 (pi. 12, fig. 57); 3814-3881 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: In 1938 Statz figured two specimens with the name Oreodites crypto- lineatus with no accompanying description invalidating his later use of the name in 1944. Family GYRINIDAE Palaeogyrinus strigatus Schlechtendal, 1894 STATZ, 1938b as fig. 22. STATZ, 1939 as pi. 15, figs. 69-72. 5 figured specimens 3882 (Statz, 1938b, .fig. 22; MISSING); 3883 (Statz, 1939, pi. 15, figs. 67-68; MISSING); 3884 (Statz, 1939, pi. 15, fig. 69); 3885 (Statz, 1939, pi. 15, fig. 70); 3886 (Statz, 1939, pi. 15, figs. 71-72); Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 51 Family HYDROPHILIDAE Berosus capitatus STATZ, 1939:62, pi. 17, figs. 77-78. Holotype 3887; Oligocene; Rott, Rheinland, Germany. Berosus morticinus (Heyden, 1866) STATZ, 1938b as fig. 24. STATZ, 1939 as pi. 16, figs. 74 and 76. 3 figured specimens 3888 (Statz, 1938b, fig. 24); 3889 (Statz, 1939, fig. 74); 3890 (Statz, 1939, fig. 76); Oligocene; Rott, Rheinland, Germany. Cymbiododyta ? austerus STATZ, 1939:66, pi. 19, fig. 88. Holotype 3891; Oligocene; Rott, Rheinland, Germany. Hydrophilus fraternus Heyden, 1859 STATZ, 1939 as pi. 19, figs. 85-87. 2 figured specimens 3892 (fig. 85); 3893 (fig. 86-87); Oligocene; Rott, Rheinland, Germany. Hydrophilus neptunus (Heyden, 1866) STATZ, 1939 as pi. 18, figs. 82-84. 2 figured specimens 3894 (figs. 82-83); 3895 (fig. 84); Oligocene; Rott, Rheinland, Germany. Hydrous ? ebeninus STATZ, 1939:64. Syntype 3896 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: There is a figured (pi. 18, fig. 81) syntype in the Kastenholz col- lection. Hydrous rottensis STATZ, 1938b as fig. 23 (both plate and counterplate figured); (nomen nudum). STATZ, 1939:63, pi. 17, figs. 79-80. Holotype 3897; Oligocene; Rott, Rheinland, Germany. NOTE: In 1938 Statz figured a specimen with the name Hydrous rottensis with on accompanying description invalidating his later use of the name in 1944. Paracymus excitatus (Heyden, 1866) STATZ, 1938b as fig. 25. STATZ, 1939 as pi. 19, fig. 89; pi. 20, figs. 90-91. 4 figured specimens 3898 (Statz, 1938, fig. 25; MISSING); 3899 (Statz, 1939, pi. 19, fig. 89); 3900 (Statz, 1939, pi. 20, fig. 90); 3901 (Statz, 1939, pi. 20, fig. 91); Oligocene; Rott, Rheinland, Germany. Eamily LUCANIDAE ^Ceruchites hahnei STATZ, 1952:5, pi. l,fig. 4. Holotype 3902; Oligocene; Rott, Rheinland, Germany. Eamily MORDELLIDAE Mordella indata STATZ, 1952:3, pi. 1, figs. 1-2. 2 syntypes 3903 (fig. 1); 3904 (fig. 2); Oligocene; Rott, Rheinland, Germany. Mordella nigrapilosa STATZ, 1952:4, pi. 1, fig. 3. Holotype 3905; Oligocene; Rott, Rheinland, Germany. 52 Contributions in Science No. 250 Family PTILIIDAE Ptilium tertiarium MORION in STATZ and MORION, 1937:8, figs. 1-4. Molotype 3906; Oligocene; Rott, Rheinland, Germany. Family SCARABAEIDAE Aphodiiis helvolus STATZ, 1952:6, pi. 1, fig. 6. Molotype 3907; Oligocene; Rott, Rheinland, Germany. Canthon practicolus vetustus PIERCE, 1946a: 122 (unfigured). Molotype 2592; paratype 2593; Pleistocene; Pit 81, Rancho La Brea, Los Angeles, Los Angeles County, California. Canthon simplex antiqiius PIERCE, 1946a: 120, pi. 10, figs. 1-3. Molotype 2594 (pi. 10, figs. 2-3); 4 paratypes 2595 (pi. 10, fig. 1); paratype 2596-2598 (unfigured); Pleistocene; Pit A, Rancho La Brea, Los Angeles, Los Angeles County, California. Copris pristinus PIERCE, 1946a: 124, pi. 1 1, figs. 4-9; pi. 13, figs. 18-19. Molotype 2850 (pi. 11, figs. 4-9); paratype (pi. 13, fig. 19); 39 paratypes, 2850 and 2853-2878 and 4329-4333 from Pit A; and 2851 and 2879-2883 and 4328 from Pit 16; 2884-2886 from Pit 13-183; all Pleistocene; Rancho La Brea, Los Angeles, Los Angeles County, California. Goet rapes ? sp. a STATZ, 1952 as pi. 2, fig. 2. Figured specimen 3908; Oligocene; Rott, Rheinland, Germany. Geotrupes ? sp. b STATZ, 1952 as pi. 2, fig. 3. Figured specimen 3909; Oligocene; Rott, Rheinland, Germany. Geotrupes rottensis STATZ, 1952:7, pi. 2, fig. 1. Molotype 3910; Oligocene; Rott, Rheinland, Germany. Maladera ? spinitibialis STATZ, 1952:9, pi. 2, fig. 4. Molotype 3911; Oligocene; Rott, Rheinland, Germany. Melolontha ? sp. STATZ, 1952 as pi. 2, fig. 6. Figured specimen 3912; Oligocene; Rott, Rheinland, Germany. Onthophagus everestae PIERCE, 1946a: 131, pi. 12, figs. 15-17. Molotype 3057 (pi. 12, figs. 15-16); paratype 3058 (pi. 12, fig. 17); Pleisto- cene; Pit 81, Rancho La Brea, Los Angeles, Los Angeles County, California. Onthophagus mulleri STATZ, 1952:8, pi. 2, fig. 5. Molotype 3913; Oligocene; Rott, Rheinland, Germany. ■^Palaeocopris labreae PIERCE, 1946a: 130, pi. 11, figs. 10-13. Molotype 3059 (figs. 10-11); 4 paratypes 3060 (fig. 12); 3061 (fig. 13); 3062 (fig. 14); 3063 (unfigured); Pleistocene; Pit 81, Rancho La Brea, Los Angeles, Los Angeles County, California. 1973 Additional Type Specimens 53 Serica kanakoffi PIERCE, 1946b: 132, pi. 14, figs. 1-2. Holotype 3071; Pleistocene; Pit 81, Rancho La Brea, Los Angeles, Los Angeles County, California. Family SILPHIDAE Nicrophorus guttuhis labreae PIERCE, 1949:63, pi. 9, fig. 7; pi. 10, fig. 9. 136 syntypes 2950 (pi. 9, fig. 7); 2951 (pi. 10, fig. 9); 2952-2979 and 4334- 4375 (unfigured); Pleistocene; 2950-2979 and 4334-4353 from Pit A; 2980- 2981 and 4354 from Pit B; 2982-3044 and 4355 from Pit X; 3045 from Pit 28; 3046 from Pit 37; 4356-4361 from Pit 81; 4362-4374 from Pit Bliss 29; 4375 from an unknown pit, all are from Rancho La Brea, Los Angeles, Los Angeles County, California. Nicrophorus giittulus punctostriatus PIERCE, 1949:66 (unfigured). Holotype 3047; Pleistocene; Pit A, Rancho La Brea, Los Angeles, Los Angeles County, California. Nicrophorus investigator alpha PIERCE, 1949:67, pi. 10, fig. 13. 5 syntypes 3048 (pi. 10, fig. 13); 3049-3052 (unfigured); Pleistocene; Pit A, Rancho La Brea, Los Angeles, Los Angeles County, California. Nicrophorus investigator latifrons PIERCE, 1949:67, pi. 10, fig. 14. Holotype 3053; Pleistocene; Rancho La Brea, Los Angeles, Los Angeles County, California. Nicrophorus mckittricki PIERCE, 1949, pi. 10, fig. 11. Holotype 3054; Pleistocene; Site 3, McKittrick, Kern County, California. Nicrophorus obtusiscutellum PIERCE, 1949: 67, pi. 10, fig. 12. Holotype 3055; Pleistocene; Pit A, Rancho La Brea, Los Angeles, Los Angeles County, California. Family TENEBRIONIDAE A psena labreae PIERCE, 1954b:98, pi. 23, figs. 6-7. Holotype 2588 (pi. 23, figs. 6-7); 2598 paratype (unfigured); Pleistocene; Pit B, Rancho La Brea, Los Angeles, Los Angeles County, California. Coniontis abdominalis caseyi PIERCE, 1954c: 145 (unfigured). Holotype 2599; 108 paratypes 2600-2707; Pleistocene; 2599-2620 from Pit A; 2621-2625 from Pit B; 2626-2640 from Pit C; 2641-2663 from Pit X; 2664- 2707 from Pit Bliss 29; all are from Rancho La Brea, Los Angeles, Los Angeles County, California. Coniontis abdominalis fragmans PIERCE, 1954c: 148, pi. 42, fig. 2. Holotype 2708, Pleistocene; Rancho La Brea, Los Angeles, Los Angeles County, California. 54 Contributions in Science No. 250 Coniontis abdominalis labreae PIERCE, 1954c: 146, pi. 41, figs. 1-2; pi. 42, figs. 3 and 10-13. Holotype 2709 (pi. 41, fig. 1); 41 paratypes 2710 (pi. 42, figs. 10-13); 2711- 2747; Pleistocene; 2709-2746 from Pit Bliss 29; 2747 from Pit 28; all are from Rancho La Brea, Los Angeles, Los Angeles County, California. NOTE: I am unable to match the remaining drawings and paratypes. Coniontis blissi PIERCE, 1954c: 149, pi. 42, fig. 16. Holotype 2748 (pi. 42, fig. 16); 13 paratypes, 2749-2761; Pleistocene; 2748 and 2759-2761 from Pit Bliss 29; 2749 from Pit A; 2750-2754 from Pit B, 2755-2757 from Pit X; 2758 from Pit 9; all are from Rancho La Brea, Los Angeles, Los Angeles County, California. Coniontis pectoralis interrupta PIERCE, 1954c: 154, pi. 42, figs. 15-15a. Holotype 2763 (MISSING); 12 paratypes 2753-2764 (MISSING); Pleisto- cene; 2753 from PitA; 2754-2758 from Pit B; 2759-2760 from Pit X; 2761 from Pit 9; 2762-2764 from Pit Bliss 29; all from Rancho La Brea, Los Angeles, Los Angeles County, California. Coniontis pectoralis paraeUiptica PIERCE, 1954c: 153, pi. 42, fig. 7. Holotype 2775 (pi. 42, fig. 7); 12 paratypes 2776-2787; Pleistocene; Rancho La Brea, Los Angeles, Los Angeles County, California. Coniontis remnans PIERCE, 1954c: 155, pi. 41, fip. 4-6. Holotype 2788; Pleistocene; Pit B, Rancho La Brea, Los Angeles, Los Angeles County, California. Coniontis trisiis alpha PIERCE, 1954c: 148, pi. 42, fig. 1. Holotype 2798 (pi. 42, fig. 1); 56 paratypes 2790-2845; Pleistocene; Rancho La Brea, Los Angeles, Los Angeles County, California. Coniontis tristis asphalti PIERCE, 1954c: 149, pi. 42, fig. 6. HOLOTYPE 2846; paratype 2847; Pleistocene; Pit 9, Rancho La Brea, Los Angeles, Los Angeles County, California. NOTE: I am unable to match the specimens with the drawing. Coniontis tristis latigula PIERCE, 1954c: 149, pi. 42, fig. 5. Holotype 2848; paratype 2849; Pleistocene; Pit Bliss 29, Rancho La Brea, Los Angeles, Los Angeles County, California. NOTE: I am unable to match the specimens with the drawing. Parasida mckittricki PIERCE, 1954a:43, pi. 8, figs. 4-5. Holotype 3064 (pi. 8, figs. 4-5); 5 paratypes 3065-3069 (unfigured); Pleisto- cene; 3064-3065 from Site 4; 3066-3069 from Site 3; all from McKittrick, Kern County, California. 1973 Additional Type Specimens 55 Order HYMENOPTERA Family APIDAE Synapis dormitans (Heyden, 1862) STATZ, 1931 as fig. 9. ALFONSUS, 1932:275 (unnumbered figure). WATSON, 1933:48 (unnumbered figure). STATZ, 1934a as fig. 2. STATZ, 1941a:101, 105, 108, 112 (unnumbered figures). FURST, 1959, pi. 13 (unnumbered figure). 5 figured specimens 3914 (Statz, 1931, fig. 5); 3915 (Statz, 1934a, figs. 1, 5-6; Statz, 1941a: 108); 3916 (Statz, 1931, fig. 9; Alfonsus, 1932:275; Watson, 1933:48; Statz, 1934a, fig 2; Statz, 1941a: 10; Furst, 1959:10; Furst, 1959:13); 3917 (Statz, 1941a: 105; MISSING); 3918 (Statz, 1941a: 112; MISSING); Oligocene; Rott, Rheinland, Germany. Synapis henshawi (Cockerell, 1907) STATZ, 1934a as figs. 5-6. STATZ, 1941a: 103 (unnumbered). Figured specimen 3919 (MISSING); Oligocene; Rott, Rheinland, Germany. Synapis kaschkei STATZ, 193 1:50, figs. 10-11. STATZ, 1934a as fig. 3. STATZ, 1941a: 109, bottom fig. (unnumbered). Holotype 3920 (Statz, 1931, fig. 10-11); figured specimen 3921 (Statz, 1934a, fig. 3; 1941a, pi. 109, bottom fig.); Oligocene; Rott, Rheinland, Germany. Xylocopa friesei STATZ, 1936b:284, pi. 13, fig. 37. Holotype 3922; Oligocene; Rott, Rheinland, Germany. Family BETHYLIDAE Epyris ? tennellus STATZ, 1938d: 107, pi. 14, fig. 46. Holotype 3923; Oligocene; Rott, Rheinland, Germany. Family BRACONIDAE Alysia latifrons STATZ, 1936b:279, pi. 9, fig. 27. Holotype 3924; Oligocene; Rott, Rheinland, Germany. Apanteles concinna STATZ, 1938d:83, pi. 8, fig. 8. Holotype 3925; Oligocene; Rott, Rheinland, Germany. Apanteles macrophthalmiis STATZ, 1938d:82, pi. 8, fig. 7. Holotype 3926; Oligocene; Rott, Rheinland, Germany. Aspicolpus longicornis STATZ, 1936b:277, pi. 8, fig. 25. Holotype 3927; Oligocene; Rott, Rheinland, Germany. Aspilota ? stigmalineata STATZ, 1938d:88, pi. 9, fig. 16. Holotype 3928; Oligocene; Rott, Rheinland, Germany. Blacus crassicapitatus STATZ, 1938d:81, pi. 7, fig. 5. Holotype 3929; Oligocene; Rott, Rheinland, Germany. 56 Contributions in Science No. 250 Cantharoctonus bruesii STATZ, 1936b:275, pi. 7, fig. 22 Holotype 3930 (MISSING); Oligocene; Rott, Rheinland, Germany. Clinocentrus rottensis STATZ, 1936b:276, pi. 8, fig. 23. Holotype 3931; Oligocene; Rott, Rheinland, Germany. Doryctes conjectus STATZ, 1938d:86, pi. 9, fig. 13. Holotype 3932; Oligocene; Rott, Rheinland, Germany. Doryctes longulus STATZ, 1938d:86, pi. 9, fig. 12. Holotype 3933; Oligocene; Rott, Rheinland, Germany. Doryctes rotiindatus STATZ, 1938d:85, pi. 8, fig. 11, Holotype 3934; Oligocene; Rott, Rheinland, Germany. Hecabolus gladiator STATZ, 1936b: 274, pi. 7, fig. 20. Holotype 3935; Oligocene; Rott, Rheinland, Germany. “Meteorus” longicornis STATZ, 1938d:81,pl. 8, fig. 6. Holotype 3936; Oligocene; Rott, Rheinland, Germany. *Palaeoblacus aculeatus STATZ, 1936b:277, pi. 8, fig. 24. Holotype 3937; Oligocene, Rott, Rheinland, Germany. Pentapleura ? filicornis STATZ, 1938d:87, pi. 9, fig. 15. Holotype 3938; Oligocene; Rott, Rheinland, Germany. Pentapleura longipennis STATZ, 1936b:278, pi. 9, fig. 26. Holotype 3939; Oligocene; Rott, Rheinland, Germany. “Rhogas” festivus STATZ, 1938d:87, pi. 9, fig. 14. Holotype 3940; Oligocene; Rott, Rheinland, Germany. Spathius longicornis STATZ, 1938d:84, pi. 8, fig. 10. Holotype 3941; Oligocene; Rott, Rheinland, Germany. Spathius ? macroradialis STATZ, 1938d:84, pi. 8, fig. 9. Holotype 3942; Oligocene; Rott, Rheinland, Germany. Spathius pedicularis STATZ, 1936b:275,pl. 7, fig. 21. Holotype 3943; Oligocene; Rott, Rheipjand, Germany. Tanycarpa interstitialis STATZ, 1936b:279, pi. 9, fig. 28. Holotype 3944; Oligocene; Rott, Rheinland, Germany. Family DIAPRIIDAE Diapriites insignicornis STATZ, 1938d:104, pi. 14, fig. 41. Holotype 3945; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 57 Diapriites minimus STATZ, 1938d:105, pi. 14, fig. 42. Holotype 3946; Oligocene; Rott, Rheinland, Germany. Pantoclis margaritaceus STATZ, 1938d:103,pl. 13, fig. 39. Holotype 3947; Oligocene; Rott, Rheinland, Germany. Family ENCYRTIDAE “Eiwyrtus” clavicornis STATZ, 1938d:102,.pl. 13, fig. 37. Holotype 3948, Oligocene; Rott, Rheinland, Germany. Family EULOPHIDAE “Eulophus” elegantulus STATZ, 1938d:100, pi. 13, fig. 35. 3 syntypes 3949 (pi. 13, fig. 35); 3950-3951 (unfigured); Oligocene; Rott, Rheinland, Germany. “Eulophus” mundus STATZ, 1938d:101, pi. 13, fig. 36. Holotype 3952; Oligocene; Rott, Rheinland, Germany. Family FIGITIDAE Figites ? planus STATZ, 1938d:90,pl. 10, fig. 19. Holotype 3953; Oligocene; Rott, Rheinland, Germany. Figites ? rotundatus STATZ, 1938d:89, pi. 10, fig. 18. Holotype 3954; Oligocene; Rott, Rheinland, Germany. Figites ? spiniger STATZ, 1938d:89, pi. 10, fig. 17. Holotype 3955; Oligocene; Rott, Rheinland, Germany. Family HELORIDAE Helorus festivus STATZ, 1938d:102, pi. 13, fig. 38. Holotype 3956; Oligocene; Rott, Rheinland, Germany. Family ICHNEUMONIDAE “Compople.x” parvulus STATZ, 1938d:79,pl. 7, fig. 3. Holotype 3957; Oligocene; Rott, Rheinland, Germany. Crypt us ? morleyi (Meunier, 1923) STATZ, 1936b as pi. 3, fig. 9. Figured specimen 3958 (MISSING); Oligocene; Rott, Rheinland, Germany. Hemiteles hirsuta STATZ, 1936b:270, pi. 5, fig. 13. Holotype 3959; Oligocene; Rott. Rheinland, Germany. Microcryptus terebrator STATZ, 1936b: 269, pi. 4, fig. 11. Holotype 3960; Oligocene; Rott, Rheinland, Germany. 58 Contributions in Science No. 250 Orthopelma curvitibialis STATZ, 1936b:272,pl. 6, fig. 17. Holotype 3961; Oligocene; Rott, Rheinland, Germany. Phygadeuon crassicornis STATZ, 1936b:269. Holotype 3962; Oligocene; Rott, Rheinland, Germany. NOTE: The figured (pi. 4, fig. 12) coimterplate is in the Kastenholz col- lection. Pimpla cyclostigmata STATZ, 1936b:270, pi. 5, fig. 14. , Holotype 3963; Oligocene; Rott, Rheinland, Germany. Polysphincta statzi (Meunier, 1923) STATZ, 1936baspl. 5, fig. 15. Figured specimen 3964; Oligocene; Rott, Rheinland, Germany. Protarchus antiquus STATZ, 1936b:273, pi. 6, fig. 18. Holotype 3965; Oligocene; Rott, Rheinland, Germany. Stenomacrus obliquus STATZ, 1936b:273, pi. 6, fig. 19. Holotype 3966; Oligocene; Rott, Rheinland, Germany. Family MEGACHILIDAE Anthidium mortuum (Meunier, 1920a) STATZ, 1936b as pi. 13, fig. 38. Figured specimen 3967 (MISSING); Oligocene; Rott, Rheinland, Germany. Family PLATYGASTERIDAE Platygasterites femoralis STATZ, 1938d:106, pi. 14, fig. 43. Holotype 3968; Oligocene; Rott, Rheinland, Germany. Platygasterites spinosa STATZ, 1938d:106, pi. 14, fig. 44. Holotype 3969; Oligocene; Rott, Rheinland, Germany. Family PROCTOTRUYPIDAE Psilomma pulchellus STATZ, 1938d:104, pi. 13, fig. 40. 2 syntypes 3970 (pi. 13, fig. 40); 3971 (unfigured); Oligocene; Rott, Rheinland, Germany. NOTE: There is an additional unfigured syntype in the Kastenholz collection. Family PSAMMOCHARIDAE Priocnemis aertsi STATZ, 1936b:283, pi. 12, fig. 35. Holotype 3972; Oligocene; Rott, Rheinland, Germany. Priocnemis wettweri STATZ, 1938d:108, pi. 14, fig. 47. Holotype 3973; Oligocene; Rott, Rheinland, Germany. Psammochares depressa STATZ, 1936b:283, pi. 12, fig. 34. Holotype 3974; Oligocene; Rott, Rheinland, Germany. 1973 Additional Type Specimens 59 Family PTEROMALIDAE “Pteromalus” abdominalis STATZ, 1938d:95, pi. 11, fig. 25. Holotype 3975; Oligocene; Rott, Rheinland, Germany. “Pteromalus'' aerosus STATZ, 1938d:98, pi. 12, fig. 30. Holotype 3976; Oligocene; Rott, Rheinland, Germany. “Pteromalus” atomus STATZ, 1938d:100,pl. 12, fig. 33. Holotype 3977; Oligocene; Rott, Rheinland, Germany. “Pteromalus” atra STATZ, 1938d:97, pi. 12, fig. 28. Holotype 3978; Oligocene; Rott, Rheinland, Germany. “Pteromalus” clavicornis STATZ, 1938d:94, pi. 11, fig. 24. Holotype 3979; Oligocene; Rott, Rheinland, Germany. “Pteromalus” crassicapitatus STATZ, 1938d:94, pi. 11, fig. 23. Holotype 3980; Oligocene; Rott, Rheinland, Germany. “Pteromalus” defossus STATZ, 1938d:92, pi. 10, fig. 21. Holotype 3981; Oligocene; Rott, Rheinland, Germany. “Pteromalus” hirtipes STATZ, 1938d:96,pl. 11, fig. 27. Holotype 3982; Oligocene; Rott, Rheinland, Germany. “Pteromalus” latipennatus STATZ, 1938d:93, pi. 11, fig. 22. Holotype 3983; Oligocene; Rott, Rheinland, Germany. “Pteromalus” longicornis STATZ, 1938d:99, pi. 12, fig. 32. Holotype 3984; Oligocene; Rott, Rheinland, Germany. “Pteromalus” pulchellus STATZ, 1938d:91, pi. 10, fig. 20. 3 syntypes 3985, (pi. 10, fig. 20); 3986-3987 (unfigured); Oligocene; Rott, Rheinland, Germany. “Pteromalus” pygmaeolus STATZ, 1938d:10(), pi. 12, fig. 34. Holotype 3988; Oligocene; Rott, Rheinland, Germany. “Pteromalus” rectispinus STATZ, 1938d:99, pi. 12, fig. 31. Holotype 3989; Oligocene; Rott, Rheinland, Germany. “Pteromalus” rottensis STATZ, 1938d:96,pl. 11, fig. 26. 7 syntypes 3990 (pi. 11, fig. 26); 3991-3996 (unfigured); Oligocene; Rott, Rheinland, Germany. “Pteromalus” venustus STATZ, 1938d:98, pi. 12, fig. 29. Holotype 3997; Oligocene; Rott, Rheinland, Germany. Family SCELIONIDAE Scelionites capitatus STATZ, 1938d:107, pi. 14, fig. 45. Holotype 3998; Oligocene; Rott, Rheinland, Germany. 60 Contributions in Science No. 250 Family SPHECIDAE Nysson rottensis MEUNIER, 1915:212, pi. 22, fig. 1. Holotype 4327; Oligocene; Roll, Rheinland, Germany. Sphex obscura STATZ, 1936b:284, pi. 12, fig. 36. Holotype 3999; Oligocene; Rott, Rheinland, Germany. Family TENTHREDINIDAE Holocampa e ns I ini STATZ, 1936b:266, pi. 3, fig. 7. Holotype 4000; Oligocene; Rott, Rheinland, Germany. Lencempria oligocaenica (Meunier, 1923) STATZ, 1936b as pi. 3, fig. 8. Figured specimen 4001 (MISSING); Oligocene; Rott, Rheinland, Germany. Tenthrado gracilis STATZ, 1936b:266, pi. 2, fig. 6. Holotype 4023, Oligocene; Rott. Rheinland, Germany. Family TRIGONAIJDAE “T rigonalys” bischoffi STATZ, 1938d:77, pl.6;pl. 7, fig. 1. Holotype 4002; Oligocene; Rott, Rheinland, Germany. Family VESPIDAE Polistes signata STATZ, 1936b:282, pi. 11, fig. 33. Holotype 4003; Oligocene; Rott, Rheinland, Germany. Vespa bilineal a STATZ, 1936b:281, pi. 10, fig. 31. 4 syntypes 4004 (pi. 10, fig. 31); 4005-4007 (iinfigured); Oligocene; Rott, Rheinland, Germany. Vespa cordifera STATZ, 1936b:280,pl. 10, fig. 30. Holotype 4008; Oligocene; Rott, Rheinland, Germany. Vespa nigra STATZ, 1936b:282, pi. 11, fig. 32. Holotype 4009; Oligocene; Rott, Rheinland, Germany. Family XYELIDAE Pleroneura graciosa (Meunier, 1920a) STATZ, 1936b as pi. l,fig. 1. Figured specimen 4010; Oligocene; Rott, Rheinland, Germany. Xyela angustipennis STATZ, 1936b:264, pi. 1, fig. 3. Holotype 4011 (pi. 1, fig. 3); 3 paratypes 4012-4014 (unfigured); Oligocene; Rott, Rheinland, Germany. Xyela latipennis STATZ, 1936b:263, pi. 1, fig. 2. Holotype 4015 (pi. 1, fig. 2); 4 paratypes 4016-4019 (unfigured); Oligocene; Rott. Rheinland, Germany. 1973 Additional Type Specimens 61 Xyela magna STATZ, 1936b:264, pi. 2, fig. 4. Holotype 4020; Oligocene; Rott, Rheinland, Germany. LITERATURE CITED Alfonsus, E. C. 1932. A fossil bee from long ago. Amer. Bee J. 72(7) :275. Anderson, F. M. 1902. Cretaceous deposits of the Pacific coast. Proc. Cal. Acad. Sci. 3rd Ser. (Geology) 2(1): 1-154. Arnold, R. 1907. New characteristic species of fossil mollusks from the oil-bearing Tertiary formations of southern California. Proc. U.S. Nat. Mus. 32(1545): 525-546. . 1908. Descriptions of new Cretaceous and Tertiary fossils from the Santa Cruz Mountains, California. Proc. U.S. Nat. Mus. 34(1617): 345-390. Bartsch, P. 1911. The recent and fossil mollusks of the genus Bittium from the west coast of America. Proc. U.S. Nat. Mus. 40(1 826): 383-41 4. Berry, S. S. 1940. New Mollusca from the Pleistocene of San Pedro, California — I. Bulls, of Amer. Paleontol. 25(94A) : 149-164. . 1941. New Mollusca from the Pleistocene of San Pedro, California — II. Bulls, of Amer. Paleontol. 27(101): 3-18. Born, I. 1778. Index Rerum Naturalium Musei Caesarei Vindobonensis. Part I: Testacea. Vienna, xlii + 458. Carpenter, F. M. 1968. The affinities of the genus Sobobapteron Pierce. Bull. So. Calif. Acad. Sci. 67(4) :263-265. Carpenter, P. P. 1855. Descriptions of (supposed) new species and varieties of shells from the Californian and west Mexican coasts principally in the col- lection of Hugh Cuming, Esq. Proc. Zool. Soc. London 23:228-235. . 1857. Catalogue of the collection of Mazatlan shells in the British Museum: collected by Frederick Reigen. London (British Museum) pp. i-xii + 1-552. . 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Rept. Brit. Assoc. Adv. Sci. for 1863, p. 517-686. Carson, C. M. 1926. New molluscan species from the Californian Pliocene. Bull. So. Calif. Acad. Sci. 25(2):49-62. Clark, B. L. 1925. Pelecypoda from the marine Oligocene of western North America. Univ. of Calif. Publ., Bull. Dept. Geol. Sci. 15(4) :69-136. Clark, B. L. ^and A. P. Woodford. 1927. The geology and paleontology of the type section of the Meganos formation (lower middle Eocene) of Cali- fornia. Univ. of Calif. Publ., Bull. Dept. Geol. Sci. 17(2) : 63-142. Cockerell, T. D. A. 1907. A fossil honey-bee. Entomologist 40(533) : 227-229. Conrad, T. A. 1834. Descriptions of nev/ Tertiary fossils from the southern States. J. Acad. Nat. Sci. Phila. 7:130-157. . 1837. Descriptions of marine shells from Upper California, collected by Thomas Nuttall, Esq. J. Acad. Nat. Sci. Phila. 7(2):227-268. 62 Contributions in Science No. 250 . 1849. Descriptions of new fresh water and marine shells. Proc. Acad. Nat. Sci. Phila. 4:152-157. — — . 1855. Descriptions of fossil shells from the Eocene and Miocene forma- tions of California. Appendix VI. U.S. House of Representatives, Document 129. . 1857. Report on the paleontology of the survey. 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Diagnoses of new species of marine bivalve mollusks from the northwest coast of America in the United States National Museum. Proc. U.S. Nat. Mus. 52(2183):393-417. . 1919. Descriptions of new species of Mollusca from the north Pacific Ocean in the collection of the United States National Museum. Proc. U.S. Nat. Mus. 56(2295) :293-371. Dana, J. D. 1847. Descriptions of fossil shells, Amer. J. Sci. 54:151-160. Davis, C. H. 1913. New species from the Santa Lucia Mountains, California with a discussion of the Jurassic age of the slates at Slate’s Springs. J. Geol. 21(5):- 453-458. Deichmuller, J. V. 1881. Fossile insecten aus dem diomeenschiefer von kutschlin bei Bilin, Bohmen. Nova Acta Acad. Caesar. Leopoldino-Davolinae Ger- manicae Naturae Curiosorum 42(6) :295-331. De Koninck, L. G. 1887. Ann. Musee Royal d’Hist. Nat. Belg. 14(6). Deshayes, G. P. 1839. Nouvelles especes de mollusques, provenant des cotes de la Californie, du Mexique, du Kamtschatka et de la Nouvelle-Zelande. 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Geol. 20(5): 3 1-42. 1973 Additional Type Specimens 63 Finlay, H. J. 1927. New specific names for Austral Mollusca. New Zealand Inst. Trans, and Proc. 57:488-533. ^ Furst, P. T. 1959. The Oligocene world of George Stratz. Pacific Discovery 12(6):11-19. . 1970. The aqualung spider-birth to death under water. Terra 9(2):22-31. Gabb, W. M. 1864. Cretaceous fossils. Paleontology of California. 1:1-197, Gale, H. R. 1928. West coast species of Hinnites. Trans. San Diego Soc. Nat. Hist. 5(9):91-94. Gould, A. A. 1850. {Descriptions of shells from the United States Exploring Expedition}. Proc. Boston Soc. Nat. Hist. 3:275-278. Grant, U. S. and H. R. Gale. 1931. Catalogue of the marine Pliocene and Pleisto- cene Mollusca of California and adjacent regions. Mem. San Diego Soc. Nat. Hist. 1:1-1036. Hagen, H. A. 1863. Neuroptern aus der braunkohle von Rott im Siebengebirge. Palaeontographica 10:247-269. Hanna, G. D. 1926. Paleontology of Coyote Mountain, Imperial County, Cali- fornia. Proc. Calif. 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Bibioniden aus der Rheinischen braunkohle von Rott. Palaeontogra- phica 14:19-30. . 1866. Kafer und Polypen aus der braunkhole des Siebengebirges. Palaeon- tographica 15:135, . 1870. Fossil Dipteren aus der braunkohle von Rott im Siebengebirge. Palaeontographica 17:249. International Commission on Zoological Nomenclature. 1964. International code of zoological nomenclature, adopted by the XV International Congress of Zoology, Inst. Trust Zool. Nomenclature, London, pp. 1-176. Landois, H. 1895. Die Riesenammonitne von Seppenrade, Pachy discus Zittel Seppenradensis H. Landois. Dreiundzwanzigster jahresbericht des West- falischen Provinzial-Vereins fiir Wissenschaft un kunst fur 1894/1895:99-108. Lea, I. 1833. Contributions to Geology. Philadelphia, pp. 1-227. Maa, T. C. 1966. Redescription of the fossil Ornithomya rottensis (Statz) (Diptera: Hippoboscidae). Pacific Insects Monog. 10:3-9. Matsumoto, T. and Y. Kanie. 1967, Ainoceras, a new heteromorph ammonoid genus from the Upper Cretaceous of Hokkaido. (Studies of the Cretaceous ammonites from Hokkaido and Saghalien — XVI). Mem. Fac. Sci., Kyushu Univ., Ser. D, Geology, 18(2): 349-359. 64 Contributions in Science No. 250 Meek, F. B. 1864. Descriptions of new organic remains from the Cretaceous rocks of Vancouver’s Island. Trans. Albany Inst. 4:37-49. Menzies, R. J. 1951. Pleistocene Brachyura from the Los Angeles area: Cancridae. J. Paleontol. 25(2): 165-170. Merriam, C. W. 1941. Fossil turritellas from the Pacific coast region of North America. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 26(1): 1-214. Meunier, F. 1914. Nouvelles recherches sur quelques insectes du Sannoisien d’Aix- en- Provence. Bull. Soc. Geol. France 4 ser. 14:187-198. . 1915a. Uber einige fossile insekten aus deh Braunkohlenischichten (Aqui tanien) von Rott (Siebengebirge). Zs. D. Ges. Berlin 67:205-217. . 1915b. Nouvelles recherches sur quelques insecta des Platrieres D’Aix en Provence. Verhandelingen der Koninklijke Akademie Van Wetenschappente Amsterdam 18(5): 1-17. . 1920. Quelques insectes de I’Aquitanien de Rott, Sept-Monts (Prusse rhenane). P. Ak. Amsterdam 22:891-898. . 1923. Sur quelques insectes de I’Aquitanien de Rott, Sept-Monts, Rhenanie). Misc. Ent. Castanet-Tolosan 26:82-87. Meyer, H. 1859. Micropsalis papyracea aus der Rheinischen braunkohle. Palaeon- tographica 8:18-21. Morris, J. 1845. Descriptions of fossils, in Strezelecki, physical description of New South Wales and Van Dieman’s Land, p. 378. Mount, J. D. 1971. A dragonfly nymph from the Miocene of San Pedro, Cali- fornia. Bull. So. Calif. Paleontol. Soc. 3(11): 3-4. Nomland, J. O. 1916. Corals from Cretaceous and Tertiary of California and Oregon. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 9(5): 59-76. Oldroyd, T. S. 1925. The fossils of the lower San Pedro fauna of the Nob Hill cut, San Pedro, California. Proc. U.S. Nat. Mus. 65(22): 1-39. Osmont, V. C. 1905. Areas of the California Neocene Univ. Calif. Publ., Bull. Dept. Geol. Sci. 4(4): 89-100. Pack, R. W. 1909. Notes on echinoids from the Tertiary of California. Univ. Calif. Publ., Bull. Geol. Sci. 5:275-283 Petrunkevitch, A. 1955. Arachinida. In R. C. Moore ed., Treatise on Invertebrate Paleontology, Part P, Arthropoda 2. Geol. Soc. Amer. and Univ. Kans. Press, pp. 42-162. Pierce, W. D. 1944a. Fossil Arthropods of California. 2. Description of a Lower Miocene fossil carabid beetle, with a discussion of Coleopterous elytra. Bull. So. Calif Acad. Sci. 43(l):4-9. . 1944b. Fossil Arthropods of California. 3. Description of a deep well Lower Miocene carabid beetle. Bull. So. Calif, Acad. Sci. 43(1): 10-12. . 1944c. Fossil Arthropods of California. 4. Two interesting Orthopteroids from diatomaceous deposits. Bull. So. Calif. Acad. Sci. 43(1): 12-17. . 1945a. Fossil Arthropods of California. 5a. A crystallized millipede from volcanic rock in a well. Bull. So. Calif. Acad. Sci. 44(1): 1-2. . 1945b. Fossil Arthropods of California. 6. Two new fossils from the Upper Miocene of the Puente Hills. Bull. So. Calif. Acad. Sci. 44(1): 3-6. . 1945c. Fossil Arthropods of California. 7. A fossil whiptail scorpion from Cabrillo Beach. Bull. So. Calif. Acad. Sci. 44(1) :7-8. . 1945d. Fossil Arthropods of California. 8. A case of Pleistocene myiasis from the La Brea Pits; Bull. So. Calif. Acad. Sci. 44(1): 8-9. . 1946a. Fossil Arthropods of California. 11. Description of dung beetles (Scarabaeidae) of the tar pits. Bull. So. Calif. Acad. Sci. 45(3) : 1 19-131 . 1973 Additional Type Specimens 65 . 1946b. Fossil Arthropods of California. 12. Description of a sericine beetle from the tar pits. Bull. So. Calif. Acad. Sci. 54(3): 131-132. . 1948a. Fossil Arthropods from British Columbia. 3. A chermid wing from interglacial lignite. Bull. So. Calif. Acad. Sci. 47(l):42-44. . 1948b. Fossil Arthropods of California. 15. Some Hemiptera from the McKittrick’asphalt field. Bull. So. Calif. Acad. Sci. 47(l):21-33. . 1948c. Fossil Arthropods from California. 16. The carabid genus Elaphrus in the asphalt deposits. Bull. So. Calif. Acad. Sci. 47(2): 53-55. . 1949. Fossil Arthropods of California. 17. The silphid burying beetles in the asphalt deposits. Bull. So. Calif. Acad. Sci. 48(2): 55-70. . 1950a. Fossil Arthropods of British Columbia. 6. Fossil spider silk. Bull. So. Calif. Acad. Sci. 49(1): 3. . 1950b. Fossil Arthropods of British Columbia. 7, A carabid from Lynn Creek lignite. Bull. So. Calif. Acad. Sci. 49(1) :4-7. . 1950c. Fossil Arthropods of British Columbia. 8. The genus Donacia in British Columbia lignite. Bull. So. Calif. Acad. Sci. 49(1) :7-9. . 1950d. Fossil Arthropods from onyx marble. 2. New pedipalpids from onyx marble. Bull. So. Calif. Acad. Sci. 49(3) : 102-104. . 1950e. Fossil Arthropods from onyx marble. 3. A primitive Thysanuran from onyx marble. Bull. So. Calif. Acad. Sci. 49(3): 104. . 1951a. Fossil Arthropods from onyx-marble. 7. The fossil pedipalpi from Bonner Quarry. Bull. So. Calif. Acad. Sci. 50(1): 38-40. . 1951b. Fossil Arthropods from onyx-marble. 8. A fossil millipede from Bonner Quarry. Bull. So. Calif. Acad. Sci. 50(l):41-43. . 1951c. Fossil Arthropods from onyx-marble. 9. Fossil primitive insects from onyx-marble. Bulk So. Calif. Acad. Sci. 50(1): 44-49. . 1954a. Fossil Arthropods of California. 18. The Tenebrionidae-Tentyriinae of the asphalt deposits. Bull. So. Calif. Acad. Sci. 53(1): 35-45. . 1954b. Fossil Arthropods of California. 19. The Tenebrionidae-Scaurinae of asphalt deposits. Bull. So. Calif. Acad. Sci. 53(2):93-98. . 1954c. Fossil Arthropods of California. 20. The Tenebrionidae-Coniontinae of the asphalt deposits. Bull. So. Calif. Acad. Sci. 53(3): 142-156. . 1958. Fossil Arthropods of California. 21. Termites from Calico Mountains nodules. Bull. So. Calif. Acad. Sci. 57(1): 13-24. . 1960. Fossil Arthropods of California. 23. Silicified insects in Miocene nodules from the Calico Mountains. Bull. So. Calif. Acad. Sci. 59(l):40-49. . 1963. Fossil Arthropods of California. 25. Silicified leafhoppers from California mountains nodules. Bull. So. Calif. Acad. Sci. 62(2): 69-82. . 1964. Three new types of invertebrates extracted from Miocene petroli- ferous nodules. Bull. So. Calif. Acad. Sci. 63(2): 8 1-85. . 1965. Fossil Arthropods of California. 26. Three new fossil insect sites in California. Bull. So. Calif. Acad. Sci. 64(3): 157-162. Pierce, W. D. and J. Gibron. 1962. Fossil Arthropods of California. 24. Some unusual fossil Arthropods from the Calico Mountains nodules. Bull. So. Calif. Acad. Sci. 61(3): 143-151. Popenoe, W. P. 1937. Upper Cretaceous Mollusca from southern California. J. Paleontol. 11 (5): 379-402. Popenoe, W. P. and W. A. Findlay. 1933. Transposed hinge structures in lamellibranchs. Trans. San Diego Soc. Nat. Hist. 7(26) : 299-3 18. Randall, J. W. 1839. Catalogue of the Crustacea brought by Thomas Nuttall and J. K. Townsend, from the west coast of North America and the Sandwich Islands. Acad. Nat. Sci. Phila. 8(1) : 106-147. 66 Contributions in Science No. 250 Rathbun, M. J. 1900. Synopsis of North American invertebrates, VII, the Cyclo- metopous or Cancroid crabs of North America. Amer. Naturalist, 34:131-143. . 1926. The fossil stalk-eyed Crustacea of the Pacific slope of North Ameri- ca. U.S.Nat. Mus. Bull. 138. Rheinhart, P. W. 1937. Three new species of the pelecypod family Arcidae from the Pliocene of California. J. Paleontol. 11(3): 181-185. . 1943. Mesozoic and Cenozoic Arcidae from the Pacific slope of North America. Geol. Soc. Amer. Special Papers No. 47. Roding, P. F. 1798. Museum Boltenianum. Hamburg (J. C. Trappii) pp. i-vii + 109. Schenck, H. G. 1943. Acila princeps, a new Upper Cretaceous pelecypod from California. J. Paleontol. 17(l):60-68. Schlechtendal, D. H. R. 1894, Beitrage zur kenntnis fossiler insekten aus dem braunkohlengebirge von Rott am Siebnegebirge. Abhl. Ges. Halle 20:197-228. . 1898. Fine fossil Naucoris-Art von Rott. Zeitschr. Naturw. 71:417-425. Schlotheim, E. F. 1816. Beitrage zur naturgeschichte der Verteinerungen in Geog- nonstischer Hinsicht. Acad, der Wissenchaft zu Miinchen, Denkschriften, Mathematische-physikalische Klasse, 6:27. Sleeper, E. L. 1968. A new fossil weevil from Nevada (Coleoptera: Curculionidae). Bull. So. Calif. Acad. Sci. 67(3): 196-198. Smith, C. T. 1945. The biostratigraphy of Glycymeris veatchii in California. J, Paleontol. 19(l):35-44. Stanton, R. J. 1966. Megafauna of the Upper Miocene, Castaic Formation, Los Angeles County, California. J. Paleontol. 40(l):21-40. Stanton, T, W. 1896. The faunal relations of the Eocene and Upper Cretaceous on the Pacific coast. U.S. Geol. Surv. Ann. Rept. 17, vol. 1, pp. 1005-1060. Statz, G. 1930. Drei neue Insektenarten aus dem Teriar von Rott am Siebengebirge. Wissenschafflichen Mitteilungen des Vereins fiir Natur- und Heimatkunde in Kohna. Rh. 1(2): 10-14. . 1931. Eine neue Bienenart aus Rott am Siebengebirge. Wissenchnfflichen Mitteilungen des Vereins fiir Natur- und Heimatkunde in Koln a. Rh. 1(2):39- 60. . 1934a. Neue Beobachtungen iiber fossile Bienen aus dem Tertiar von Rott am Siebengebirge. Archiv fiir Bienenkunde 15(1): 1-10. 10 fig. . 1934b. Neue Tipulidenfunde aus den Braunkohlenschiefern von Rott am Siebengebirge. Wissenscafflichen Mitteilungen des Vereins fur Natur- und Heimatkunde in Koln a. Rh. 1(3):90-106, 13 figs. . 1936a. U^ber neue Funde von Neuropteren, Panorpaten und Trichopteren aus den Tertiaren Schiefern von Rott am Siebengebirge. Decheniana 93:208- 255, 28 figs. . 1936b. Ueber alte und neue fossile Hymenopterenfunde aus dem Ter- tiaren Ablagerungen von Rott am Siebengebirge. Decheniana 93:256-304, 5 pis., 15 figs. . 1937. Uber fossile niedere Wassertiere aus dem Tertiar von Rott am Siebengebirge. Die Natur am Niederrheim 13(1): 1-16, 23 figs. . 1938a. Die Tertiarschichten von Rott und ihre Wasserfauna. Heimatblatter des Siegkreises 14(2): 18-30, figs. 1-12. . 1938b. Die Tertiarschichten von Rott und ihre Wasserfauna. Heimatblatter des Siegkreises 14(3): 56-68, figs. 13-32. . 1938c. Fiinf neue fossile Cerambyciden-Arten aus den Mitteloligocanen Ablagerungen von Rott am Siebengebirge. Entomoligische Blatter 34(4): 173- 179, 1 pi. 1973 Additional Type Specimens 67 . 1938d. Neue Funde parasitischer hymenoptern aus dem Tertiar von Rott am Siebengebirge. Decheniana Abt. A, 98(1) :71-144, 9 pis. . 1939. Geradfliiger und Wasserkafer der Oligocanen Abla^erungen von Rott. Decheniana Abt. A, 99:1-102, 20 pis. . 1940a. Rheinifche Termiten! Rheindifcher Naturfreund 1:1-12, figs. 1-11. . 1940b. Neue Dipteren (Brachycera et Cyclorhapha) aus dem Oberoligazan von Rott. Palaeontographica Abt. A, 91 : 120-174, 9 pis., 1 fig. . 1941a. Bienen, Blatter un Bliiten aus dem Rheinischen Braunkohlenwald. Rheinische Heimatpflege 13:100-112, 14 figs. . 1941b. Plagegifter aus dem rheinifchen Braunkohlertwalde. Rheinifcher Naturfreund 5(1) : 1-16, 19 figs. . 1941c. Termiten aus dem rheinischen Braunkohlenwald. Die Umschau 1941(45):7-11, 10 figs. . 1942a. Versteinertes Liebesleben im Reiche der Insekten. Die Umschau 1942(32):n-4}, 8 figs. . 1942b. Bienen, Blatter und Bliiten aus dem Rheinischen Braunkohlenwald. Dtsch. Imkerfuher 16(1-2): 1-6, 22 figs. . 1943. Neue Dipteren (Nematocera) aus dem Oberoligocan von Rott. I. Familie: Bibionidae (Haarmucken). Palaeontographica Abt. A, 95:1-65, 12 pis. . 1944a. Neue Dipteren (Nematocera) aus dem Oberoligozan von Rott. IF Familie: Fungivoridae (Pilzmiicken) . Palaeontographica Abt. A, 95:67-92, 7 pis. . 1944b. Neue Dipteren (Nematocera) aus dem Oberoligozan von Rott. III. Familie: Limnobiidae (Stelzmiicken). IV. Familie: Tipulidae (Schnaken). “ V. Familie: Culicidae (Stechmiicken). Palaeontographica Abt. A, 95:93-120, 5 pis. . 1944c. Neue Dipteren (Nematocera) aus dem Oberoligozan von Rott. VL Familie: Tendipedidae (Zuck-oder Schwarmiicken). VIL Familie: Helei- dae (Gnitzen). VIII. Familie: Lycoriidae (Trauermucken). Palaeontographica Abt. A, 95:123-188, 17 pis. . 1950a. Cicadariae (Zikaden) aus den Oberoligocanen Ablagerungen >/on Rott. Palaeontographica Abt. A, 98:1-46. . 1950b. Alte und neu Hydrocorisae (Wasserwanzen) aus dem Oberoligocan von Rott. Palaeontographica Abt. A, 98:47-96. . 1952. Fossile Mordellidae und Lamellicornia (Coieoptera) aus dem Oberoligozan von Rott. Palaeontographica Abt. A, 102: 1-18. Statz, G. and A. Horion. 1937, Ein Fossiler Ptiliidenfund aus den mitteloligocanen Ablagerungen von Rott am Siebengebirge. Entomoligische Blatten 33(1):8-10. Statz, G. and E. Wagner. 1950. Geocorisae (Landwanzen) aus dem Oberloigocanen Ablagerungen von Rott. Palaeontographica, Abt. A, 98:97-136. Stearns, R. E, C. 1890. Descriptions of new west American land, freshwater, and marine shells, with notes and comments. Proc. U.S. Nat. Mus. 13:205-225. Stehli, F. G. 1955. A new Devonian terebratuloid brachiopod with preserved color pattern. J. Paleontol. 29(5) : 868-870. . 1956. Dielasma and its external homeomorph Beecheria. J. Paleontol. 30(2):299-302. . 1961a. New terebratuloid genera from Australia, J. Paleontol. 35(3) :451- 456. . 1961b. New genera of Upper Paleozoic terebratuloids. J. Paleontol. 35(3):- 457-466. 68 Contributions in Science No. 250 Waring, C. A. 1917. Stratigraphic and faunal relations of the Martinez to the Chico and Tejon of southern California. Proc. Calif. Acad. Sci. 4th Series, 7(4):41-124. Watson, L. R. 1933. As I see the old bee. Amer. Bee J. 73(2):48-49. White, C. A. 1885. On new Cretaceous fossils from California Bull. U.S. Geol. Surv. No. 22, p. 65-84. Wiedey, L. W. 1928. Notes on the Vaqueros and Temblor Formations of the California Miocene with descriptions of new species. Trans. San Diego Soc. Nat. Hist. 5(10):95-182. Willett, G. 1937. Additions to knowledge of the fossil invertebrate fauna of California. Bull. So. Calif. Acad. Sci. 36(2):61-64. Wilson, E. C. 1970. Conocardium langenheimi sp. n. (Mollusca: Bivalvia) in the Lower Permian Series of the McCloud Limestone, northern California. Los Angeles Co. Mus,, Contrib. Sci. 184:1-14. Wilson, E. C. and D. E. Bing. 1970. Type specimens of fossil invertebrata in the Los Angeles County Museum of Natural History, exclusive of paleoentomology. Los Angeles Co. Mus., Contrib. Sci. 181 : 1-20. Woodring, W. P. 1930. Upper Eocene orbitoid Foraminifera from the western Santa Ynez range, California, and their stratigraphic significance. Trans. San Diego Soc. Nat. Hist. 6(4) : 145-170. . 1931. A Miocene Haliotis from southern California. Paleontol. 5(1): 34-39. . 1942. Marine Miocene mollusks from Cajon Pass, California. J. Paleontol. 16(l):78-83. Woodring, W. P. and M. N. Bramlette. 1950. Geology and Paleontology of the Santa Maria District, California. Geol. Surv. Prof. Paper 222, pp. 1-185 -t- iv. Woodring, W. P., M. N. Bramlette, and W. S. Kew. 1946. Geology and paleon- tology of Palos Verdes Hills, California. U.S. Geol. Surv. Prof. Paper 207, pp. 1-112. Woodring, W. P., R. Stewart, and R. W. Richards. 1940. Geology of the Kettleman Hills oil field, California. U.S. Geol. Surv. Prof. Paper 195, pp. 1-170 + v. 1973 Additional Type Specimens 69 Species abbreviatipennis, Culicoides, p, 37 aberrans, Deltocephalus, p. 28 aberrans, Leia, p. 42 abdominalis caseyi, Coniontis, p. 53 abdominalis, Culicoides, p. 37 abdominalis fragmans, Coniontis, p. 53 abdominalis, Gyponites, p. 29 abdominalis labreae, Coniontis, p. 54 abdominalis, Pteromalus, p. 59 aculeatus, Palaeoblacus, p, 56 adustus, Termes, p. 21 aeneus, Syrphus, p. 46 aerosa. Bibio, p. 32 aerosus, Pteromalus, p. 59 aertsi, Priocnemis, p. 58 aestimata, Penthetria, p. 33 aethiops, Termes, p. 21 albuminosa, Marginella, p. 13 alpha, Coniontis tristis, p. 54 alpha, Nicrophorus investigator, p. 53 amabilis, Sciapus, p. 40 amicula, Lycoria, p. 43 ampliventris, Thamnotettis, p. 30 amygdala, Fletcherina, p. 4 anae, Glycymeris veatchii, p. 6 andersoni, Lyria, p. 14 andersoni mulleri, Turritella, p. 12 angulata, Calicantharus fortis, p, 13 angustipennis, Boletina, p. 41 angustipennis, Thamnotettix, p. 30 angustipennis, Xyela, p. 60 Anomalosipho, p. 14 antennaria, Lycoria, p. 44 antennatus, Calocoris, p. 24 anthracinus, Chlorops, p. 39 anthracinus, Thamnotettix, p. 30 Antiplanes, p. 14 antiqua, Argyoneta, p. 16 antiqua, Leuctra, p. 19 antiqua, Limnobia, p. 47 antiquus, Canthon simplex, p. 52 antiquus, Protarchus, p. 58 aquatilis, Oligosaldina, p. 26 arizonae, Onycholepisma, p. 18 arnoldi, Bittium, p. 1 1 ashleyi, Lirofusus, p. 13 asphalti, Coniontis tristis, p. 54 asphalti, Ranatra, p. 25 aster, Actinocyclina, p. 2 atomus, Pteromalus, p. 59 atomus, Termes, p. 21 atra, Opetia, p. 43 atra, Pterom^us, p. 59 atra. Sepsis, p, 45 atra, Usia, p. 35 atratus, Culicoides, p. 37 aulica, Lycoria, p. 44 auripes. Bibio, p. 32 austera, Lycoria, p. 44 austera, Serromyia, p, 38 austerus, Bythoscopus septulus, p. 28 austerus, Culicoides, p. 37 austerus, Cymbidodyta, p. 51 austerus, Raglius, p. 24 badia, Notonecta (Paranecta), p. 25 barbarensis woodfordi, Mitromorpha, p. 14 bauckhorni, Leria, p. 41 bauckhorni, Ulmeriella, p. 20 Beecheria^^p. 4 bella, Lycoria, p.*44 bellarugosa, Inoperna, p. 6 beringianus, Panomya, p. 9 bessomi, Ranatra, p, 25 beta, Lima, p, 6 bi angulata, Psammotreta, p. 9 Bibio, p. 32 bicolor, Culicoides, p. 37 bicolora, Lycoria, p. 44 bicolorata, Sepsis, p. 45 bifurcatus, Brachidontes, p. 6 bilineata, Vespa, p. 60 bischoffi, Trigon^ys, p. 60 blanda. Drosophila, p. 40 blandus, Temnostethus, p. 22 blandus, Termes, p. 21 blissi, Coniontis, p. 54 bonneri, Onychothelyphonus, p. 16 bouata, Tipula, p. 48 bouatia, Velia, p. 23 boulderana, Acteon, p. 15 branneri. Cancer, p. 17 brevis. Turcica imperialis, p. 1 1 broili, Timorina, p. 4 bruesii, Cantharoctonus, p, 56 brunnescens, Atrichopogon, p. 35 brunnescens, Boletina, p. 41 calico, Parastylotermes, p. 2 1 calico, Protochlorotettix, p. 27 capitata, Tettigoniella, p. 29 capitatus, Berosus, p. 51 Contributions in Science 70 No. 250 capitatus, Scelionites, p. 59 carinata, Liotia, p. 10 carpenteri, Jaton, p. 13 carrizoensis, Anadara, p. 5 caseyi, Coniontis abdominalis, p. 53 cellolusa, Celithemis, p. 19 Ceratopogon, p. 36 Cercopis, p. 27 ceres, Libellula, p. 19 chacei, Moniliopsis, p. 14 Cixius, p. 28 clairvillei lynni, Elaphrus, p. 49 clavicornis, Encyrtus, p. 57 clavicornis, Pteromalus, p. 59 clypealis, Thamnotettix, p. 30 Coccinella, p. 49 colorata, Serromyia, p, 38 comosella, Bibio, p. 32 compta. Bibio, p. 32 compta, Lycoria, p. 44 comstocki, Protohepialus, p. 3 1 concentrica, Astarte, p. 7 concinna, Apanteles, p. 55 concinnus, Termes, p. 21 concolor, Manota, p. 42 condoni, Pecten, p. 7 conjectus, Doryctes, p. 56 contractulus, Termes, p. 22 cooperi, Turritella, p. 12 cor, Trinacria, p. 6 cordifera, Vespa, p. 60 cordovae, Donacia(Donacia), p. 49 cordovae, Draycotia, p. 27 Coreide, p. 22 Corixa, p. 22 corrugatus, Clisocolus, p. 8 cowperi, Pecten, p. 7 cras^icapitatus, Blacus, p. 55 crassicapitatus, Pteromalus, p. 59 crassicornis, Phygadeuon, p. 58 crassiplicatus, Hinnites multirugosus, p.7 cristobalensis, Pecten, p. 7 cryptocephala, Lycoria, p. 44 cryptolineatus, Oreodites, p. 50 Culicoides, p. 36 curvitibialis, Orthopelma, p. 58 Cyclostigmata, Pimpla, p. 58 cynbaeformis, Gilledia, p. 4 davidae, Bembidion (Ochthedromus), p. 48 decoratus, Helmis, p. 50 decoratus, Raglius, p. 24 defossus, Pteromalus, p. 59 delta, Isocardia, p. 8 demessa, Acila, p. 5 Dentalium, p. 15 depressa, Psammochares, p. 58 Dielasma, p. 3 Dinidorine, p. 26 discus, Periploma, p. 10 dormescens, Oscinella, p. 39 dorminans, Pelopia, p. 38 dormitans, Synapis, p. 55 Dosinia, p. 9 draycoti, Donacia (Donacia), p. 49 Dytiscites, p. 50 ebeninus. Hydrous, p. 5 1 elegans, Drosophila, p. 40 elegantula, Phryganea, p. 3 1 elongatulus, Culicoides, p. 37 elegantulus, Eulophus, p. 57 elisabethae, Myiolepta, p. 46 elongata, Nyctibora, p. 19 elongatum, Dielasma, p. 3 enigmatica, Pararotifera, p. 3 enslini, Hoplocampa, p. 60 Eristalis, p. 46 Eulalia, p. 45 everestae, Bembidion, p. 48 everestae, Onthophagus, p. 52 excitatus, Paracymus, p, 5 1 excellenta, Neuronia, p, 3 1 exhumata, Leia, p. 42 exigua, Plecia, p. 34 exilis, Lycoria, p. 44 expositia, Plecia, p. 34 femoralis, Batrachomorphus, p. 28 femoral is, Platygasterites, p. 58 femuralis, Lycoria, p. 44 fenestrata, Iselica, p. 14 fernandoensis, Astrodapsis, p. 16 festivus, Helorus, p. 57 festivus, Rhogas, p. 56 filicornis, Pentapleura, p. 56 fisheri, Onychomachilis, p. 18 fortjs angulata, Calicantharus, p. 13 fortivenosa, Paropia, p. 29 fossilis, Daphinia, p. 17 fossilis, Daphnia, p. 17 fossilis, Mycomyia, p. 42 fosteri, Exaeretoptera, p. 19 foveatus, Elaphrus ruscarius, p. 49 fragmans, Coniontis abdominalis, p. 53 1973 Additional Type Specimens 71 fraternus, Hydrophilus, p. 5 1 freya, Turrit ell a, p. 12 friesei, Xylocopa, p. 55 fulvescens, Rhamphomyia, p. 41 fulvescens, Tabanus, p. 46 galeana, Mitromorpha, p. 14 gamagei, Mioaphelenchus, p. 3 gamma, Crassatella, p. 8 gastonensis, Tivela, p. 9 Geotrupes, p. 52 Gerris, p. 23 gibroni, Miochlorotettix, p. 27 gigantea, Glycymeris, p. 5 glabellus, Ectobius, p. 19 gladiator, Hecabolus, p. 56 goeckei, Idiocerus, p. 29 goetghebueir, Stilibezzia, p. 38 gracilenta, Lycoria, p. 44 gracilentus, Thamnotettix, p, 30 gracilior, Culicoides, p. 37 gracilior, Lycoria, p. 44 gracilior, Oncopygius, p. 40 gracilis, Panorpa, p, 30 gracilis, Tenthredo, p. 60 graciosa, Boletina, p. 42 graciosa, Pleroneura, p. 60 gradata, Astraea, p. 1 1 grandaeva, Eotermes, p. 2 1 grandaeva, Plecia, p. 34 granti, Crassatella, p. 8 grandis, Lestes, p. 18 grandis, Pandora, p. 10 grossa, Plecia, p. 34 guttulus labreae, Nicrophorus, p. 53 guttulus punctostriatus, Nicrophorus, P. 53 hadleyi, Thelyphonus, p. 16 hahnei, Ceruchites, p. 5 1 hakei, Pecten, p. 7 hannibali, Dendrophyllia, p, 3 harfordus, Pecten, p. 7 hastata, Polyxena, p. 43 haupti, Oligogypona, p. 29 heerii, Hilara, p. 41 helvolus, Aphodius, p. 52 henshawi, Synapis, p. 55 Hermione, p. 46 Heterocordylus, p. 25 Heterogasterine, p. 24 heydeni. Soevenia, p. 26 hirsuta, Hemiteles, p. 57 hirtipes, Pteromalus, p. 59 hochi, Agabus, p. 50 hodgei, Chlamys, p. 7 holmgreni, Termes, p. 22 horioni, Limnobia, p. 47 howardae, Protochrysomyia, p. 35 hypogaea, Plecia, p. 34 Idiocerus, p. 29 ignobilis. Sepsis, p. 45 imperialis brevis. Turcica, p. 1 1 imperialis, Plecia, p. 34 indata, Mordella, p. 5 1 inezana, Turritella, p. 12 inf era, Turritella uvasana, p. 12 insignicornis, Diapriites, p. 56 interrupta, Coniontis pectoralis, p. 54 interstitialis, Tanycarypa, p. 56 inversa, Fletcherina, p. 4 investigator alpha, Nicorphorus, p.53 investigator latifrons, Nicrophorus, P. 53 iota, Turritella, p. 12 jondani. Cancer, p. 17 jubata, Notonecta, p. 25 jucunda, Lycoria, p. 44 jucundus, Culicoides, p. 37 juliae, Miomesamia, p. 27 juliae, Palaeosminthurus, p. 17 kamuy, Ainoceras, p. 15 kanakoffi, Kelletia, p. l3 kanakoffi, Serica, p. 53 kaschei, Synapis, p. 55 kastenholzi, Epistrophe, p. 46 kathrynae, Rhyncolus, p. 50 kirkbyae, Sobobapteron, p. 18 kirkbyi, Miochlorotettix, p. 27 kuhni, Mycomyia, p. 42 labreae, Apsena, p. 53 labreae, Coniontis abdominalis, p. 54 labreae, Nicrophorus guttulus, p. 53 labreae, Palaeocopris, p. 52 langenheimi, Conocardium, p. 4 lapidaria nigra, Plecia, p. 34 lapidaria, Plecia, p. 34 lasti, Parajulus, p. 17 laticornis, Lycoria, p. 44 latifrons, Alysia, p. 55 latifrons, Nicrophorus investigator, p. 53 latigula, Coniontis tristis, p, 54 latipennata, Rhamphomyia, p. 41 latipennatus, Pteromalus, p. 59 72 Contributions in Science No. 250 latipennis, Xyela, p. 60 latisternum, Calcoschizomus, p. 16 laurae, Recticulitermes, p. 21 lengersdorfi, Lycoria, p. 44 lepida, Dziedzickia, p. 42 lepisma, Acmaea, p. 10 Lestes, p. 18 lewisii, Gyrineum, p. 13 Libellulidae, p. 19 liliputanus, Culicoides, p. 37 Limnophila, p. 47 lithophila, Phryganea, p. 3 1 Iloydi, Protosegestes, p. 20 lomaensis, Haliotis, p, 10 longa, Lycoria, p. 45 longicornis, Aspicolpus, p. 55 longicornis, Boletina, p. 42 ‘ longicornis, Metorus, p. 56 longicornis, Pteromalus, p. 59 longicornis, Spathius, p. 56 longipennis, Bezzia, p. 36 longipennis, Leptura, p. 49 longipennis, Pentapleura, p. 56 longipoda, Phthinia, p. 43 iongulus, Doryctes, p. 56 luctuosa, Plecia, p. 34 lugens, Lycoria, p. 45 lunatus, Bythoscopus, p. 28 Lycoria, p. 43 Lygaeide, p. 23 lygaeus, Sehirus, p. 23 lynni, Donacia (Donacia), p, 49 lynni, Elaphrus clairvillei, p, 49 M acorn a, p. 8 macrophthalmus, Apanteles, p. 55 macroradialis, Spathius, p. 56 macroschisma, Pododesmus, p. 6 maculatus, Hydroporus, p, 50 maculatuosa, Panorpa, p. 30 maculipennis, Deltocephalus, p. 28 maculosa, Maclarenella, p. 3 maculosus, Deltocephalus, p, 28 madens, Crocistethus, p. 23 magna, Lirularia, p. 1 1 magna, Xyela, p. 61 magnificus, Anaesthetis, p. 49 magnoculus rousei, Gnathamitermes, p. 22 major, Glycymeris veatchii, p. 6 margaritaceus, Pantoclis, p. 57 martinii, Mansonia, p. 39 matilijaensis, Astropecten, p. 15 maxima, Tipula, p. 48 mckittricki, Nicrophorus, p. 53 mckittricki, Parasida, p. 54 meganosensis, Glycymeris, p. 5 melanopa, Lycoria, p. 45 Melolontha, p. 52 meuniere, Phryganea, p. 3 1 michalki, Rhinocoris, p. 26 mmimus, Diaprites, p. 57 miocaenica, Trichocera, p. 48 miocenae, Gibronea, p. 3 miocenae, Rhyssematus, p. 50 minuscula, Lycoria, p. 45 minutus, Dolichopus, p. 40 montereyana, Anadara, p. 5 moorei, Cyprimeria, p. 9 morleyi, Cryptus, p. 57 morticinus, Eerosus, p, 51 mortuum, Anthidium, p. 58 mulleri, Onthophagus, p. 52 mulleri, Turritella andersoni, p. 12 multipunctatus, Hydroporus, p. 50 multirugosus crassiplicatus, Hinnites, p.7 munda, Gonomyia, p. 47 mundus, Eulophus, p. 57 Mycomyia, p. 42 neptunus, Hydrophilus, p. 51 Nerita, p, 10 nervosus, Deltocephalus, p. 28 nigra, Penthetria, p. 33 nigra, Plecia lapidaria, p. 34 nigrapilosa, Mordella, p. 5 1 nitra, Vespa, p. 60 nuculiformis, Crassinella, p. 8 nuttalli, Sanguinolaria, p. 8 obesus, Culicoides, p. 37 obliquus, Stenomacrus, p. 58 obscura, Pelopia, p. 38 obscura, Sphex, p. 60 obscuratus, Culicoides, p. 37 obtusiscutellum, Nicrophorus, p. 53 ocalana, Operculina, p. 2 ocoyana. Ficus, p. 12 ocoyana topangaensis, Turritella, p. 12 oligocaenica, Aethalion, p, 27 oligocaenica, Leucempria, p. 60 oligocaenica, Lygus, p. 25 oligocaenica, Rhabdomastix, p. 48 oligocaenicus, Oncopygius, p. 40 onychis, Onychocampodea, p. 17 onychis, Parajulus, p. 17 orcutti, Coralliochama, p. 4 1973 Additional Type Specimens 73 oregonensis, Psephaea, p. 14 Orthocladius, p. 38 ossa, Turritella, p. 12 ovalipennata, Lycoria, p. 45 ovalis, Oligopenthimia, p. 29 packardi, Ampullina, p, 12 pagasti, Erioptera, p. 47 pagasti, Pelopia, p. 38 palaea, Haliotis, p. 10 palaemon, Pelopia, p. 39 paludosus, Sehirus, p. 23 papyracea, Nicropsalis, p. 17 paraelliptica, Coniontis pectoralis, p. 54 paucicostata, Homalopoma, p. 1 1 parva, Venercardia, p. 8 parvulus, Campoplex, p. 57 pectoralis interrupta, Coniontis, p. 54 pectoralis, Macropsis, p. 29 pectoralis paraelliptica, Coniontis, p. 54 pedicularis, Spathius, p. 56 pelidua, Dryomyza, p. 41 pelidua, Lycoria, p. 45 pelidua, Opomyza, p, 43 Pelopia, p. 38 Pelopiina, p. 39 pennata, Lycoria, p. 45 pennata, Plecia, p, 34 pennatus, Deltocephalus, p. 29 pentadiscalia, Stratiomyia, p. 46 Pentatomine, p. 26 Penthetria, p. 33 percarus, Pecten, p. 7 perlaminosa, Mercenaria, p. 9 petreficata, Clusiodes, p. 39 petrorum, Saperda, p. 49 petrothauma, Astraea, p. 1 1 peusi, Tipula, p. 48 philippi, Plecia, p. 34 Phryganea, p. 31 picata, Chilosa, p. 46 picatus, Cydnus, p. 23 pinguis, Plecia, p. 35 pilosa, Docosia, p. 42 pilosella, Ditomyia, p. 42 Pitar, p. 9 planus, Figites, p. 57 plateosus, Euxcelis, p. 29 Plecia, p. 33 praeclara, Nothochrysa, p. 30 praticolus vetustus, Canthon, p. 52 primaeva, Notonecta, p. 26 primitivus, Plioprojapyx, p. 18 princeps, Crepidula, p. 1 1 pristinus, Copris, p, 52 problem aticum, Fungivorites, p. 42 Procladius, p. 39 productus. Cancer, p. 17 pronota, Gyponites, p. 29 pronotumalis, Oligoidiocerus, p. 29 proserpina, Plecia, p. 35 pseudoillota, Barbatia, p. 5 psila, Discocylina, p. 2 pterotrichia, Limnophila, p. 47 pulchellus, Psilomma, p. 58 pulchellus, Pteromalus, p. 59 pulchellus, Raglius, p. 24 punctata, Scopeuma, p. 43 punctatus, Drymus, p. 24 punctiventris, Syromastes, p. 22 punctostriatus, Nicrophorus guttulus, p. 53 pusillus, Pionosomus, p. 24 pygmaea, Asindulun, p. 41 pygmaeolus, Pteromalus, p. 59 rectispinus, Pteromalus, p, 59 reddingensis, Glycymeris veatchii, p. 6 regina, Calva, p. 9 reisingeri, Mycomyia, p; 42 remnans, Coniontis, p. 54 rhenana, Anthomyza, p. 32 rhenana, Blaberites, p. 19 rhenana, Corixa, p. 23 rhenana, Delphax, p. 28 rhenana, Oligosaldina, p, 26 rhenana, Plecia, p. 35 rhenana, Polyxena (Cordyla), p. 43 rhenanus, Calotermes, p, 2 1 riguus, Trapezonotus, p, 25 robusta, Limnophila, p. 47 robustus, Bythoscopus, p. 28 robustus, Deltocephalus, p. 29 rohli. Bibio, p, 32 rottensis. Anopheles, p. 39 rottensis. Bibio, p. 32 rottensis, Calotermes, p. 20 rottensis, Clinocentrus, p. 56 rottensis, Deltocephalus, p. 29 rottensis, Dicranoptycha, p. 47 rottensis, Ectrepesthoneura, p. 42 rottensis, Geotrupes, p, 52 rottensis, riyalesthes, p. 28 rottensis. Hydrous, p. 5 1 rottensis, Ischnodemus, p. 24 74 Contributions in Science No. 250 rottensis, Liogryllus, p. 19 rottensis, Naucoris, p. 25 rottensis, Nysson, p. 60 rottensis, Oligosaldina, p. 26 rottensis, Ornithoponus, p. 41 rottensis, Panorpa, p. 3 1 rottensis, Penthetria, p. 33 rottensis, Poecilocoris, p. 27 rottensis, Pteromalus, p. 59 rottensis, Rhamphomyia, p. 4 1 rottensis, Sciapus, p. 40 rottensis, Tetrops, p. 49 rottensis, Trepidaria, p. 48 rottensis, Ula, p. 48 rottensis, Ulmeriella, p. 20 rotundatus, Doryctes, p. 56 rotundatus, Figites, p. 57 rousei, Gnathamitermes magnoculus, p. 22 rousei, Miotroctes, p. 19 ruscarius foveatus, Elaphrus, p. 49 ryshkoffi, Cryptotermes, p. 21 sack!, Temnostoma, p. 46 santacruzana, Turcicula, p. 1 1 santamariensis. Area, p. 5 satyrus, Plecia, p. 35 scarificata, Tivela, p. 9 schmidti, Onychojapyx, p. 18 scita, Penthetria, p. 33 scitus, Dolichopus, p. 40 Scuteilerine, p. 27 seppenradensis, Pachydiscus, p. 15 septemmontanus, Dendroleon, p. 30 sepultus austerus, Bythoscopus, p. 28 septultus, Bythoscopus, p. 28 Serromyia, p. 37 setaria, Norellisoma, p. 43 seymouri, Salticus, p. 16 shamanensis, Thais, p. 14 signata, Polistes, p. 60 simplex antiquus, Canthon, p. 52 sisquocensis. Area, p. 5 Sophia, Haplocnemia, p. 49 spadicea. Bibio, p. 32 spicata, Oliva, p. 13 Spilomya, p. 46 spinifera, Empis, p. 41 spiniger, Figites, p. 57 spinitibialis, Deltocephalus, p. 29 spinitibialis, Maladera, p. 52 spinitibialis, Sehirus, p. 23 spinofemorata, Serromyia, p. 38 spinosa, Platygasterites, p. 58 spinosus, Dolichopus, p. 40 Spondylus, p. 7 Spongostylum, p. 35 stantoni, Macrocallista, p. 9 statzi, Polysphincta, p. 58 statzi, Scolopostethus, p. 24 stearnsii, Pecten, p. 7 stewarti, Turritella uvasana, p. 13 stigmalineata, Aspilota, p. 55 stigmaticus, Chrysopiius, p. 43 stocki, Aphelophlebodes, p. 18 stocki, Helminthoglypta, p. 15 Stratiomya, p. 46 striatus, Trapezonotus, p. 25 strigatus, Palaeogyrinus, p. 50 sturmi, Plecia, p. 35 stygia, Plecia, p. 35 subgibbosa, Eucrassatella, p. 8 submersa, Phryganea, p. 31 subobsoleta, Leptothyra, p. 1 1 tarsalis, Paraclia, p. 47 tenellus, Epyris, p. 55 tenelius, Xanthochlorus, p. 40 tenera, Helius, p. 47 tenuipennis, Culicoides, p. 37 tenuis, Calcibacunculus, p. 20 tenuis, Dactylolabis, p. 47 terebrator, Microcryptus, p, 57 tertiaria, Phalacrotophora, p. 43 tertiarium, Ptilium, p. 52 Tettigoniidarum, p. 20 texana, Lowenstamia, p, 4 Thamnotettix, p. 29 thienemanni, Pelopia, p. 39 thoracica, Lycoria, p. 45 Thrips, p. 30 tibialis, Pyrrhocoris, p. 26 Tipula, p. 48 topangaensis, Turritella ocoyana, p. 12 trapezoides, Thracia, p. 10 tridesmia, Clathurella, p. 14 tristis alpha, Coniontis, p. 54 tristis asphalti, Coniontis, p. 54 tristis latigula, Coniontis, p. 54 trochiformis, Trochita, p. 1 1 tropica, Penthetria, p. 33 Tropidia, p. 46 umbonata, Empis, p. 41 umbonata, Limnophila, p. 47 umbonata, Macrocera, p. 42 umbonata, Mycomyia, p. 42 1973 Additional Type Specimens 75 unicolor, Mycomyia, p. 42 univestis, Chilacis, p. 24 ustus, Thammotettix, p. 30 uvasana infera, Turritella, p. 12 uvasana stewarti, Turritella, p. 13 vancouverensis. Area, p. 5 Vancouver!, Notiophilus, p. 49 varivestita, Mansonia, p. 40 veatchii anae, Glycymeris, p. 6 veatchii, Glycymeris, p. 6 veatchii major, Glycymeris, p. 6 veatchii reddingensis, Glycymeris, p. 6 ventricosa, Venericardia, p. 8 venturana, Henricia, p. 16 venusta, Opomyza, p. 43 venustus, Oncopygius, p. 40 venustus, Pteromalus, p. 59 veterana, Limnophila, p. 48 veterana, Plecia, p. 35 veterana, Stilobezzia, p. 38 vetustus, Canthon praticolus, p. 52 vidua, Corixa, p. 23 vulpina. Bibio, p. 32 vulpina, Plecia, p. 35 wagneri, Lygaeosoma, p. 24 washingtonianus, Viviparus, p. 13 weigandi, Elephantomyia, p. 47 weissmanae, Phlepsius, p. 27 wettweri, Proicnemis, p. 58 winnertzi, Plecia, p. 35 weylandi, Lycoria, p. 45 woodfordi, Mitromorpha barbarensis, p. 14 zeta, Flaventia, p. 9 Accepted for publication July 24, 1972 Ill I I I X* ■s I I I i NUMBER 251 NOVEMBER 9, 1973 0'7 f 73 2 i,5// SYSTEMATICS OF THE MARMORATUS GROUP OF THE FROG GENUS LEPTODACTYLUS (AMPHIBIA, LEPTODACTYLIDAE) By W. Ronald Heyer CONTRIBUTIONS IN SCI6NCE NATURAL HISTORY MUSEU M • LOS ANGELES COUNTY CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Natural History Museum of Los Angeles County. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientific institutions and scien- tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries should be directed to Virginia D. Miller, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS 1 Manuscripts for CONTRIBUTIONS IN SCIENCE may be in any field of Life | or Earth Sciences. Acceptance of papers will be determined by the amount and char- i acter of new information. Although priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the collections of the Muse- ^ um, other technical papers will be considered. All manuscripts must be recommend- ' ed for consideration by the curator in charge of the proper section or by the editorial board. Manuscripts must conform to those specifications listed below and will be ex- ' amined for suitability by an Editorial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indi- i cate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.— (1) The 1972 CBE Style Manual, third edition i ( AIBS) is to be followed in preparation of copy. (2) Double space entire manuscript. : (3) Footnotes should be avoided if possible. Acknowledgments as footnotes will not , ^ be accepted. (4) Place all tables on separate pages. (5) Figure legends and unavoid- v | able footnotes must be typed on separate sheets. Several of one kind may be placed on > i a sheet. (6) An abstract must be included for all papers. This will be published at the head of each paper. (7) A Spanish summary is required for all manuscripts dealing . with Latin American subjects. Summaries in other languages are not required but are ^ strongly recommended. Summaries will be published at the end of the paper. (8) A | ■ diagnosis must accompany any newly proposed taxon. (9) Submit two copies of Isi manuscript. I * ILLUSTRATIONS. — All illustrations, including maps and photographs, will be » referred to as figures. All illustrations should be of sufficient clarity and in the proper ^ ! proportions for reduction to CONTRIBUTIONS page size. Consult the 1972 CBE * Style Manual, third edition (AIBS) in preparing illustration and legend copy for J I style. Submit only illustrations made with permanent ink and glossy photographic | prints of good contrast. Original illustrations and art work will be returned after the | manuscript has been published. f , PROOF. — Authors will be sent galley proof which should be corrected and ; • returned promptly. Any changes or alterations, other than typographical corrections, ; : will be billed to the author. One hundred copies of each paper will be given free to ( each author or divided equally among multiple authors. Orders for additional copies | j must be sent to the Editor at the time corrected galley proof is returned. Appropriate r ! order forms will be included with the galley proof. 1 1 Virginia D. Miller | : Editor I SYSTEMATICS OF THE MARMORATUS GROUP OF THE FROG GENUS LEPTODACTYLUS (AMPHIBIA, LEPTODACTYLIDAE)i By W. Ronald Heyer^ Abstract: The geographic distribution patterns of eight characters of size, shape, texture, and color pattern were examined for members of the Marmoratus species group of the frog genus Leptodactylus . Analysis of the geographic distribution of character states indicates that the group is com- posed of five species. Leptodactylus bokermanni, new species, has nonex- panded toe tips and occurs in the forests of southeast Brasil. Leptodactylus hylaedactylus has nonexpanded or slightly expanded toe tips and inhabits the forests of greater Amazonia and southwestern Brasil. Leptodactylus andreae has toe disks and occurs in the Orinocan and greater Amazonian forests. Leptodactylus marmoratus has toe disks and occurs in the forests of southeastern Brasil. Leptodactylus martinezi has nonexpanded toe tips and inhabits the dry forests of central Brasil. Each species is diagnosed and described, and a key is provided. It is hypothesized that the foam nest in the Marmoratus group and that of the other species groups in the genus Leptodactylus evolved in response to different selective forces. In the case of the other species groups, the foam nest has evolved in reponse to arid conditions with sporadic heavy rain. In these situations, rainfall can be used in a predictive manner to ensure reproductive success. In the Marmoratus group, rainfall may not be a good predictor of reproductive success. Terrestrial organisms have a relatively stable reproductive environment in wet tropical forests, how- ever. The foam nest in the Marmoratus group apparently has developed as a response to selection for terrestriality. Introduction This paper is the second of a projected series on the systematics of the species groups of the genus Leptodactylus. The first treated the Melanonotus group (Heyer, 1970a); the present paper treats members of the Marmoratus group as previously defined by Heyer (1968). The members of this group have been poorly understood, both with respect to how many species comprise the group and to what the relation- ships of the Marmoratus group are to the other species groups within the genus. This paper is addressed to the first problem. Acknowledgments I thank the following institutions and curators for sending specimens, informa- tion, and location of obscure localities (museum abbreviations as used in text in parentheses): American Museum of Natural History, New York, Charles W. Myers and Richard G. Zweifel (AMNH); Zoologische Sammlung des Bayerischen Staates, ^Review Committee for this Contribution Robert L. Bezy William E. Duellman Jay M. Savage John W. Wright ^Research Associate, Section of Herpetology, Natural History Museum of Los Angeles County; and Biology Department, Pacific Lutheran University, Tacoma, Washington 98447. (Present address: National Museum of Natural History, Washington, D. C. 20560.) 1 2 Contributions in Science No. 251 '■ I Munich, U. Gruber and W. Hellmich; British Museum (Natural History), London, ; Alice G. C. Grandison, (BMNH); Carnegie Museum, Pittsburgh, Clarence J. McCoy, | Jr., (CM); Field Museum of Natural History, Chicago, Hymen Marx, (FMNH); Naturhistoriska Museet, Goteborg, Birgitta Hansson, (Ba); Museum of Comparative Zoology, Harvard University, Cambridge, Ernest E. Williams, (MCZ); Museum of Natural History, University of Kansas, Lawrence, William E. Duellman, (KU); Natural History Museum of Los Angeles County, Los Angeles, Robert L. Bezy and John W. Wright, (LACM); Museum of Zoology, University of Michigan, Ann Arbor, Michael C. Devine and Charles F. Walker, (UMMZ); Academy of Natural Sciences, Philadelphia, James E. Bohlke and Edmund Malnate, (ANSP); University of Puerto || Rico, Mayaguez, Juan A. Rivero, (UPR); Rijksmuseum van Natuurlijke Historie, Leiden, Marinus S. Hoogmoed, (RM); Museu de Zoologia da Universidade de Sao | Paulo, Paulo E. Vanzolini, (DZ); Natur Museum Senckenberg, Frankfurt, Robert Mertens; United States National Museum, Washington, D.C., the late James A. Peters, (GOV, JAP, USNM); Naturhistorisches Museum, Vienna, Josef Eiselt, ( Vien- i na). Werner C.A. Bokermann (WCAB) and Jean Lescure (LG) allowed me to examine specimens from their private collections. For critically reading the manuscript, I thank Avelino Barrio, Centro Nacional de Investigaciones lologicas, Buenos Aires, Werner C.A. Bokermann, Sao Paulo, and Jay M. Savage, University of Southern California. Wendy Enger, Pacific Lutheran I University, executed figures 2, 5, 7, 12, 26, and 28. The reviewers of this paper do not all concur that the methods of analysis used are the most appropriate. That decision, as well as all interpretations, remains solely mine. Support from National Science Foundation grant GB-27280 is gratefully acknow- ledged. Methods and Materials Over 1300 preserved specimens form the basis of this analysis. Early in the study, it became clear that the problem could best be treated in the manner used by Gans (1959, 1966). The analysis was done in three stages; in the first, the following were recorded: 1) Sex. Individuals were determined to be either male, female, or juvenile. Males were determined by the presence of vocal slits. Females were determined by presence of eggs visible through the ventral body wall, or by size if males were present from the same locality. If there was any question, the individual was scored as a juvenile. 2) Size. The snout-vent length was measured to the nearest 0. 1 mm with vernier calipers. 3) Snout shape. A ratio was determined which in selected samples reflects the differences between blunt and pointed snout shapes. A specimen was lined up under a grid-type ocular micrometer in a dissecting microscope so that one line traversed the distance between the anterior corners of the eyes. This distance was read and used as the denominator. The numerator was determined by reading the distance at right angles from the grid line connecting the anterior eye corners to the tip of the snout. The larger the value of the ratio, the longer the snout. 4) Toe tip shape. Four categories of toe tip shape were recognized, ranging from 1973 Systematics of Marmoratus Group 3 A B C D Figure 1. Toe tip character states, see text. state A, the tips pointed, to state D, the tips with large, distinct disks (Fig. 1). In this character only, intermediate categories were used, such as B-C or C-D. 5) Dorsal texture. Four character states were recognized: A, dorsum smooth; B, Dorsum with white tipped tubercles; C, dorsum with ridges or warts; D, dorsum with dorsolateral folds and white tipped tubercles. The method of preservation affects this character. Many of the specimens scored as having state A certainly had one of the other states in life. Because of this, the variation of dorsal texture was not analyzed in detail, but the range of variation is described in the individual species accounts in a later section. 6) Dorsal pattern. Six different character states were recognized (Fig. 2). Here, Figure 2. Dorsal pattern character states, see text. 4 Contributions in Science No. 251 as in the other characters which were broken down into arbitrary states (with the exception of toe disks), a specimen with an intermediate category was placed into the category it most resembled. 7) Dorsolateral stripes. Three states were recognized: A, no dorsolateral stripes; B, light, narrow, dorsolateral stripes from the eye to the inguinal region; C, light, broad, conspicuous dorsolateral stripes from the eye to the inguinal region. 8) Light mid-dorsal stripe. Seven states were recognized: A, no mid-dorsal stripe; B, a mid-dorsal pin stripe from above the vent to about midbody; C, a pin stripe from above the vent to the snout; D, a stripe wider above the vent, tapering to a point not beyond the sacrum; E, a broad stripe from above the vent to the tip of the snout; F, a pin stripe from above the vent not extending beyond the sacrum; G, a wide stripe in the sacral region only. The specimens were examined locality by locality. As the first stage of analysis was being done, the specimens from each locality were examined to determine if there were one or more distinctive forms; that is, specimens exhibiting consistent discon- tinuous variation in two or more characters. Localities of presumed instances of sympatry of two distinctive types were noted. Detailed comparison of the specimens from the presumed sites of sympatry formed part of the second stage of analysis. This part of the analysis, termed “the recognition of non-dimensional species” by Cans (1966), centers about study of sympatric occurrences. Distinctive specimens from the same locality are examined in detail. Samples from surrounding localities are then compared with and determined to be one or the other distinctive forms from the site of sympatry. This was done separately for each of seven large areas. The third stage of analysis, termed, “the recognition of species in space” by Gans (1966), involves detailed analyses of similar forms over the entire geographic range. Decisions are made as to how many species are represented based on this third stage of analysis. This methodology is nothing more than standardized common sense as histori- cally used in systematics. It is not a phenetic analysis in the numerical taxonomy sense, but rather a “classical” taxonomic approach. The reason for utilizing the methods outlined by Gans (1959, 1966) is that it allows one to keep track of variability when large numbers of specimens over wide geographic areas are being studied. The Recognition of “Non-Dimensional’ ’ Species Seven large areas were delineated to form the study units for the recognition of “non-dimensional” species. Each area was determined on the basis of at least one locality where two forms occurred together, proximity of other localities from which specimens originated, amount of material, and intuition. The situation for each area follows. Area A — This area consists of lands of north draining rivers. A single instance of sympatry was found (Surinam; Nickerie, Sipaliwini, Fig. 3, number 1). The forms are distinctive when closely compared (Fig. 4). The forms are designated as I and II and differ as follows. Form 1(11 specimens) is more robust than form II (4 specimens); form I has either a uniform dorsum or wide saddlelike dark blotches, form II has more or less distinct small spots dorsally; 1 specimen of form I has type C dorsolateral 1973 Systematics of Marmoratus Group 5 Figure 3. The seven study areas for recognition of “non-dimensional” species. Numbers indi- cate localities where more than one form is represented in collections. stripes, none of the form II specimens has dorsolateral stripes; the snout of form I males is sharply pointed in profile, the snout of form II males is rounded in profile; the toe tips of form I specimens are either type C or D, the toe tips of form II specimens are type B, or type B almost type C; the dorsum is smooth in all form I specimens, the dorsum in a single form II specimen is warty. Specimens from all localities within Area A were readily allocated as either form I or II using the Sipaliwini specimens as standards. Area B — This area covers the Brasilian territory and states of Amapa, Mato Grosso, Para, and Goias which are drained by the Amazon or Tocantins river systems. Instances of sympatry are at Amapa; Serra do Navio 220-300 m, Serra do Veado, 290-310 m (Fig. 3, number 2), Mazagao (Fig. 3, number 3); Para; Belem (Fig. 3, number 4), IPEAN, 3 km E Belem (Fig. 3, number 5), and Cachimbo (Fig. 3, 6 Contributions in Science Figure 4. Frogs from locality one of Fig. 3. Left — dorsal views; upper two sj?ecimens are form I individuals, lower two form II. Right — ventral views; upper two specimens are form I individu- als, lower two form II. number 6). At each site except Cachimbo, there are two forms, which are again i designated as I and II. Here and in the other major areas, designation of I forms i' indicates that the I form of Area A is most similar to the I form of Area B, but ' relationships among all the I forms from all areas constitutes the analysis of the species in space of the next section. The sympatric I forms of Area B (143 specimens) differ from the II forms of Area B (17 specimens) as follows: the body shape of form I specimens is slightly more robust than in form II specimens, but the differences are not as pronounced as in Area A; the dorsal patterns of form I specimens are more uniform, the dorsal patterns of form II specimens are more mottled with a tendency toward | spotting; the venters of form I specimens are darker than in form II specimens; some form I individuals have broad, light dorsolateral stripes, no form II individuals have such stripes; no form I specimens have B or C category light mid-dorsal stripes, while some form II individuals do; the toe tips of form I individuals are C or D category | (Fig. 1), the toe tips of form II individuals are either B or C category (Fig. 1) — in the C cases of both forms the disks are distinctive in that the C category disk of form I is flattened while the C category disk of form II is rounded (Fig. 5). At Cachimbo, two distinctive forms were recognized. The first corresponds to the II form of Area B. The second corresponds to the F dorsal pattern of Figure 2 and is designated form III. This is the only site where more than one distinctive form is known to occur in which both forms have no disk development of the toes. The specimens from all other localities from within Area B. were readily designated as being either form I, II or III. The form I specimens from Para, Jacareacanga have notably large toe disks. Area C — This area includes samples from the Brasilian state of Amazonas. In- stances of sympatry were found at Itapiranga (Fig. 3, number 7) and Ducke Reserve, near Manaus (Fig. 3, number 8). At both sites, the different forms consisted of forms I and II. The differences between form I (7 specimens) and form II 1973 Systematics of Marmoratus Group 7 Figure 5. A, C category of toe disk of form I individuals. B, C category of toe disk of form II individuals. (5 specimens) at the two sites of sympatry are; some form I individuals have light, broad dorsolateral stripes, no form II individuals have such stripes; no form I speci- mens have B or E category mid-dorsal light stripes, some form II specimens do; form I individuals have C or D category toe tips, form II individuals have B category toe tips. Individuals from all other localities within Area C were readily distinguished as either I or II forms using the specimens from localities 7 and 8 (Fig. 3) as standards with the exception of the specimens available from Manaus. These 4 particular individuals are so desiccated that proper allocation is not possible. Area D — This area includes samples from the Brasilian state and territory of Acre and Rondonia. Two sites of sympatry were identified: Rondonia, Porto Velho (Fig. 3, number 9) and Igarape Marmelo (Fig. 3, number 10). At Porto Velho, two forms were present, I (4 specimens) and II (9 specimens) which differ as follows: two form I individuals have broad, light, dorsolateral stripes, no form II individuals have such stripes; no form I individual has an E category light mid-dorsal stripe, one form II individual does; form I individuals have C or D category toe tips with flattened disks (Fig. 5A), form II individuals have category B or C toe tips, if category C, the disks are rounded (Fig. 5B). Specimens from other localities within Area D were readily identified as either form I or form II using the Porto Velho specimens as standards with the exception of the specimens from Igarape Marmelo. Igarape Marmelo is the only locality where intermediate individuals between form I and form II types were iden- tified for any of the areas. Three individuals are clearly the same as form I individuals from the other localities within Area D with respect to toe disk development. Seven individuals are clearly the same as form II individuals from other Area D localities in toe tip development and in some individuals having B, C, and E categories of mid- dorsal stripe development (Fig. 6). Two individuals are intermediate with respect to toe tip development. In addition, the dorsal patterns are complex and not easily described in the total sample, but the form I specimens tend towards uniformity, the form II specimens tend toward spotting, while the two individuals intermediate in toe disk development appear to be a combination of both form I and II patterns (Fig. 6). Area E — This area includes Amazonian localities from Colombia, Ecuador, and Peru. Two instances of sympatry were noted: Peru; Loreto, Iparia (Fig. 3, number 11) and Peru; Pasco, Iscozazin Valley, Pan de Azucar (Fig. 3, number 12). Seven form I 8 Contributions in Science No. 251 Figure 6. Specimens from Igarape Marmelo, Territorio do Rondonia, Brasil. Specimens on left of upper (dorsal view) and lower (ventral view) figures are form I individuals; specimens on right are form II individuals, specimens in middle are intermediates. 1973 Systematics of Marmoratus Group 9 individuals differ from nine form II individuals at the two localities in the following: form I individuals have broader heads than form II individuals; the dorsal surfaces of form I individuals are uniform except for a posterior continuation of the dark interorbi- tal blotch, the dorsal surfaces of form II individuals are fuzzy-spotted; no form I individuals have category B light mid-dorsal stripes, some form II individuals have the B category stripe; the ventral surfaces of the thigh and calf are darker in form I individuals, lighter in form II individuals (Fig. 7), the toe tips of form I individuals are category C, form II individuals are category B. Individuals from all other localities in Area E were readily categorized as either I or II forms using the specimens from the two sites of sympatry as standards. In this area, dorsal and particularly ventral pattern proved more useful than toe tip shape in separating the two forms. Area F — This area includes all samples from Bolivia. One instance of sympatry was noted, Bolivia; Santa Cruz, Buenavista (Fig. 3, number 13). Differences between the form I specimens (31) and form II specimens (39) are: dorsal surfaces of the form I specimens are more or less uniform in pattern, the dorsal surfaces of the form II individuals are blotched, not quite forming distinct spots; some form I individuals have broad, light dorsolateral stripes, no form II individuals have such stripes; no form I individuals have B or C category light mid-dorsal stripes, some form II individuals do; the snouts of form I males are pointed in profile, in form II males the snout profiles are rounded; form I specimens have C or D category toe tips, most form II specimens have category B toe tips, a few specimens have toe tips approaching the C category; two form I individuals have dorsolateral folds, no form II individuals have dorsolateral folds. All other samples from Area F were readily categorized as either form I or II using the Buenavista individuals as standards with the exception of a single individual from Santa Cruz, Rio Ichilo, 34 km S Boca Chapare. In this instance, determination was made on the basis of dorsal pattern as the toe tips were not clearly either B or C category. Area G— Area G covers all lands drained by rivers that empty to the east or south of Brasil and southern South America. A single instance of sympatry was found: Figure 7. Ventral patterns of frogs from area E. A, form II individual, B, form I individual. 10 Contributions in Science No. 251 Brasil; Rio de Janeiro, Teresopolis (Fig. 3, number 14). The specimens from this locality are poorly preserved, making detailed comparisons impossible. One specimen from this series clearly has the C type category toe tips while the other nine specimens clearly have the B type category; the single form I individual also has broad, light dorsolateral stripes, none of the form II individuals have such stripes. Specimens from most other localities were categorized as either I or II forms, but specimens from several localities were difficult to allocate. First of all it was evident that more than one type of II form was involved. The form II individuals from the coastal states of Guanabara, Minas Gerais, Rio de Janeiro, and Sao Paulo differ from form II individu- als from Goias and Mato Grosso. The latter specimens are more similar to the form II individuals from Amazonas and Para. None of the coastal form II individuals have a dorsal pattern of distinct spots nor do any individuals have B, C, or E category light mid-dorsal stripes. Further, some individual specimens in series from Guanabara; Rio de Janeiro, Tijuca, and Cosme have very poorly developed disks but seem to be form I specimens. The most difficult situation to interpret at this point is some of the speci- mens from Parana. The form I individuals from the southern part of the state of Sao Paulo and the state of Santa Catarina are quite small and have well-developed disks. Most of the Parana specimens are larger with no disk development at all. These same specimens have broad, light dorsolateral stripes, which throughout all the other study areas were found only in form I individuals. Because of the broad, light dorsolateral stripes, the specimens from Parana are included in with the form I specimens for further analysis in the next section. Discussion At this point it is appropriate to ask whether at each site of sympatry, the different forms represent a single species polymorphic system, or the different forms represent distinct species. To help reach this decision, presentation of the only mating call recordings is necessary. Werner Bokermann kindly sent me audiospectograms of four individuals of the Marmoratus group from recordings he took in the field (Figs. 26, 28). One audiospectogram is from a form I individual (Fig. 28), the other three were recorded from form II individuals. All recordings are based on allopatric specimens. I interpret the audiospectograms as representative of two call types, correlating with the different morphological forms. Detailed descriptions of the calls will be presented elsewhere. Levins (1968) discussed a theoretical situation involving polymorphism within a single species which could account for the situation presented here. In Levins’ terms, a species with a concave fitness set living in a coarse grained environment will be polymorphic, and the morphs might be mistaken by systematists for closely related species. Briefly, this would mean that two distinctive environments occur in such a pattern that an individual is likely to spend its entire lifetime in a patch of a single kind. The alternative environments are very different compared to individual tolerance, so the population strategy is to have two morphs, each adapted to one environmental type. In practice. Levins states that such organisms demonstrating this strategy would most likely be small organisms with reduced mobility. If frogs demonstrate the pattern 1973 Systematics of Marmoratus Group 11 proposed by Levins, then members of the Marmoratus group are likely candidates. The system described by Levins implies that the polymorphism is controlled by very few genes. At most of the sites of sympatry, two forms were recognized, I and II, which differed in dorsal ground pattern, development of dorsolateral stripes, development of mid-dorsal stripes, development of toe disks, and apparently mating calls. One would have to assume either that several groups of genes or a few pleiotropic genes are involved. It is probably most reasonable to assume that several different groups of genes are controlling the differences between form I and II individuals. This in turn suggests that form I is a species distinct from form II in each of the major study areas. Form III is very distinct from either I or II so that it probably is representative of a distinct breeding unit. The one exception to the discussion involves the specimens from Brasil: Rondonia; Igarape Marmelo. This was the only locality where inter- mediates between form I and form II individuals were recognized. Due to the distinc- tiveness of the forms at all of the other localities, I interpret the situation at Igarape Marmelo to represent hybridization between two closely related species. The results of this portion of the analysis are the recognition of three “non-dimensional” species correlating to forms I, II, and III as recognized above. The next analytic procedure will attempt to determine whether all species I populations from each major study area represent the same species and whether all species II populations from each major study area represent the same species. THE RECOGNITION OF SPECIES IN SPACE Small samples are available for species III which are rather uniform. No further analysis of variation is presented for this species. The character states for the I and II forms were plotted separately. Plotting was done on map overlays so any geographic trends could be noticed. Samples within a 150 km circle diameter area were combined in cases where no differences were noted among the samples during prior examination. For numerical data, when individual samples consisted of 10 or more individuals, the standard error was determined for that sample. For male and female sizes, the sample mean and maximum were plotted for each locality. Such trends are not necessarily statistically significant, but may indicate differences. The reasoning here is that any populational variation may be helpful in determining the status of the populations. The determination of the status of the populations depends on a value judgment of all known trends and variables. That is, the results of this analysis do not in themselves provide any answers, but they do provide data from which more intelligent inferences may be drawn. Analysis of Form II Specimens Male size — Form II males range in size from 19.4-26.2 mm S-VL (snout- vent length). There were only three samples which had 10 or more males: These are from French Guiana, Bolivia, and the state of Sao Paulo, Brasil. The samples from French Guiana and Brasil differ by one standard error. The overall trend demonstrates a slight difference in size between the Peruvian and Bolivian samples, the Peruvian samples having the largest individuals examined (Fig. 8). The smallest males examined come 12 Contributions in Science No. 251 23. T 2h.6 2h.O 23.5- 2h.J 24.5- 24.7 22.0- 26.2 22.1 24.9- 25.6 22.4 / 26.1 23.4 / 23.8 20.1 21.2 22.8 20.6- 20.6 23.6 22.9- 2U.0 2k. 0- 21+.1 22.1 21.8+.81 22.9 N 23.1- 23.9 22.6f.6 23.4 22.2- 23.0 22.1 22.0- 23.0 22.2 24.2 23.0 23.7+.9 25.1 23.4 23.5 Figure 8. Relative geographic distribution of male size of form II individuals. Single value indicates length of single individual; one value immediately below another indicates the upper value is the mean based on 2 to 9 individuals, the lower value is the maximum; the addition of one standard error indicates a sample size of 10 or more. See text for further explanation. from three different Brasilian localities: Para, near Belem, Rondonia, Porto Velho, and Rio de Janeiro, Teresopolis (Fig. 8). Female size — Form II females range in size from approximately 20 mm to 31.0 mm S-VL. Four localities were analyzed with 10 or more individuals: French Guiana, 22.5 (mean) ± 1.5 (one standard error) mm, Peru; Departamento de Huanuco, 23.9 ± 2.0 mm, Bolivia; Departamento de Beni, 22.7 ±1.8 mm and Bolivia; Estado do Santa Cruz, 23.2 ± 1.8 mm. The only noticeable trend is that the largest individuals come from the states of Fluanuco and Pasco in Peru and from Peru: Loreto; Suhuaya, near Contamana on Rio Ucayali. Male snout ratio — Three locality samples of 10 or more individuals were availa- ble: French Guiana, Bolivia, and from the state of Sao Paulo, Brasil (Fig. 9). There are no statistical differences among these samples. The only noticeable geographic pattern is that males with high snout ratios are found in central Mato Grosso and southern Goias, Brasil (Fig. 9). Female snout ratio — The female snout ratio is consistently smaller than the male snout ratio at any individual locality. The four localities adequate for statistical consid- erations are: French Guiana, Cayenne and surrounding area, .68 (mean) ± .04 (one standard error), Bolivia, Beni, Huachi and surrounding area, .67 ± .06, Bolivia; Santa 1973 Systematics of Marmoratus Group 13 .69 .67 .68 .W 72 .7^ .82 .74 .60 .70 .74 .74 .83 .73+. 05 .77 .74 .74 .76 .74 N .74 Figure 9. Relative geographic distribution of male snout ratios of form II individuals. Individual values are where a single specimen is represented; a horizontal bar above the index value indicates a mean value of 2 to 9 individuals represented; a horizontal bar and a plus or minus value is the mean value and one standard error for 10 or more individuals from a given locality. See text for further explanation. Cruz, Buenavista, .67 ± .04, and Peru; department of Huanuco samples, .64 ± .06. No geographic patterns are evident. Dorsal pattern — There are no absolute differences in patterns, but certain trends are clear. Almost all of the populations north of Amazonia have a large percentage of B category pattern, as do certain Brasilian populations in the territory of Acre and the states of Mato Grosso and Goias (Fig. 10). Remaining populations are characterized by a preponderance of E category patterns. A single individual from near Belem was categorized as D. Dorsolateral stripes — Populations from the Brasilian territory of Amapa, French Guiana, Surinam, southern Guyana, and the localities of around Feijo, Acre, Brasil; Rio Perene, Junin, Peru; and Belo Horizonte, Minas Gerais, Brasil have a high incidence of B category individuals (Fig. 11). Two northernmost populations have C category individuals. In these instances, the stripes begin at the shoulder, not the eye 14 Contributions in Science No. 251 E-50 C E B-12 C-35 E-53 C-17 E E-83 B-33 E-67 C-33 D-ll E-56 C C-50 E-50 *C-8 TT in *E-81* E-lOO E E-lOO C-33 *C-20 E-67 *E-77 ® *C-36 *E-6U E-lOO © *b-6 *C-35 *E-59 E C-50 ®C *E-100 E-50 E E C-17 E-83 E-lOO Figure 10. Relative geographic distribution of dorsal color pattern of form II individuals. Single letters indicate the state of a single specimen, letters followed by percentages indicate the distribu- tion of states among 2 to 9 individuals, letters preceded by an asterisk and followed by percen- tages indicate the distribution of states among 10 or more individuals from a given region. See text for further explanation. (Fig. 12A). All other populations have a preponderance of individuals lacking dor- solateral stripes (Fig. 11). Mid-dorsal stripe — The clearest difference is between the populations from the Brasilian states of Minas Gerais, Rio de Janeiro, Sao Paulo and all other populations. No individuals from these three states have B or C category mid-dorsal stripes. The D category is also more abundant in populations from these three states than elsewhere (Fig. 13). Populations from the Brasilian territory of Amapa, French Guiana, Surinam, Guyana, and Venezuela do not have any individuals with E or G category stripes; category E or G individuals occur only in Amazonian populations (Fig. 13). Toe tip development — Populations from Venezuela, the Brasilian state and terri- tory of Acre and Rondonia, the mouth of the Amazon, and Belo Horizonte, Minas Gerais, Brasil differ from all other populations in having C category individuals (Fig. 14). Populations from the state of Sao Paulo, Brasil at Anhembi and Campos do Jordao are distinctive in having a preponderance of A category individuals (Fig. 14). Discussion of Taxonomic Decisions Combining the above distributions of character states, three transitional zones are apparent: 1) the mouth of the Amazon region, especially the Brasilian state of Amapa, 1973 Systematics of Marmoratus Group 15 A-lOO *A-80 *B-20 A-T5 B-25 A-lOO *A-T9 *B-21 A-l'i B-C-25 A-50 B-50 A A-67 B-33 A A A-lOO A-lOO A-lOO *A-100 A-lOO A- 100 A A-lOO A- 100 A-lOO A-89 B-11 A-lOO A-lOO A- 100 *A-92 *B-8 A-lOO A A-lOO *A-9T *B-3 A-lOO A A A *A-90 *B-10 A A A A- 100 *A-100 A-lOO A A-lOO Figure 11. Relative geographic distribution of dorsolateral stripes of form 11 individuals. See legend of Fig. 10 and text for explanation. separating the lands draining north from the Amazonian drainage; 2) a broad zone of western Brasil bordering Peru, Bolivia, and Paraguay; 3) the Brasilian states of Guanabara, Minas Gerais, Rio de Janeiro, and Sao Paulo. Direct comparison of specimens from French Guiana, and the Brasilian territory and state of Amapa and Para indicates that a single species is involved. Morphologi- cally, the specimens are similar. There does appear to be a slight break in color pattern in the territory of Amapa, where the northern populations have many individuals with small spots on the dorsums, and the southern populations have individuals with larger, blotchier spots, or no distinct spots at all. I prefer not to recognize these differences taxonomically at present, because there are so few individuals available from the critical area of Amapa and the differences are not striking in the material at hand. The northern populations do differ from other populations with respect to details of dorsal pattern, dorsolateral stripes, and mid-dorsal stripes. These differences are here recog- nized in an informal manner (Fig. 24). Direct comparison of individuals from the Brasilian territory of Rondonia, the states of Acre, Goias, Mato Grosso, and from Bolivia do not indicate any clear cut differences. The situation, in fact, is rather confusing. The samples vary in degree of toe tip expansion, morphology, and dorsal pattern. Morphologically, the sample from Mato Grosso, Urucum, has an individual with a broader head (DZ 9204) than other individuals from the same sample. In other respects, the sample is uniform in charac- teristics. The single individual from Goias, Jatai (WCAB 15139) has a more pointed 16 Contributions in Science No. 251 Figure 12. Category C dorsolateral stripes. A, form II specimen; B, form I specimen from greater Amazonia; C, form I specimen from southeast Brasil. A A-50 C-50 *A-T7 „ *B-8 ^ *F-15 A- 12 B-25 D-12 F-38 G-12 A-50 B-50 A- 35 B-65 A-50 B-17 C-IT F-17 A-78 D-11 E-11 A- 33 B-33 E-17 F-17 C-50 E-50 B-50 C-50 A-33 B-33 G-33 *A-6 *B-17 *C-67 »F-9 *A-67 *B-lU »C-17 *F-3 A-33 : B-67 B-33 *A-83 C-33 C-22 A »B-lt F-12 *C-1 *F-12 A-62 A- 100 B-12 C-12 F-12 B-IOC A-lOO A A-33 F-67 B-33 C-33 D-33 A-20 B-50 C-10 D-10 G-iO B-50 C-20 F-20 Figure 13. Relative geographic distribution of mid-dorsal stripes of form II individuals. See legend of Fig. 10 and text for explanation. head than any of the other individuals. The single example from Vila Murtinho, Mato Grosso, next to Paraguay, has distinctive toe disks. The disks are not clearly of the I or II forms, that is rounded or flattened (Fig. 5). On the basis of the single faded specimen (UMMZ 76081), it is in fact difficult to tell whether it is a form I or II individual. The specimen is tentatively assigned to form II until fresh material has been collected. Pattern wise, the specimens from Tarauaca and Placido de Castro, Acre, Chapada dos Guimaraes, Mato Grosso, and Bolivia are similar in having individuals with small discrete dark spots more or less arranged in four longitudinal rows. The specimens from Feijo, Acre and the specimens from Urucum, Mato Grosso are similar in that some of the spots are coalesced along the mid-line to form short, narrow bands. 1973 Systematics of Marmoratus Group 17 Figure 14. Relative geographic distribution of toe tip development of form II individuals. See legend of Fig. 10 and text for explanation. The specimens from Jatai, Goias and Sto. Antonio Leverger, Mato Grosso are similar in having many indistinct dark spots not arranged in any pattern. The specimens from Nhambiquara and Guajara-Assu Falls, Rondonia are too poorly preserved to determine the dorsal patterns. The overall pattern from this region is a mosaic. Few individuals are available from this region and there are no consistent, clear cut differences separat- ing groups of populations. I prefer to treat informally the series of populations as distinctive until further work elucidates the relationships among these populations (Fig. 24). It should be kept in mind that the populations of frogs found along western Brasil are more or less distinctive from frogs of the northern Amazon, Peru, and Bolivia, but also, the individual populations within western Brasil are distinctive among themselves. Direct comparison of individuals from the Brasilian states of Guanabara, Minas Gerais, Rio de Janeiro, Sao Paulo, Goias, and Mato Grosso indicates that two distinc- tive types are represented. The specimens from Guanabara, Minas Gerais, Rio de Janeiro, and Sao Paulo differ from other specimens in that the appearance of many of the specimens is Pseudis-WkQ, that is, with dorsally directed eyes and more or less triangular bodies. The mid-dorsal stripes are distinctive as noted above. Only two larval samples are known for any members of the Marmoratus group. Both belong to 18 Contributions in Science No. 251 Figure 15. Relative geographic distribution of male size of form I individuals. See legend of Fig. 8 and text for explanation. group II individuals. One sample is from French Guiana, Regina, the other from Sao Paulo, Campos do Jordao. The two tadpole lots are strikingly different. The larvae from Regina lack spiracles and denticles in the tooth rows (Fig. 25), whereas the larvae from Campos do Jordao have spiracles and well-developed denticulate tooth rows (Fig. 23). These differences are certainly indicative of species differences. Life history information may well clarify the status of the populations lying between the extremes of Regina and Campos do Jordao. I consider the form II populations from Minas Gerais, Rio de Janeiro, and Sao Paulo to represent a distinct species from all other form II populations (Fig. 24). Within the southeast coastal Brasilian species, the specimens from Belo Horizonte, Minas Gerais are the most distinctive. None of the individuals have distinct, small dark spots, many individuals are finely mottled, a pattern not seen elsewhere, and some individuals have C category disks. Analysis of Form I Specimens Male size — Form I males range in size from 16.2 to 27.0 mm. There are six samples with 10 or more males for which standard errors were calculated: one from Ecuador, one from Bolivia, and two each from the Brasilian states of Rio de Janeiro and Sao Paulo. The sample from around Teresopolis, Rio de Janeiro differs from the Ecuadorian and Bolivian samples by more than one standard error, the sample from Ilha Sao Sebastiao also differs from the Ecuadorian and Bolivian samples by more than one standard error. None of the other samples differ significantly (Fig. 15). There is 1973 Systematics of Marmoratus Group 19 2k. h- 24.7 23.1 237^- 25.9 20.1 22.4- 24.3 21.6 21.0- 22.0 22.3- 23. T 21.8+. 8 22.9 25.7 N 21.4 20.5 22.4 22.5+1.2 18.3 Figure 16. Relative geographic distribution of female size of form 1 individuals. See legend of Fig. 8 and text for explanation. no distinctive geographic pattern in male size: the largest individuals are found in the western Amazon regions of Ecuador, Peru, Bolivia, and Brasil; the smallest individu- als are from the state of Santa Catarina, Brasil (Fig. 15). Female size — Form I females range in size from about 18 to 28.9 mm. Ten samples are represented by 10 or more individuals for which standard errors were determined: 1) Provincia de Napo, Ecuador, 2) Provincia de Pastaza, Ecuador, 3) vicinity of Benjamin Constant, Amazonas, Brasil, 4) vicinity of Belem, Para, Brasil, 5) Buenavista, Santa Cruz, Bolivia, 6) vicinity of Teresopolis, Rio de Janeiro, Brasil, 7) vicinity of Rio de Janeiro, Guanabara, Brasil, 8) Ilha Sao Sebastiao, Sao Paulo, Brasil, 9) vicinity of Sao Paulo, Sao Paulo, 10) Humboldt, Santa Catarina, Brasil. Sample 10 differs by more than one standard error from samples 1, 2, 3, 5, 6; sample 8 from samples 1, 2, 5; sample 4 from samples 1, 2, 5; sample 1 from sample 6. None of the other combinations differ significantly. No distinct patterns are evident in the geographic distribution ofTemale size: the largest individuals (S-VL^26.0 mm) are found in the western Amazonian regions of Colombia, Ecuador, and Peru, and the isolated localities of Sipaliwini, Nickerie, Surinam; Buenavista, Santa Cruz, Bolivia; Alcatrazes Is., Sao Paulo, Brasil; Porto de Cima, Morretes, Parana, Brasil; the smal- lest individuals are found in the state of Santa Catarina, Brasil (Fig. 16). 20 Contributions in Science No. 251 E-lOO C-25 E-T5 ^E-100 *E-100 ® E-lOO E-lOO *E-100 E-lOO E *E-100 E E E-lOO E-lOO E-lOO / ^ I *C-28 ) E \ *E-56 / *C-21 *E-T9 C-20 E-80 E-lOO E-lOO E *C-4 *E-96 E *C-UO *e-6o / 'c-67 E-33 B-IT C-50 E-34 C-33 E-6t '^C-11 \ B-50 C-50 ; *D-1 \ *E-8£ *A-2 *B-5 *C-10 *E-83 *C-l!+ *E-86 ° C-28 E-lOO p_22 E-50 Figure 17. Relative geographic distribution of dorsal color pattern of form I individuals. See legend of Fig. 10 and text for explanation. Male snout ratio — Six localities are represented by 10 or more males for which standard errors were determined on the snout ratios: 1) Provincia de Pastaza, Ecuador, .70 (mean) ± .06 (one standard error); 2) Buenavista, Santa Cruz, Bolivia, .72 ± .03; 3) vicinity of Teresopolis, Rio de Janeiro, Brasil, .71 ± .04; 4) vicinity of Rio de Janeiro, Guanabara, Brasil, .73 ± .04; 5) Ilha Sao Sebastiao, Sao Paulo, Brasil, .68 ± .05; 6) vicinity of Sao Paulo, Sao Paulo, Brasil, .72 ± .04. No geographic patterns or trends were evident when the data were plotted on a map. Female snout ratio — The female snout ratio is consistently smaller than the male snout ratio at any locality represented by at least two specimens of each sex. Ten samples are represented by 10 or more individuals for which standard errors were determined: 1) Provincia de Napo, Ecuador, .64 ± .06, 2) Provincia de Pastaza, Ecuador, .65 ± .05, 3) vicinity of Benjamin Constant, Amazonas, Brasil, .69 ± .07, 4) vicinity of Belem, Para, Brasil, .65 ± .04, 5) Buenavista, Santa Cruz, Bolivia, .67 ± .04, 6) vicinity of Teresopolis, Rio de Janeiro, Brasil, .70 ± .04, 7) vicinity of Rio de Janeiro, Guanabara, Brasil, .69 ± .05, 8) Ilha Sao Sebastiao, Sao Paulo, Brasil, .67 ± .04, 9) vicinity of Sao Paulo, Sao Paulo, Brasil, .67 ± .06, 10) Humboldt, Santa Catarina, Brasil, .73 ± .03. Sample 10 differs by 1 standard error or more from 1973 Systematics of Marmoratus Group 21 A- 100 A A- 100 A-83 C-IT *A-8T *C-13 A-100 *A-90 *C-10 A- 100 *A-81 *0-19 *A-79 *0-21 A A-100 *A-69 *0-31 0 A-6T 0-33 A-100 A 0-100 *A-8T *0-13 A-IT 0-83 *A-81 V *0-18 J *C-19 ^ A *A-93 *0-7 A-67 0-33 Figure 18. Relative geographic distribution of dorsolateral stripes of form 1 individuals. See legend of Fig. 10 and text for explanation. samples 1, 2, and 4. Other than the samples of frogs from Humboldt having long snouts, no geographic trends were apparent when the ratios were plotted on a map. Dorsal pattern — Most individuals are either C or E category with respect to their dorsal patterns. The B category is found in a few individuals from Estiron, Ampiacu River, Loreto, Peru, and in scattered populations in the Brasilian states of Rio de Janeiro, Sao Paulo, and Parana (Fig. 17). The A category is found in a low frequency in the northern populations and from around Teresopolis, Rio de Janeiro, Brasil (Fig, 17). The D category is found in low or moderate frequency in the northern populations, Ilha Sao Sebastiao, Sao Paulo, Brasil, and the state of Santa Catarina, Brasil (Fig. 17). Dorsolateral stripes — Most populations have around a 20% frequency of indi- viduals with C category stripes (Fig. 18). The only populations in which the C categ- ory was absent in samples of 10 or more individuals are from the western Amazon regions of Ecuador and Peru (Fig. 18). Low frequencies of B category individuals are found in central and southern Amazonian Ecuador, southern Venezuela, southern Guyana, around Rio de Janeiro, Guanabara, Brasil, and Itapetininga, Sao Paulo, Brasil (Fig. 18). Mid-dorsal stripe — Most samples have individuals with A, D, and F category 22 Contributions in Science No. 251 A-lOO A-25 D-50 F-25 *A-12 *D-63 *F-13 *A-T1 *D-21 *F-8 A-33 D-33 F-33 A- 100 *A-21 F *D-51 *F-28 A A F-lOO A A-6o D-40 N *A-5 *D-85 *F-10 A-33 D-50 F-17 D-lOO d-66 F-33 D *A-T1 »D-23 *F-6 A-50 D-25 *A-32 *d-6o *F-8 A-6T D-33 A-6T D-33 D-83 F-IT A- ■100 *A-62 D-lOO *D-35 *F-3 A *A-T^ *D-26 / *A-58 / *B-8 A-6t \*DB-8 D-33 ^\^D-25 A-100 . *A-22 *d_26 *B-9 Af-30 'v*C-13l_ , *A-71\ *DB-1 \ *D-25 / A Figure 19. Relative geographic distribution of mid-dorsal stripes of form I individuals. See legend of Fig. 10 and text for explanation. stripes (Fig. 19). B category stripes are found in low to moderate frequencies in samples from the southern border of Guyana and Surinam; Villavicencio, Meta, Col- ombia; Provincia de Napo, Ecuador; Ilha de Sao Sebastiao, Sao Paulo, Brasil, around Sao Paulo, Sao Paulo, Brasil; and the state of Santa Catarina, Brasil (Fig. 19). The only C category individuals are from the state of Santa Catarina, Brasil; the only E category individuals are found in northcentral samples and are strictly correlated with the A category dorsal pattern (Fig. 19). Toe tip development — During the data taking, it was noticed that juvenile speci- mens often had poorly differentiated disks while adults from the same locality had well-developed disks. Only adults were used for this character analysis. Almost all samples are characterized by having individuals with either C or D category disks. B category individuals are from three separate areas: Sipaliwini, Nickerie, Surinam; southern Venezuela; and the state of Parana, Brasil (Fig. 20). BC category individuals are from five areas: French Guiana and east-central Surinam; Abuna, Rondonia, 1973 System ATics of M armor at us Group 23 C-50 c D-50 C-86 d-i4 c c *C-T4 *D-26 / B-20 \ I C-60 I \ D-20 J *C-T2 *D-28 C-6t D-33 *C-83 *D-17 C-6t D-33 C-100 D C C c c-100 *C-58 *D-i*2 C-33 D-6t *C-5T *D-43 D D D-lOO *BC-3 *C-64 *D-32 ( B-5o\ V D-50 ) C-IT D-83 C-33 D-6t D *C-6l *D-39 Figure 20. Relative geographic distribution of toe tip development of form I individuals. See legend of Fig. 10 and text for explanation. Brasil; Buenavista, Santa Cruz, Bolivia; the states of Rio de Janeiro and Sao Paulo, Brasil; and the state of Santa Catarina, Brasil (Fig. 20). Discussion of Taxonomic Decisions The above analyses indicate that there are three somewhat distinctive regions: 1) the region represented by the north-central samples. The samples are uniform among themselves, but differ with respect to details of dorsal pattern, dorsolateral stripes, mid-dorsal stripes, and toe disk development from the other samples; 2) the region represented by the western Amazonian samples from Colombia, Ecuador, Peru, Amazonas, Brasil, and Bolivia. The samples are basically uniform among themselves, but differ with respect to details of size and dorsolateral stripes from the other samples; 3) the region of the Brasilian states of Rio de Janeiro, Sao Paulo, Parana, and Santa Catarina. These samples differ among themselves and collectively differ from the other samples. The specimens from southern Guyana and Surinam are the most robust individu- als of those examined (e.g.. Fig. 4, form A). These individuals contrast in body shape with specimens from localities directly south along the Amazon River in the Brasilian states of Amazonas and Para. There is a continuum of body shape of specimens from southern Guyana and southern Surinam— > western and northern Surinam and French Guiana^ Territorio de Amapa, Brasil-^ Estados do Para, Brasil and eastern 24 Contributions in Science No. 251 18.5 20.4- 20.7 22.1 A-67 0-33 A-28 B-14 DB-28 Figure 21. Relative geographic distribution of four characters of form I individuals in the Brasilian states of Sao Paulo, Parana, and Santa Catarina. To the left and above lines is mainland, figures to the right of lines indicate island populations. A, Male size-See legend of Fig. 8 and text for explanation. B, Female size-See legend of Fig. 8 and text for explanation. C, Mid-dorsal stripes-See legend of Fig. 10 except that vertical bar rather than asterisks indicates a sample size of 10 or more individuals. See text for further explanation. D, Toe tip development-As for mid-dorsal stripes. Amazonas, Brasil, however. With the specimens at hand, and lack of large series from critical areas, I prefer not to recognize these differences as representative of specific differentiation. The north central specimens are distinctive in details of body shape, dorsal pattern, and toe disks, which features are recognized informally (Fig. 27). The specimens from northern draining lands of Colombia and Venezuela are more similar to specimens from Amazonian Colombia, Ecuador, and Peru than the north-central South American localities. Specimens from western Amazonian Colom- bia, Brasil, Ecuador, Peru, and Bolivia are somewhat distinctive in details of size and dorsolateral stripes; these differences together with the homogeneity of dorsal pattern, mid-dorsal stripes, and toe disk development are recognized informally (Fig. 27). BC-67 C-33 BC BC-20 1973 Systematics of Marmoratus Group 25 All specimens from the Brasilian states of Rio de Janeiro, Sao Paulo, Parana, and Santa Catarina differ from other specimens in not being as robust, most of the speci- mens having dorsolateral stripes of narrower type (Fig. 12C), B and C category mid-dorsal stripes present, and B and BC category toe disks reasonably represented. While these differences are not absolute, they appear to be consistent. There is at present, a wide zone where no form I individuals have been collected, making it difficult to assess the degree of taxonomic differentiation of the southeast Brasil forms from the Amazonian and northern South American form. To my knowledge, no comparative information on either mating call structure or larvae is available. Taking the cue from the form II analysis and for reasons discussed in the relationship section, I prefer to recognize the differences at the species level. There remains within the southeast Brasilian form I samples considerable heterogeneity, calling for a closer scrutiny of variation in the states of Sao Paulo, Parana, and Santa Catarina with respect to size, mid-dorsal stripes and toe disks. The geographic distribution of male and female size is a continuum from rela- tively small size in northern Sao Paulo to the smallest sizes in Santa Catarina with three large-sized populations from Alcatrazes Island, Sao Paulo, and Parana (Fig. 21 A,B). The geographic distribution of mid-dorsal stripes is a crude continuum from B cate- gory in Sao Paulo, leading to expression of the B and C categories in Santa Catarina. The only real break is the unique presence of the C category in Santa Catarina (Fig. 2 1C). The geographic distribution of toe disks shows that the Parana samples are unique (Fig. 21D). Taken together, the samples form an integrated series with the exception of the samples from Alcatrazes Island, Sao Paulo; Paranagua, Parana; and Porto de Cima, Morretes, Parana. The sample from Alcatrazes Island is unique only with respect to size, but in this respect it is distinctive. Gigantism is a relatively common phenomenon on islands, but neither of the island populations from Sao Sebastiao nor Buzios are as distinctively large as the Alcatrazes population. Further work is needed to determine the significance of size in the Alcatrazes population. I consider the Alcatrazes popula- tion to represent the same species as the mainland form I species, but point out the distinctive large size of the population in an informal manner (Fig. 27). The samples from Paranagua and Porto de Cima, Morretes are distinctive with repect to size and toe disk development. The larger sizes of the specimens seem to be superimposed over the general trend of decreasing size of the mainland populations (Figs. 21 A,B); the B category of disk development is unique in these two samples. Data from three indi- viduals from Volta Grande, Parana were not plotted on the maps as the specimens are faded juveniles. The 11.1 and 14.5 mm specimens have BC category disks, the 16.0 mm specimen has C category disks. These three individuals fit into the general patterns evidenced by the rest of the mainland form 1 individuals and contrast with the other individuals from Parana. As indicated previously, the decision to include the Paranagua and Porto de Cima, Morretes individuals in the form I individuals analysis was based on the fact that certain of these individuals had C category dorsolateral stripes. Direct comparison of the distinctive Parana specimens with form II specimens from Sao Paulo indicates that the only consistent difference is in the dorsolateral stripes. All specimens are of similar size, dorsal pattern, toe tip development, and 26 Contributions in Science No. 251 ventral pattern. I consider the Parana slender toed forms to represent the same species as the slender toed forms from Sao Paulo until further collections and information are gathered to indicate otherwise. The distinctiveness of the Parana form II individuals is pointed out in an informal manner, however (Fig. 24). Taxonomic Conclusions Five species are recognized. The most distinctive occurs in the arid interior of Brasil. The remaining four species form a complex of two allopatric groups of two species each. Two species are broadly sympatric over much of northern South America, Amazonia, and southwestern Brasil, the other two species are broadly sym- patric over much of southeast Brasil from Minas Gerais to Santa Catarina. Nomenclature Twelve names have been proposed previously for members of this group. In addi- tion, one name has been associated with this group from time to time. Each of the names is dealt with in the order that it was proposed. Rana pygmaea Spix, 1824. Dr. Gunther Peters (personal communication) in- formed me that the type is no longer in the Berlin collections. W. Peters (1873) examined Spix’s type material and briefly indicated that the holotype ofR. pygmaea was from Bahia and that it was a juvenile of Cystignathus pachypus. C. pachypus has been placed in the synonymy of several species by various authors, but has most commonly been placed in the synonymy ofL. ocellatus. Melin (1941: 60) suggested that/?, pygmaea may actually pertain to members of the Marmoratus group. Examina- tion of Spix’s figure of R. pygmaea could support Melin’s suggestion, although the figure is quite stylized. As the type is no longer extant, a decision must be made with the available data, the three most critical bits being the locality, the original figure, and W. Peter’s synonymy based on examination of the specimen. No recent collections of members of the Marmoratus group are from the Brasilian state of Bahia. This, together with W. Peter’s synonymy lead to the conclusion that the name does not apply to any member of the Marmoratus group but that the name applies to a member of either the Fuscus or Ocellatus groups. Adenomera marmorata Fitzinger in Steindachner, 1867. There is confusion as to exactly who authored the species. In the text of Steindachner (1867), the new genus and species is clearly attributed to Fitzinger (p. 37). The proper citation would thus appear to be Fitzinger in Steindachner. As Parker (1932) points out, however, the description does not apply to the figure. Parker suggested that 1) a mix-up of speci- mens occurred and that the specimen that Steindachner chose to have illustrated was not the one described by Fitzinger, and 2) that the name /I. marmorata be associated with the figure and the extant specimen rather than the description. As it is clear that Steindachner considered Fitzinger to be the author of Adenomera marmorata and apparently made a mistake in having the incorrect specimen figured, I prefer to use the citation Adenomera marmorata Fitzinger in Steindachner. The holotype (the figured specimen), Vienna 16453, is completely faded, but otherwise in good condition. I follow Bokermann’s (1966) suggestion that the holotype originated from around Rio 1973 Systematics of Marmoratus Group 27 de Janeiro, Brasil. The toe tips are expanded into distinct, flattened disks. Adenomera marmorata is the oldest available name for the form I species of southeast Brasil. Cystignathus hylaedactylus Cope, 1868. The holotype, ANSP 2240, is presently in a state of fair preservation. The original description was extremely brief with respect to color pattern and no distinctive pattern can now be discerned on the type. Cope indicated that the toes had “distinct dilations at the end.” Several of the toe tips of the holotype have been dissected by previous workers to examine the state of the terminal phalanges. The intact toe tips are somewhat desiccated, but are dilated. They corres- pond to the B-C state used herein. The swellings are rounded, not flattened. Cystignathus hylaedactylus Cope is the oldest available name for the form II species found in northern South America, Amazonia, and southwest Brasil. Leptodactylus diptyx Boettger, 1885. One of the five syntypes, BMNH 1947. 2.17.47, is clearly representative of the form II species from northern South America, Amazonia, and southwest Brasil. The tips of the toes are pointed, the dorsum is spotted, there are narrow lateral stripes on the sides of the body, and there is a mid-dorsal pin stripe from above the vent extending beyond the sacral region. I hereby designate BMNH 1947.2.17.47, an adult male, the lectotype of Leptodactylus diptyx Boettger. The locality of Paraguay represents the only record of any representative of the Marmoratus group from that country. Records are available close to the Paraguayan border in the state of Mato Grosso, Brasil, suggesing that the provenance of the type poses no problem. Leptodactylus glandulosus Cope, 1887. Cope indicated that the species was small, it appears as though all of the 14 specimens comprising the type series are juveniles. The toe tips are either the A or B category, some individuals have the dorsum with indistinct spots, and some individuals have a light, mid-dorsal pin stripe from above the vent extending to the snout. Leptodactylus glandulosus clearly applies to the form II species from northern South America, Amazonia, and southwest Brasil. I hereby designate ANSP 10275 as the lectotype of Leptodactylus glandulosus Cope. Leptodactylus hololius Boulenger, 1918. The holotype is in a good state of preservation. In addition to the spots on the anterior part of the head and body dorsally as described by Boulenger, indistinct spots are present more posteriorly. The tips of the toes are slightly expanded, state B., and rounded. The holotype is clearly represen- tative of the form II species from northern South America, Amazonia, and southwest Brasil. Leptodactylus nanus Miiller, 1922. Dr. U. Gruber kindly sent the available type specimens. He informed me (personal communication) that the holotype, Munich 659/1920 was lost in the war. The three extant types are in excellent condition, with expanded toe disks and type C dorsolateral stripes. The types are clearly representative of the form I species of southeast Brasil, in particular the southern populations. I hereby designate the 19.4 mm S-VL female specimen, Munich 661/1920 as the lectotype of Leptodactylus nanus Muller. Leptodactylus andreae Muller, 1923. The holotype, Munich 136/1911, was lost in the war (U. Gruber, personal communication). The remaining four types are in a good state of preservation. The types have expanded toe disks (type D), and one has category D dorsal pattern. The 21.1 mm S-VL specimen, Munich 145/191 1 , is hereby 28 Contributions in Science No. 251 designated as the lectotype of Leptodactylus andreae Muller (the specimen was not dissected, but apparently is a juvenile female). Leptodactylus andreae is the oldest available name for the form I species of northern South America including Amazonia. Leptodactylus minutus Noble, 1923. The holotype is in a good state of preserva- tion. The dorsum is spotted, there is a C category mid-dorsal stripe, and the toes are B category, all of which associate the holotype with the form II species from northern South America, Amazonia, and southwest Brasil. Leptodactylus trivittatus Lutz, 1926. A series of syntypes at the United States National Museum are in fair to poor states of preservation. Only one of the specimens, a juvenile, has the pattern on which the name was based. For this reason, USNM 96943, a juvenile, is hereby designated as the lectotype of the species. The toe disks are C-D category and flattened. The name applies to the form I species of southeast Brasil. Leptodactylus rugosus Melin, 1941. Lutz and Kloss (1952) pointed out that L. rugosus Melin was preoccupied by L. rugosus Noble and renamed Melin’s species as L. melini. The holotype has category B toe tips which are rounded. The dorsal pattern is faded but may have been spotted. The sides are warty (as are many individuals of this group) and apparently this is the reason for the original specific name. The dorsum is smooth. The name clearly applies to the form II species of northern South America, Amazonia, and southwest Brasil. Leptodactylus poeppigi Melin, 1941. The toe tips are B category, the dorsum has irregular spots and a B category mid-dorsal light pin stripe, all of which indicates the name applies to the form II species of northern South America, Amazonia, and south- west Brazil. Leptodactylus martinezi Bokermann, 1956. The types were included in the analysis and were part of the form III species of central Brasil. In summary, the form I species of southeast Brasil has been described three times, A. marmorata is the oldest name. The form I species of northern South America and Amazonia has been described once as L. andreae. The form II species of northern South America, Amazonia, and southwest Brasil has been described seven times, C. hylaedactylus is the oldest name. The form III species has been described once asL. martinezi. One species remains, which apparently has never been described. This new species is described in the next section. Species Accounts The information from the previous analyses is summarized in the accounts. Addi- tional information was taken from a series of 10 males and 10 females of each species representing the extremes of geographic range within each species except for L. martinezi, for which all available adult specimens were examined. The snout-vent length and percentages of S-VL are mean values which are followed by one standard error except for maleL. martinezi. The methods are the same as those used previously (Heyer, 1970b). Adult members of the Melanonotus and Ocellatus species groups have distinct lateral fringes distinguishing them from members of the Marmoratus group which have free toes. Adult members of the Pentadactylus species group are moderate to large 1973 Systematics of Marmoratus Group 29 sized frogs (greater than 60 mm S-VL), the males have thumb spines, differentiating them from the small sized (smaller than 31 mm S-VL) members of the Marmoratus group in which the males lack thumb spines. Members of the Marmoratus group can be differentiated from members of the Fuscus group as follows (characteristics for mem- bers of the Marmoratus group in parentheses): dorsolateral folds usually present (al- most always absent), a light longitudinal stripe often on posterior face of thigh (ab- sent), first finger usually much longer than second (about equal), moderate in size, 25-65 mm S-VL (small, never exceeding 31 mm S-VL). Leptodactylus andreae Muller Leptodactylus andreae Muller, 1923:41-43 (Type locality, Brasil: Para; Peix- eboi. Lectotype Zool. Sammlung Bayr. Staates, hert. nr. 145/1911, juvenile female). Diagnosis — The only other species in the Marmoratus species group in addition to L. andreae with flattened toe disks is L. marmoratus. Some individuals of L. andreae have a striped pattern dorsally, no L. marmoratus have dorsal longitudinal stripes, almost noL. andreae have a light mid-dorsal pin stripe extending from above the vent to beyond the sacral region, someL. marmoratus have such stripes extending to the shoulder region or to the snout. Summary of characteristics — Adults: Dorsal outline of snout rounded, subovoid, or subelliptical in females, subovoid, subelliptical, or pointed in males; snout rounded in profile in females, rounded or usually rounded-acute in males; female snout ratio .643 ± .060, male snout ratio .703 ± .056, canthus rostralis indistinct; loreal slightly concave or obtuse in cross section; tympanum distinct, equal to or just greater than Vi diameter of eye; vocal sac single, internal or very slight folds under angle of jaw to forearm; vomerine teeth in transverse (usually) or very slightly obtuse series posterior to choanae; head length 38.8 ± 1.8% S-VL; head width narrower than length, 35.4 ± 1.9% S-VL; interorbital distance 9.1 ± 0.8% S-VL; length of first finger equal to, just greater than, or just lesser than length of second finger; dorsum smooth or with ridges, warts, or rarely tuberculate dorsolateral folds, males usually with white-tipped tubercles scattered over posterior dorsum, females almost always lacking such tuber- cles; no distinctive glands or two small round glands on either side of anus on hind- limb; toes usually with distinct, flattened disks (Figs. 1, C, D; 5, A); toes free; subarticular tubercles pronounced; no metatarsal fold; weak, sinuous tarsal fold ex- tending V2-% length of tarsus, usually smooth or with series of white-tipped tubercles; lower surface of tarsus and sole of foot almost smooth to profused with white-tipped tubercles; female size 24.1 ± 2.0 mm S-VL, 28.9 mm maximum, male size 23.3 ± 1.5 mm S-VL, 27.0 mm maximum; femur 42.1 ± 3.8% S-VL; tibia equal to or longer than femur, 45.8 ± 1 .9% S-VL; foot longer than femur, equal to or longer than tibia, 48.2 ± 2.4% S-VL; dorsal pattern most commonly uniform (Fig. 2E), or often with indistinct marks (Fig. 2C) or distinct blotches (Fig. 2D), rarely with distinct lines (Fig. 2A), most individuals lacking light dorsolateral stripes, a few with narrow stripes, several with well defined light stripes from eye to groin (Fig. 12B), narrow light mid-dorsal stripe absent or extending from anus to sacral region or rarely to middle of back, broad mid-dorsal stripe from snout to anus present only in the indi- viduals with striped dorsums (Fig. 2A), center of throat and belly immaculate, ventral 30 Contributions in Science No. 251 borders of chin, thigh, and tibia (or entire tibia) mottled, some males with dark vocal | folds; posterior thigh finely mottled to uniform. | Geographic variation — Populations from northern South America (Guyana, i Surinam, French Guiana and part of Amapa, Brasil) are rather uniform with respect to morphology and color pattern. They differ in detail from all other populations in having the dorsal pattern of three light broad longitudinal stripes (A category) rep- resented at a low frequency, the blotched dorsal pattern represented in moderate frequency, a low frequency of narrow dorsolateral stripes not reaching the eye (cate- gory B), and the pin stripe mid-dorsal stripe (B category) is present at a low frequency. Populations from the western Amazon (Colombia, Ecuador, Peru, Amazonas-Brasil, Bolivia) are also rather uniform with respect to morphology and color pattern. They differ in detail from other populations in being larger and having a very low frequency of broad light dorsolateral stripes (C category). Remaining populations do not demon- strate any geographic trends in morphology or color pattern. Distribution (Fig. 27) — Range: North, central, and westen South America east of the Andes. Known elevational range, sea level to 1200 m. , Localities — BOLIVIA. Santa Cruz: Buenavista, 500 m, MCZ 15583, UMMZ 64031 B-C, J, 64032B 66483A-F, 66484A-E, 66493A-D, F, 66494C, 66504A-G, ' 66542,66611. BRASIL. Amapa: Mazagao, WCAB 19184-5; Serra do Navio, LACM 44281, I 44726-53, WCAB 2317-8, 3623, 18169, 18224, 19979, 35245-8. Amazonas: Codajaz, DZ 28325; Ducke reserve, near Manaus, KU 129938-41; Ilha Terra Nova, in Amazon River, LACM 44714, 44716; Itapiranga, DZ 27755, 27757-8; Lago de January, island next to Channel Pixuna, LACM 44719, 44721-3; Livramento, region, AMNH 44782-6; Tapuruquara, WCAB 8768-9; Teresina, 3 hrs downstream Leticia, LACM 50179-97. Para: Belem, 20 m, DZ 24999-25000, LACM 44792-8, USNM i 154062-65; IPEAN, 3 km E Belem, KU 127338, 127340-52, 127354-5, 12357, 127359-64, 127366-9, 127371-5, 127377, 127380-90, 127392-4, 128268-71; Jacareacanga, DZ 24949, WCAB 45483, 45508-12; Rio Mapuera (at equator), 80 m, AMNH 80026; Sudam Floral Reserve, 74 km SE Santarem, KU 129934-7. Ronddnia: Abuha, WCAB 10050; Igarape Marmelo, WCAB 9852, 9856, 9858; Porto Velho and vicinity, UMMZ 64119A-C, E. COLOMBIA. Amazonas: Rio Apaporis, USNM 144843. Caqueta: Tres Es- quinas, Rio Arteguaza, WCAB 40641-3; Villa Maria, USNM 147038. Cordoba; Tierra Alta, FMNH 61802, 61805. Meta; Villavicencio, 500 m, FMNH 30812, 81795-8, USNM 144844-6. Vichada; Anaben, UPR 102. ECUADOR. Moruna-Santiago: trail between Copal and Mendez, 580-910 m, USNM 192705; trail between Plan Grande and Rosario, between Mendez and Limon (Gral. Plaza), 910-1130 m, USNM 192706-9; Sucua and environs, 820 m, USNM 1926.92-9; Rio Yuquipa, nr. Macas, USNM 192710. Napo: Lago Agrio, KU 126243-5; Loreto, 400 m, JAP 9216; Puerto Libre, Rio Aguarico, 570 m, KU 119355-86; 2 km W Puerto Napo, Hacienda of George Kiederle, on S bank of Rio Napo, 470 m, USNM 192703; San Jose Viejo de Sumaco, JAP 9215; Santa Caecilia, 340 m, KU 109161-2, 111426-8, 119344-5, 119348-9, 119352-4 UMMZ 129285(2); 1 mi NE Tena, E of Rio Masahualli, 490 m, USNM 192701-2. Pastaza: Arajuno, 537 1973 Systematics of Marmoratus Group 31 m, USNM 192712-25; Canelos, 530 m, KU 1 19324-42; Caveceras del Ri'o Bobonaza, 670 m, USNM 192685-6, 192689-91; Vh km downstream from Caveceras, 650 m, USNM 192687-8; Moreta, below Montalvo, Rio Bobonaza, JAP 9219, 3 km SSE Puyo, 975 m, USNM 192683-4; 5 km SSE Puyo, Hacienda Madrid, 975 m, USNM 192681-2; Headwaters of Rio Capahuari, JAP 9209-10; Rio Conambo, JAP 9220-1; Rio Pindo, 190 m, USNM 192726; Rio Puyo at Puyo, 970 m, USNM 192680; Rio Rutuno, trib. of Rio Bobanaza, JAP 9214; Rio Sandalias, trib. of Rio Puyo, Rio Bobonaza, USNM 192711, WCAB 35526-7; Rio Solis, headwaters of Rio Bobonaza, JAP 9211-2; Rio Villano, 300-400 m, GOV 9523, JAP 9213, 9217; 2 km S. Shell Mera, 970 m, USNM 192700; Veracruz (± 10 km E Puyo), 1000 m, KU 119323, USNM 192704. FRENCH GUIANA. Cabassou (Remire), LG 358-9; 6 km SSE Cayenne, Mont. Cabassou, LACM 44592; trail between Flueve Approuague and summit Mont Tortue, LACM 44607-11; Maturi, LG 555-6; Montagne des Chevaux, LG 397-8; Regina, 25 m, LACM 44597-9, 44612-6; Remire, LG 789; Lower Riviere Matarony, upstream from Matarony, LACM 44593-6, 44600-5, 44617-23; Rochambeau, LG 711, 724, 733, 741-5; Sant Tortue, Orstom Camp, LACM 44606, 44669; trail from Sophie to La Greve, MCZ 44559. GUYANA. Essequibo: N of Acarahy Mts W of New River, KU 69714; Shudikar-wan, 200-300 m, AMNH 46264, 87877-85, 46268-9, 46272-3, 80028, 87886-8. PERU. Loreto: Estiron, Ampiacu River, 200 m, DZ 24284-90, 24838-41, 24843, 24847, 25491; Iparia, MCZ 75033, 75035-8; Iquitcs, AMNH 43471; Itaya River, 150 m, AMNH 42175; Lago Mirano, mouth of Rio Napo (at Rio Amazonas), AMNH 43155, 43161, 43168, 43184, 43190; Pampa Hermosa (Cushabatay), 176 m, AMNH 42048; Rio Tamaya, 150 m, AMNH 43363, 43423-4; Tibi Playa (above mo. Rio Ucayali), 150 m, AMNH 42785; Pasco; Iscozazin Valley, Pan de Azucar, 380 m, LACM 40654, 40657; San Martin; Achinamisa, below Chasuta on Rio Huallaga, AMNH 42595. SURINAM. Marowijne: Anapaike Village, Lawa River, DZ 24778-80; Lawa River; DZ 24774-6. Nickerie: Kaysergebergte, RM 16751(2); Post Gonini, Coeroeni Rivier, RM 16739(2), 16740(4); Post Tigrie, New Rivier, RM 16741; Sipaliwini, RM 16724, 16727, 16745, 16749(7), 16753. Suriname: Berlijn, RM 16742. VENEZUELA. Amazonas: Base of Mt. Duida, UPR 3125-9; Pico Cunoto, Mt. Duida, UPR 3130; Mt. Marahuaca, 1200 m, UPR 103. Barinas: Palma Sola, UMMZ 55550. Leptodactylus bokermanni, new species Figures 22, 23 Holotype — UMMZ 104257, an adult male from Paranagua, Estado do Parana, Brasil. Elevation 30 m. Paratopotype — WCAB 22787. Paratypes — BRASIL. Minas Gerais; Agua Limpa, USNM 96997; Belo Horizonte, UMMZ 109990 (2 specimens), WCAB 7236-42; Po^os de Caldas, 1000 m, WCAB 18735; Serra da Piedade, Caete, DZ 25071. Parana: Porto de Cima, Morretes, 32 Contributions in Science No. 251 O 3 OO ' oo8° Figure 23. Lateral view and mouthparts of larva of Leptodactylus bokermanni. Upper line equals 1 cm, lower line equals 1 mm. DZ 15781-2, 24292. Rio de Janeiro: Niterof, AMNH 20278, 87876; Saco Sao Fracisco, USNM 99122; Teresopolis, Guapi, USNM 97681, 97683-90. Sdo Paulo: Anhembi, 400 m, WCAB 30860-1; Botucatu, DZ 6388; Campos do Jordao, WCAB 30035, 30097-102, 34462 (larvae), 34463-76, 36957-60, 37628-9; Iguape, DZ 508; Limeira, DZ 9638; Piquete, DZ 530; Sta. Branca, DZ 25445. Diagnosis — The other species of the Marmoratus group in addition to L. boker- manni with toe tips not developed into flattened disks are L. hylaedactylus and L. Q UBi I qO Figure 22. Dorsal and ventral views of a paratype oi Leptodactylus bokermanni, WCAB 30035, from Campos do Jordao, Sao Paulo, Brasil. Specimen is 24.3 mm S-VL. 1973 System ATics of Marmoratus Group 33 martinezi. Leptodactylus bokermanni does not have four longitudinal rows of symmet- ric dark spots as does L. martinezi. No individuals of L. bokermanni have a light mid-dorsal stripe extending from above the vent to the middle of the body or to the snout, many L. hylaedactylus do; L. bokermanni larvae have a spiracle and well- developed denticle rows, L. hylaedactylus larvae lack a spiracle and denticle rows. Description of Holotype — Dorsal outline of snout subelliptical; snout profile rounded; canthus rostralis indistinct; loreal slightly concave in cross section; tym- panum distinct, greatest diameter V2 eye diameter; vomerine teeth in short transverse series, posterior to choanae; vocal slits elongate, paired; vocal sac single, vocal fold from jaw to forearm on each side; finger lengths in order of decreasing size first finger just shorter than second; large ovoid inner metacarpal tuber- cle smaller than ovoid outer metacarpal tubercle; head and shoulders smooth, sacral region and upper surfaces of tibia profused with white-tipped tubercles; angle of jaw and sides with diffuse glandular development, compact, white oval glands on either side of the anus on hind limbs; glandular fold from eye, over tympanum, to shoulder; ventral surfaces smooth, belly fold distinct; toe tips slightly expanded, state B; toes without fringe; subarticular tubercles moderately developed; distinct, oval inner metatarsal tubercle larger than rounded, distinct outer metatarsal tubercle; tarsal fold not continuous with inner metatarsal tubercle, extending % distance of tarsus; line of white-tipped tubercles in place of metatarsal fold; lower surface of tarsus and sole of foot profused with white-tipped tubercles. Snout-vent, 24.0 mm; head length, 9.0 mm; head width 7.9 mm; interorbital distance, 2.2 rnm; greatest diameter of tympanum, 1.3 mm; diameter of eye, 2.7 mm; eye-nostril distance, 2.0 mm; femur, 9.7 mm; tibia, 10.4 mm; foot, 12.3 mm. Dorsum cream, in preservative (probably faded), with tan markings, interorbital bar extended into a triangle posteriorly, posterior tip of interorbital triangle contiguous with a chevron in shoulder region, dorsolateral dark stripes from shoulder to groin, other tan markings in sacral region; upper surfaces of limbs striped, supratympanic fold dark; throat profused with melanophores, other ventral surfaces appearing im- maculate; posterior surface of thigh mottled. Summary of characteristics — Adults: Dorsal outline of snout rounded, subovoid, or subelliptical in females, subovoid, subelliptical, or pointed in males; snout profile rounded in females, rounded or rounded-acute in males; female snout ratio .702 ± .046, male snout ratio .717 ± .050; canthus rostralis indistinct; loreal very slightly concave or obtuse in cross section; tympanum distinct, greatest diameter equal to or just greater than V2 eye diameter; male vocal slits elongate, very slightly oblique to jaw or parallel to jaw; vocal sac single, internal or a slight fold along margin of jaw to forearm; vomerine teeth in transverse series, posterior to choanae; head length 38.2 ± 1.7% S-VL; head width less than length, 34.0 ± 1.4% S-VL; interorbital distance 8.9 ± 0.8% S-VL; length of first finger equal to or just greater than length of second finger; dorsum smooth or with ridges or warts, males often with white-tipped tubercles posteriorly, females rarely with white-tipped tubercles posteriorly, sometimes appear- ing glandular over much of body, with no distinct glands or some males with distinct, small, ovoid glands on either side of anus on hindlimb; toes commonly pointed or slightly swollen at tips (Fig. lA, B), but when disked, disks rounded, never flattened 34 Contributions in Science ' No. 251 Figure 24. Distribution of Leptodactylus bokermanni (triangles) andL. hylaedactylus (circles). Solid circles indicate 1) a northern homogeneous population and 2) distinctive heterogeneous grouping of localities. Open triangles indicate distinctive southern populations ofL. bokermanni. See text for further details. (Fig. 5B); toes free; subarticular tubercles moderately developed; line of tubercles forming a metatarsal fold or absent; weak, sinuous or almost straight, tuberculate, tarsal fold extending % to % length of tarsus; lower tarsus and sole of foot scattered or profused with white tipped tubercles; female size 24.2 ± 2.5 mm S-VL, 27.6 mm maximum, male size 23.2 ± 1.2 rnm S-VL, 25.1 mm maximum; femur 41.0 ± 2.6% S-VL; tibia equal to or longer than femur, 44.0 ±1.7% S-VL; foot longer than femur or tibia, 50.2 ± 2.1% S-VL; dorsal pattern commonly uniform (Fig. 2E) or with indistinct marks (Fig. 2C), rarely with distinct spots (Fig. 2B), most individuals lacking dorsolateral stripes, rare individuals with narrow or broad light stripes from eye to groin, narrow, light mid-dorsal stripe from anus not extending past sacral region or absent; venter immaculate or scattered with melanophores, not distinctly mottled; posterior face of thigh mottled to uniform. 1973 Systematics of Marmoratus Group 35 Larvae: Nostril nearer tip of snout than eye; distance between nares about equal to or just less than interorbital distance; eye moderate, 10.5 ± 1.2% body length; mouth- parts subterminal; oral papillae in single row; oral disk entire; oral disk width 27.0 ± 1.8% body length; anterior oral papilla gap 77.8 ± 6.6% oral disk width; tooth row formula or denticles in posteriormost row weakly or not developed; beak mod- erately developed, spiracle sinistral, halfway up body, % distance from tip of snout to anus; dorsal fin origin at body-tail juncture; tail height just greater than body height; tail tip elongate, rounded to pointed; anal tube median; dorsum dark, uniform gray to brown, in preservative; venter with profusion of melanophores on throat region, no melanophores on belly or anal tube; few or no melanophores on ventral tail fin, rest of tail mottled; total length largest specimen, stage 30, 13.5 mm; body length 31.4 ± 1.0% total length. Geographic variation — Specimens from Guanabara, Minas Gerais, Rio de Janeiro and Sao Paulo differ from other specimens in having more dorsally directed eyes and triangular shaped bodies. Within this area, the northernmost specimens from Belo Horizonte, Minas Gerais are distinctive in that none of the individuals have distinct, small dark spots, many individuals are finely mottled, and some specimens have C category disks. The southernmost demes from the state of Parana are distinctive from the remaining populations in that many of the individuals have broad, light dorsolateral stripes. Distribution (Fig. 24) — Range: Southeastern Brasil from Minas Gerais to Parana, sea level to 1500 m. Etymology — The species is named for Werner C. A. Bokermann in recognition of his contributions to the study of frogs of the genus Leptodactylus . Leptodactylus hylaedactylus (Cope) Figure 25, 26 Cystignathus hylaedactylus Cope, 1868:115 (Type locality. Probably Peru, Napo or upper Maranon. Holotype ANSP 2240, male). Leptodactylus diptyx Boettger, 1885:32-33 (Type locality, Paraguay. Lectotype BMNH 1947:2.17.47, male). Leptodactylus glandulosus Cope, 1887:52-53 (Type locality, Brasil: Mato Grosso, Chupada, 30 mi NE Cuyata, nr. headwaters of Xingu. Lectotype ANSP 10275, juvenile). Leptodactylus hololius Boulenger, 1918:430-431 (Type locality, Peru: Rio Maranon, Pebas. Holotype BMNH 1915.3.9.13, female). Leptodactylus minutus Nobel, 1923:295-297 (Type locality, British Guiana, Bartica District. Holotype AMNH A- 13495, female). Leptodactylus poeppigi Melin, 1941:59-60, Fig. 33 (Type locality, Peru: San Martin; Roque. Holotype Ba. Ex. 507, female). Leptodactylus melini Lutz and Kloss, 1952:639-640 [Substitute name for Leptodactylus rugosus Melin, 1941:58-59, Fig. 32] (Type locality, Brasil: Amazonas; vicinity of Manaos. Holotype Ba. Ex. 506). 36 Contributions in Science No. 251 Diagnosis — The other species of the Marmoratus group in addition to L. hylaedactylus with toe tips not developed into flattened disks areL. bokermanni andL. L martinezi. L. hylaedactylus does not have four longitudinal rows of symmetric dark [ spots as does L. martinezi. Many individuals of L. hylaedactylus have a mid-dorsal |j stripe extending from above the anus to the middle of the body or the snout, no L. j bokermanni do; L. hylaedactylus larvae lack a spiracle and denticles, L. bokermanni larvae have a spiracle and denticle rows. Summary of characteristics — Adults: Dorsal outline of snout subovoid or subel- liptical in females, subelliptical or pointed in males; snout rounded in profile in females, rounded or rounded-acute in males; female snout ratio .672 ± .052, male snout ratio .723 ± .065; canthus rostralis indistinct; loreal very slightly concave or obtuse in cross section; tympanum distinct, greatest diameter equal to, just less than, or just greater than Vi diameter of eye; male vocal slits elongate, very slightly oblique to jaw or parallel to jaw; vocal sac single, internal, or a fold along margin of jaw to forearm; vomerine teeth in obtuse to transverse (usually) series, posterior to choanae; head length 38.0 ± 1.6% S-VL; head width less than length, 34.3 ± 1.4% S-VL; interorbital distance 8.8 ± .8% S-VL; length of the first finger equal to, just greater | than, or just less than length of second finger; dorsum smooth or with ridges and warts, j rarely with dorsolateral folds, most males with white-tipped tubercles profused over | posterior dorsum, females without such tubercles; females without distinctive glands, males lacking distinctive glands or with paired, small oval glands at angle of jaw, just i in back of axilla, in groin, and on either side of anus on hindlimb; toes usually with tips I expanded, but not distinctly disklike (Fig. IB), some toes lacking expanded tips (Fig. lA) or with distinct disks (Fig. 1C), if with distinct disks, rounded, not flattened (Fig. 5B); toes free; subarticular tubercles moderately developed or pronounced; line of tubercles forming a metatarsal fold or absent; weak, sinuous or almost straight tarsal fold extending % to % length of tarsus, smooth or with white-tipped tubercles; lower tarsus and sole of foot scattered or profused with white-tipped tubercles; female size 23.2 ± 2.1 mm S-VL, 31.0 mm maximum, male size 22.7 ± 1.5, 26.2 mm max- imum; femur 41.0 ± 2.7% S-VL; tibia equal to or (usually) greater than femur, 44.6 ±2.3% S-VL; foot longer than tibia or femur, 50.0 ± 3.4% S-VL; dorsal pattern commonly uniform (Fig. 2E), with indistinct marks (Fig. 2C), or distinctly spotted (Fig. 2B) rarely blotched (Fig. 2D), most individuals lacking light dorsolateral stripes, some with a narrow, light dorsolateral stripe, rarely with broad, light dorsolateral stripes, light mid-dorsal pin stripe from snout to above anus to absent, rarely with a broad light stripe from tip of snout to above anus to sacral region; throat and belly immaculate or scattered with melanophores, ventral edges of throat, belly, femur, tibia (or entire surface) scattered with melanophores, usually not distinctly mottled, some males with black vocal folds; posterior face of thigh mottled to uniform. Larvae: Nostril midway between eye and tip of snout or nearer snout; distance between nares equal to interorbital distance; eye large, diameter 15.2 ± 1.6% body length; mouthparts subterminal; oral papillae in single row; oral disk entire; oral disk width 18.0 ± 1 .6% body length; anterior oral papilla gap 82.5 ± 5.8% oral disk width; no denticles on tooth rows; beak weakly developed; no spiracle; dorsal fin origin at body-tail juncture, or origin on tail, posterior to body-tail juncture; tail height less 1973 Systematics of Marmoratus Group 37 o ^ ^OQOO OOOo oo Figure 25. Lateral view and mouthparts of larva of Leptodactylus hylaedactylus . Upper line equals 1 cm, lower line equals 1 mm. than, equal to, or greater than body height; tail tip elongate and rounded to pointed; anal tube median; dorsum with uniform pattern, gray to brown in preservative; venter with suffusion of melanophores on throat region, few melanophores on belly, no melanophores on anal tube; no melanophores on tail fins, scattered melanophores on upper two thirds of tail musculature; total length largest specimen, stage 34, 13.2 mm, stage 40, 12.6 mm; body length 36.7 ± 3.1% total length. Geographic variation — The northernmost populations (Venezuela, Guyana, Surinam, French Guiana, and Amapa (part), Brasil) are distinctive from the remaining populations in having a high frequency of dorsal spotting (B category), a high fre- quency of narrow dorsolateral stripes that do not reach the eye (B category), and no instances of broad mid-dorsal stripes (E or G category). Andean slope populations from Peru and Bolivia are relatively homogeneous and are not distinctive from other populations. Populations from the Brasilian state and territory of Acre and Rondonia and from the state of Mato Grosso are distinctive both from other samples and among themselves (see analysis of form II specimens). Distribution (Fig. 24) — Range: The species occurs from northeastern South America, central and southern Amazonia and south central Brasil. The known eleva- tional range is sea level to 500 m. Dr. Avelino Barrio has recently collected material of the Marmoratus group from northern Argentina and kindly allowed me to examine the specimens. Morphologically, they are L. hylaedactylus. Dr. Barrio is preparing a paper on the specimens describing their morphology, mating call, ecology, and karyotype. Localities — BOLIVIA. Beni: Upper Beni, below Huachi, UMMZ 74817; Cachuela Esperanza, 150 m, UMMZ 67738; Huachi, 250 m, MCZ 12900-1, 15578-81, UMMZ 64110-1, 64114; Ivon, UMMZ 74818; Lake Rogoagua, 200 m, UMMZ 64115; Puerto Almacen, 260 m, AMNH 72380; 4-5 km from confluence of Rio Blanco and Rio Itenez, AMNH 79093; Rio Marmore, nr. Guajara-Mirim, 160 m. 38 Contributions in Science No. 251 Figure 26. Audiospectrograms of L. hylaedactylus . Upper call from specimen WCAB 15091, Brasil, Mato Grosso, Sto. Antonio Le verger, temperature 29°C; middle call from specimen WCAB 15139, Brasil, Goias, Jatai, temperature 26°C; lower call probably from specimen WCAB 15550, Brasil, Mato Grosso, Chapada dos Guimaraes, temperature 19°C. 1973 Systematics of Marmoratus Group 39 CM 2665; Trinidad, AMNH 79091-2. Cochabamba: Sta. Elena, UMMZ 64112. La Paz: Lower Bopi, UMMZ 64113. Santa Cruz: Buenavista, 500 m, AMNH 34012, 34016, 34072, MCZ 12898-9, 15582, 15584, UMMZ 64031A, D-I, K-P, 64032A, 64033A-C, 64034, 64035A, B, 64036A-E, 66476, 66493E, G, 66494A, B, D, 66501. BRASIL. Acre: Feijo, 249 m, WCAB 1263; Placido de Castro, DZ 6521, 6522, 6524; Tarauaca, WCAB 1387, 1389-90, 2513, 2855. Amapd: Mazagao, WCAB 19136-7, 35966; Oiapoque River, MCZ 25722, USNM 11511-2, 115514, WCAB 5614; Rio Tracajatuba, WCAB 19006; Serra do Navio, 220-300 m, WCAB 35243-4. Amazonas: Benjamin Constant, DZ 24890-92, 24893-6; Ducke Reserve, near Manaus, KU 129930-3; Itapiranga, DZ 27756; Manaus, AMNH 51756-9 (specimens referred to this species, although poor condition precludes positive identification); Ponta Negra, Negro River, DZ 24879-84; Puraquequara River (mouth) affl. Amazonas, DZ 24875-6. Golds: Jatai, WCAB 15139. Mato Grosso: Chapada dos Guimaraes, 800 m, WCAB 15550; Porto Murtinho, UMMZ 76081 (specimen referred to this species, although poor condition precludes positive identification); Sto. Antonio Leverger, 100 m, WCAB 15091-2; Urucum, S. of Corumba, FMNH 9180-1, 9203-4, 9237. Pard: Belem, 20 m, WCAB 2014; Cachimbo, 500-600 m, DZ 21938; IPEAN, 3 km E Belem, KU 127339, 127353, 127356, 127358, 127365, 127370, 127376, 127378-9, 127391; Obidos, KU 129929. Pernambuco: Recife, DZ 25027. Rondonia: GuajaraAssu Falls, FMNH 64238; Igarape Marmelo, WCAB 9846-51, 9854-5, 9859; Nhambiquara, WCAB 13279; Porto Velho and vicinity, 60 m, DZ 16350-1, UMMZ 64119D, F, WCAB 10709, 31721-3, 34103. COLOMBIA. Amazonas: Leticia, 275 m, KU 124743. FRENCH GUIANA. Cacao, LG 124; Cayenne, sea level, LACM 44280, LG 34; Crique Gabrielle, LG 126; Regina, 25 m, LACM 42082-3 (larvae), 44282-378; Roura, LG 877. GUYANA. Demerara: Dunoon, 20 m, UMMZ 52506. Essequibo: Kartabo, 10 m, AMNH 39671, 39696; Marudi, 250 m, AMNH 46565-8 + 4; Membaru Creek, upper Mazaruni River, UMMZ 85148-9; Yarikita River, mouth and Haul Over, UMMZ 83581-2. PERU. Amazonas: Mamayacu, Rio Cenipa (trib. of Marahon), AMNH 42414. Huanuco: Divisoria, FMNH 56318; Hac. Pampyacu, MCZ 22810-9, 82199-204, UMMZ 83093(4); Tingo Maria, WCAB 36498. Junin: Rio Perene, MCZ 22878-81. Loreto: Cumaria, AMNH 42799; Igarape Champuia, Curanua Riv., DZ 10343-8; Iparia, MCZ 75032, 75034; Marahon, mouth of Rio Pastaza, 150 m. AMNH 42228; Parinari (Marahon), AMNH 43437; Pucallpa, Rio Ucayali, FMNH 56321; Rio Pisque, AMNH 43548, 43561; San Regis, on Rio Marahon, AMNH 42059, 42487; Suhuaya, nr. Contamana on Rio Ucayali, AMNH 42075. Pasco: Iscozazin Valley, Pan de Azucar, 380 m, LACM 40651-53, 40655-6, 40658-9; Oxapampa, Nevati Mission, USNM 166769-70. SURINAM. Marowijne: Langaman Kondre, DZ 24757. Nickerie: Sipaliwini, RM 16719, 16721, 16723, 16755. Suriname: Brownsweg, RM 16734; Lelydorp, RM 16731; Paramaribo, AMNH 77464, MCZ 17925, RM 16732-3, 16735, 16738. VENEZUELA. Monagas: Caripito, ± 50 m, AMNH 70654-64, USNM 117090-1. 40 Contributions in Science No. 251 martinezi (squares). Solid circles indicate two homogeneous population units ofL. andreae, one along the Andes, the other of North Central South America. Open triangle indicates distinctive island population of L. marmoratus. Leptodactylus marmoratus (Fitzinger in Steindachner) Figure 28 Adenomera marmorata Fitzinger in Steindachner, 1867:37, Taf. Ill, Figs. 5-8 (Type locality, Brasil. Holotype Vienna 16453, male). Leptodactylus nanus MUller, 1922: 168-179, Figs. 4-6 (Type locality, Rio Novo, Santa Catarina, Brasil. Lectotype Munich 661/1920, a female). Leptodactylus trivittatus Lutz, 1926:151, Plate 32, Figs. 14, 15 (Type locality, Campo Belo, Alto da Serra De Cubatao, Brasil. Lectotype USNM 96943, a juvenile). Diagnosis — The only other species in the Marmoratus species group in addition 1973 Systematics of Marmoratus Group 41 Figure 28. Audiospectrogram ofL. marmoratus, Brasil, Guanabara, Tijuca, temperature 22°C. No specimen. to L. marmoratus with flattened toe disks is L. andreae. No individuals of L. mar- moratus have dorsal longitudinal stripes, someL. andreae have such stripes; someL. marmoratus have light mid-dorsal pin stripes extending from above the anus to the snout or shoulder region, almost noL. andreae have the pin stripe, if present, extend- ing beyond the sacral region. Summary of characteristics — Adults: Dorsal outline of snout nearly semicircular, rounded, subovoid, or subelliptical in females, subovoid, subelliptical, or pointed in males; snout rounded in profile in females, rounded or rounded-acute in males; female snout ratio .687 ± .046, male snout ratio .712 ± .052; canthus rostralis indistinct; loreal slightly concave or obtuse in cross section; tympanum distinct, equal to or just less than V2 diameter of eye; male vocal slits elongate, slightly oblique to jaw or parallel to jaw, vocal sac single, internal, or a fold along margin of jaw to forearm; vomerine teeth in transverse (usually) or slightly obtuse series posterior to choanae; head length 38.4 ± 2.2% S-VL; head width less than length, 33.6 ± 1.1% S-VL; interorbital distance 9.0 ± 0.8% S-VL; length of first finger equal to, just greater than, or just less than length of second finger; dorsum smooth or with white-tipped tubercles on posterior portion in individuals of both sexes, ridges or warts, or rarely with distinct dorsolateral folds; small, round or oval paired glands present at angle of jaw and on either side of anus on hind limb present or absent; toes usually with distinct, flattened disks (Figs. 1C, D; 5 A); toes free; subarticular tubercles moderately de- veloped; weak line of small white-tipped tubercles forming a metatarsal fold or absent; weak, almost straight tarsal fold extending % — % length of tarsus with series of very small tubercles; lower surface of tarsus and sole of foot profused with white-tipped tubercles; female size 21.9 ± 1.7 mm S-VL, 27.9 mm maximum, male size 20.8 ± 1.8 mm S-VL, 26.0 mm maximum; femur 40.4 ± 3.0% S-VL; tibia equal to or (usually) longer than femur, 44.2 ± 2.2% S-VL; foot longer than either femur or tibia, 49.5 ± 2.7% S-VL; dorsal pattern commonly uniform (Fig. 2E) or with indistinct marks (Fig. 2C), rarely distinctly spotted (Fig. 2B) or blotched (Fig. 2D), very rarely lined (Fig. 2A), most individuals lacking light dorsolateral stripes, a few with narrow stripes, several with well defined light stripes from eye to groin (Fig. 12C), light 42 Contributions in Science No. 251 mid-dorsal stripe ranging from present from tip of snout to anus to absent; throat scattered with melanophores or center of throat and belly immaculate, ventral edges of throat, belly, femur, and tibia mottled; posterior face of thigh mottled to uniform. Geographic variation — Mainland populations appear to form a north-south cline with respect to size and mid-dorsal stripes. The larger individuals are in the north (maximum male S-VL about 24 mm, female S-VL about 25 mm) the smallest indi- viduals are in the south (maximum male S-VL about 19.5 mm, female S-VL about 20 mm). No individuals from the north (Guanabara, Rio de Janeiro) have light mid-dorsal pin stripes extending beyond the sacral region anteriorly. In Sao Paulo, some individu- als have light mid-dorsal pin stripes extending to midbody (B category). In Santa Catarina some individuals have B category mid-dorsal stripes as well as some indi- viduals having a continuous pin stripe from the snout to the vent (C category). The deme on Alcatrazes Island is distinctively larger than any other island or mainland population (male S-VL to 26 mm, female S-VL to 27.9 mm). Distribution (Fig. 27) — Range: Southeast Brasil from the states of Guanabara and Rio de Janeiro to the state of Santa Catarina. Known elevational range, sea level to 850 m. Localities — BRASIL. Guanabara: Rio de Janeiro, Cosme, USNM 119000-3; Jacarepagua, USNM 97472-85; Sumre, 850 m, WCAB 12984; Tijuca, 400 m, AMNH j 36257, MCZ 15846, UMMZ 68792, 104275, 104278, USNM 81132, 96300-2 + 10, , WCAB 14342, 16641, 16643-4, 16646, 16932, 17357, 18943, ?>056A. Parana: Volta Grande, USNM 125505, \2553\-2. Rio de Janeiro: Angra dos Reis, USNM 70591-2, 96519; Barro Branco, UMMZ 104263, USNM 133009; Grande Island, DZ 25101; Guanabara, DZ 7, 8, 13, 22, 27, 35, 38, 148; Itatiaia National Park, DZ 4130, Vienna 4490; Petropolis, USNM 96417, 96430, Vienna 15802, 19413:1, 2; Pico de Tijuca, USNM 133010; Represa Rio Grande, Guanabara, 50 m, WCAB 7589-643, 9380, 22727, 14120-21; Rio de Janeiro, USNM 97235; Teresopolis, USNM 97682. Santa Catarina: Rio Humboldt, AMNH 15569-70, FMNH 6473, MCZ 8841-2, 11680, 19510-9, 12903, 82205-10, USNM 66583, 118177-8; Rio Vermelho, WCAB 3416-7, 5105; Sao Bento do Sul, WCAB 6104. Sao Paulo: Alcatrazes Island, DZ 24163, 24172, 24175, 24177, 24182, 24205-6, 24231-2; Alta da Serra de Cubatao, USNM 97858; Boracea, USNM 129175; Buzios Island, DZ 24140; Caraguatatuba, DZ 24291; Cubatao, 800 m, DZ495, 1972, WCAB 2425-6; Est. Biol. Boraceia, Salesopolis, DZ 2733, 2745-6, 3373, 3375, 24293; Itanhaem, DZ 626, 10931; Itapetininga, DZ 25430-1; Paranapiacaba, 800 m, DZ 10783-4, WCAB 6581, 6849, 11957; Piassa- guera, DZ 771, 914, 2041, 2042, 2043; Rio Grande, DZ 658; Sao Paulo, 800 m, DZ 4218, 6386, 6469, 9372, 10578-80, 15662, WCAB 45104-6; Sao Sebastiao Island, 50 m, CM 33439, DZ 6408-19, 24276-83, FMNH 67257-60, KU 74217-21, WCAB 4907-8, 8210-20, 8222-6, 8821-30, 8898-900, 9389, 13643-53; Sao Vicente, 5 m, DZ 4155, WCAB 7316. Leptodactylus martinezi Bokermann Leptodactylus martinezi Bokermann, 1956: 37-40, Figs. 1-4 (Type locality Cachimbo, Para, Brasil. Holotype WCAB 71, male). Diagnosis — Leptodactylus martinezi is the only species in the Marmoratus 1973 Systematics of Marmoratus Group 43 species group that has four symmetrically arranged rows of longitudinal spots. The dark spots of other species, if present, are never thus arranged. Summary of characteristics — Adults: Dorsal outline of snout subovoid or subel- liptical in females, subelliptical or pointed in males; snout profile rounded; female snout ratio .736 ± .071, male snout ratio .95; canthus rostralis indistinct; loreal very slightly concave or obtuse in cross section; tympanum distinct, greatest diameter equal to or just less than Vi diameter of eye; male vocal slits elongate, very slightly oblique to jaw; vocal sac single, internal; vomerine teeth in transverse series, posterior to choanae; head length 39.4 ± 1.6% S-VL; head width less than length, 32.8 ± 2.1% S-VL; interorbital distance 8. 1 ± .9% S-VL; length of first finger equal to or just greater than length of second finger; dorsum smooth or warty, white-tipped tubercles posteriorly in a few individuals of both sexes; much of dorsal and lateral surfaces glandular appearing, but no distinct glands; toe tips pointed or slightly swollen (Fig. lA, B); toes free; subarticular tubercles moderately developed or pronounced; line of white-tipped tubercles forming a metatarsal fold or absent; tarsal fold weak, sinuous or almost straight, extending % to % length of tarsus, tuberculate; female size 22.3 ± 1.1 mm S-VL, 24.5 mm maximum, male size 23. 1 mm, 23.4 mm maximum; femur 39.3 ± 3.6% S-VL; tibia longer than femur, 46.0 ± 1.2% 5-VL; foot longer than tibia, 50.8 ± 1.8% S-VL; dorsal pattern of symmetrically arranged distinct dark spots in four longitudinal rows, light, thin mid-dorsal stripe from tip of snout to above anus, no light dorsolateral stripes; ventral surfaces scattered or profused with melanophores, not mottled; posterior face of thigh mottled. Distribution (Fig. 27) — Range: L. martinezi is found in Central Brasil. The known elevational range is 200-600 m. Localities — BRASIL. Goids: Sta. Isabel, Bananal Island, DZ 25325. Mato Grosso: Mato Verde, DZ 25320-1, 25242-9, 25322-4; Sao Domingos, Rio das Mortes, 200 m, DZ 25216. Para: Cachimbo, 500-600 m WCAB 71-2. An Artificial Key to the Adults of Members of the Marmoratus Group The key is provided only as an aid to identification. Because of the subtle differ- ences separating the species, extensive use of the analysis section must be used for verification. lA. Dorsal pattern of four longitudinal rows of symmetrically arranged dark spots L. martinezi IB. Spots, if present on dorsum, never symmetrically arranged in four rows ... 2 2A. Toe tips pointed or expanded; if disked, disks are round, not flattened .3 2B. Toe tips expanded into flattened disks 4 3A. No individuals with a light mid-dorsal stripe extending from above the anus to the middle of the body; southeast Brasil L. bokermanni 3B. Many individuals with light mid-dorsal stripes extending from above the anus to the middle of the body or the snout; northeastern South America, central and southern Amazonia, south central Brasil L. hylaedactylus 44 Contributions in Science No. 251 4A. Some individuals with a striped dorsal pattern; almost no individuals with light mid-dorsal pin stripe extending from above the anus to beyond the sacral region; north, central, and western South America east of the Andes L. andreae 4B. No individuals with striped dorsal patterns, some individuals with light mid-dorsal pin stripe extending from above the anus to beyond the sacral region; southeast Brasil L. marmoratus RELATIONSHIPS One member of the Marmoratus group is clearly distinctive from the others: L. martinezi. The remaining four species form a closely related complex. The working hypothesis of the previous analysis was that the allopatric populations form species pairs: L. hylaedactyhis and bokermanni; andreae and marmoratus . The distribution of toe tip character states supports this arrangement (Fig. 29D). The distributions of other character states agree in part only or not at all, however: 1) Dorsal pattern. L. boker- manni and hylaedactylus are similar in sharing states B, C, and E, L. andreae and marmoratus are similar in sharing states C, D, and E. State A is found in some L. andreae, only oneL. marmoratus examined approaches state A (Fig. 29A). State B is extremely rare inL. andreae, more common in marmoratus (Fig. 29D). More interest- ing is that no L. bokermanni have state D patterns, while state D is rarely present in hylaedactylus (Fig. 29E). 2) Dorsolateral states. All four species have states A and B. Leptodactylus hylaedactylus does not have the typical state C category. The few light dorsolateral stripes that are present inL. hylaedactylus extend from the shoulder region to the groin. Leptodactylus bokermanni, andreae, nnd marmoratus have the C state (Fig. 29C). 3) Mid-dorsal stripes. Leptodactylus bokermanni and andreae share state A, D, and F, L. hylaedactylus and marmoratus share states A, B, C, D, and F (Fig. 29B). The composite of all the characters is a mosaic which does not provide any clear indications of relationships among the four species. In the section, recognition of “non-dimensional” species, a single instance of apparent hybridization was noted between L. hylaedactylus and L. andreae. Hybridization data are difficult to interpret. In fact, one could interpret the data as either implying thatL. hylaedactylus andL. andreae were the most closely related species pair or the most distantly related species pair. If the former were true, then the four species would represent an interesting case of parallel speciation involving similar adaptive types. Further speculation should await the availability of more data, particularly from life histories and karyotypes. Distribution The forests of southeast Brasil contain two species, L. bokermanni and marmoratus . Two species are found in Amazonian and Orinocoan Forests, L. andreae and hylaedactylus. It appears thatL. andreae is confined to these forest regions. The only exception is a single possibly erroneous record from forest west of the Andes. Leptodactylus hylaedactylus also occurs in the drier forests of southern Bolivia and Mato 1973 Systematics of Marmoratus Group 45 B andreae hylaedactylus marmoratus bokermanni bokermanni andreae hylaedactylus marmoratiis D Figure 29. Phenetic character trees for four species of the Marmoratus group. Also see text. Grosso, but the species is definitely excluded from the Chaco. A single specimen from Recife, Pernambuco, Brasil is frustrating. As far as habitat is concerned, the species should occur there. But why there should be only a single record available from the east Brasilian forest strip seems odd. The region has been visited by collectors and other species of Leptodactylus are known from there (Heyer, 1970a). The only species with a distribution confined to dry forests is L. martinezi in central Brasil. Leptodactylus andreae andL. marmoratus are found in two types of habitat: primary forest and house gardens (Marty Crump and William E. Duellman, personal communication. Lutz, 1947:248). These two types of habitats suggest that the frogs are limited by soil moisture. The soil must be sufficiently moist so that larvae do not desiccate within the chambers. The soil must not be flooded, however, as flooded soil would not permit the construction and maintenance of the chambers. It seems probable that the members of the Marmoratus group evolved in primary forests and have secondarily invaded gardens. Refugia — Comparison of the patterns of differentiation in the Marmoratus group with results of Vanzolini and Williams’ (1970) analysis of the Anolis chrysolepis species group indicates some striking parallels. Vanzolini and Williams demonstrated the existence of four core areas within the geographic distribution of A. chrysolepis. These core areas contain populations that are rather uniform, but the regions between core areas have heterogeneous populations. They reason that the core represented forest refugia during dry Pleistocene periods, the populations within each refugium became homogeneous and with more mesic climates, such as now, the species have spread out from the refugia. 46 Contributions in Science No. 251 Before comparing results of core area distributions, it is instructive to compare and contrast the distributions and evolutionary origins of the Anolis chrysolepis and i| Marmoratus species groups. Anolis chrysolepis occurs throughout northern Venezuela. ^ Members of the Marmoratus group are absent over much of northern Venezuela. The | Marmoratus group does not provide a test for Vanzolini and Williams’ core area III [j which covers this region. At the other extreme, Anolis chrysolepis does not occur in | southeast Brasil, so no information from that study is available for comparison. What | remains for comparison is Vanzolini and Williams’ core areas I, III, and IV. Members | of the Anolis chrysolepis group are relatively recent invaders of South America, with I their evolutionary origins in Middle America and Mexico. Members of the Mar- moratus species group are strictly South American, probably having undergone their total evolutionary history in situ. The distribution patterns of figures 24 and 27 are readily comparable with Vanzo- lini and Williams’ core areas. Leptodactylus andreae (Fig. 27) demonstrates the pres- | ence of two core areas, corresponding with core areas I (Surinam, French Guiana, and j Amapa, Brasil) and III (east Andean slopes of Colombia, Ecuador, and Peru) of Vanzolini and Williams. Leptodactylus hylaedactylus (Fig. 24) demonstrates clearly a core area corresponding with core area I of Vanzolini and Williams. The other region delimited in figure 24 is a heterogeneous area, which by default leaves the east Andean slope populations a homogeneous group, although it is not particularly distinctive in i any regard. As a homogeneous assemblage, the east Andean slope populations ofL. hylaedactylus correspond to core area III of Vanzolini and Williams. Vanzolini and Williams suggested that their core area III was actually a composite of three smaller core areas which have merged. The combined distributions of L. andreae and L. hylaedactylus substantiate that hypothesis. The absence of L. hylaedactylus from Ecuador and the east Andean slopes of Colombia is easiest explained if L. hylaedac- tylus was never present in the northern refugia within core area III. The distribution of L. martinezi coincides with core area IV of Vanzolini and Williams. The core areas deduced from detailed geographic analyses of the Anolis chrysolepis and Marmoratus groups coincide almost too perfectly. The validity of the core areas is attested to by the fact that organisms with different evolutionary origins have similar core area patterns. The critical similarity between the two groups of organisms is that they are both forest groups. Vuilleumier (1971), in reviewing the Pleistocene changes in fauna and flora of South America noted the relative goodness of fit between Haffer’s (1969) proposed forest refugia based on bird studies and Vanzolini and Williams’ (1970) proposed refugia. The results of this study are consistent with Vanzolini and Williams’ results and differ from Haffer’s in the same way as Vanzolini and Williams’ differed. Vuil- leumier (1971) also indicated that Muller (1968) had proposed a forest refugium in Southeast Brasil, the Serra do Mar refugium. Leptodactylus wagneri: A contrast — In a previous study (Heyer, 1970a), analysis of the geographic distribution of certain character states within L. wagneri, which is distributed throughout the same region as the Anolis chrysolepis group and along east coastal Brasil, did not have any patterns indicative of core areas or refugia. Rather, the only characters with significant geographic variation were clinal in nature. Members of 1973 Systematics of Marmoratus Group 47 the Marmoratus group differ fromL. wagneri in one significant ecological way. While the Marmoratus group members are forest forms, L. wagneri is more of a clearing or ecotonal species (Heyer and Beilin, 1973). Leptodactylus wagneri is associated with forests in its distribution pattern, but within the general forested regions it is found in clearings, or to be more precise, in the most arid microhabitats within the forests (Heyer and Beilin, 1973). ThusL. wagneri by the nature of its ecology should not be restricted to forest refugia during xeric periods, but in fact, its population should be maximum at such times providing the climate is not too extreme. Ecological and Evolutionary Implications Ecological data are scarce for all species. Heyer and Silverstone (1969) indicated that in French Guiana, L. hylaedactylus and andreae (reported as marmoratus) were ecologically distinct. Collections from the localities where L. bokermanni and marmoratus or L. hylaedactylus andL. andreae have been taken together are sugges- tive that ecological differences are found thoughout the ranges of the species pairs. With a single exception, one species is common, the other rare at sympatric sites. This may well be due to the bias of individual collectors, collecting one habitat type more thoroughly than another. Lutz (1947) has discussed the terrestrial life history of L. marmoratus (as L. nanus and trivittatus) . The eggs are placed in terrestrial chambers, hatch, and metamorphose without leaving the chamber. The yolk stores are sufficient to carry the larvae through metamorphosis without the necessity of feeding. Unfortunately, Lutz did not comment on the morphology of the larvae. Heyer and Silverstone in describing the larvae ofL. hylaedactylus (1969) suggested i\\2H hylaedactylus probably do not feed as larvae as in marmoratus because the larvae had large yolk stores, degenerate mouthparts, and no spiracle. The larvae of L. bokermanni appear to have functional mouthparts (although the denticles are at best weakly developed) and a spiracle is present. There are fewer yolk stores in the stage 30 larvae ofL. bokermanni than in the stage 40 larvae of L. hylaedactylus. In the L. bokermanni larvae the yolk is visible inside the few large coils of the intestine. The fact that the intestine is composed of few large coils filled with yolk at stage 30 is indicative that the individuals are still being nourished by the yolk. Later staged larvae may have to utilize additional food re- sources, however. A female L. bokermanni, WCAB 34465, had about 18 large eggs in the body. An attempt was made to determine the variation in clutch size from the museum materials, but it proved impractical. It is obvious that a female starts with 40 or 50 ova of small size. Some ova are reabsorbed while others grow and it is not possible to determine where the process stops in the preserved specimens. Exact data on clutch size will have to be gathered from nests in the field. The greatest number of large eggs examined was 18 in the L. bokermanni female mentioned above. The smallest number of large eggs examined was about 5 in a single female. There appears to be a considerable range in clutch size in the group. The species with the largest clutches may have larvae that are not as terrestrially adapted as the species with smaller clutch sizes (and concomitant larger individual egg size). At any rate, the presence of the less specialized L. bokermanni larvae as compared to L. hylaedactylus larvae 48 Contributions in Science No. 251 supports the earlier hypothesis (Heyer, 1969) that evolution towards terrestriality has occurred independently in the Marmoratus group on one hand and the other species groups of Leptodactylus on the other. I have previously suggested that the foam nest as found in members of the genus Leptodactylus has been in major response to arid conditions. This hypothesis is sup- ported by the ecology of such forms as L. wagneri and L. mystaceus which breed in temporary ponds in environments where more stable ponds also occur (Heyer and Beilin, 1973). The breeding biology of the Marmoratus group has been a reponse to veiy different environmental conditions. In the tropical rainforest (sensu lato) the temporary pond or stream environments are less predictable than in tropical dry forests (sensu lato). In dry forests, the rains may be sporadic, but usually occur at the same time of the year and the frogs are seemingly able to gauge the intensity of the rains, breeding at times of sufficient rainfall. In the rainforests the rains are more random and the frogs are not able to use them in a predictive manner. The terrestrial environment is more or less stable with respect to moisture throughout the year; if frogs were to become terrestrial, they could maintain breeding populations throughout the year. This is apparently what members of the Marmoratus group have accomplished through the mechanism of the foam nest. If these hypotheses are true, the foam nest of the Marmoratus group has evolved in response to mesic, unpredictable environments while the foam nest of the other Leptodactylus groups has evolved in response to xeric but predictable environments. Leptodactylus andreae-hylaedactylus and bokermanni-marmoratus occur sympat- rically over wide geographic areas. But there have doubtless been a series of many, separate encounters among the species pairs that occurred at the deme level. It is at the deme level, of course, where character displacement would operate to build premating isolating mechanisms between the species. Examination of the specimens from the several localities where two species occur together indicates that the displacement has not affected the same characters in the same ways. For example, at some localities there is a definite size difference between the species, but at other localities there does not appear to be a size difference. At some localities the toe disk development is distinctly different, at others it is not as nearly pronounced. In actuality, the mor- phological differences between the species pairs are so slight that they have been overlooked previously. One would expect the mating calls would be one of the first characters to diverge to maintain the species distinctness. But the divergence of mating calls has doubtless occurred many different times with many different demes with the recurrent pattern of expansion and contraction of ranges. Thus one might expect that the mating calls of the species pairs are distinct wherever the species occur sympatri- cally, but the characteristics of the mating calls within a species may demonstrate a complicated pattern when examined geographically. Resumen Se examinaron la distribucion geografica de ocho caracteristicas de tamano, forma, textura de la superficie de la piel, y coloracion de las especies del grupo marmoratus del genero anuro Leptodactylus . Una analisis de la distribucion geografica 1973 Systematics of Marmoratus Group 49 I de las caracteristicas supracitadas indica que el grupo abarca cinco especies. En i| Leptodactylus bokennanni, una nueva especie de los bosques del sureste del Brasil, las I puntas de los dedos no son dilatadas. En Leptodactylus hylaedactylus, que habita los bosques de Amazonia y el suroeste del Brasil, las puntas de los dedos no son dilatadas |j o son escasamente dilatadas. En Leptodactylus andreae, que se encuentra en los j bosques de Amazonia y la region del Orinoco, las puntas de los dedos son dilatadas en I la forma de discos. En Leptodactylus marmoratus, que se encuentra en los bosques : del sureste del Brasil, las puntas de los dedos son dilatadas en la forma de discos. I En Leptodactylus martinezi, que habita los bosques secos de la parte central del Brasil, i las puntas de los dedos no son dilatadas. Hay una diagnosis y descripcion de cada j especie y una clave para la identificacion. Se presenta la hipotesis de que el nido j de espuma del grupo marmoratus y el de los otros grupos del genero Leptodactylus se desarrollaron como resulta de distintas fuerzas de seleccion natural. En los otros grupos, el nido de espuma se desarrollo como resulta de un clima arido con fuertes Iluvias esporadicas. En tal clima, se puedeasegurar el exito de la reproduccion mediante utilizar la Iluvia como pronostico de condiciones favorables. En cambio en el grupo marmoratus es posible que la Iluvia no sea un buen pronostico de exito en la reproduccion. El ambiente de animales terrestres en bosques tropicales pluviales es relativamente estable en cuanto a la reproduccion. En el grupo marmoratus, el nido de espuma aparentemente se desarrollo como resulta de seleccion natural a favor de un modo de vida terrestre. Literature Cited Boettger, O. 1885. Liste von Reptilien und Batrachiem aus Paraguay. Z. Natur. 58:1-36. Bokermann, W. C. a. 1956. Sobre una nueva especie de Leptodactylus del Brasil (Amphibia). Neotropica 2:37^0. 1966. Lista anotada das localidades tipo de Anfibios brasileiros. Service de Documentagao, Sao Paulo. 183. pp. Boulenger, G. a. 1918. Descriptions of new South American batrachians. Ann. Mag. Nat. Hist. 9(2):427-433. Cope, E. D. 1868. An examination of the Reptilia and Batrachia obtained by the Orton Expedi- t tion to Ecuador and the upper Amazon, with notes on other species. Proc. Acad. Natur. Sci., Philadelphia, 1868:96-140. 1887. Synopsis of the Batrachia and Reptilia obtained by H. H. Smith, in the province of Mato Grosso, Brazil. Proc. Amer. Phil. Soc. 24:44—60. Gans, C. 1959. A taxonomic revision of the African snake genus Dasypeltis (Reptilia: Ser- pentes). Ann. Mus. Roy. Congo Beige. Sci. Zool. 74:1-237 and 13 plates. 1966. Studies on amphisbaenids (Amphisbaenia, Reptilia) 3. The small species from southern South America commonly identified as Amphisbaena darwini. Bull. Amer. Mus. Natur. Hist. 134:185-260. Haeeer, j. 1969. Speciation in Amazonian forest birds. Science 165:131-137. Heyer, W. R. 1968. Biosystematic studies on the frog genus Leptodactylus . Ph.D. Dissertation, Univ. Southern Calif. 234 pp. 1969. The adaptive ecology of the species groups of the genus Leptodactylus (Amphibia, Leptodactylidae). Evolution 23:421^28. 1970a. Studies on the frogs of the genus Leptodactylus (Amphibia: Leptodactylidae). VI. Biosystematics of the Melanonotus group. Los Angeles Co. Mus., Contrib. Sci. 191:1^8. 1970b. Studies on the genus Leptodactylus (Amphibia: Leptodactylidae). II. Diagnosis and distribution of Xhe Leptodactylus of Costa Rica. Rev. Biol. Tropical 16:171-205. 50 Contributions in Science No. 251 , AND M.S. Bellin. 1973. Ecological notes on five sympatrie Leptodactylus (Amphibia; Leptodactylidae) from Ecuador. Herpetologica 29:66-72. , AND P. A. SiLVERSTONE. 1969. The larva of the frog Leptodactylus hylaedactylus (Leptodactylidae). Fieldiana, Zool. 51:141-145. Levins, R. 1968. Evolution in changing environments, some theoretical explorations. Monogr. Population Biol., Princeton, 2:1-120. Lutz, A. 1926. Observagoes sobre Batrachios brasileiros. Parte 1: O Genero Leptodactylus Fitzinger. Mem. Inst. Oswaldo Cruz 19:139-157. Lutz, B. 1947. Trends towards non-aquatic and direct development in frogs. Copeia 1947:242-252. , AND G. R. Kloss. 1952. Anfibios anuros do alto Solimoes e Rio Negro. Apontamentos sobre algumas formas e suas vicariantes. Mem. Inst. Oswaldo Cruz 50:625-678. Melin, D. 1941. Contributions to the knowledge of the amphibia of South America. Gdteborgs Kungl. Vetenskaps och Vitterhets samhalles handlingar sjatte foljden. Ser. B 1:1-71. Muller, L. 1922. Uber eine Sammlung Froschlurche von Sta. Catharina nebst Beschreibung zweier neuer Arten. Blatter Aquarien-und Terrarien-Kunde 33:167-171. 1923. Neue oder seltene Reptilien und Batrachier der Zoologischen Sammlung des bayr. Staates. Zool. Anz. 57:39-54. Muller, P. 1968. Die Herpetofauna der Insel von Sao Sebastiao (Brazilien). Saarbriicker. 68 pp. Noble, G. K. 1923. New Batrachians from the tropical research station British Guiana. Zoologica 14:289-299. Parker, H. W. 1932. The systematic status of some frogs in the Vienna Museum. Ann. Mag. Nat. Hist. 10 (10):341-344. Peters, W. 1872. fiber die von Spix in Brasilien gesammelten Bratrachier des Konigl. Naturalienkabinets zu Miinchen. Monatsber. K.P. Akad. Wiss. Berlin, 18:196-227. Spix, J. B. 1824. Animalia nova sive species novae Testudineum et Ranarum, quas in itenere per Brasiliam, annis 1817-1820. Monaco. 53 pp. and 22 plates. Steindachner, F. 1867. Reise der osterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859. Amphibien. Kaiserlich-Koniglichen staatsdruckerei Wien. 70 pp. Vanzolini, P. E., and E. E. Williams. 1970. South American anoles: the geographic differen- tiation and evolution of XfioAnolis chrysolepis species group (Sauria, Iguanidae). Arq. Zool., Sao Paulo, 19:1-298. VuiLLEUMEiR, B. S. 1971. Pleistocene changes in the fauna and flora of South America. Science 173:771-780. Accepted for publication August 1, 1973 so 1.1 5 NUMBER 252 DECEMBER 21, 1973 SUBSPECIES OF THE SEA OTTER, ENHYDRA LUTRIS By Aryan I. Roest CONTRIBUTIONS IN SCICNCE NATURAL HISTORY MUSEUM • LOS ANGELES COUNTY SERIAL PUBLICATIONS OF THE !* NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Prior to November 30, 1973, publications of the Natural History Museum have appeailj under various formats — Leaflet Series, Museum Graphic, Science Series, Study Guides, C(i tributions in Science, Contributions in History, Science Bulletins, unnumbered catalogs [ exhibitions, and other miscellaneous publications. The Museum now publishes the follow | serials at irregular intervals as CONTRIBUTIONS IN SCIENCE, HISTORY BULLETII * SCIENCE BULLETINS, EDUCATION SERIES, HISTORY SERIES, and SCIENCE j ’ RIES. The Contributions are short papers of octavo size. The Bulletins are longer, comp hensive papers of quarto size. The Series are papers of variable lengths of quarto or lar | size. Papers in each serial are numbered separately and consecutively. | CONTRIBUTIONS IN SCIENCE contain articles in the earth and life sciences, present || results of original research. Emphasis is intended principally for papers allied to biosystemsS research, but other subjects and review-oriented ones will be considered. Number 1 was issi l on January 23, 1957. Contributions must be not less than 8 nor exceed 72 printed pages. | INSTRUCTIONS FOR AUTHORS , Acceptance of manuscripts will be determined by the significance of new information. Prior/j will be given to manuscripts by staff members. All manuscripts must be recommended by t I curator in charge of each discipline or by the Editorial Board. Manuscripts must conform\ the specifications listed below. They will be examined for suitability by the Editorial Bodi and will include review by specialists outside the Museum. | Authors must adhere to the International Code of Nomenclature of Bacteria and Virus j International Code of Botanical Nomenclature, and International Code of Zoological Nomei j clature, including their respective recommendations. Further, authors proposing new taxa? | a CONTRIBUTIONS IN SCIENCE must indicate that all primary types have been deposit j in an appropriate scientific institution of their choice and must cite that institution by nan| MANUSCRIPT FORM. — (1) In preparation of copy follow the 1972 CBE Style Manui ' third edition (AIBS), Chapters 5 and 6. (2) Footnotes should be avoided; acknowledgmeiljl as footnotes will not be accepted. (3) An informative abstract must be included for all pape|i (4) A Spanish summary is required for all manuscripts dealing with Latin American subjeql:l Summaries in other languages are not required but are strongly recommended. (5) A difftill ential diagnosis must accompany any newly proposed taxon. (6) Submit two copies of man j script. ILLUSTRATIONS.— All illustrations, including maps and photographs, will be referred . as figures. All illustrations should be of sufficient clarity and in proper proportions for redu j tion to CONTRIBUTIONS page size. In preparing illustrations and legends consult the 19'l CBE Style Manual, third edition (AIBS), Chapter 5. Submit only illustrations made wi permanent ink and glossy photographic prints of good contrast. Submit duplicate copies • all illustrations. Original illustrations will be returned after the manuscript has been publishe PROOF. — Authors will be sent galley proof which should be corrected and returned prompt! No changes or alterations, other than typesetting corrections, will be allowed unless paid 1 author. Requests for reprints may be placed through the Editor. Virginia D. Miller, Ediu All communications concerning CONTRIBUTIONS IN SCIENCE should be sent to tl < Editor, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, L( Angeles, California 90007 SUBSPECIES OF THE SEA OTTER, EN HYDRA LUTRIS^ By Aryan I. Roest^ Abstract: Two subspecies of Enhydra lutris are recog- nized: E. 1. lutris, found from the Commander Islands through the Aleutians to Prince William Sound, and with a disjunct popu- lation along the California coast; E. 1. gracilis, occurring in the Kurile Islands and extreme southern Kamchatka. Although the California population has some distinctive features, it represents the southern extreme of a dine which originally involved a con- tinuous distribution of otters along the entire Pacific coast of North America. The sea otter (Enhydra lutris) originally ranged along 6000 miles of the Asian and North American coastline, from northern Hokkaido to Baja Cali- fornia. Intensive hunting, for the fur trade, brought the species to the verge of extinction by the middle of the 19th century. In spite of its economic im- portance during this period, few specimens were obtained for scientific col- lections. Only recently have sufficient specimens become available to permit detailed taxonomic studies. Three subspecies of Enhydra lutris have been described, all based on relatively small samples. The sea otters of central Kamchatka, the Comman- der Islands, and the Aleutians are considered the nominate race, E. /. lutris (Linnaeus) by Barabash-Nikiforov (1947), Miller and Kellogg (1955), and Hall and Kelson, (1959). Barabash-Nikiforov (1947) re-established E. /. gracilis (Bechstein), on the basis of five specimens from Cape Lopatka, Kam- chatka. Merriam (1904) based his description of E. 1. nereis on a single skull from the coast of California. Scheffer and Wilke (1950) questioned the valid- ity of E. /. nereis after examining eight skulls from Washington, Oregon, and California; their conclusions were ignored by several later authors. Both of the two preceding references mention a size difference between Alaskan and Californian skulls, but present little data. The size of Aleutian otters was also discussed by Scheffer (1951), Lensink (1962), and Kenyon (1969). Miller and Kellogg (1955) recognized both E. 1. lutris (‘Kamchatka to western Aleu- tian Islands, Alaska’) and E. 1. nereis (‘Washington, Oregon, and California coasts’). Hall and Kelson (1959) also recognized both subspecies, but indi- cated that E. 1. lutris occurred along the Alaskan coast as far south as British Columbia, as well as in the Aleutians. Roest (1971) considered E. 1. nereis to be a valid form during the earlier stages of the present study. ^Review Committee for this Contribution Karl W. Kenyon G. Victor Morejohn Donald R. Patten ^Department of Biological Sciences, California Polytechnic State University, San Luis Obispo, California 93407; and Research Associate in Mammalogy, Natural History Museum of Los Angeles County, Los Angeles, California 90007 1 Contributions in Science No. 252 |. Full protection has allowed sea otter populations to increase during the ji past 50 years, and adequate series of specimens from Alaska and California! li are now available for study. The present work re-examines the status of sub-B species in the sea otter after a study of 267 Alaskan and Californian skulls,® ip all adults. Data from most of these specimens was subjected to modern sta- p tistical analyses. Alaskan specimens came from Amchitka and Adak Islands, in the Aleu- '( tians, and from the Alaska Peninsula, southern Alaska, and Prince William « Sound (the latter regions are collectively termed southwestern Alaska below). i Methods A skull was considered adult if the basioccipital-basisphenoid suture was H closed, since this suture closes at an age of about three years (Lensink, 1962). || Twenty-four measurements were taken on each skull, in millimeters. Total f length of many specimens was available, in centimeters, and was also used in I preliminary analyses. Weight, in pounds, was known for a number of Alaskan ; specimens, largely from animals weighed shortly after being collected. Only j; a few weights were available for California animals, mostly from animals | found dead on the beach for various undetermined periods of time. Since | weight is a characteristic which depends greatly on the condition of the animal I prior to death (Kenyon, 1969), it was not used in the analyses. However, j available weight data are presented in Table 1 . i Table 1. Weights of sea otters from four localities, in pounds Males Females Locality N Mean Range N Mean Range Adak 17 72.4 59-89 43 47.0 35-74 Amchitka 2 69.0 65-73 23 47.0 38-55 S. W. Alaska 12 76.9 47-96 1 49.0 — California 5 56.0 38-74 9 44.4 27-63 Stepwise Discriminant Analysis was used to analyze the data, the specific ; program being BMD07M, revised July 24, 1969 (Dixon, 1970). Preliminary , analyses indicated that the variables which discriminated (were statistically || significant) between different groups of specimens included frontal notch, projecting coronoid, nasal length, and condylobasal length. These four meas- Ji urements were selected for further analysis because they provided large F j values (Table 2) and were of potential taxonomic value. Due to the distinct differences in size between males and females, reflected particularly in con- i dylobasal length, sexes were analyzed separately. Total lengths were also || considered separately, but were excluded from the final analyses when it was 8 1973 Subspecies of the Sea Otter 3 Table 2 Statistical significance of the four variables submitted together for Stepwise Discriminant Analysis (BMD07M) ; sexes were analyzed separately Degrees of F value Variable N Freedom at step 1 MALES: Notch 106 3 102 52.5813 Nasal length 106 3 102 35.9030 Coronoid process 106 3 102 34.8241 Condylobasal length 106 3 102 21.1130 FEMALES: Coronoid process 142 3 138 30.2828 Nasal length 142 3 138 18.7986 Notch 142 3 138 17.3719 Condylobasal length 142 3 138 5.7635 learned that there were some differences in the manner in which they were taken. Each of these variables is discussed below. Frontal notch.— A distinct constriction in the frontal bones just anterior to the braincase was noted in most Alaskan skulls (Fig. 1). Skulls were classi- fied as to whether a notch was present (which was assigned a value of 3 for statistical analysis), only a suggestion of a notch was present (assigned a value of 2), or complete absence of a notch (value of 1). Table 3 indicates the occurrence of a frontal notch in four populations of sea otters. Projecting coronoid. —Merriam (1904) considered a backward sloping coronoid process a characteristic of E. 1. nereis, Scheffer and Wilke (1950) felt this distinction was of little value, since they found both sloping and up- right processes in their California sample. In the present study it was observed that in mandibles which had a strongly sloping coronoid process, the tip of the process projected posteriorly past the base of the condyloid process, and the posterior edge of the coronoid process was concave in outline. Some man- dibles had a coronoid process which extended posteriorly only as far as the base of the condyloid process; in these the posterior edge of the coronoid process was relatively straight. A third group had upright coronoid processes which did not project posteriorly at all, and in which the posterior edge of the process was convex (Fig. 2). Values were assigned to these variations as fol- lows: projecting coronoid— 3; coronoid equal to condyloid process— 2; upright coronoid— 1. The numbers and percentages of specimens which possessed, or lacked, projecting coronoid processes are presented in Table 4. Nasal length.— Length of the internasal suture line. Even in mature speci- mens the internasal suture line can usually be discerned. Table 5 presents data on the nasal lengths found in four different sea otter populations. Condylobasal length.— 'Distance from the anterior edge of the premaxilla to a line connecting the most posterior projection of the occipital condyles. Condylobasal lengths from five populations are presented in Table 6. 4 Contributions in Science No. 252 1973 Subspecies of the Sea Otter 5 Table 3 Occurrence of a frontal notch in sea otters from four localities. M/F indicates males/females Present Small Absent Locality N Number (M/F) % Number (M/F) % Number (M/F) % Adak 60 44 (12/32) 73.5 12 (5/7) 20.0 4 (0/4) 6.7 Amchitka 111 88 (33/55) 79.3 16 (2/14) 14.4 7 (3/4) 6.3 S.W. Alaska 41 5 (3/2) 12.2 25 (17/8) 61.0 11 (8/3) 26.8 California 43 3 (1/2) 7.0 7 (2/5) 16.2 33 (23/10) 77.0 Total: 255 Table 4 Projection of coronoid processes in sea otters from four localities. M/F indicates males/females Projects Equal No Projection Locality N Number (M/F) % Number (M/F) % Number (M/F) % Adak 60 11 (5/6) 18.3 6 (1/5) 10.0 43 (11/32) 71.6 Amchika 111 6 (2/4) 5.4 14 (9/5) 12.6 91 (27/64) 82.0 S.W. Alaska 37 11 (7/4) 29.7 25 (18/7) 67.5 1 (0/1) 2.7 California 43 34 (22/12) 79.0 7 (3/4) 16.2 2 (1/1) 4.7 Total: 25l Total length.— Length from the tip of the nose to the tip of the tail. The total lengths of specimens from four different regions are presented in Table 7. Total lengths of Alaskan and Californian otters in Table 7 are not directly comparable, since most of those from Alaskan specimens are curvilinear, while those from Californian animals are standard lengths. The difference in the two methods of measuring is probably not large, but there is little evidence to indicate its magnitude. One Alaskan specimen (AFG SO 72-13) was measured both ways; it had a curvilinear length of 76 cm and a standard length of 70 cm. Since male Californian otters have standard lengths which are about 20 cm shorter than the curvilinear lengths of male Alaskans, and the standard lengths of female Californians are 1 1 cm shorter than the curvilinear lengths of female Figure 1. Frontal notch. Upper — typical Aleutian skull with a distinct notch clearly visible in the narrowest part of the frontal region. Middle — Aleutian skull with a suggestion of a notch. Lower — typical Californian skull without a frontal notch; the frontal bones flare smoothly into the cranial region. All three skulls are from adult males. Note also the longer internasal suture of the California specimen (lower). 6 Contributions in Science No. 252 Figure 2. Coronoid process. Upper — typical Aleutian mandible with highest point of coronoid process anterior to the base of the condyloid process; posterior edge of coronoid process slightly convex. Middle — Aleutian mandible with suggestion of posterior projection of the coronoid process; posterior edge of coronoid process is essentially straight. Lower — typical Californian mandible with coronoid process projecting backward; posterior edge of coronoid process concave. Alaskans, a size difference between otters from the two regions probably exists. It is not as great, however, as the data in Table 7 suggests. During the early part of this study, very few specimens were available from southwestern Alaska. Comparisons of otters from the Aleutian Islands with those from California led to the conclusion that California otters could be readily distinguished from the Aleutian form, and hence deserved subspe- 1973 Subspecies of the Sea Otter 7 Table 5 Nasal length in sea otters from four localities, in millimeters Males Females Locality N Mean ± S.D. Range N Mean ± S.D. Range Adak 17 17.5 ± 1.16 15.0-19.6 43 16.9 + 1.64 13.2-20.8 Amchitka 38 16.1 ± 1.90 11.2-19.7 73 16.2 + 1.65 11.6-20.4 S.W. Alaska 24 19.6 ± 1.35 16.6-22.0 10 18.3 + 1.21 16.4-20.0 California 27 19.1 ± 1.04 16.7-21.1 16 19.0 + 0.98 17.2-20.8 Totals: 106 142 Table 6 Condylobasal length in sea otters from five localities, in millimeters. Commander Island data from Stroganov (1962) Males Females Locality N Mean + S.D. Range N Mean + S.D. Range Commander Ids. 17 136.5 130-140 1 125.4 120-128 Amchitka 38 135.5 ± 3.29 128-144 73 128.5 ± 3.69 115-136 Adak 17 137.8 + 3.71 129-144 43 128.2 + 3.83 121-136 S.W. Alaska 24 136.2 + 2.75 130-142 10 125.6 + 2.32 122-134 California 27 130.4 3.90 124-136 16 125.0 ± 2.61 119-130 Table 7 Total length of sea otters from four localities, in centimeters. Measurements of Alaskan otters are curvilinear lengths; those of Californian otters are standard lengths. Curvilinear lengths may be 5 to 15 cm longer than standard lengths Males Females Locality N Mean + S.D. Range N Mean + S.D. Range Adak 17 146.5 + 6.85 135-153 43 131.8 ± 6.08 118-145 Amchitka 2 146.0 ± 2.83 144-148 23 131.2 ± 4.99 121-140 S.W. Alaska 12 145.6 7.08 131-161 3 121.3 + 6.13 117-130 California 11 127.1 + 6.19 117-137 11 118.7 ± 6.03 106-127 Totals: 42 80 cific recognition (Roest, 1971). Subsequently additional material from south- western Alaska became available in November, 1972 and has been included in the final analyses presented here. Barabash-Nikiforov (1947), in discussing subspecies in the sea otter, presented some data in terms of percentages of the total length of the skull. Among the measurements he used were condylobasal length, zygomatic width 8 Contributions in Science No. 252 (width across zygomatic arches), mastoid width (width between mastoid processes), infraorbital width (width between infraorbital foramina), and postorbital width (width between postorbital, or supraorbital, processes). These measurements were also taken on specimens examined in this study. To permit comparisons, all measurements were recalculated as percentages of condylobasal length. Condylobasal length was used instead of total skull length in order to eliminate variation due to the size of the lambdoidal crest in older specimens. The results of these calculations are presented in Table 8. Table 8 Skull ratios of sea otters from various localities, expressed as percentages of con- dylobasal length. ZW = zygomatic width; MW = mastoid width; 10 = infraorbital foramen width; PO = postorbital width; CBL m condylobasal length. Russian local- ities and “Aleutians” are data recalculated from Barabash-Nikiforov, 1947 Locality N Range ZW/CBL N Range MW/CBL N Range lO/CBL N Range PO/CBL S. Kamchatka (C. Lopatka) 3 85.9-86.5 4 80.5-83.5 4 37.6-38.8 4 39.9-41.7 Mednyi Island 4 74.7-77.6 4 71.8-74.7 5 32.2-34.8 5 30.2-36.3 Bering Island 1 75.2 1 75.2 1 35.3 1 36.1 Commander Id. 2 71.5-72.0 1 72.0 2 32.8-34.6 2 32.4-33.6 “Aleutians” 5 74.7-80.6 5 73.1-78.0 5 32.2-34.6 5 29.4-35.2 Adak 10 74.3-83.2 10 68.5-76.5 10 32.4-35.3 10 31.6-35.1 S.W. Alaska 18 74.3-80.4 18 72.8-78.8 18 31.2-35.2 18 30.9-37.3 California 18 73.3-80.0 18 67.5-77.5 18 31.2-34.8 18 29.1-34.9 Results and Discussion In all analyses, specimens from Amchitka and Adak Islands, in the Aleu- tians, are not significantly different. This is shown well on the canonical graphs produced by the discriminant analysis program, and reproduced as Figures 3 and 4 (males and females, respectively). Also shown is the fact that Califor- nia specimens differ from Aleutian animals, but specimens from southwest- ern Alaska are clearly intermediate. Further indication of the relationship between these sea otter populations is presented in Table 9, which duplicates the discriminant function assignment of individual specimens to specific popu- lations. Otters from southwestern Alaska are similar to those from the Aleutians in condylobasal length, total length, and weight (Tables 6, 7, 1). They are similar to Californian otters in nasal length (Table 5), and are intermediate between the other two populations in their display of frontal notches and coronoid processes. Examination of the particular combination of characters in skulls from specific localities in southwestern Alaska indicates there may be a slight gradient from more Aleutian features near Umnak Island and along the north shore of the Alaska Peninsula (3 specimens show primarily 1973 Subspecies of the Sea Otter 9 Figure 3. Canonical graph produced by BMD07M Program for male sea otters. Open circle = specimen from Amchitka, dotted circle = Adak, half black circle = southwestern Alaska, solid back circle = California. Abscissa is first canonical axis, ordinate is second. Alaskan features, 6 are intermediate, and 1 could be considered Californian) toward more Californian features in Prince William Sound (2 Alaskan, 2 intermediate, 12 Californian). The ratios of skull widths to condylobasal lengths in Alaskan and Cali- fornian otters are essentially the same as those of otters from the Commander Islands (Table 8). Condylobasal lengths of Commander Island otters are similar to those from Alaskan animals (Table 6). Barabash-Nikiforov (1947) did not mention frontal notches or coronoid projections, but illustrated skulls from the Commander Islands (Figs. 5, 16, 18) which show a notch in three skulls and a possible notch in a fourth. Projecting coronoids are illustrated by Ognev (1931: Figs. 134, 135), and an ‘equal’ projection by Barabash-Niki- forov (1947: Fig. 20). Novikov (1956: Fig. 149) and Stroganov (1962: Fig. 72) both illustrate skulls with a frontal notch and mandibles which lack pro- jecting coronoids. The available data indicates that otters from the Com- mander Islands are essentially similar to those from Alaska, as previously sug- gested by Barabash-Nikiforov (1947). 10 Contributions in Science No. 252 Figure 4. Canonical graph produced by BMD07M Program for female sea otters. Open circle = specimen from Amchitka, dotted circle == Adak, half black circle = southwestern Alaska, solid black circle = California. Abscissa is first canonical axis, ordinate is second. In addition to skull differences, total length is also greater in Alaskan sea otters. Table 7 indicates that these northern animals are probably about 10 cm longer than those from California, even allowing for the differences gen- erated by taking curvilinear instead of standard length measurements. They are also larger than those from the Commander Islands; Barabash-Nikiforov (1947) indicates a maximum length of only about 144 cm, with the average of adult specimens ranging between 101 and 130 cm, the oldest animals being the largest. However, he does not differentiate between males and females, which undoubtedly resulted in a lower average figure. Among Alaskan and Californian specimens, the size difference is essentially parallel in both sexes, and is reflected also in the weight data (Table 1) and the condylobasal lengths (Table 6). Barabash-Nikiforov (1947) indicates that the fur of E. /. lutris is most commonly dark brown, ranging to nearly black, although occasionally nearly 1973 Subspecies of the Sea Otter 11 Table 9 Discriminant function assignment of specimens: four skull variables and four otter populations. Specimen from N Assigned to: Adak Amchitka S.W. Alaska California MALES: Adak 17 10 5 2 Amchitka 38 8 27 3 — S.W. Alaska 24 2 — 20 2 California 27 — — 5 22 FEMALES: Adak 43 19 16 7 1 Amchitka 73 18 49 6 — S.W. Alaska 10 2 — 6 2 California 16 — — 3 13 Combining the sexes in the above assignments, and calculating percentages: BOTH SEXES: Adak 60 29 (48%) 21 (35%) 9 (15%) 1 (2%) Amchitka S.W. 111 26 (23%) 76 (69%) 9 (8%) — Alaska 34 4 (12%) — 26 (76%) 4 (12%) California 43 — — 8 (19%) 35 (81%) orange individuals are taken. Alaskan otters are also dark, but those from California are most commonly medium brown in color. The fur is grayish at the base, as can be easily determined by parting the hairs to examine the underfur. Kenyon (1969) quotes observations by Victor Scheffer that the pelage is “light smoky gray near the skin, darkening gradually to smoky brown” at the tips. Over 40 fresh pelts of California otters were examined dur- ing the present investigation, as well as several tanned skins, and in all cases the underfur was essentially grayish, with a suggestion of brownish or even silvery color, becoming darker at the tips. In summary, sea otters from the Commander Islands, the Aleutians, southwestern Alaska, and California are all part of a single population which varies from large, dark, short-nasaled forms in the north to smaller, lighter, longer-nasaled forms in the south. This suggests an example of Bergmann’s Rule (larger forms of a species in the northern parts of its range). Specimens from near the two ends of this dine (Aleutian Islands and California) can be distinguished from each other on the basis of differences in size, nasal length, shape of coronoid process, and presence or absence of a frontal notch. Merriam (1904) compared specimens from the ends of this dine, and de- scribed E. 1. nereis as a distinct, southern form. Roest (1971) followed suit on the basis of many more specimens from California and the Aleutians. As has happened in the course of studying geographic variation in other species, the subsequent examination of specimens from intervening regions established the existence of intergrade populations. In this case, the intermediate speci- 12 Contributions in Science No. 252 Figure 5. Distribution of the sea otter, Enhydra lutris. Original distribution in- cludes all areas within the dashed lines. Present distribution indicated by stippling; note small, disjunct population of E. L lutris along the California coast. E. /. gra- cilis is currently restricted to the Kurile Islands and extreme southern Kamchatka. The circled x’s indicate the type localities of the two forms. mens are from southwestern Alaska, which still leaves a distributional gap of 2000 miles southward to California. Examination of otter remains from Indian shell mounds (middens) or similar deposits, from localities between Prince William Sound and Monterey Bay, may provide additional data to sup- port the concept of a dine. For example, Grinnell, Dixon, and Linsdale ( 1937 : Fig. 109) illustrate a skull from a shell mound in Oakland, California, which shows a notch (Aleutian feature) and has nasals over 18 mm long (Califor- nian feature). Sea otters from southern Kamchatka (Cape Lopatka) have wider skulls than do those from other localities in the North Pacific (Table 8). Since no specimens of this population were personally examined, and none have been illustrated in the Russian literature, it is not known whether they possess any of the cranial features noted in Alaskan and Californian otters. Barabash- Nikiforov (1947) indicated that they are smaller in size than typical E. 1. lutris, darker in color, and had a distinct reddish tint to the underfur: “the corresponding hair in the Kamchatkan sea otter is a cinnamon hue through- out the entire length, approximating orange or pinkish-cinnamon in tone.” He considered these animals to be a distinct subspecies, E. 1. gracilis, occurring not only in southern Kamchatka but also in the Kurile Islands. Since most authors have considered the range of the sea otter to be continuous from the Kuriles to Baja California, it would be desirable to study the variation of Kurile sea otters to determine whether their broad heads, small size, and reddish underfur are involved in a dine along the western side of the Pacific. Figure 5 indicates geographical ranges. Some of the differences among various sea otter populations around the rim of the northern Pacific may be due to the fact that all existing otter popu- lations are descendants of small relict groups of survivors from the early fur- trading days. Kenyon (1969) indicates that nine such groups survived into the twentieth century. Each of these groups was reduced to very small num- 1973 Subspecies of the Sea Otter 13 bers by the turn of the century, and each may have possessed, by chance, a unique combination of characteristics which had previously occurred in a more general way throughout the entire range of the species. If such a local combination of characters did occur in a restricted group of animals, then their descendants would be likely to retain those features— the ‘Founder Effect.’ This possibility, combined with the complete geographic and genetic isolation of the California population, may have been instrumental in bringing about the smaller size, lighter color, and longer nasals so prevalent in modern speci- mens from California. At the present time there appear to be only two forms of sea otter that merit subspecific recognition. These are discussed below. Subspecies Accounts 1. Enhydra lutris lutris (Linnaeus) \15S.Mustela lutris Linnaeus, Systema naturae, ed. 10, 1:45. Type locality, Kamchatka, U.S.S.R. nil. Lutra marina Erxleben, Systema regni animalis . . . p. 445. Type locality, unknown. 1816. Pusa orientalis Oken, Lehrbuch der Naturgeschichte. 3, abt. 2:986. 1827. Lutra stelleri Lesson, Manuel de mammalogie, ou histoire naturelle des Mammiferes, p. 156. Type locality, Kamchatka. 1843. Enhydra lutris, Gray. List of the . , . Mammalia in the . . . British Museum, p. 72. 1880. Enhydris marina Brandt, Beobacht. uber die verschied. Kleider Seeotter, “Melang, Biolog. Bullet, de I’Academ. Imp. de St. Petersb.,” 9:15. nM. Enhydris lutris, True, Proc. U.S. Nat. Mus., 7 (App., Circ. 29): 609. 1898. Latax lutris Stejneger, The Asiatic fur-seal islands and the fur seal industry, in The fur seals and fur seal islands of the North Pacific Ocean, by David Starr Jordan, U.S. Treasury Dept. Doc. 2017, p. 4:29. 1922. Enhydra lutris kamschatica Dybowski, Arch. Tow. Nauk. Lwow. 1:350. 1924. Enhydra lutris lutris Miller, List of North Amer. Recent Mamm. (1923), U.S. Nat. Mus. Bull. 128:131. Type /ocu/i7y.— Kamchatka, U.S.S.R. Should probably be restricted to the east central coast of Kamchatka, opposite the Commander Islands. Distribution.— CuTXQniXy ranging from the Commander Islands eastward through the Aleutian Islands to the Alaska Peninsula; along the north coast of the Alaska Peninsula to at least Port Eleiden; along the south shore of the Alaska Peninsula, Kodiak Island, the Kenai Peninsula, and throughout Prince William Sound to the vicinity of Kayak Island and Cape Suckling; along the 14 Contributions in Science No. 252 California coast from Monterey Bay south to Morro Bay. Formerly distributed as a continuous population from the central Kamchatka coast eastward through the Aleutians and along the southern coast of Alaska, British Colum- bia, Washington, Oregon, California, and Baja California to at least Morro Hermoso (27°32' N. lat.) (Kenyon, 1969). Descriptive rem«rA:5.— Zygomatic and mastoid widths less than 80% of the condylobasal length; infraorbital and postorbital widths less than 37% of the condylobasal length. Underfur grayish. This form varies clinally from one end of its range to the other. North- ern animals are large, while those from California are distinctly smaller (Tables 1, 6, 8). Individual adult males have reached a maximum weight of 46.25 kg (102 lbs.; Schneider, personal communication), although most are between 27 and 36 kg. Maximum length reported is 166 cm curvilinear length (Schneider, personal communication). Nasal lengths vary from about 11 mm in Aleutian specimens to over 21 mm in California animals. The California population can be distinguished from the Aleutian popu- lation, but not from the population in southwestern Alaska. California speci- mens are smaller than Aleutian animals, have nasal lengths usually over 18 mm, lack a frontal notch, and have coronoid processes which project back- wards at an angle. When the fur of specimens of this form is parted, the underfur thus ex- posed is characteristically grayish. Typical pelts from northern animals are dark brown to nearly black, while California specimens are usually medium brown. In the list of specimens examined, the following abbreviations have been used to identify collections; ADFG— Alaska Department of Fish and Game, Anchorage, Alaska; CAS— California Academy of Sciences, San Francisco, California; CFG— California Department of Fish and Game (specimens at California Polytechnic State University and at Moss Landing Marine Labora- tories); CPSU— California Polytechnic State University, San Luis Obispo, California; HSU— Humboldt State University, Areata, California; LLU— Loma Linda University, Loma Linda, California; MLML— Moss Landing Marine Laboratories, Moss Landing, California; MML— Marine Mammal Labora- tories, Bureau of Sport Fisheries and Wildlife, Seattle, Washington; MVZ— Museum of Vertebrate Zoology, Berkeley, California; PGMNH— Pacific Grove Museum of Natural History, Pacific Grove, California; SBMNH— Santa Barbara Museum of Natural History, Santa Barbara, California; SDMNH— San Diego Museum of Natural History, San Diego, California; SSU— Sacramento State University, Sacramento, California. Adult specimens examined. — (267) RUSSIA: Medny Island: MML — 1 speci- men, no number. ALASKA: Amchitka Island: ADFG-SO-67-305, 349, 351, 364, 365, 368, 380, 387, 390, 391, 394, 407, 411, 436, 443, 445, 446, 450, 455, 456, 461, 478, 485, 490, 492, 497, 498, 500; HSU — 3 specimens, no numbers; MML — KWK 59-5, 59-6, 59-16, 59-21, 59-23, 59-34, 59-38, 59-39, 59-40, 59-43, 59-48, 59-49, Subspecies of the Sea Otter 15 1973 I 59-56, 59-57, 59-64, 59-67, 59-69, 59-71, 59-80, 59-84, 59-90, 59-91, 59-92, 59-94, I 59-96, 59-98, 59-100, 59-118, 59-130, 59-134, 60-4, 62-7, 62-11, 62-21, 62-24, 62-47, I 62-50, 62-72, 62-77, 62-78, 62-80, 62-89, 62-94, 62-105, 62-111, 62-118, 62-121, , 62-124, 62-131, 62-135, 62-136, 62-138, 62-145, 62-147, 62-152, 62-156, 62-159, ii 62-175, 62-176, 62-184, 62-211, 62-226, 62-268, 62-272, 16-56, 17-56, 19-56, 39-56, i' 44-56, 46-56, 52-56, 59-56, 70-56, 71-56, 72-56, 73-56, AIWR 4-53, 6-53, 7-53, 9-53, j 16-53, 32-53, 33-53, 48-53, 50-53, 54-53, 63-53, 75-53, J-53, K-53, L-53, X-53, 8-54, ! 42-54, 66-54, 71-54, 37-55, 27-56, JEB 63-80, 63-273, 130-1949, 133-1949; MVZ , 113396; Adak Island: ADFG SO-67-5, 7, 9, 11, 17, 23, 26, 27, 28, 29, 34, 39, 44, I 63, 70, 77, 79, 85, 86, 88, 98, 104, 106, 107, 108, 114, 116, 118, 131, 133, 135, 177, :l 242, 260, 261, 272, 278, 281, 293; MLML SO-67-22, 195; St. Paul Island, Pribilof ! Islands: ADFG SO-72-7; Amak Island: MML, 1 specimen, no number; Port Moller: I ADFG SO-67-511; Ilnik: ADFG SO-71-3, 4, 5, 6; Port Heiden: ADFG SO-71-2, I 10, 72-1; Sanak Island: MML KWK 60-15; Sandman Reefs: ADFG CJL 1-58; Popof I Island: ADFG SO-69-4; Nagai Island: MML KWK 60-12, 60-13, 60-14; Simeonof ! Island: MML KWK 60-5, 60-6, 60-8, 60-10; Tugidak Island: ADFG SO-66-1, 68-22; I English Bay: ADFG SO-68-6, 72-5; Prince William Sound: ADFG 1 specimen, no I number, CJL 1-56, 2-56, No. 1, No. 2, No. 3, 2, 61, 80, 1500, 1501, SO-72-10, 72-12, I 72-15, 72-16, 72-17, 72-19, 72-20. CALIFORNIA: Monterey: CFG 150-69, 155-69, 175-70, LLU 1 specimen, no number, SBMNH 225, SDMNH 18865; Hopkins Marine Station: CFG SO- 134-70, 189-70; Point Pinos: CFG SO- 18 1-70; Asilomar: CFG SO-249-72, PGMNH 2436A, 2439A; Seventeen-mile Drive: MLML 412; Carmel: CFG SO-98-68, 1 specimen, no number; Point Lobos: CFG SO- 134-69, 190-69, MVZ 114550, 119829; Bixby Creek: MVZ 84812; Point Sur: CAS 637, MLML 1225; Gorda: MVZ 116213; San Luis Obispo County: MVZ 116611; San Carpoforo Creek: CFG SO-141-70; Cambria: CFG SO-160-69, 186-70; Cayucos: CFG SO-248-72; Morro Bay: CFG SO-1 17-69, 159-69, 163-70, CPSU M-263, M-983, MLML 0-3; Montana de Oro State Park: CFG SO- 187-70; No specific locality: CAS 4428, MLML SO-5, 0-7, 494, 495, MVZ 123192, 123194, 123196, 123197, 123198, 123199, SSU 1202, SDMNH 16371. 2. Enhydra lutris gracilis Bechstein 1799. Lutra gracilis Bechstein, Thomas Pennant’s allgemeine Uebersicht der vierfussigen Thiere . . ., v. 2, p. 248. Type locality “Staatenland” (= southernmost of the Kurile Islands: see Hollister, 1921). 1931. Enhydra lutris, Ognev, Mammals of eastern Europe and northern Asia, 2:402. 1947. Enhydra lutris gracilis, Barabash -Nikiforov, The sea otter, p. 20. Type locality:— Kurile Islands; possibly Kunashir Island. Distribution:— The northern Kurile Islands and the southern tip of Kam- chatka. Formerly found along the coasts of southern Sakhalin and northern Honshu and Hokkaido, and the southern Kuriles (Barabash-Nikiforov, 1947). Descriptive remarks :—SmdX\er than northern specimens of E. 1. lutris, although about the same size as specimens from California. The largest meas- urement reported is a condylobasal length of 133.6 mm. This form is dis- tinguished from E. 1. lutris by its broad head (Table 8). The zygomatic and mastoid widths are greater than 80% of the condylobasal length, and the 16 Contributions in Science No. 252 infraorbital foramen and postorbital widths are over 37% of the condylobasal length. Another distinctive feature is the reddish tone of the underfur. Specimens examined:— Nont. The above description and remarks are based entirely upon the discussion by Barabash-Nikiforov (1947). Key to Subspecies Differences between the two subspecies of Enhydra lutris are summarized in the following key: lA. Zygomatic width and mastoid width less than 80% of condylobasal length; infraorbital foramen width and postorbital width less than 37% of condylobasal length; underfur grayish: . E.l. lutris IB. Zygomatic width and mastoid width more than 80% of condylobasal length; infraorbital foramen width and postorbital width more than 37% of condylobasal length; underfur reddish: E.l. gracilis Acknowledgments I thank the following institutions and individuals for their courtesies and cooperation: Marine Mammal Laboratories, Bureau of Sport Fisheries and Wildlife, Seattle, Wash, (and particularly Karl W. Kenyon); Alaska Depart- ment of Fish and Game, Anchorage, Alaska (particularly Karl B. Schneider) ; California Department of Fish and Game, Marine Resources, Monterey, Calif, (particularly Mel Odemar and Paul Wild); California Academy of Sciences, San Francisco, Calif.; Natural History Museum of Los Angeles Co., Los Angeles, Calif, (particularly Donald Patten); Pacific Grove Museum of Natural His- tory, Pacific Grove, Calif.; San Diego Museum of Natural History, San Diego, Calif.; Santa Barbara Museum of Natural History, Santa Barbara, Calif.; Museum of Vertebrate Zoology, Berkeley, Calif.; Humboldt State University, Areata, Calif.; Moss Landing Marine Laboratories, Moss Landing, Calif.; Sacramento State University, Sacramento, Calif.; Loma Linda University, Loma Linda, Calif. I also thank Luther Bertrando, Richard Pimentel, and Nancy Mosman for their assistance with the statistical analyses. 1973 Subspecies of the Sea Otter 17 Literature Cited Barabash-Nikiforov, I. I. 1947. The Sea Otter. Translated from the Russian by the Israel Program for Scientific Translations, Jerusalem, 1962. 227 pp. Dixon, W. J. (ed.). 1970. BMD Biomedical Computer Programs. Univ. Calif. Publ. in Automatic Computation, No. 2. Univ. Calif. Press, Berkeley. 599 pp. Grinnell, J., J. S. Dixon, and J. M. Linsdale. 1937. Fur-bearing Mammals of California. Univ. Calif. Press, Berkeley. 2 vols. 777 pp. Hall, E. Raymond, and K. R. Nelson. 1959. The Mammals of North America. Ronald Press, New York. 2 vols. 1083 pp. Hollister, N. 1921. Review of Pohle, H., Die Unterfamilie der Lutrinae. J. Mam- mal. 2(3):177-178. Kenyon, K. W. 1969. The Sea Otter in the Eastern Pacific Ocean. North American Fauna Series, No. 68. 352 pp. Lensink, C. j. 1962. The history and status of sea otters in Alaska. Ph.D. thesis, Purdue University, Univ. Microfilms, Ann Arbor. 188 pp. Merriam, C. H. 1904. A new sea otter from southern California. Proc. Biol. Soc. Wash. 17:159-160. Miller, G. S. and R. Kellogg. 1955. List of North American Recent Mammals. U.S. Nat. Mus. Bull. 205. 954 pp. Novikov, G. A. 1956. Carnivorous Mammals of the Fauna of the USSR. Translated from the Russian by the Israel Program for Scientific Translations, Jerusalem, 1962. 284 pp. Ognev, S. I. 1931. Mammals of Eastern Europe and Northern Asia, v. 2, Carnivora (Fissipedia). Translated from the Russian by the Israel Program for Scientific Translations, Jerusalem, 1962. 590 pp. Roest, a. I. 1971. A systematic study of the sea otter (Enhydra liitris). Proc. 8th Ann. Conf. Biol. Sonar and Div. Mamm., Biol. Sonar Lab., Mar. Mammal. Study Cent., SRI, Menlo Park, pp. 133-135. Scheffer, V. B. 1951. Measurements of sea otters from western Alaska. J. Mam- mal. 32:10-14. AND F. Wilke. 1950. Validity of the subspecies Enhydra lutris nereis, the southern sea otter. J. Wash. Acad. Sci. 40:269-272. Stroganov, S. U. 1962. Carnivorous Mammals of Siberia. 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