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DC^ _ XOiiiS^ Q ^ i: X^OlllST^ O Ml’^NVINOSHilWS S3 I ava a n'^LI B RAR I ES^SM!THSONIAN“^}NSTITUTlON NOlini r- , 2 r- 2 r- 2 ° 2 CD CD I) - 4 LOS — ANGBLES COUNTY MUSEUM CONTRIBUTIONS IN SCIENCE DECEMBER 31, 1971 13 C TABLE OF CONTENTS and AUTHOR INDEX 1971 Nos. 206-218 Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 2 CONTRIBUTIONS IN SCIENCE TABLE OF CONTENTS No. 206. The biology of Anthophora (Micranthophora) peritomae Cockerell, by Philip F. Torchio. 14 pp., 13 figs. January 5, 1971. No. 207. Trombiculid mites of the genus Microtrombicula (Acarina) from Costa No, 208, Rica, by James P, Webb, Jr. and Richard B. Loomis. 15 pp. , 4 figs. February 4, 1971. The role of introgressive hybridization in the development of Salvia: section Audibertia (Labiatae), by William A. Emboden, Jr. 15 pp. , 4 figs, February 4, 1971. No, 209, Five new species of Acmaeodera (Coleoptera: Buprestidae) from the southwestern United States and Baja California, by Richard L. Westcott. 19 pp. , 15 figs. February 17, 1971. No. 210. Observations on the biology of land crabs and their burrow associates on the Kenya coast, by Charles L, Hogue and Donald B. Bright. 10 pp. , 5 figs. February 17, 1971. No. 2U. The birds of the lowlands of Bwamba, Toro Province, Uganda, by Herbert Friedmann and John G. Williams. 70 pp. , 1 fig. April 7, 1971. No. 212. Notes on the bat flies of Costa Rica (Diptera: Nycteribiidae), by B. V. Peterson. 8 pp, , 2 figs, March 31, 1971, No. 213. Type specimens of Recent mollusks in the Los Angeles County Museum of Natural History, by Gale Sphon. 37 pp. , 21 figs. May 27, 1971. No. 214. Studies on California ants. 6. Three new species of Myrmecocystus (Hymenoptera: Formicidae), by Roy R. Snelling. 16 pp. , 3 figs. May 28, 1971. No. 215. Status of certain frogs of the genus Colostethus, with descriptions of new species, by Philip A. Silverstone. 8 pp. , 3 figs, June 9, 1971, TABLE OF CONTENTS 3 No. 216. Phylogenetic relationships of certain Neotropical toads with the description of a new genus (Anura : Bufonidae), by Linda Trueb. 40 pp. , 11 figs. July 8, 1971. No. 217. Pliocene avian remains from Baja California, by Hildegarde Howard. 17 pp. , 2 figs. November 12, 1971. No. 218. A distributional checklist of the amphibians of Honduras, by John R. Meyer and Larry David Wilson. 47 pp., 2 figs. December 29, 1971. 4 CONTRIBUTIONS IN SCIENCE AUTHOR INDEX Bright, Donald B. No. 210 Emboden, William A. , Jr. No. 208 Friedmann, Herbert No. 211 Hogue, Charles L. No. 210 Howard, Hildegarde No. 217 Loomis, Richard B. No. 207 Meyer, John R. No, 218 Peterson, B. V. No. 212 Silverstone, Philip A. No. 215 Snelling, Roy R. No. 214 Sphon, Gale G. No. 213 Torchio, Philip F. No. 206 Trueb, Linda No. 216 Webb, James P, , Jr. No. 207 Westcott, Richard L. No. 209 Williams, John G. No. 211 Wilson, Larry David No. 218 LOS ANGELES COUNTY MUSEUM CONTRIBUTIONS IN SCIENCE Dumber 206 January 5, 1971 501, 73 THE BIOLOGY OF ANTHOPHORA (MICRANTHOPHORA) PERITOMAE COCKERELL (HYMENOPTERA: APOIDEA, ANTHOPHORIDAE) By Philip F. Torchio Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. Robert J. Lavenberg Managing Editor INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 BIOLOGY OF ANTHOPHORA (MICRANTHOPHORA) PERITOMAE COCKERELL (HYMENOPTERA: APOTDEA, ANTHOPHORIDAEO By Philip F. Torchio“ Abstract: The biology of Anthophora (Micranthophora) peritomae Cockerell is described and compared with that of A. (M.) flexipes Cresson. Several habits of A. peritomae not ob- served previously within the genus include: salivary material secreted by larvae; cell refurbishment; and use of communal nest entrances. Two parasitoids (Zacosmia maciilata (Cresson) and An- thrax limatiilus artemisia Marston) were found to attack immatures of A. peritomae. Observations on the biology of Z. maculata are included. The fungus, Ascosphaera apis (Maassen ex Claussen) Olive and Spiltoir, which is the causal agent of chalk brood disease of Apis mellifera Linnaeus immatures, grew only on feces within cells. The biology of the Anthophora subgenus, Micranthophora, has not been studied extensively. Hicks (1934) reported some biological features of A. (M.) curta Provancher and was the first to associate the parasitic bee, Zacos- mia maculata (Cresson), with curta. Torchio and Youssef (1968) described the biologies of A. (M.) flexipes Cresson and Z. maculata and the immature stages of Z. maculata. This paper describes the biology of A. (M.) peritomae Cockerell and makes comparisons with the biology of A. flexipes. New obser- vations on the biology of Z. maculata are also included. Acknowledgments I would like to thank G. E. Bohart (Leader, Wild Bee Pollination In- vestigations, ARS, USDA, Logan, Utah), R. W. Thorp (Department of Entomology, University of California, Davis), and W. P. Stephen (Depart- ment of Entomology, Oregon State University, Corvallis) for reviewing the manuscript. Howard Potter, an Agricultural Research Technician stationed at this laboratory, was responsible for the photographs. Habitat Anthophora peritomae was found nesting in an eroded, vertical embank- ment adjacent to Bear River, two miles east of Cornish, Cache County, Utah, during September and October, 1968. This embankment, formed by sedimen- tation of the river and subsequent erosion, was composed of coarse-grained, pebble-free, tightly-packed greyish sand. It measured 4.5 m in height and iln cooperation with Utah Agricultural Experiment Station. ^Entomology Research Division, Agricultural Research Service, Logan, Utah 84321; and Research Associate in Entomology, Los Angeles County Museum of Natural History. 1 2 Contributions in Science No. 206 18.3 m long, was devoid of plant growth and curved from a south to an east exposure. Dense aggregations of nests (more than 30 per square meter) were found in limited areas between the 0.5 and 2-m levels on the south and southeast slopes which were exposed to the early morning sun. Most entrance holes were shaded from the sun by 2:00 pm Mountain Standard Time (MST). On clear days, flight occurred whenever the temperature rose above 24° C (usually from about 9:30 am to 6 pm), but little flight took place during cloudy weather though temperatures often rose above 24° C. Nest architecture Entrance Hole: The structure, dimensions, and construction of entrance holes were similar to those of A . flexipes, which nested in the vertical embank- ments near Moab, Utah (Torchio & Youssef, 1968). Both species also em- ployed the same method (rapid flicking motions with the hind legs) to remove soil from the nest. Anthophora peritomae, however, initiated con- struction of new nest burrows throughout the daily flight period and directed them into the substrate at various angles. Because nests of A. flexipes found near Cornish were constructed in flat or gently sloping sand surfaces, they were surmounted by ellipsoidal tumuli formed by excavated sand flicked backwards from the nest entrances. Burrows: Burrows constructed by both species were similar: They had the same diameter and their walls were not lined with secreted materials. Also, the lateral burrows were comparable in length, and each was partially plugged a short distance above the cells with loose, dry sand excavated from new burrows or from one or more areas along existing ones. The main burrows and entrance holes remained unplugged during and after nesting activities. The nesting site of A. peritomae was completely honeycombed with nests constructed in previous seasons. As a consequence, even the most care- ful excavations were inadequate to trace the branches associated with each nest. The problem was compounded when I found that the bees commonly used communal entrances. However, once I devised a method to establish the number of females using a communal entrance, it was possible to de- cipher the architecture of individual nests branching from the common entrance cavity. To observe those bees using a communal nest and to distinguish between them, I marked each bee (without disturbing it) in such a way that I could recognize it for several days. Small spots of lacquer or oil-based pigments painted on the mesonotum or apical terga were unsatisfactory: some bees died within minutes after application; others became disoriented; and many quickly removed the markings. In contrast, bees dusted with fluorescent powder were apparently unaffected. This material was applied by blowing a small quantity of powder through a soda straw directly onto a bee hovering 1971 The Biology of Anthophora Peritomae Cockerell 3 in front of her nest entrance. The powder, which adhered to body hairs, was quite visible on most individuals for at least four days. Seven communal nests were observed in this study, and each occupant of each nest was marked with a different colored powder. The number of females per nest was small (minimum, three; maximum, six), and the nest architecture was variable. In four nests, a large communal chamber was excavated immediately behind the common entrance hole, and each burrow leading away from it represented an individual nest. Each nest was normally composed of a main burrow and several lateral burrows, each originating at a particular point along the terminal half of the main burrow. The lateral burrows meandered for variable distances until they terminated in cells. The entrances to the communal chambers were normally enlarged gradually during the nesting period. When nesting was completed, they were nearly as wide as the chambers. A fifth nest lacked a communal chamber, and each burrow originating along the entrance burrow represented an individual’s nest which normally possessed its own lateral burrow system. The entrance hole and burrow, however, were not enlarged as one might expect. The other two nests studied were multi-entranced and lacked communal chambers. One of these nests, occupied by four bees, had three entrance holes; the other, which possessed four entrances, housed three bees. These multi-entranced nests were more than likely developed when burrowing bees accidentally broke into existing nest burrows of other solitary nesting bees. Subsequently, these drilled-into burrows became interconnecting, communal nests. Occasionally, burrowing bees would happen onto inactive nests excavated in previous years, where- upon they often, but not always, incorporated them into their nest archi- tecture. When the entrance of a single-entrance nest containing five females was plugged with cellucotton, each returning bee hovered directly in front of it for a short period. Then she darted about searching the surrounding bank for five to 10 seconds before she returned to her original hovering position. After several minutes, she flew away and then returned a few minutes later to repeat the same pattern. After an hour, one bee landed on the plug and tried to pull it loose. Soon, two others joined her, and the three (sometimes working together and sometimes individually) removed the plug in 3 hr., 52 sec. and re-established communal nesting. The fourth female reoriented to an abandoned nest several meters away, and the fifth bee exca- vated a new nest 30 m away. A check of the five bees four days later proved that the forced reorientation was permanent. Bees inhabiting nests with more than one entrance also varied in their behavior. Normally, one entrance hole was entered by two or more bees and the others were each utilized by one bee. These bees continued to enter and exit through the same holes even after the main or a lateral burrow was partially exposed by the burrowing activities of another bee or by my exca- 4 Contributions in Science No. 206 vations. If an entrance used by one female was plugged while she was in the nest, she quickly located another exit hole, oriented to the new entrance by hovering in front of it for a few seconds, and then flew in a lateral zigzag and figure eight pattern in front of and directly away from it. Subsequently, she used the new entrance exclusively. Conversely, a bee returning to a nest whose entrance was plugged while she was away was unable to locate alterna- tive entrances and abandoned her nest. Those bees utilizing communal en- trances associated with multi-entranced nests seemed to be strongly oriented to their entrances. If a communal entrance was plugged while the bees were in the nest, they sometimes drilled another hole as close as possible to the original. At other times, bees escaped from other entrances in the nest but then returned to the blocked entrance and tried to unplug it. If they could not remove the plug, they usually excavated another entrance near the plugged one. However, in one case, two bees abandoned their nests and started new solitary nests 40 and 48 cm distant. , Although plugging the entrances with cellucotton disturbed the orien- tation of the bees, carving the entrances or applying fluorescent powder did not. Also, careful observations revealed that most nests were occasionally entered by bees from other nests. Some of these visitors deposited one or two loads of pollen. Two such bees, marked as they left the alien nest, were subsequently discovered nesting within a 45 cm radius. Cells: Cells of A. peritomae and flexipes were similar in shape, place- ment at the terminus of burrows, extractability from surrounding soil, attributes of the cell lining, and details of the cell cap. Although both species commonly constructed a single cell at the terminus of each branch burrow, peritomae placed two to four cells in linear series or in loose clusters at the terminus of nearly 20 per cent of the burrows studied. Cells in linear series were usually separated from each other by a plug of loose sand 2 to 3 mm deep, but I found one series where cells were joined. Both species placed cells at various declinations ranging from subhorizontal to perpen- dicular. The area immediately in front of most cells of A. peritomae was hol- lowed, resembling the basal area of typical cells (Fig. 1). The inner surface of these antechambers was lined with small, uniform soil particles similar to the soil lining found in cells, but it lacked the secreted, waxlike lining characteristic of true cells. Although this antechamber (8 mm long by 6 mm wide) was completed during cell construction, its function was not determined. Antechambers were reduced or lacking in front of cells constructed in linear series. Cells of A. peritomae did not vary in size ( 10 by 8 mm), shape, or structure. Construction of cells was begun when the terminus of the burrow was carved into a crude cell with dimensions slightly larger than those of the completed cell. The final cell walls were formed by lining the roughed-in cell with small soil particles of a relatively uniform size extracted from other 1971 The Biology of Anthophora Peritomae Cockerell 5 areas within the nest. The cell was then coated with a thin lining of a trans- parent, waxlike secretion. Cell cap construction was not observed, but com- pleted caps were lined with a waxlike material and were nearly identical to those constructed by A. flexipes. Figures 1-4. Anthophora peritomae. Fig. 1. Longitudinal section of cell and un- lined antechamber. Fig. 2. Basal area of cell with arrows designating the 2 waxed linings. Fig. 3. Cell constructed across open space. Arrow points to cell cap. Fig. 4. First instar larva nearly free of its egg chorion. During excavations of A. peritomae nests, one cell was found with two wax linings separated by a thin layer of soil (Fig. 2). Thus, peritomae re- furbishes and relines abandoned cells in some circumstances. This time-saving habit is, surprisingly, not often practiced by bees except for some Apidae and some Megachilidae. Another cell (Fig. 3) was found positioned across the diameter of a burrow, suggesting that the species can construct cells across unsupported spaces. Provisioning and development Nest provisioning by A. peritomae (Fig. 13) was studied from Sep- tember 5 through September 10, 1968. Similar studies of A. flexipes (Fig. 12) were made a few weeks earlier at the nearly flat-surfaced sandy area one mile to the north. A. peritomae required an average time of 5 hr, 50 min to provision a cell: this included the deposition of 20 pollen loads in a period of 1 hr, 50 sec (an average of 3 min/load); and a foraging period of 4 hr 6 Contributions in Science No. 206 (an average of 12 min, 37 sec/ trip). A. flexipes deposited 18 pollen loads/cell (Fig. 12) in 1 hr, 18 min (an average of 4 min, 20 sec/ load); and the foraging period was 1 hr, 27 min, 30 sec (an average of 6 min, 44 sec/ trip). Thus, the two species were very similar in their provisioning behavior at Cornish in spite of the dissimilarity in their nesting habitats. The bottom three-eighths to one-half of each A. peritomae cell was filled with a dark yellow, pastelike provision. Samples of pollen taken from 10 cells proved it to be Grindelia squarrosa (Pursh) Dunal (determined by G. E. Bohart) , which was the predominant flowering species near the nesting site. Anthophora peritomae, unlike A. flexipes, stored pollen as a nearly dry, asymmetrical ball. When the provision was nearly complete, liquid (probably regurgitated nectar) was added until the mass became a semiliquid that filled the bottom half of the cell. The surface of the completed provision was tacky to the touch, but it was not covered with a layer of nectar. The pungent odor characteristic of anthophorine cells was present in the cells of both species. The egg of A. peritomae was dull white, opaque, weakly arched, and measured 3.5 mm long by 0.9 mm wide. It was deposited on the surface of the provision and was attached to it only by its tips which were slightly em- bedded (Fig. 5). The egg was always positioned with its narrowed anterior tip (0.5 mm wide) less than 1.0 mm from the cell wall and with its broadened, posterior tip (0.9 mm wide) resting on or near the center of the provision. The position of A. flexipes eggs as described by Torchio and Youssef ( 1968: 292-293) should be reversed: the narrowed, anterior tip of each egg faced the cell wall and the broadened, posterior tip rested near the center of the provision’s surface. The egg was turgid when deposited, and its surface, though minutely reticulated, was slightly reflective. As the embryo matured, this turgidity and reflectivity were gradually lost. The first instar completely freed itself from the egg chorion 68 to 74 hours after the egg was deposited. Emergence was signaled by the splitting of the chorion along the pleural region directly above the spiracular line of the embryo. Within 10 hours, the chorion was completely separated between the anterior and posterior spiracles, and, during the eleventh hour, it split transversely between the anterior pair of spiracles. Also, the posterior areas of the chorion separated rapidly until the four pos- terior abdominal segments of the first instar larva were exposed. The pleural region of the chorion then separated anteriorly above the prothorax of the larva and finally above the lateral and posterior margins of its head capsule. The first instar was completely freed from its chorion (Fig. 4) within 12 hours after the chorion began to separate from the head capsule. Subsequently, the chorion was represented by two chorionic strips lying on top of each other on the surface of the provision directly behind the larva. The first instar encircled the outer edge of the provision as it fed (Fig. 8), and it usually completed one circle before molting. The second and third instars followed the same pathway and consumed sufficient quantities of 1971 The Biology of Anthophora Peritomae Cockerell 7 Figures 5-9. Anthophora peritomae. Fig. 5. Position of egg on provision within cell. Fig. 6. Dissected cell exposing anal excreta smeared on cell wall and partially covered by a few sausage-shaped fecal particles. Fig. 7. Fecal particles with one having a transparent thread extruding from it. Fig. 8. First instar encircling provi- sion as it feeds. Egg chorion visible on central surface of provision. Fig. 9. Last instar forming a cone-shaped provision as it encircles and consumes the outer edges. pollen to furrow the outer edge of the provision, leaving a central pillar. The last instar enlarged and deepened the furrow as it fed until the provision was shaped into a conelike structure whose sides sloped nearly 60 degrees (Fig. 9). The larva then lifted its posterior abdominal segments until its anus touched the cell wall immediately below the cell cap and deposited a light gray, granular material through its anus. This material, as seen through the translucent cuticle, was stored in a pair of convoluted tubules within the terminal body segment. Each tubule emptied into the colon immediately behind the zone filled with feces. Within 43 minutes, the larva had smeared the material (probably Malpighian excreta) across a limited area adjacent to the cell cap by oscillating its terminal abdominal segment. The thin coating dried rapidly into a brittle scale which remained firmly attached to the cell wall (Fig. 6). 8 Contributions in Science No. 206 As the feeding larva was completing deposition of the anal excretion, it began to deposit a salivary material as a rapidly drying, shiny, transparent thread. This material was secreted onto the provision continuously for 5 to 9 hours. Eventually, the larva ingested these threads as it consumed its pro- vision. Since it made no attempt to construct a cocoon, the function of the salivary secretion is not understood. Possibly, it serves a nutritive role, or it may merely represent the vestige of a cocoon-spinning habit. When the larva had voided its anal excreta and salivary secretion, it rapidly consumed the remaining provision by two methods of feeding: most larvae fed upon the surface of the provision until it was consumed; others continued to encircle and deepen the furrow around the outer edge of the provision until it was shaped into a parallel-sided column (Fig. 10). Then the larva consumed the basal area of the column (Fig. 11) until it rested with its terga at the bottom of the cell beneath the provision. Eventually, the head and anus of the C-shaped larva faced the cell cap with the medial abdominal terga resting on the bottom of the cell and the provision resting on the medial abdominal sterna. At this juncture, the subspherical food remnant was rapidly consumed. Defecation was usually begun about 8 min after the larva deposited its anal excreta. The first fecal particles were deposited adjacent to or covering this layer (Fig. 6). The sausage-shaped fecal pellets measured 0.75 to 1.10 Figures 10-11. Anthophora peritomae. Fig. 10. Last instar larva encircling the parallel-sided provision shaped by the feeding larva. Fig. 1 1 . Position of last instar larva as it begins to consume basal area of provision. Some fecal particles have been deposited on the cell wall, and one is extruding from the anus of the larva. 1971 The Biology of Anthophora Peritomae Cockerell 9 mm long and 0.5 mm wide. They were subcylindrical, but both tips were dorsoventrally flattened and blunted apically. Also, each tip was provided with one to three thin, transparent threads which interconnected the pellets. At least one thread was continuous as it traversed the core of the fecal chain. Several partially extruded fecal particles were gently extracted from the anus of various larvae. In each case, a transparent thread which entered the posterior tip of the fecal particle was also pulled from the anus (Fig. 7). The material resembled a strand of molten glass when it was first exposed to air, but it dried quickly into a brittle, shiny thread. The thread varied in length (9 to 18 mm) and was incorporated into each fecal particle prior to its emission. Defecation was not continuous. At times, three or four pellets were deposited within a 2-minute period and then an hour or more of inactivity followed before additional pellets were deposited. Defecation was completed approximately three days after it was initiated. Anthophora peritomae, unlike A. flexipes, deposited unappressed fecal particles on the cell walls, but like A. flexipes, it did not normally smear feces over the inner face of the cell cap. The postdefecating larva of A. peritomae quickly changed in appear- ance: it became semiflaccid; its integument turned an opaque, cream color; and it assumed a nonmotile, C-shaped, overwintering position. Seventy such prepupae collected from the nest site on October 8, 1968, were stored in the laboratory at room temperature (21-24°C). Although none were cold-condi- tioned, six adults emerged in late December and 24 more emerged by May 1. The remaining 40 larvae were then cold-conditioned throughout the month of May at 5-7 °C, and most of these emerged as adults during the third week of June. The sex ratio was 1:1. In the field, A. peritomae overwinters as a prepupa. Pupation occurs during mid-August, and most adults have emerged by September first. Associates Two parasitoids (a melectine bee, Zacosmia maculata (Cresson) and the bombyliid fly. Anthrax limatulus artemisia Marston) were reared from cells of Anthophora peritomae. Four planidia of Anthrax /. artemisia were recovered from three of several hundred host cells excavated during the nesting season. One active planidium in each of two cells was crawling on the provision adjacent to the host cell. In the third cell, two planidia were found, one crawling over the surface of the provision and the other crawling back and forth on top of the first-instar host larva. Also, on October 13, 1968, 68 host cells were collected from the nest site and transferred to the laboratory where they were stored at room temperature (21-24°C). On December 29, 1968, these cells were opened and five were found to contain Anthrax 1. artemisia (one mature larva and four adults). 10 Contributions in Science No. 206 The same 68 host cells also included three containing Z. maculata (two prepupae and one adult). Various stages of Z. maculata were also found in 36 additional cells excavated in 1968. The biology agreed with that reported by Torchio and Youssef (1968), but additional details of its ethology were observed. ( 1 ) In one cell, an egg chorion of the parasite was found attached to the cell wall immediately below the cap. A scar visible only on the inner face of this cap indicated that the parasite had drilled a hole through it, attached an egg to the cell wall rather than to the cap, and then repaired the cap. (2) In one cell, the first-instar parasite punctured the host egg, migrated a short distance along the surface of the provision, and then ingested a small quantity of it before molting. The head capsule below the antennae was immersed in the provision during feeding. (3) The second and subsequent instars, like those of the host, consumed the outer edge of the provision, but they periodically meandered across the surface of the provision and con- sumed the developing central pillar. (4) Also, territoriality was evidenced by three male Z. maculata which were observed perched on plants of inland salt grass [Distichlis stricta (Torr.) Rybd.] 6 to 9 m in front of the nesting embankment. Two were dusted with marking powder, and each was subse- quently found perched on plants growing over a particular square meter surface during various periods through four consecutive days. Each time another Z. maculata flew over an occupied plant, the “owner” took flight and grasped the second bee, causing both to fall. Then, the “owner” immedi- ately assumed a copulatory position and used his antennae to stroke those of the invader. Coition was not observed because every invader proved to be a male. When pebbles approaching the size of an adult Z. maculata were tossed near a perched male, he flew up, examined the intruding object, and returned to his perch. A fungus, Ascosphaera apis (Maassen ex Claussen) Olive and Spiltoir, was found in most cells of Anthophora peritomae, and hyphae were visible on feces of the host bees within 24 hours after defecation was completed. Forty-eight hours later, a solid mat of mycelia and nutriocytes covered the feces. Some fecal particles were embedded in this mycelial mat, and each particle became shrunken, collapsed, and brittle. In Europe, Ascosphaera apis sometimes attacks larvae of Apis mellifera L. and causes an infection known as chalk brood disease (Maassen, 1916). Recently chalk brood disease has been reported from honey bee colonies in the United States. Melville and Dade (1944) and Clout (1956) were the first to report the incidence of Ascosphaera apis on wild bees {Megachile sp. and Chalico- doma sp., respectively). The fungus infests the bee larva after it attacks the pollen provision, according to Clout. Baker and Torchio (1968), reported the first incidence of this fungus from the New World in cells of two species of wild bees. Megachile (Megachile) inermis Provancher and Anthophora (Anthophora) pacifica pacifica Cresson. They associated the fungus with the MINUTES 1971 The Biology of Anthophora Peritomae Cockerell 11 sdnoH Figure 12. Nesting activities of Anthophora flexipes during cell provisioning. MINUTES 12 Contributions in Science No. 206 SdnOH Figure 13. Nesting activities of Anthophora peritomae during cell provisioning. 1971 The Biology of Anthophora Peritomae Cockerell 13 feces in both species but did not recover infested larvae. In Anthophora peritomae, the fungus did not appear in cells parasitized by Zacosmia nor in cells in which the host larva died before or during defecation. Apparently, Ascosphaera apis in the New World does not infect the larvae of bees other than honey bees. Discussion In Table I the contrasting features in the bionomics of Anthophora peritomae and A. fiexipes are outlined. Of the 13 items, numbers 1 through 5 are probably of specific significance; the others are more likely expressions of habits imposed on the bee by the niches in which nesting occurred. Items 6 and 7, for example, may simply reflect the different host plants available to each species. Anthophora peritomae visited Grindelia squarrosa, whereas A. fiexipes visited several hosts growing in abundance near its nest sites. Perhaps the moistness of the provision and the presence or absence of a nectar layer may be correlated with the host plants rather than with the species of bee. Apparently, Z. maculata survives equally well on either a solid- or liquid- surfaced provision. Numbers 8 through 13 may simply reflect a more crowded condition at the nest site of A. peritomae. Table I Ethological differences between A nthophora peritomae and Anthophora fiexipes ( + = yes; — = no) A. fiexipes A. peritomae 1. Area immediately above each cell carved into a pocket lined with small soil particles + 2. Anal excreta smeared on cell wall before defecation - + 3. Feces deposited as a continuous, thin layer covering cell walls + — 4. A transparent thread traverses the central core of each fecal particle and interconnects them -(?) + 5. Salivary material secreted - + 6. Pollen initially stored in a dry state - + 7. Surface of provision covered with a nectar layer + - 8. Initiation of nesting restricted to morning - 9. Nest entrances enter substrate at a particular angle + - 10. Lateral burrows in each nest mostly parallel + - 11. Some old burrows refurbished - + 12. Some old cells refurbished - + 13. Some nests communal — + 14 Contributions in Science No. 206 A, peritomae demonstrates the following biological habits heretofore unnoticed within the genus: salivary material is secreted by the larvae; old cells are refurbished; and communal nest entrances are used. Provisioning of cells by more than one individual appears to be accidental. Literature Cited Baker, G. M., and P, F. Torchio. 1968. New records of Ascosphaera apis from North America. Mycologia 60: 189-190. Clout, G. A. 1956. Chalk brood and hunch back flies. Bee Craft 38: 135. Hicks, C. H. 1934. Some reared insect parasites and their hosts. Univ. Color. Studies 21: 265-271. Maassen, a. 1916. tJber Bienenkrankheiten. Mitt. Biol. Anst. Land.-Fortsn. 16: 51-58. Melville, R., and H. A. Dade. 1944. Chalk brood attacking a wild bee. Nature 153: 112. Torchio, P. F., and N. N. Youssef. 1968. The biology of Anthophora (Micrantho- phora) fiexipes and its cleptoparasite, Zacosmia maculata, including a descrip- tion of the immature stages of the parasite (Hymenoptera: Apoidea, Antho- phoridae). Jour. Kans. Ent. Soc. 41; 289-302. Accepted for publication September 8, 1970. Printed in Los Angeles, California by Continental Gra LOS CONTRIBUTIONS m'lli IN SCIENCE ANGELES COUNTY MUSEUM JMBER 207 February 4, 1971 TROMBICULID MITES OF THE GENUS MICROTROMBICULA (ACARINA) FROM COSTA RICA By James P. Webb, Jr. and Richard B. Loomis Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 summai-y 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 TROMBICULID MITES OF THE GENUS MICROTROMBICULA (ACARINA) FROM COSTA RICA James P. Webb, Jr.^ and Richard B. Loomis^ Abstract: Five species of Microtrombicula are reported for the first time from Costa Rica: M. perplexa, new species, type host Liomys salvini, from Guanacaste Province, Costa Rica and from Nicaragua; M. starretti, new species, type host Myotis nigricans, from Heredia Province, Costa Rica; M. sturnirae, new species, type host Sturnira lilium, from Chinandega, Nicaragua, Costa Rica, and Mexico; M. boneti (Hoffmann) NEW COMBI- NATION (Trombicula tibbettsi Brennan and White, NEW SYNONYMY) from bats of southern United States to Trinidad; and M. carmenae (Brennan and Jones) NEW COMBINATION, from bats of southern Mexico to Trinidad. A key to the larvae is included. Introduction This is the second report of trombiculid mites from Costa Rica obtained by field parties of the Los Angeles County Museum of Natural History. Geest and Loomis ( 1968) listed 13 species, 9 of which were new, of Pseudoschoen- gastia from approximately 291 of more than 500 examined rodents, taken between January 1962 and December 1964. They also discussed the biotic districts and provinces and the 24 localities where larvae of Pseudoschoen- gastia had been found. Described below are five species of Microtrombicula, including three new species and two new combinations, all reported from Costa Rica for the first time. Four species are recorded from bats and the fifth is known only from the heteromyid rodent, Liomys salvini (Thomas). In addition to the Costa Rican specimens, larvae are reported from southern United States, Mexico, Nicaragua, Panama and Trinidad. Acknowledgments Sincere gratitude is extended to Dr. Fred S. Truxal, Dr. Richard S. Casebeer, Julius C. Geest, Dr. Charles L. Hogue, David G. Marqua, Dr. Charles A. McLaughlin, Dr. Jay M. Savage, Dr. Andrew Starrett, Robert C. Stephens, and others who assisted in the recovery of chiggers in Costa Rica under the auspices of the Los Angeles County Museum of Natural History (LACM). The field studies and initial chigger research were supported by the United States Army Medical Research and Development Command Grant DA-MD-49-193-63-G94, F. S. Truxal and C. A. McLaughlin, Principal Investigators. Recent studies were supported by the U.S. Public Health Service ^Dept. Biol., California State College, Long Beach, California 90801. ^Dept. Biol., California State College, Long Beach, California 90801; and Research Associate, Los Angeles County Museum of Natural History. 1 2 Contributions in Science No. 207 Grant AI-03407 from the National Institute of Allergy and Infectious Dis- eases, Richard B. Loomis, Principal Investigator. In addition, we thank Dr. James M. Brennan of the Rocky Mountain Laboratory (RML), Hamilton, Montana, who graciously supplied paratypes and referred material. We are indebted to Dr. J. Knox Jones, Jr. and others of The University of Kansas (KU) for specimens from Nicaragua obtained under United States Army Medical Research and Development Command Grant DA-49-1 93-MD- 2215. Finally, appreciation is expressed to Lee C. Spath (formerly of California State College, Long Beach) for the preparation of the excellent illustrations, and to Lynell K. Tanigoshi (University of California at Riverside) for pro- viding us with the initial studies of M. perplexa, new species. Accounts of the Taxa The description of each new species is based upon the holotype, aug- mented by paratypes. Previously named species are redescribed on the basis of specimens examined. The terminology generally follows Wharton et al. (1951), Wharton and Fuller (1952), and Audy (1954). All measurements are in microns. A key to the species is included. Except for the specimens from the Rocky Mountain Lab (RML), the specimens from Costa Rica, including all holotypes, are in the Los Angeles County Museum of Natural History (LACM), and the other larvae are in the chigger research collection at California State College, Long Beach. Para- types will be given to the chigger research collection, California State College, Long Beach; Rocky Mountain Laboratory, Hamilton, Montana; United States National Museum; Institute of Acarology, Ohio State University; The B. P. Bishop Museum, Honolulu, Hawaii; and to other appropriate institutions and individuals. Subclass ACARINA Order Acariformes Suborder Prostigmata Family Trombiculidae Subfamily Trombiculinae Microtrombicula Ev/'mg, 1950 Microtrombicula: Webb and Loomis 1970:655 (Diagnosis of genus) Species known from Costa Rica: Trombicula boneti Hoffmann ( = T. tibbettsi Brennan and White); T. carmenae Brennan and Jones, Microtrom- bicula perplexa, new species; M. starretti, new species and M. sturnirae, new species. 1971 Trombiculid Mites from Costa Rica 3 Diagnosis of Costa Rican species. Pretarsala II present; palpotarsus with six branched and nude setae; palpotibial claw trifurcate or bifurcate; 2-3 genualae I; 2-3 pairs of sternal setae; coxa III with 1-4 setae; galeala nude or branched; sensilla flagelliform or with shaft slightly expanded; scutum mod- erately punctate; and tarsus III usually with proximal whorl of 3 setae (1 nude or nearly nude dorsal mastitarsala and 2 ventral branched setae) and with media whorl usually of 5 branched setae. Remarks. Subgenera have not been included as none of these species clearly belongs to any of the recognized categories. Microtrombicula perplexa, new species Figures 1, 4 Types. Larvae: Holotype and 95 paratypes from 7.5 km S Liberia, 125 m, Guanacaste Province, Costa Rica, off Liomys salvini Thomas, Sal- vin’s spiny pocket mouse, original number 0-3296 (LACM 26609), taken 8 August 1964 by C. L. Hogue, R. C. Stephens and J. C. Geest. Diagnosis. Larva similar to M. crossleyi (Loomis) and M. trisetica (Loomis and Crossley) in having coxa III with multiple setae (2-4); sternal setae 2-2-2; and palpotibial claw bifurcate, and differing from both in having 2 rather than 3 genualae I, sensilla with shaft slightly expanded instead of flagelliform, mastitarsala III branched rather than nude, and eyes absent. Description of holotype (with mean and range of 14 types, holotype and 13 paratypes, in parentheses, unless otherwise noted). Body slightly engorged, 220 by 155, eyes absent. Dorsal setae 2-10-10-10-8-8 + 20, total 68; measurements of humeral seta 31, anterior dorsal seta 21, posterior dorsal seta 38. Ventral setae 2-2-2 (sternals) 6-4-6 + 30, total 52; measurements of anterosternal seta 20, posterosternal seta 20, posterior ventral seta 38. Scutum: Subpentagonal, moderately punctate; sensilla slightly expanded with numerous setules along entire length. Scutal measurements: AW 59 (61, 59-62; 13), PW 73 (76.5, 73-80), SB 23 (24, 23-25), ASB 24 (24, 22-26), PSB 28 (26, 24-29), AP 19 (19, 16-22), AM 25 (27, 25-30), AL 26 (26, 23-29; 13), PL 28 (29, 28-31), S47 (48, 32-52; 12). Gnathosoma: Cheliceral blade with small tricuspid cap and elongate, pointed dorsal projection; cheliceral base and capitular sternum with moder- ate punctuation. Galeala nude. Palpal setae B/B/BNB (palpotibial setae with few branches); palpotarsus with 1 nude and 5 branched setae, and tarsala 6; palpotibial claw bifurcate. Leg I, 2 genualae, tarsala 14 (14, 13-16) ; leg II, tarsala 13 (13, 13-14), proximal microtarsala, and pretarsala; leg III, coxa with 2-4 (usually 3) branched setae, and mastitarsala with several branches. Leg measurements: I, 190 (203, 190-218); II, 182 (177, 166-186); III, 199 (192, 182-204); total, 571 (575, 550-598). 4 Contributions in Science No. 207 Figure 1. Microtrombicula perplexa, new species. A. Scutum and eyes. B. Dorsal aspect of gnathosoma with chelicera, cheliceral base, galeala, and tibial setae. C. Ventral aspect of palpal tibia and tarsus. D. Coxa III, with branched setae. E. Leg I, with specialized setae (measurements in microns) and bases of branched setae of the three distal segments. F. leg II; as above. G. Leg III; as above. H. Representative body setae; PD, posterior dorsal; 1 St, first sternal; H, humeral. 1971 Trombiculid Mites from Costa Rica 5 Remarks. Examination of 183 larvae revealed the following variation in the number of setae on coxa III: 2-2 setae (11), 2-3 (42), 3-3 (124), and 3-4 setae (6 larvae). This is the first species of Microtrombicula recorded from a heteromyid host. The specific name refers to the original problem of generic placement. Geographic distribution. Known from Guanacaste Province, Costa Rica and Boaco, Matagalpa, and Rivas, Nicaragua. Specimens examined (195), all from Liomys salvini. COSTA RICA. GUANACASTE PROVINCE: 8.3 km N Liberia, 3 Aug. 1964 (1); 5 km N, 4 km W Liberia (Finca Coyo^ar), 4 Aug. 1964 (1); 7.5 km S Liberia, 8 Aug. 1964 (9 L. salvinr, holotype + 182). NICARAGUA. BOACO: 17 km N. 15 km E Boaco, 9 Aug. 1967 (1). MATAGALPA: 1 km NE Esquipu- las, 16 March 1968 (6). RIVAS: 2 km N, 3 km E Merida, 7 April 1968 (3). Microtrombicula starretti, new species Figures 2, 4 Type. Larva. Holotype from 5 km SE Los Cartagos, Heredia Province, Costa Rica, off Myotis nigricans, black myotis, original numbers 0-2336 to 0-2350, lot (LACM 25646-60), taken 15 August 1963, by A. G. Hollister, F. G. Thompson, and A. Starrett. Diagnosis. Larva similar to Microtrombicula carmenae (Brennan and Jones) and M. sturnirae, new species, in having two pairs of sternal setae, 3 genualae I, palpotibial claw trifurcate, and coxa III unisetose, but differing from both in having sensilla with 12 branches (2-3 branches in M. carmenae and 3-5 in M. sturnirae) and cheliceral blade with a small dorsal projection (absent in other two species). Description of holotype. Body unengorged, 295 by 195; eyes 2/2, anterior larger, lens and ocular plate distinct. Dorsal setae 2-6-6-6-4-4-2-2, total 32; measurements of humeral seta 46, anterior seta 37, posterior dorsal seta 37. Ventral setae 2-2 (sternals) 4-6-8-6 + 8, total 36; measurements of anterosternal seta 22, posterosternal seta 26, posterior ventral seta 35. Scutum: Subpentagonal, moderately punctate; sensilla flagelliform with 12 short distal branches. Scutal measurements: AW 55, PW 65, SB 17, ASB 26, PSB 27, AP 19, AM 37, AL 26, PL 46, S 56. Gnathosoma: Cheliceral blade with small tricuspid cap; cheliceral base and capitular sternum lightly punctate. Galeala nude. Palpal setae B/B/BBB; palpotarsus with 2 nude and 4 branched setae, and tarsala 6; palpotibial claw trifurcate. Leg I, 3 genualae and tarsala (13); leg II, tarsala (14) and pretarsala; leg III, coxa unisetose, and 1 nude mastitarsala. Leg measurements: I, 209; II, 193; III, 216; total, 618. 6 Contributions in Science No. 207 Figure 2. Microtrombicula starretti, new species. A. Scutum and eyes. B. Dorsal aspect of gnathosoma with chelicera, cheliceral base, galeala, and tibial setae. C. Ventral aspect with palpal tibia and tarsus. D. Representative body setae: 1 St, first sternal; 2 St, second sternal; H. humeral; PD, posterior dorsal. E. Leg I, with specialized setae (measurements in microns) and bases of branched setae of the three distal segments. F. Leg II; as above. G. Leg III; as above. 1971 Trombiculid Mites from Costa Rica 7 Ecological notes. The type host was taken near Los Cartagos (Volcan Barba) in the Montane Life Zone of the Costa Rica Highlands Biotic District (Geest and Loomis, 1968). Microtrombicula sturnime, new species Figures 3, 4 Types. Larvae : Holotype and 15 paratypes from Volcan Casita, 720 m, Hda, Bellavista, Chinandega, Nicaragua, off Sturnira liliiim parvidens, yellow-shouldered bat, original number TEL 844 (KU 106090), taken 17 July 1966 by T. E. Lawlor; and 9 paratypes from Monteverde, 1380 m, Puntarenas Province, Costa Rica, off 3 Sturnira ludovici, Anthony’s bat, original numbers 0-2836 to-2838 (LACM 26144-46) taken 14 May 1964 by F. S. Truxal, C. A. McLaughlin, and J. M. Savage. Diagnosis. Larva similar to Microtrombicula carmenae (Brennan and Jones) and M. starretti, new species, in having two pairs of sternal setae, 3 genual ae I, palpotibial claw trifurcate, and coxa III unisetose, and differing from M. carmenae in having coxa III seta on or near anterodistal margin (medial, not marginal in M. carmenae) and from M. starretti in having sensilla with 3-5 branches rather than 12 branches. Description of holotype (with mean and ranges of 8 types, holotype and 7 paratypes, in parentheses, unless otherwise noted). Body engorged, 430 by 230, eyes 2/2, distinct, nearly equal in size, separated by striae; ocular plate distinct. Dorsal setae 2-6-6-6-6 + 10, total 36; measurements of humeral seta 36, anterior dorsal seta 32, posterior dorsal seta 42. Ventral setae 2-2 (sternals) 4-4-4-8-8 + 10, total 42; measurements of anterosternal seta 30, posterior ventral seta 23. Scutum : Subpentagonal, moderately punctate; sensilla flagelliform with 3-5 distal branches. Scutal measurements : AW 36 (36, 35-38), PW 42 (44.5, 42-46), SB 13 (14, 13-14), ASB 24 (26, 24-27), PSB 21 (20, 19-21), AP 22 (23, 22-24), AM 20 (22, 20-23), AL 19 (20, 17-21), PL 34 (33.5, 31-36), S 39 (43, 39-45; 4). Gnathosoma: Cheliceral blade with small tricuspid cap; cheliceral base with light and capitular sternum with moderate punctation. Galeala nude. Palpal setae B/B/BBB; palpotarsus with 1 nude and 5 branched setae, and tarsala 7; palpotibial claw trifurcate. Leg I, 3 genualae, tarsala 13 (13, 12-15); leg II, tarsala 10 (11, 10-13), proximal microtarsala and pretarsala; leg III, coxa unisetose and mastitarsala with 1 long basal branch. Leg measurements: I, 201 (198, 181-211); II, 169 (170, 147-177); III, 202 (198, 190-203); total, 572 (566, 518-580). Remarks. Larvae from Costa Rica and Nicaragua have two branched setae on trochanter III whereas those from Mexico possess a single seta. 8 Contributions in Science No. 207 Figure 3. Microtrombicula sturnirae, new species. A. Scutum and eyes. B. Dorsal aspect of gnathosoma with chelicera, cheliceral base, galeala, and tibial setae. C. Ventral aspect of palpal tibia and tarsus. D. Representative body setae: 1 St, first sternal; PD, posterior dorsal; H, humeral. E. Leg I, with specialized setae (measure- ments in microns) and bases of branched setae of the three distal segments. F. Leg II; as above. G. Leg III; as above. 1971 Trombiculid Mites from Costa Rica 9 The larvae of M. sturnirae were recovered from the ears of Sturnira lilium parvidens taken in Nicaragua and Mexico. None of the 55 Costa Rican S. 1. parvidens yielded this species although six of 121 S. ludovici and one of 12 S. mordax possessed numerous larvae. The bat hosts were taken from April through October. Geographic distribution. Known from three provinces in Costa Rica, southern Sonora, Mexico, and western and central Nicaragua. Specimens examined (73). COSTA RICA. CARTAGO PROVINCE: Tapanti, 30 July 1963, 2 Sturnira ludovici (14). HEREDIA PROVINCE: 2 km N El Angel Falls, 18 Aug. 1964, Sturnira mordax (17). PUNTARENAS PROVINCE: Finca Los Helechales, 3 Oct. 1964, S. ludovici (2) ; Monte- verde, 14 May 1964, S. ludovici (9). MEXICO. SONORA: 13 km SSE Alamos, Rio Cuchujaqui, 10-11 April 1963, 2 Sturnira lilium (7). NICA- RAGUA. CHINANDEG A : Volcan Casita, Hda. Bellavista, 17 July 1966, S. lilium (16). CHONTALES: 1 km N, 2.5 km W Villa Somoza, 6 Aug. 1967, S. lilium (8). Microtromhicula boneti (Hoffmann), NEW COMBINATION Figure 4 Trombicula (Trombicula) boneti Hoffmann, 1952: 87, holotype from Cueva de Quintero, Tamaulipas, Mexico, host Mormoops megalophylla, Dec. 30, 1950; Brennan and Jones, 1959: 12. Eltonella (Coecicula) boneti: Vercammen-Grandjean, 1965: 44. Trombicula tibbettsi Brennan and White, 1960: 348, type from Frio Cave, Uvalde Co., Texas, host Mormoops megalophylla senicula, Nov. 17, 1956, NEW SYNONYMY; Brennan and Jones, 1960: 533; Goodwin and Greenhall, 1961 : 204; Loomis and Crossley, 1963 : 379; Brennan and Yunker, 1966: 257; Brennan, 1967: 155. Eltonella (Coecicula) tibbettsi : Vercammen-Grandjean, 1965: 44. Diagnosis. Larva similar to Microtromhicula carmenae (Brennan and Jones, M. starretti, new species, and M. sturnirae, new species, in having 3 genualae I, 2 pairs of sternal setae, and coxa III unisetose but differing from them in having palpus with pronounced lateral angulation and lacking eyes. Description of species. Based upon 2 presumed paratopotypes : Body engorged, 370 by 265; eyes absent. Dorsal setae 2-6-6-6-6-6-6-2, total 40; measurements of humeral seta (44, 45), anterior dorsal seta (38, 39), posterior dorsal seta (28, 29). Ventral seta 2-2 (sternals) 4-6-8-6-4-4-6 + 6, total 48; measurements of anterosternal seta (35, 36), posterosternal seta (22, 26), posterior ventral seta (24, 26). Scutum : Subpentagonal, posterior margin with acute angle; AM setal base below anterior line of AL setal bases, sensilla flagelliform with 2-4 long distal branches. 10 Contributions in Science No. 207 Figure 4. Localities of Microtrombicula in Costa Rica: M. boneti ■ , M. carmenae # , M. per pie xa ▼ , M. starretti ♦ , and M. sturnirae A • Scutal measurements: AW (57, 57), PW (70, 70), SB (20, 21), ASB (23, 26), PSB (26, 26), AP (19, 23), AM (36, 39), AL (27, 27), PL (38, 46), S (48, -). Gnathosoma: Cheliceral blade with medium sized tricuspid cap. Galeala stout with several branches. Palpal setae B/B/BNB; palpotarsus with 1 nude or nearly nude and 5 branched setae, and tarsala; palpotibial claw trifurcate. Leg I, 3 genualae and tarsala (17, 17); leg II, tarsala (12, 12) and pretarsala; leg III, coxa unisetose, and 1 nude mastitarsala. Leg measurements: I (197, 215); II (168, 181); III (189, 197); total (554, 593). 1971 Trombiculid Mites from Costa Rica 11 Taxonomic remarks. The types of Trombicula boneti were from Mor- moops megalophylla, a phyllostomatid bat, taken in northeastern Mexico. Trombicula tibbettsi Brennan and White (1960) was based upon larvae from Texas recovered from the same host species, as well as from several other species of bats taken in Alabama and Trinidad. Vercammen-Grandjean (1965) placed them in the genus Eltonella, subgenus Coecicula, and sepa- rated the two species on the basis of the number of body setae. A study of the original description and two presumed paratypes (RML) of T. boneti revealed 32-40 dorsal body setae and 34-38 ventral body setae. These counts overlap those of the original description and of four larvae of T. tibbettsi (RML) with dorsal setae numbering 28-34 and ventral body setae 36-42. In addition, a careful comparison of other characteristics, including the scutal shape, acute palpal angulation, lack of eyes, two pairs of sternal setae, and the 2-5 distal branches on the sensillae, has led to the conclusion that all of these larvae belong to the single species, M. boneti. Vercammen-Grandjean ( 1965) reported that the eight larvae of M. boneti from Uvalde Co., Texas had shorter legs than two larvae from San Jose Province, Costa Rica. Selected scutal measurements of the same two specimens from Costa Rica (RML) and 10 larvae from La Aduana, Sonora, Mexico, also revealed a larger size for the Costa Rican specimens. The ten larvae from Sonora, Mexico measured : AW 57, 56-58; PW 73, 71-78; SB 21, 19-22; PL 46, 43-50. The two specimens from Costa Rica measured : AW 66, 65-67; PW 88, 87-89; SB 27, 27; PL 47, 46-48. Ecological notes. Larvae of M. boneti have been recovered from the wings of natalid, phyllostomatid and vespertilionid bats. Geographic distribution. Known from Alabama and southwestern Texas, Mexico (Cape Region of Baja California, southern Sonora, Sinaloa, Tamauli- pas, southward to Chiapas), Costa Rica (San Jose Province), Panama, Curasao, and Trinidad. Specimens examined (50). COSTA RICA. SAN JOSfi PROVINCE: Rock Cave, W of Santa Ana, Desmodus rotundus (2, RML). MEXICO. BAJA CALIFORNIA SUR: 5 km SE Pescadero, 10 July 1963, Natalus stramineus (8). CHIAPAS: 3 km N Las Margaritas, 23 July 1964, D. rotundus ( 1). SONORA: 1 km SW La Aduana, 6 Aug. 1963, Leptonycteris sanborni (28) ; La Aduana, 7 Aug. 1964, Glossophaga soricina (2). SINA- LOA: 2 km SW Santa Lucia, 28 July 1963, Mormoops megalophylla (4) ; 1.5 km W Copala, 29 July 1963, Pteronotus parnellii (1). TAMAULIPAS : Cueva de Quintero, 30 Dec. 1950, M. megalophylla (2, RML). TEXAS. Uvalde Co.: Frio Cave, 22 Jan. 1956, M. megalophylla (3). Additional records. PANAMA (Brennan and Yunker, 1966). PAN- AMA: Chilibre, Chilibre Caves, 2 Aug. 1960, Pteronotus suapurensis (1). ALABAMA (Brennan and White, 1960). Conecuh Co. : Sanders Cave, 9 Aug. 1953, My Otis grisescens (1); Lauderdale Co. : Collier Cave, 27 Sept. 1953, M. grisescens (1). TEXAS (Brennan and White, 1960). Uvalde Co. : 12 Contributions in Science No. 207 Frio Cave, 17 Nov. 1956, Mormoops megalophylla senicula (3). WEST INDIES. Bahama Isl. (Brennan, 1967), New Providence: Hunt’s Cave, 22 Aug. 1949, Erophylla sezekorni (1). CURACAO (Brennan, 1967): Cave of Hato, 22 Dec. 1950, Glossophaga elongata (1). TRINIDAD (Brennan and White, 1960 and/or Brennan and Jones, 1960), St. Andrew Co.: Tamana Hill Cave, 12 June 1956, 3 Chilonycteris rubiginosa rubiginosa (16), 20 Nov. 1957, Mormoops tumidiceps (5), M. megalophylla (4). Microtrombicula carmenae (Brennan and Jones) , NEW COMBINATION Figure 4 Trombicula carmenae Brennan and Jones, 1960: 512, holotype from Em- peror Valley Zoo, Port of Spain, St. George Co., Trinidad, host Phyllos- tomus discolor, Goodwin and Greenhall, 1961 : 204; Brennan and Yunker, 1966: 253. Eltonella {El ton ell a) carmenae, Vercammen-Grandjean, 1965 : 44. Diagnosis. Larva similar to Microtrombicula starretti new species, and M. sturnirae new species, in having coxa III unisetose, 2 pairs of sternal setae, 3 genualae I, and palpotibial claw trifurcate, and differing from M. starretti in having sensilla with 2-3 branches (M. starretti with 12 branches) and from M. sturnirae in having coxa III medial, not marginal (on or near anterodistal margin in M. sturnirae). Description of species (based on the holotype after Brennan and Jones, 1960, 1 paratype, 3 referred slides from RML and specimens examined). Body partly engorged, 465 by 300; eyes 2/2, anterior lens larger or equal in size with posterior lens, no ocular plate. Dorsal setae 2-6-6-6-4-2-2, total 28; measurements (paratype) of hu- meral seta 37, anterior dorsal seta 37, posterior dorsal seta 32. Ventral setae (paratype) 2-2 (sternals) 4-2-6-6-4-4, total 30; measure- ments of anterosternal seta 22, posterosternal seta 23, anterior ventral seta 22, posterior ventral seta 29. Scutum : Subpentagonal, lightly punctate; sensilla flagelliform with 2-3 long distal branches. Scutal measurements ( 1 paratype in parentheses) : AW 48 (46), PW 60 (58), SB 18 (15), ASB 26 (24), PSB 20 (21), AP 21 (21), AM 31 (28), AL 23, (21 ), PL 36 (35), S 44 (39). Scutal measurements including the mean and range of selected samples (in parentheses) : Costa Rica (11): AW 45, 42-49; PW 55, 53-57; SB 15, 15-16; ASB 23, 22-27; PSB 23, 20-24; AP 23, 21-26; AM 30, 27-32 (10); AL 22, 20-24; PL 36, 34-39. Nicaragua (11): AW 49, 46-50; PW 58, 57-59; SB 16, 15-19; ASB 26, 23-29; PSB 25.5, 24-27; AP 25, 22-27; AM 30, 24-32 (9); AL 24, 23-27; PL 40, 47-42. Oaxaca, Mexico (10): AW 49, 46-53 (8);PW 62, 56-70 (9); SB 17, 15-19 (9); ASB 26, 24-30 (9); PSB 25, 23-28 (9); AP 25, 22-28; AM 29, 24-31 (5); AL 24, 22-27; PL 39, 34-42 (6). Gnathosoma: Cheliceral blade with small tricuspid cap; cheliceral base 1971 Trombiculid Mites from Costa Rica 13 and capitular sternum moderately punctate. Galeala nude. Palpal setae B/B/BNB (paratype exhibited lateral palpotibial seta with several small branches and ventral palpotibial seta forked); palpotarsus with 1 nude and 5 branched setae, and tarsala; palpotibial claw trifurcate. Legs (measurements of 1 paratype in parentheses): Leg I, 3 genualae and tarsala 13 (14); leg II, tarsala 1 1 ( 10) and pretarsala; leg III, coxa uni- setose, and 1 nude (sometimes with 1-3 barbs) mastitarsala. Measurements of tarsala I and II with means and extremes of 30 larvae from the bat host, Artibeus jamaicensis: Costa Rica (10): Tj 15, 13-17; Tn 11.5, 10-13. Nica- ragua (10): Ti 15, 14-16; T„ 12, 10-13. Oaxaca, Mexico (10); Tj 15, 14-16; Tn 12, 10-13. Leg measurements of 1 paratype: I, 213; II, 180; III, 212; Total, 605. Ecological notes. Specimens of M. carmenae and M. sturnirae have been taken from the same individual host and from different hosts at the same locality in Costa Rica. Larvae are recorded from all months except January, February, May and December. Geographic distribution. Known from eastern Oaxaca, Mexico, Nica- ragua, Costa Rica, Panama, and Trinidad. Specimens examined (151). COSTA RICA. CARTAGO PROVINCE: Tapanti, 30 July 1963, Sturnira hidovici (1). GUANACASTE PROVINCE: Bahia del Coco, 19 July 1962, Phyllostomus discolor (2); 9 km N Liberia, 15 July 1962, Artibeus jamaicensis (1); Playas del Coco, 20-23 July 1962, 3 A. jamaicensis (3), 3-4 April 1963, 2 A. jamaicensis (17). HEREDIA PROVINCE: 2 km N El Angel Falls, 18 Aug. 1964, Sturnira mordax (6). PUNTARENAS PROVINCE: Boca de Barranca, 1 Aug. 1962, P. discolor (1); Finca Don Nicholas, 3 km N Tambor, 14 Nov. 1964, 2 P. discolor (5); Finca Los Helechales, 4 Oct. 1964, A. jamaicensis (6); Rincon, Osa Pen- insula, 28 June 1963, Artibeus lituratus (1), 3 July 1963, A. Uturatus (5). MEXICO. OAXACA: Las Minas (8 km E Tapanatepec), 10 June 1964, 2 A. jamaicensis (21). NICARAGUA. BOACO: 14 km S Boaco, 18 July 1964, P. discolor (9). CARAZO: 3 km N, 4 km W Diriamba, 12 Aug. 1967, A. jamaicensis (1). GRANADA: Finca El Progresso, 10 km SE Guanacaste, 15 June 1966, P. discolor (15). JINOTEGA: Yali, 4 Aug. 1966, A. jamai- censis (16). MANAGUA: 2 km N Sabana Grande, 11 June 1966, 2 P. dis- color (5). NUEVA SEGOVIA: 2 km N, 1 km E Jalapa, 24-27 July 1967, 4 A. jamaicensis (21). RIVAS: 3 km N, 4 km W Sapoa, 24 June 1966, Des- modus rotundas (11). PANAMA. PANAMA: Pacora, 6 June 1961, Phyl- lostomus hastatus (1). WEST INDIES. TRINIDAD, Mayaro Co.: Point Radix, 24 April 1959, D. rotundus (1). St. Andrew Co.: Sangre Grande, 16 April 1959, D. rotundus (1). St. George Co.: Emperor Valley Zoo, Port of Spain, 14 March 1958, P. discolor (1 paratype, RML). Additional records (Brennan and Yunker, 1966). PANAMA. BOCAS DEL TORO: Rio Changena, 19 Sept. 1961, Artibeus jamaicensis (4), Sturnira ludovici ( 1 ) . 14 Contributions in Science No. 207 KEY TO THE LARVAE OF MICROTROMBICULA FROM COSTA RICA 1. Two genualae I; sternal setae 2-2-2; sensilla slightly expanded; coxa III with 2-4 setae (from Liomys salvini) M. perplexa Three genualae I; sternal setae 2-2; sensilla flagelliform; coxa III unisetose (from bats) 2 2. Palpus with pronounced lateral angle, tarsus III with medial whorl of 4 setae M. boneti Palpus without pronounced lateral angle, tarsus III with medial whorl of 5 setae 3 3. ASB > PSB, coxa III seta on anterodistal margin (from Sturnira species) M. sturnirae ASB ^ PSB, coxa III seta medial, not marginal 4 4. Sensilla with 2-3 branches, proximal mastitarsala III with 1-3 barbs M. carmenae Sensilla with 10-12 branches, proximal mastitarsala III nude (from Myotis nigricans) M. starretti Resumen Cinco especies de Microtrombicula se reportan por primera vez de Costa Rica: M. perplexa, especie nueva, tipo hospedero Liomys salvini, de la provincia de Guanacaste, Costa Rica, y de Nicaragua; M. starretti, especie nueva, tipo hospedero Myotis nigricans, de la provincia de Heredia, Costa Rica; M. sturnirae, especie nueva, tipo hospedero Sturnira lilium, de Chinan- dega, Nicaragua, Costa Rica, y Mexico; M. boneti (Hoffmann) COMBINA- CION NUEVA {Trombicula tibbettsi Brennan y White, SINONIMIA NUEVA) de los murcielagos del sur de los Estados Unidos hasta Trinidad; y M. carmenae (Brennan y Jones) COMBINACION NUEVA de los murcie- lagos de sur de Mexico hasta Trinidad. Una clave de las larvas esta incluida. Literature Cited Audy, J. R. 1954. Notes on the taxonomy of trombiculid mites, with descriptions of a new subgenus. Malaysian Parasites IX. Stud. Inst. Res. Malaya (26): 123- 170. Brennan, J. M. 1967. New records of chiggers from the West Indies. Stud. Fauna Curacao and Other Caribbean Isis. 24(95): 146-156. Brennan, J. M., and E. K. Jones. 1959. Keys to the chiggers of North America with synonymic notes and descriptions of two new genera (Acarina: Trombi- culidae). Ann. Ent. Soc. Amer. 52(1): 7- 15. Brennan, J. M., and E. K. Jones. 1960. Chiggers of Trinidad, B. W. I. (Acarina: Trombiculidae). Acarologia 2(4) : 493-540. Brennan, J. M., and J. S. White. 1960. New records and descriptions of chiggers (Acarina: Trombiculidae) on bats in Alabama. J. Parasit. 46(3) :346-350. 1971 Trombiculid Mites from Costa Rica 15 Brennan, J. M., and C. E. Yunker. 1966. The chiggers of Panama (Acarina: Trombiculidae). In Wenzel, R. L. and V. J. Tipton, eds., Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago, 111., p. 221-226. Ewing, H. E. 1950. A redescription of four genera of chigger mites, together with a description of a new genus and subgenus (Acarina, Trombiculidae). Proc. Ent. Soc. Wash. 52(6):291-299. Geest, J. C., and R. B. Loomis. 1968. Chiggers of the genus Pseudoschoengastia (Acarina: Trombiculidae) from Costa Rica. Los Angeles Co. Mus., Contrib. Sci. 150:1-49. Goodwin, G. G., and A. M. Greenhall. 1961. A review of the bats of Trinidad and Tobago. Bull. Amer. Mus. Nat. Hist. 122(3) : 191-301. Hoffmann, A. 1952. Contribuciones al conocimiento de los trombiculidos Mexi- canos. Ciencia 12(3-4): 87-94. Loomis, R. B. 1954. A new subgenus and six new species of chigger mites (genus Trombicula) from the central United States. Univ. Kans. Sci. Bull. 36(13): 919-941. Loomis, R. B., and D. A. Crossley, Jr. 1953. A new species of chigger from east- ern Kansas (Acarina: Trombiculidae). J. Kans. Ent. Soc. 26(l):32-34. Loomis, R. B., and D. A. Crossley, Jr. 1963. New species and new records of chiggers (Acarina: Trombiculidae) from Texas. Acarologia 5(3) :371-383. Vercammen-Grandjean, P. H. 1965. Revision of the genera: Eltonella Audy, 1956 and Microtrombicula Ewing, 1950, with descriptions of fifty new species and transferal of subgenus Chiroptella to genus Leptotrombidium (Acarina: Trom- biculidae). Acarologia 7 (fasc. suppl.) : 34-257. Webb, J. P., Jr., and R. B. Loomis. 1970. A new subgenus of intranasal chiggers of the genus Microtrombicula from North America and Korea. J. Med. Ent. 7(6):655-663. Wharton, G. W., and H. S. Fuller. 1952. A manual of the chiggers. Mem. Ent. Soc. Wash. (4): 1-185. Wharton, G. W., D. W. Jenkins, J. M. Brennan, H. S. Fuller, G. M. Kohls, AND C. B. Philip. 1951. The terminology and classification of trombiculid mites (Acarina: Trombiculidae). J. Parasit. 37(1): 13-22. Accepted for publication September 8, 1970 .1; LOS ANGELES COUNTY MUSEUM 1 CONTRIBUTIONS IN SCIENCE P UMBER 208 February 4, 1971 i 1 THE ROLE OF INTROGRESSIVE HYBRIDIZATION 1, j IN THE DEVELOPMENT OF SALVIA: SECTION AUDIBERTIA (LABIATAE) j |, 1 By William A. Emboden, Jr. 1 1 ;i !| I Los Angeles County Museum of Natural History • Exposition Park j Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) JDouble space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 ROLE OF INTROGRESSIVE HYBRIDIZATION IN THE DEVELOPMENT OF SALVIA: SECTION AUDIBERTIA (LABIATAE) By William A. Emboden, Jr/ Abstract: The nineteen western North American species of Salvia (section Audibertia), while occupying distinct niches, are frequently sympatric throughout a part of their range. Hybridization within the section is not uncommon and blocks of characters found in closely related species suggest hybrid origins. Prior to this paper only one instance of introgression has been reported, and the documentation of one more pair of introgressed species suggests that introgression is of greater importance than was previously realized, and may have played an even more significant role in the formation of the plexus as it is known today. Degrees of genetic exchange within the section are diagrammed. The integrity of subsection Echinosphace is vindicated by a corrected chromosome count for one of the species which was miscounted and another member which was imperfectly known cytologically. Further chromosome counts of subsection Pycno- sphace substantiate thirteen as the gametic chromosome number; previous reports had varied considerably. Introduction The role which hybridization has played in the development of the North American species of Salvia: section Audibertia, will never be fully comprehended, for we have almost no fossil record for this derivative of the Madro-Tertiary geoflora. Studies by Epling (1938), Munz (1927), Epling, Lewis, and Raven (1962), and Emboden (1964 and 1967) all point to hybridization as one of the most important factors in the evolution of Salvia as it occurs in the western United States and Baja California. The arguments rest largely upon morphological and palynological data. Beyond simple hybridization, there is introgression, defined by Anderson in 1953 as, “the concept of infiltration of germplasm of one species into that of another as a consequence of hybridization and repeated backcrossing.” This definition is refined from the less eloquent enunciation by Anderson and Hubricht who originated the concept in 1938. In his 1938 monograph of section Audibertia, Epling pointed out that this section of the genus Salvia is confined primarily to arid portions of the southwest. Most members are components of either the Larrea-Franseria shrub formation of the Colorado desert or the related Artemesia californica- Salvia shrub formation of the coastal plain. Since distributions were well ^Research Associate, Los Angeles County Museum of Natural History; Professor of Biology, San Fernando Valley State College, Los Angeles, California 91324. 1 2 Contributions in Science No. 208 mapped by Epling, one need only note the addition of S. chionopeplica and its range to this section of the monograph. Salvia chionopeplica was dis- covered by Wiggins in 1931, thirty-six miles east of Rasario (Stanford University type specimen 5300). Epling (1940) agreed to its inclusion in subsection Jepsonia of Audibertia. This subsectional disposition was later vindicated by chromosome counts made by Epling, Lewis, and Raven (1962). Of the nineteen species of Salvia in this section, seven are conspicuous localized elements of the Larrea-Franseria association, five are important elements of the Artemisia calif ornica-Salvia formation, and the remaining seven are ubiquitous, or are disjunct elements of one of the previously mentioned groups; all, presumably, are derivatives of the Madro-Tertiary geoflora. This section makes its greatest impression as an element of the coastal sage formation. It was Epling (1938) who first suggested that most members of section Audibertia have arisen through hybridization. This he based upon the frequency with which natural hybrids occur, especially in instances of sym- patric distributions of species of subsections Jepsonia and Munzia. Com- munication with Dr. Philip Munz has established that the report of a hybrid between Salvia carduacea and S. mellifera (1959): was a misreading of Epling’s monograph (1938) and that he knows of no such hybrid. This eliminates the inexplicable link between subsections Echinosphace and Parishiella. Floral morphology in section Audibertia, comprised of nineteen species is more variable than that of section Calosphace (circa five hundred species) and one sees a progressive change from one extreme to the other, especially in the morphology of the androecium. A careful study of the overall morphology of Audibertia suggests a series of interchanges of blocks of characters that could only have arisen through hybridization and subsequent introgression. Epling would relate section Audibertia to subgenus Calosphace, but the examples which he uses, S. clinopodioides and S. axillaris, are most unlikely progenitors for this western plexus. Recent evidence for the antiquity of the genus in north temperate regions perhaps as early as the Miocene (Emboden, 1964) argues against making comparisons between two species of more southern distribution and an entire assemblage of nineteen species which could be assigned subgeneric status. This coupled with the Pleistocene glaciation and progressive aridity of the southwest might in itself be argument against making such contemporary species comparisons. However, recent evidence from Crowell (1962 and 1968) on paleogeographical coordinations suggests that the coastal block west of the San Andreas fault has been trans- ported northward along the coast since the Early Miocene by a distance of perhaps 175 miles. Such geographical, climatic, and paleogeographical co- ordinations make speculation on specific ancestors of section Audibertia seem quite inconsequential. The significance of this sort of data is that it helps explain some of the opportunities which occurred in the past for hybridization 1971 Introgression in Salvia 3 of now disjunct entities, some of which still retain the capacity to exchange genes. Contemporary Aspects of Hybridization A summary of known hybridization within section Audibertia is presented in Figure 1 and is fairly representative of the extent of hybrid- ization which may be realized. All but S. mohavensis, S. pachyphylla, and S. clinopodioides have been grown in the gardens and greenhouses of San Fernando Valley State College at Northridge, California, and repeated reciprocal crosses have been attempted between all of these species. It is possible to make crosses between plants which are seasonally isolated by altering amounts of water and temperatures, both being important factors in floral initiation. Most of the crosses obtained had previously been reported to occur in nature or had been found growing in botanical gardens when allopatric species were grown in proximity to each other. It is obvious, both in morphology and breeding behavior, that subsections Greenostachys and Echinosphace are isolated from the other subsections, and the only con- nection between S. columbariae of subsection Pycnosphace and the other subsections is the infrequent hybrid with S. meUifera of subsection Parishiella. As a generalization, it may be said that whenever members of subsections Jepsonia and Parishiella have sympatric distribution patterns, it is likely that hybrids will be found, especially in instances of a disturbed environment. Prior to this paper, only one documented instance of introgressive hybrid- ization has been found in this section and that is between S. apiana and S. meUifera. So important is this latter instance of introgression that it drew the attention of Epling (1947), Anderson and Anderson (1954), Webb and Carlquist (1964), and Emboden (1964, 1967, and 1969). Since the 1949 publication by Anderson of Introgressive Hybridization, this phenomenon has continued to be one of the most controversial aspects of evolution among higher plants, and to a lesser extent among the cryptogams and lower animals. It is especially noteworthy, therefore, that an unpublished instance of introgression in this section of one of the largest genera of flowering plants be documented. Chromosomes, Cytology, and Subsectional Disposition Before discussing the particulars of introgression in section Audibertia, it is necessary to clarify some cytological phenomena that are in error or imperfectly known. Epling, Lewis, and Raven (1962) did a great deal to clarify the unfortunate miscounts published by Carlson and Stuart (1936), Stewart (1939), and Delestaing (1954), and added seven unpublished counts. Most counts were made from somatic material, especially roottips, because of the difficulties attendant with fixation of the oily buds. Of the nine species from which meiotic counts were taken by Epling et al., none showed meiotic irregularities. All members of section Jepsonia and Parishiella, of which there are fourteen species, have a gametic number equal to fifteen. Of the collections of S. columbariae, the only member of subsection Pycno- 4 Contributions in Science No. 208 sphace, the gametic number was thirteen. Since this species hybridizes with S. mellifera, having a gametic number of fifteen, it seemed probable that more than one number might occur in this species. Earlier reports of gametic numbers of sixteen, fourteen, and eight were questioned by Epling et al. with the following comment, “. . . the possibility of more than one chromo- some number in this species cannot be excluded. However, the known errors in determining chromosome numbers for other species leads us to question the validity of reports other than n equals thirteen for this species.” Sub- sequently, I have sampled four widely disjunct populations of S. columbariae and have obtained meiotic counts of thirteen in all cases (vouchers L.A. 7678-76881). Whereas subsection Echinosphace has been characterized by Epling et al. as being “chromosomally and morphologically the most variable within Audibertia,” it seems likely that the variable morphology is due to con- temporary selective pressure, for when grown under uniform greenhouse conditions, all of these members show a striking degree of morphological similarity except for the annual habit of S. carduacea. As for chromosomal dissimilarity, I must take exception. The material of S. greatae collected by Lewis was not amenable to meiotic preparations and consequently the examination of three premeiotic divisions in another tissue of a bud too immature for meiotic divisions led to a report of circa 30 as a somatic count. Subsequently, I have had the opportunity to observe a prepared slide of root- tips of S. greatae (L.A. 77004) from a plant growing in a canyon north of Hidden Springs in the Orocopia Mountains of Riverside County, California, and this material revealed several cells with a somatic number of 32; this then is consistent with other members of subsection Echinosphace which have chromosome numbers built upon a gametic number of 1 6. Epling et al. report a gametic count of circa 32 from a single plant of S. funerea horn Hole in the Rock Spring, Inyo County, California. The authors state, “. . . the probability exists that it was an autotetraploid but aberrant for the species as a whole.” Since there were frequent quadrivalents in the chromo- somal configurations of this plant (L.A. 76882), it was thought to be an autotetraploid. Although, as these authors note, “genomes of Salvia species of section Audibertia are structurally similar, and quadrivalents might be expected in an allotetraploid.” In order to clarify this enigmatic count, I collected flowering material from several plants of S. funerea growing in Titus Canyon of Death Valley, one mile west of Klare Spring. Meiotic prepa- rations of this material revealed a maximum configuration of 22 pairs and five alternate disjunctive rings of four chromosomes, substantiating the earlier count of 64 as a somatic count and 32 as a meiotic number. Anaphase was normal in every way, and the pollen was 97.5 per cent viable as stained with lactophenol blue. The question of allopolyploidy versus autopolyploidy remains unresolved. Further cytological work as well as an attempt to re- construct the putative allopolyploid from extant species might provide a 1971 Introgression in Salvia 5 solution. It does not seem likely that such crosses can be made, as they have already been attempted, and the possibility of reconstructing an autopolyploid from colchicine treatment of any member of this subsection does not seem promising. These additional counts do argue strongly for the unity of sub- section Echinosphace as established by Epling in 1938, in contrast to the later statement (as quoted previously) concerning chromosomal dissimilarity. The only apparent chromosomal dissimilarity is the size relationships; S. carduacea, an annual of semiarid regions, has much larger chromosomes than the other species in this subsection. The most questionable subsectional disposition is that of Jepsonia and Parishiella, the members of which show no chromosomal dissimilarity in size or number, and freely exchange genes when found in sympatric dis- tributions. The morphology is quite similar among these species with the exception of the conspicuous and highly colored bracts of S. pachyphylla and a few of the subspecies of S. dorii (formerly S. carnosa). It is interesting to note that palynological size relationships do not as readily distinguish subsections Jepsonia and Parishiella as the others (Emboden, 1964). Introgression The observation of introgression dates back to the early eighteenth century, for Lawson (1714) hints at introgression in describing the natural history of Carolina. Subsequently papers have appeared which suggest that this phenomenon is common to most families of flowering plants, and in- cludes such woody members as the pines, spruces, oaks, junipers, birches, etc. Some of this data has paleohistorical import as well. Both fossil leaves and pollen suggest paleohistorical introgression of the oaks, Quercus macro- car pa and Q. gambelii in the Early and Late Pleistocene (Maze, 1968). Since the 1949 publication of Anderson, hundreds of papers have appeared in which introgression figures as a major feature of evolution and speciation. It is not the purpose of this paper to present a bibliography of such studies, but since this plethora of information has accrued, a synoptical bibliography on introgression would be in order. This aspect of hybridization, once thought to be restricted to a few plant species, has now been found in animal genera as well. It includes such genera as: Drosophila (Van Valen, 1969; Pipkin, 1968); Rotaria (Pefler, 1956); Hyla (Meehan, 1966); Bufo (Guttman, 1968); Cyanocitta (Brown, 1963), to mention but a few. It is to be anticipated that greater numbers of animal genera, including the vertebrates, will soon figure in introgression studies. Like polyploidy, the occurrence of this phenomena, has been neglected by zoologists until recently. A lesser frequency among animals is to be expected due to behavioral factors, ease of migration, frequently shorter life spans, and generally less physiological plasticity. It has not been emphasized by botanists that introgression is probably much more frequent among perennials than among annuals, for the conditions which permit the establishment of introgressants are often ephemeral, but 6 Contributions in Science No. 208 once established, the effect upon gene flow is considerable and is operative for several successive years. The role of introgression in evolution is multifaceted and certainly con- troversial. It is not of equal import in all taxa in which it is known to occur, but it is of some consequence to authenticate those instances in which it does occur or has occurred with some frequency. Hulten (1956) has presented an impressive body of evidence substantiating introgression to be the primary agent responsible for the variation encountered in the Cerastium alpinum complex, and this would seem to obtain for the genus Elymus (Brown and Pratt, 1960), Tucker’s (1970) study of the Quercus undulata complex is the synopsis of a long-term study and indicates that introgression and trans- gressive segregation account for the variation patterns in this group. These are but a few examples of studies in which introgression is a major feature of evolution. With respect to the genus Salvia in western North America, the inter- change of blocks of characters among the species, noted by Epling (1938) were attributed to hybridization. It would be more appropriate to say that this variation could come about only through hybridization followed by subsequent backcrossing, or in a word— introgression! Much of the variation in Audibertia may be accounted for historically by post Pleistocene dis- ruption resulting in the creation of new habitats (in Anderson’s terms, the “hybrid habitat”) which could be exploited by hybrid entities or introgres- sants. Such a postulate would help to account for the extant gene flow between contemporary Audibertias (Figure 1). Reproductive isolation has rarely been achieved in this section of the genus except in instances where distri- bution patterns indicate relictual populations which have doubtless had long histories of isolation, or where populations are geographically disjunct. While Figure I indicates known gene flow in Audibertia to date, further studies may indicate that this hybridization is more extensive than it is now assumed to be. To date the only introgression recorded for this section has been that of Salvia apiana x S. mellifera. This paper presents one more example of contemporary introgression, that of S. apiana x S. leucophylla. Introgression between SALVIA APIANA and SALVIA LEUCOPHYLLA In Los Angeles County, Topanga Canyon Boulevard winds through a canyon of coastal chaparral vegetation which includes S. apiana, S. leuco- phylla, S. mellifera, S. spathacea, and S. columbariae as well as all of the other elements which typify this association. In several areas S. apiana and S. mellifera are sympatric and FI hybrids are relatively frequent. Hybrids between S. mellifera x S. leucophylla and S. apiana x leucophylla are less often encountered, both for reason of less frequent appearance, and less frequent populational contact. One site along the boulevard, just 3.7 miles north of the town of Topanga is of especial interest, for here S. apiana and 1971 Introgression in Salvia 7 SALVIA ; SECTION AUDIBERTIA Figure 1. Breeding behavior in Salvia: section Audibertia. Known crosses between species in this section are indicated as follows: Hybrids which have been found in nature Hybrids which have been made by deliberate pollination— —Intro- gression which has been found in nature hmi S. leucophylla are sympatric in an area which still bears the scars of a previous roadcut, the disturbances resulting from the laying of underground telephone cables at some earlier date, and constant erosion of the de- composing granite-shale hillside resulting from construction of the present boulevard. The site has so many edaphic disturbances coupled with light differences in the form of a northwest slope and a southwest slope that it presents an ideal environment for introgression (Figure 2). In this area, it 8 Contributions in Science No. 208 Flat land ^ Barbed C J leucophylla ^ wire S. apiana and Unused intragressants raad Figure 2. Site of introgression between S. apiana x S. leucophylla 3.7 miles north of Topanga. 1971 Introgression in Salvia 9 is easy to see occasional FI hybrids which show such a degree of heterosis that they appear monstrous in overall appearance; this coupled with the pink flower color as opposed to the violet-pink of S. leucophylla and the white of S. apiana as well as a host of more cryptic characters makes them readily distinguishable (Figure 3). Ascertaining the degree of gene infiltration into one or the other of the two parental species is not so easily observed. As Anderson has repeatedly pointed out, introgression is most significant when it is least observable. Inspection of the population revealed the pollinators to be primarily social bees such as Apis and several species of the bumblebees, Bombus. Since Grant and Grant (1964) have pointed out that social bees are not effective pollinators of S. apiana for reason of the widely spaced anthers and acentric position of the style, I could discount these as an ethological factor bringing about any appreciable infiltration of S. leucophylla genes into S. apiana or back to the FI hybrid. There is limited potential for social bees to carry pollen between S. leucophylla and the FI hybrid, the limitation being the distance between anthers on the FI and the slightly recurved style. Bombus species, on the other hand, are capable of effectively pollinating both parental species and hybrid recombinants by virtue of their size. Carpenter bees are so infrequent in the area as to be a negligible factor, and I have not had occasion to witness pollination by humming-birds or night-visiting moths. It is obvious that the greatest amount of pollination and the most efficacious pollination involves Bombus. A superficial assay of introgression in this area leads one to suspect that the hybrid population is introgressed in the direction of S. apiana, partly because of the great morphological variation in the S. apiana growing there with respect to compaction of verticils, pink dots covering the lower lip, violet striations on the stem, and a great difference in the number of verticils and overall size of the plants. Much greater uniformity is encountered in the S. leucophylla population. The apiana-\[\iQ variants, as indicated in Figure 3 are found predominantly on the west facing and southwest-facing slopes which are also occupied by some typical S. apiana. Gentle slopes, washes and flatlands are more favorable to the growth of S. leucophylla, although they do invade the northwest-facing slopes in this area. Salvia apiana and S. apianaA\kt individuals, can occupy steep, eroding, sun-baked hillsides with little competition from any other plants. In analyzing the population it was desirable to collect representative specimens (LACM Herbarium 70578-70622) and to prepare a scatter diagram (Figure 4) to clarify the occurrence of introgression. Two characters which exhibited measurable extremes in character states were the distance between verticils and the breadth of the lower lip; these were selected as ordinate and abscissa. Other characters could have been used to portray the same phenomenon, although perhaps not as effectively. Having positioned the metroglyphs utilizing these two characters, the other five characters chosen 10 Contributions in Science No. 208 U Figure 3. Hybridization between Salvia apiana x Salvia leucophylla showing the upper portion of the inflorescences and examples of each flower. A. S. leucophylla; B. FI hybrid between S. leucophylla x S. Apiana found in nature; C. S. apiana. 1971 Introgression in Salvia 11 were reduced to three states and were indicated as whiskers on the metro- glyphs. The resultant scatter diagram (Figure 4) shows the introgressants to be apiana-\\\iQ in most instances, as was anticipated by a casual perusal of the population. Parental types are easily distinguished in this diagram and show a distributional cohesion which further vindicates this mode of pre- sentation. Variation in the hybrids is considerable and conspicuous which suggests that introgression at this site is not of great antiquity. Another interesting feature of the area is the lack of real intermediacy of the habitat. Studies which I made previously (1964 and 1967) are indicative that the “hybrid habitat” of Anderson consists primarily of a disturbed area in prox- imity to the parent which is experiencing introgression. Unpublished data which I have on soil types in instances of introgression between S. apiana and S. mellifera indicate that introgressants are supported by a soil type which is similar to that of\S. mellifera in humus, mineral and moisture content, and light exposure whidh is also similar to S. mellifera. A thorough study of the nature of the “hybrid habitat” in instances of introgression is in order. The ability of an introgressant to compete satisfactorily with a parental species which has presumably reached an adaptive peak for that area is difficult to explain, except in instances where the “hybrid habitat” is an area which is reconstituted edaphically and climatically. Observations on the effects of heterosis would seem to indicate that the hybrid vigor conferred through this process of massive gene flow might be the key to the establishment and perseverance of recombinants. It should also be stressed that in most instances involving introgression the plants in question are perennials; the critical period in their development is the seedling stage of growth, and once established it is not difficult for the plant to persist and compete. Frequently the introgressants and apparently “good” parental types are found growing adjacent to one another in what would seem to be an identical habitat; more frequently, however, the introgressants occupy a niche, or a series of micro- niches, which is not totally characteristic of either of the parental species. Taxonomic Aspects of Introgression in Audibertia An outstanding feature of the two aforementioned populations which involve introgression with S. apiana is the apparent compaction of the loose paniculate inflorescence of S. apiana into symmetrical compressed cymes which approximate those of the species from which genetic infiltration is taking place. This compacted inflorescence of S. apiana has been recognized as a variety of S. apiana by Munz (1927) on the basis of a collection made at Morongo Wash in the Colorado Desert in Riverside County, California. He characterizes this variety as having, “Panicles condensed, spicate, branches being reduced, and appressed; otherwise much as in {S. apiana) typica.” His diagnosis of this compact form is correct, and I believe that his impli- cation that S. vaseyi is in some way involved is also correct. However, I have noted that when S. apiana is found hybridizing with S. mellifera or 12 Contributions in Science No. 208 COROUA COLOR INFLORESENCE FORM POSITION OF STYLE CONFORMATION OF COROLLA LENGTH OF UPPER LIP Verticils compact i Straight Thrysis with perpendicular labia Slighty branched Curved slightly Thrysis intermediate White Vettieels branched Curved outside of the corolla Thrysis geniculate Figure 4. Scatter diagram indicating an introgressed population of S. apiana x S. leucophylla. 1971 Introgression in Salvia 13 S. leucophylla, numerous members of the S. apiana population exhibit com- pact verticils. I suggest that this is due to historical or contemporaneous introgression with these other species. Further support of such a hypothesis comes from a careful analysis of other features of the compact variety. For example, when introgression with S. melUfera or S. leucophylla is involved the degree of verticilate compaction in S. apiana is likely to be greater and the cymose condition is less prominent. When the introgression is with S. melUfera, the compact form of S. apiana usually exhibits blue-violet dots over the labia; when the compaction is due to gene infiltration from S. leuco- phylla, the dots on the labia of the compact variety are violet-pink. An extension of this study will be to seek out contemporary instances of intro- gression between S. apiana and S. vaseyi and present further evidence in support of this contention. I question the taxonomic status of a variety whose characteristic aspects are derived independently from three different species of a section in dis- junct areas. The compact form is by no means localized. It has been collected from various extensions of the range of S. apiana and is, I believe, but one aspect of introgression, either historically based or through present gene flow. The importance of introgression in general has been well summarized by Anderson; its impact on evolution has yet to be clearly understood. Mas- sive short term gene flow between two sympatric species certainly has the advantage of increasing the range of the introgressant by the exploitation of newly opened habitats. It is a source of variability which is of a magnitude considerably greater than that of mutation, and whereas the variability in- volved in mutation is fortuitous, the variability introduced via introgression is far from chance, but represents recombinations, or blocks of characters, which have a history of being at a selective advantage in a habitat not altogether dissimilar to that occupied by the introgressant. This has doubtless led to the preservation of taxa during periods of disruption. In section Audi- bertia, introgression may be the primary source of species which constitute the section at this time. Whereas most flowering plants and animals produce sterile or partially sterile FI hybrids, in section Audibertia the hybrids en- countered are quite fertile, averaging around 75 per cent good pollen in the FI (as determined by staining with lactophenol blue). Epiing (1947) records 50 per cent of the nutlets developing in hybrids between S. apiana and S. melUfera, and 2.0 per cent of these being viable. He refers to the degree of fertility as, “sufficient to permit frequent localized hybrid swarms.” If the effect of introgression is sufficient to preserve frequent hybrid swarms, while not swamping parental species, it permits rapid exploitation of new environ- ments by new recombinants as such niches become available. This would indicate that the effect of introgression approximates that of allopolyploidy, and in short terms, exceeds that of mutation. Its importance can only be minimized by the limited instances in which we can cite this feature of 14 Contributions in Science No. 208 evolution as an active force; these examples are growing, and a new per- spective will soon be in order. With reference to the S. apiana x S. leucophylla introgressants discussed in this paper, it is one more piece in a complex jigsaw that will lead to an understanding of one of the most important elements of the vegetation of western North America, as well as contributing to a clearer understanding of the magnitude of role of introgression in the process of evolution. Acknowledgments I extend my thanks to Mrs. Doris Hoover for calling my attention to paleogeographic data, to Mrs. Frances Runyan for Figure 2, to Miss Mary Butler for Figure 3, and to Mr. Joe Nakanishi for Figures 1 and 4. Literature Cited Anderson, E. 1949. Introgressive Hybridization. John Wiley and Sons, London. 1953. Introgressive Hybridization. Cambridge Phil. Soc. Bull. Rev. 28:280-307. , and B. Anderson. 1954. Introgression of Salvia apiana and Salvia mellifera. Ann. Mo. Bot. Card. 41:329-338. Brown, J. 1963. Ecogeographical variation and introgression in an avian visual signal. Evolution 17:23-29. Brown, W., and G. Pratt. 1960. Hybridization and introgression in the grass genus Elymus. Amer. Jour. Bot. 47:669-767. Carlson, E. M., and B. C. Stuart. 1936. Development of spores and gametophytes in certain New World species of Salvia. New Phytol. 35: 1-49. Crowell, J. 1962. Displacement along the San Andreas Fault. Calif. Geol. Soc. Am. Spec. Paper. 61 p. 1968. The California coast ranges. U. M. R. Jour. No. 1 (April) : 133-155. Delestaing, N. 1954. Contribution a I’etude cytologique du genre Salvia. Rev. Cytol. et Biol. Veg. 15:195-236. Emboden, W. a., Jr. 1964. Pollen morphology of the genus Salvia, section Audi- bertia. Pollen et Spores 6:527-536. 1969. Detection of palynological introgression in Salvia (Labiatae). Los Angeles Co. Mus. Contrib. Sci. 156:1-10. , and H. Lewis. 1967. Terpenes as taxonomic characters in Salvia: section Audibertia. Brittonia 19:152-160. Epling, C. 1938. The California Salvias. A review of Salvia, section Audibertia. Ann. Mo. Bot. Gard. 25:95-188. 1940. Supplementary notes on Salvia: Audibertia. Ann. Mo. Bot. Gard. 27:259-262. 1947. Natural hybridization of Salvia apiana and S. mellifera. Evolution 1:69-78. , H. Lewis, and P. Raven. 1962. Chromosomes of Salvia: section Audi- bertia. Aliso 5:217-221. Grant, K., and V. Grant. 1964. Mechanical isolation of Salvia apiana and Salvia mellifera. Evolution 18:196-212. 1971 Introgression in Salvia 15 Guttman, S. 1968. Transferrin and hemoglobin polymorphism, hybridization and introgression in two African tree frogs, Bufo regularis and B. rangeri. Comp. Biochem. and Phys. 23:871-877. Hulten, J, 1956. The Cerastium alpinum complex. Sv. Bot. Tidskr. 50:434-456. Lawson, J. 1714. A History of Carolina. London. Maze, J. 1968. Past hybridization between Quercus macrocarpa and Quercus gambelii. Brittonia 20:321-333. Mechan, j. 1966. Introgressive hybridization of tree frogs. Evolution 14:445-457. Munz, P. a. 1927. The southern California species of Salvia (including Ramona). Bull. So. Calif. Acad. Sci. 26:17-29. 1959. A California Flora. University of California Press, Berkeley. Peeler, B. 1956. Introgression in planktonic Rotaria. Evolution 10:246-261. Pipkin, S. 1968. Introgression between two closely related species of Drosophila in Panama. Evolution 22:140-156. Stewart, W. S. 1939. Chromosome numbers of California salvias. Am. Jour. Bot. 26:730-732. Tucker, J. M. 1970. Studies in the Quercus undulata complex IV. The contribution of Quercus harvardii. Am. Jour. Bot. 57:71-84. Van Valen, L. 1963. Introgression in laboratory populations of Drosophila per- similus and D. pseudoobscura. Heredity 18:205-214. Webb, A., and S. Carlquist. 1964. Leaf anatomy as an indicator of Salvia apiana X S. mellifera introgression. Aliso 5:437-499. Accepted for publication November 24, 1970 LOS ANGELES j COUNTY MUSEUM CONTRIBUTIONS IN SCIENCE I^JUMBER 209 February 17, 1971 1 FIVE NEW SPECIES OF ACMAEODERA (COLEOPTERA: BUPRESTIDAE) FROM THE SOUTHWESTERN UNITED STATES AND BAJA CALIFORNIA By Richard L. Westcott Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 FIVE NEW SPECIES OF ACMAEODERA (COLEOPTERA: BUPRESTIDAE) FROM THE SOUTHV'HSTERN UNITED STATES AND BAJA CALIFORNIA" By Richard L. Westcott^ Abstract: Five new species of the genus Acmaeodera are described: A. recticolloides (southern Baja California del Norte to San Diego County, California), A. panamintensis (mountains of southwestern Nevada and adjacent California), A. atactos- pilota (central Nevada and southwestern Utah to southern Cali- fornia), A. verityi (Colorado Desert, California), and A. mojavei (southern Arizona to southern California). Confusion of the last three species, both in collections and in the literature, with A. pubiventris Horn and A. lanata Horn is clarified. Con- siderable emphasis is placed on the use of the female genitalia as a taxonomic aid. Pertinent bionomical information is given. Introduction Extensive collecting and study in recent years by entomologists in the western United States has revealed many new species of this interesting and widespread genus of wood-boring beetle. The five species described herein belong to the group “Sinuatae,” as defined by Horn (1878), which inhabits, for the most part, the desert and chaparral of southern California and adja- cent regions. Two of the species have been recently discovered. The other species have been known but confused^ both in collections and in the litera- ture, with Acmaeodera lanata Horn and A. pubiventris Horn. Acknowledgments I wish to express my deep gratitude to Dr. W. F. Barr, University of Idaho, for his role as major professor in the course of this study and for the loan of specimens, both personal and those in his care. Special thanks go to Dr. G. H. Nelson, Kansas City, Missouri, and Mr. D. S. Verity, Santa Monica, California, for use of their collections which have made it possible to greatly enlarge the scope of this study. For additional material appreciation is ex- pressed to the following individuals and institutions: R. C. Bechtel (Nevada State Department of Agriculture), F. M. Beer (Oregon State University), M. A. Cazier (Arizona State University), H. F. Howden (Canada Depart- ment of Agriculture), H. B. Leech (California Academy of Sciences), Alodified from part of a thesis submitted in partial fulfillment for the degree of Master of Science, Department of Entomology, University of Idaho, Moscow, Idaho. ^Research Assistant, Los Angeles County Museum of Natural History; and Plant Division, Oregon Department of Agriculture, Salem, Oregon 97310. 1 2 Contributions in Science No. 209 D. L. Mays (University of Florida), F. H. Parker (Globe, Arizona), V. M. Tanner (Brigham Young University), J. W. Tilden (San Jose State College), F. S. Truxal (Los Angeles County Museum of Natural History), American Museum of Natural History, California Department of Agriculture, and the University of California ( Berkeley, Davis, and Riverside). My sincere thanks go to Mr. Ken Gray, Portland, Oregon, for his photographic work. In listing type material the author has found it convenient, in most instances, to use abbreviations for collector’s names as follows : R. P. Allen (RPA) W. F. Barr (WFB) R. C. Bechtel (RCB) F. M. Beer (FMB) R. M. Bohart (RMB) M. A. Cazier (MAC) F. C. Clark (FCC) H. E. Cott (HEC) H. K. Court (HKC) J. H. Davidson (JHD) J. M. Davidson (JMD) D. W. Davis (DWD) A. Fenyes (AF) D. L. Flaherty (DLF) A. R. Gittins (ARG) T. R. Haig (TRH) J. C. Hall (JCH) S. M. Hogue (SMH) M. E. Irwin (MEI) A. E. Lewis (AEL) E. G. Linsley (EGL) G. P. Mackenzie (GPM) C. D. MacNeill (CDM) J. W. MacSwain (JWM) W. R. M. Mason (WRM) D. L. Mays (DLM) A. E. Meier (AM) A. S. Menke (ASM) A. T. McClay (ATM) A. E. Michelbacher (AEM) D. R. Miller (DRM) G. H. Nelson (GHN) K. T. Nelson (KTN) F. W. Nunenmacher (FWN) F. D. Parker (FDP) W. M. Pearce (WMP) R. L. Penrose (RLP) W. Reeves (WR) W. R. Richards (WRR) E. S. Ross (ESR) E. I. Schlinger (EIS) W. E. Simonds (WES) R. F. Smith (RFS) M. D. Snelling (MDS) L. A. Stange (LAS) O. H. Sweezy (OHS) V. M. Tanner (VMT) J. W. Tilden (JWT) P. C. Ting (PCT) F. S. Truxal (FST) R. L. Usinger (RLU) E. P. Van Duzee (EPV) E. C. Van Dyke (ECV) D. S. Verity (DSV) R. L. Westcott (RLW) B. E. White (BEW) Discussion of Taxonomic Characters In the species descriptions to follow, elytral markings when median or submedian are referred to as discal, and when lateral are termed submarginal, even though some may reach the margin. The markings are additionally referred to in accordance to their arrangement from anterior to posterior. Varying degrees of coalescence may be exhibited. A marking at the humeral 1971 Five New Species of Acmaeodera 3 angle is always referred to as the humeral spot. Usually it is yellow, but it may be brownish. The humeral interval, located between the umbone and the lateral margin, is used in species descriptions, but due to its variability is not employed as a uniformly reliable diagnostic character. The colored hairs on the pronotum and elytra range from very light coppery to fuscous and the lighter shades may be difficult to detect. The genitalia have seldom been used in classifying Acmaeodera, but in most cases it is necessary to see them in order to differentiate the sexes. I have found that the female genitalia have several important characters of taxo- nomic value. Tanner (1927) made a study of the female genitalia of Coleop- tera, and I have followed his terminology. The female genital apparatus of Acmaeodera is largely membranous. These membranous areas apparently are of little taxonomic value. The membranes bear setae (particularly apically) which might be significant, but are much too easily lost during mounting to be reliable. The characters of greatest importance are the form, size and arrangement (particularly the proportionate lengths) of the sclerotized struc- tures. These structures consist of the proctiger, styli, and the baculi of the proctiger, valvifers, and coxites. The membranous portions of the coxites near the styli are flared. White ( 1939) incorrectly referred to the coxites as the subgenital plate. With additional study, characters which may prove of value are a length to width ratio of the distal portion of the ovipositor and the ratio of the distance between the styli to the length of a stylus. The male genital apparatus is rather simple, of the modified trilobed type, and affords few characters of use in species separation. The only charac- ter of any value found in this study is the shape of the distal end of the penis, however the apex is sharply attenuate and in mounting may be easily broken behind the actual opening of the ejaculatory duct. The five species described herein will key out, under “Acmaeoderae Sinuatae,” either to couplet 10 or 1 1 according to Fall (1899). Difficulty arises in the interpretation of density of vestiture of the ventral surface and this cannot now, in my opinion, be adequately used as presented by Fall. Any attempt to define the species described herein on this basis would be of little value without a complete revision of the group “Sinuatae.” In the discussion to follow the species will be related to Fall’s key as a matter of formality, since it is the only key to the North American Acmaeo- dera. However, in my opinion, this is of very limited value due to the large number of species described since and the many species yet to be named. Acmaeodera recticolloides, new species Figures 1, 10 FEMALE: Medium-sized, moderately robust. Head and pronotum black with slight bronzy luster. Elytra black, shining, with a discal row of five and a marginal-submarginal row of six to seven light yellow spots, apical two-fifths with reddish markings laterally, first discal spot immediately behind front 4 Contributions in Science No. 209 margin, median spots coalesced into a fascia which does not reach as far mesad as other discal spots, humeral spot present. Head densely clothed with long, erect, silky white hairs, interspersed with dark hairs dorsally, coarsely, densely punctate, reticulate. Pronotum slightly narrower than elytra, 1.87 times as wide as long, widest at base; surface with a feeble, median, basal depression and a slight depression in front of lateral pit, densely clothed with long, erect, silky white hairs on basal half and sides, and with dark coppery brown hairs on apical half, disc coarsely, densely punctate, punctures becoming coarser and denser towards sides; lateral margins arcuate, entire, more arcuate on apical half which is flanged and visible from above. Elytra widest just behind umbone; sides subparallel for basal three-fifths then gradually rounded to apex; lateral margins serrate on apical two-fifths; surface moderately clothed with long, erect, fuscous hairs which are inter- spersed with long, silky white hairs, basal three-fifths with recumbent to subrecumbent silky white hairs at sides; strial punctures medium to coarse, becoming confused at base, discal striae paired, at least on basal three-fifths, outer striae with coarser punctures, interstrial punctures medium, shallow; humeral interval moderately developed, not elevated in front. Ventral surface with prosternal margin trisinuate, with a moderately developed rounded prominence on either side of middle; abdomen moderately clothed with long, subrecumbent to suberect, silky white hairs at middle becoming dense at sides, punctures medium-sized and moderately placed at middle becoming denser and slightly larger laterally, punctures of first visible sternite much coarser anterolaterally, second visible sternite distinctly more densely punctate at middle than first, last visible sternite with a well-developed, rounded, subapical plate. Female genitalia as in Fig. 10. Length: 9.1 mm. Width: 3.3 mm. TYPE MATERIAL AND LOCATION: Holotype female. La Zapopita, Valle de Trinidad, Baja California del Norte, Mexico, April 16, 1962 (G. H. and K. T. Nelson), collected on Ephedra californica, deposited in the Cali- fornia Academy of Sciences. One hundred seventy paratypes, same locality as holotype, April 16-18, 1962 (GHN, KTN, DSV, RLW); 4 mi W La Zapopita, April 16, 1961 (FST). Seven, 5 mi SE and 15 mi NW Desengano, Baja California, Mexico, June 6, 1966 (DSV). One, 4 mi W Jacumba, San Diego Co., California, June 5, 1957 (DSV). Four, Boulevard, San Diego Co., California, June 1, 1968 (DSV). Paratypes in the collections of the Los Angeles County Museum of Natural History, United States National Museum, J. M. Davidson, G. H. Nelson, D. S. Verity, W. F. Barr, and the author. DISTRIBUTION : Southern Baja California del Norte, Mexico to extreme southern California. BIONOMICS: This species was taken at the type locality and at Boulevard, San Diego Co., California, resting on or flying over Ephedra californica 1971 Five New Species of Acmaeodera 5 Figures 1-9. Dorsal view of adult Acmaeodera selected to show general form and elytral maculation. Fig. 1, recticolloides, new species, type locality. Fig. 2, recti- collis Fall, 2 mi S Holbrook, Ariz. Fig. 3, panamintensis, new species, type locality. Fig. 4, pubiventris Horn, Independence, Calif. Fig. 5, lanata Horn, Wildrose Can., Panamint Mts., Calif. Fig. 6, atactospilota, new species, Lone Pine Campground, Calif. Fig. 7, verityi, new species, 5 mi NW Palm Springs, Calif. Fig. 8, verityi, Snow Creek, Calif. Fig. 9, mojavei, new species, 6-9 mi NE Red Mt., Calif. 6 Contributions in Science No. 209 Watson. The habits of related Ephe dr a-inhdhitmg species, plus the fact that A. recticolloides was taken on no other plant in the area, indicate that E. cali- fornica is its host. Several specimens were collected on Lotus scoparius (Nuttall in Torrey and Gray) Ottley near Desengano, Baja California del Norte. In northern Baja California del Norte this beetle inhabits an area reminiscent of the moister Sonoran Desert element of Arizona, yet represent- ing the beginning of a transition toward chaparral. Several species of Acmae- odera commonly associated with chaparral have been taken in this area. There is an abundance of mesquite and the vegetation is quite dense in places. In contrast, the related A. recticoUis Fall inhabits a wide range far to the east of A. recticolloides from the sparsely vegetated plateau regions of northeastern Arizona to the desert regions of Big Bend National Park, Texas. It, too, is associated with Ephedra. VARIATION : This species exhibits slight variation. It ranges from 7.7 to 10.7 mm in length and from 2.7 to 4.0 mm in width. The basal discal spot of each elytron was found in all but two specimens. However, it varies in size and is on rare occasion coalesced with the spot behind it. The median discal spots of each side are almost always coalesced into a fascia. The reddish apical spots of each elytron are narrow, usually three in number, and may be variously coalesced with each other or with adjacent yellow spots. Occasionally the color is a faint pinkish which is difficult to detect if only one or two spots are present. It is of interest that this species can, in most cases, be sexed by examining the form of the subapical plate. In the male it is usually broadly rounded or truncate behind. The female has the plate more narrowly rounded, usually giving it a triangular appearance. DIAGNOSIS AND COMPARISON : Acmaeodera recticolloides will key into the couplet containing A. lanata and A. pubiventris or to A. recticoUis in Fall (1899). The presence of dark hairs on the elytral surface and reddish mark- ings on the elytral apices will readily separate A. recticolloides from A. lanata and A. pubiventris. It appears most closely related to A. recticoUis and may be readily distinguished from that species by its blacker ground color and abun- dant dark hairs on the elytra. Acmaeodera recticolloides averages smaller in size and is less robust. The lateral reddish markings are smaller and less pro- nounced. The second visible abdominal sternite of A. recticolloides is dis- tinctly more densely punctate at the middle than the first and the punctures appear equal in size, while in A. recticoUis the difference in density is much less pronounced and the punctures of the second visible sternite are finer. Acmaeodera recticolloides might be confused with some of the chaparral- inhabiting species of California, but the form and color of its markings, coupled with the much denser vestiture beneath should serve to readily separate it. This species is named for its superficial resemblance to A. recticoUis. 1971 Five New Species of Acmaeodera 7 Acmaeodera atactospilota, new species Figures 6, 13 FEMALE: Medium-sized, moderately robust. Head and pronotum black, with coppery reflections near front margin and laterally on pronotum. Each elytron black with purplish reflections, markings yellowish, those of disc vermiculate and approximating or meeting a submarginal, longitudinal, broken row of small spots which extends nearly to apex, humeral spot present. Head densely clothed with moderately long, semirecumbent, silky white hairs, coarsely, densely punctate, reticulate, and with a very well-developed, median, dorsal carina. Pronotum as wide as elytra, 1.79 times as wide as long, widest behind middle, wider at base than apex; surface with a shallow median depression in front of base and a slight depression in front of basal pits, basal two-thirds of disc, sides, and extreme front densely clothed with long, subrecumbent, silky white hair, apical third of disc with long, erect, coppery brown hairs, disc coarsely, densely punctate, punctures becoming much coarser and denser towards sides; lateral margins arcuate, entire, prominent apically, not visible from above. Elytra widest at base; sides subparallel for basal two-thirds, then moderately converging to apex; lateral margins serrate on apical two-fifths; surface moderately clothed with long, erect to recumbent, silky white hair, inner discal striae more widely separated than outer striae and with punctures small to large, punctures of outer discal striae coarser, deeper, and denser, interstrial punctures shallow, small to medium-sized; humeral interval feebly developed, not elevated to level of front margin. Ventral surface with prosternal margin trisinuate, with a shallowly rounded prominence on either side of middle; abdomen moderately clothed with long, recumbent or subrecumbent, silky white hair at middle of first visi- ble sternite, becoming dense elsewhere, most dense at sides, punctures mod- erately placed at middle of first visible sternite, becoming coarser and denser toward sides, second visible sternite more finely, distinctly more densely punc- tate at middle than first, last visible sternite with a large, well-developed, widely rounded, subapical plate. Female genitalia as in Fig. 13. Length: 9.3 mm. Width: 3.2 mm. TYPE MATERIAL AND LOCATION: Holotype female, near Littlerock, Los Angeles County, California, May 10, 1959 (R. L. Westcott), deposited in the California Academy of Sciences. Two hundred fifty-nine paratypes from the following localities: NEVADA: Nye County: near Berlin, VI-27- 1960 (RLW); Ichthyosaur State Park, VI-21-24-1960 (RLW); Beatty, V- 3 1-1962 (FDP, LAS). Esmeralda County: Goldfield, VI-26- 1907 (FWN), III-20-1961 (FMB), dug from Ephedra. Clark County: Jean, V-28-1950 (CDM); Lee Canyon, Charleston Mtn., V-24-1940 (WR, MAC, PCT), VI- 27-1966 (WFB, RLP), VII-1-1966 (RLW), VI-17-18-1967 (WFB, RLP, SMH); Kyle Canyon, VI-27- 1947 (ATM), V-8-1964 (RLW), reared from 8 Contributions in Science No. 209 Figures 10-15. Female genitalia of Acmaeodera: Fig. 10, recticolloides; Fig. 11, panamintensis; Fig. 12, lanata; Fig. 13, atactospilota; Fig. 14, verityi; Fig. 15, mojavei. 1971 Five New Species of Acmaeodera 9 Ephedra viridis. CALIFORNIA: Inyo County: Bishop, VI-21-1929 (EPV), VI- 28-29-1950 (JWT); Big Pine, VI-17-1929 (RLU); 5 mi NE Big Pine, V-29, VI-1-1960 (DSV, RLW); 7 mi E Big Pine, VII-2-1960 (WFB); Independence, VI-11-1929 (RLU), V-25-1959 (GHN), V-29, VI-1-1960 (DSV, RLW); Lone Pine, V-19-27-1937 (ECV); 3 mi W Lone Pine, V-29- 1960 (RLW) ; Lone Pine Campground, 6 mi W Lone Pine, V-28-1960 (DSV, RLW); Lone Pine Creek, VI-7-1939 (RMB); Olancha, VI-15-1941 (GPM); Westgard Pass Plateau, V-26-1937 (AM); Batchelder Spring, VI- 16- 1960 (HKC); 8 mi W Deep Springs, VII-3-1965 (FMB); Panamint Mts., S of Emigrant Jet., 4500-5000', V-24-1959 (GHN); 16 mi N Trona, V-30-1960 (DSV). Kern County: Randsburg, V-25-1958 (DSV, RLW). San Bernardino County: 1 mi S Adelanto, VI- 1-1 962 (MEI); 2 mi SW Phelan, VII-13-1963 (DSV); Desert Springs, VIII-3-1958 (GHN), VI-11-1962 (GHN); 4 mi SE Desert Springs, VII-12-1959 (RLW); Yermo, IV-28-1949 (EGL, JWM, RFS); 25 mi E Baker, VI-24- 1948 (ECV); Windmill Station, V-28-1950 (CDM, HEC) ; Bonanza Mine, Providence Mts., 1939 (EPV). Los Angeles County: Palmdale, V-26-1957 (DSV); Littlerock, VI-8-1941 (GPM), V-28- 1955 (WES), V-26-1957 (DSV), V-30-1957 (WES), V-10-1959 (DSV, RLW); 3 mi SE Pearblossom (DSV). Paratypes deposited in the collections of the American Museum of Natural History, California Academy of Sciences, California Department of Agriculture, Los Angeles County Museum of Natural History, United States National Museum, University of California at Davis and Riverside, W. F. Barr, F. M. Beer, J. M. Davidson, G. H. Nelson, R. L. Penrose, J. W. Tilden, D. S. Verity, and the author. Forty-seven specimens from the following localities are not included as paratypes : NEVADA: Washoe County: Nixon, VI-24- 1964 (FDP). White Pine County: Ely, VI-15-1941 (GPM). Lincoln County: Pioche, VII-9-1965 (WFB, RLW), VI-25-1966 (RLW). UTAH: Washington County: Zion Canyon, VI-19-1933 (OHS). ARIZONA: Mohave County: Wolfhole, VI-3- 1941 (VMT). Yuma County: 18 mi SE Parker, V-14-1966 (MAC, JMD). CALIFORNIA: Mono County: Topaz Lake, VI-26-1957 (DLF). San Ber- nardino County: Morongo Valley, V-26-1941 (ECV), V-19-1951 (RCB), VII- 6-1959 (DSV), V-21-1961 (GHN); Joshua Tree, VI-24- 1955 (RPA). Riverside County: Banning, V-20-1941 (ECV); 3 mi N US 99 on road to Morongo Valley, VI-22- 1957 (RLW); 5 mi W Desert Hot Springs, VI-22- 1957 (DSV); North Palm Springs, VI-21, 23-1959 (GHN), VI-5- 1960 (DSV); Snow Creek, VI-1-1957 (DLM, RLW), VI-23-1957 (DSV); Palm Springs, VII- 1-1941 (BEW) ; 2 mi W Mecca, VI-10-1963 (DRM). San Diego County: 2 mi E Jacumba, VI-23-1960 (GHN). DISTRIBUTION: Central Nevada to southwestern Utah, northwestern Arizona and southern California. BIONOMICS: Two adults of A. atactospilota were reared from Ephedra viridis Coville collected in Kyle Canyon, Clark County, Nevada. This plant 10 Contributions in Science No. 209 does not occur in many of the localities where A . atactospilota is taken, there- fore the beetle probably utilizes additional species of Ephedra as hosts. Adults have been taken visiting flowers of the following plants : Sphaeralcea rusbyi eremicola (Jepson) Kearn, Viguiera reticulata Watson, Encelia farinosa Gray, Coreopsis bigelovii (Gray) Hall, Stephanomeria pauciflora (Torrey) A. Nelson, Salvia dorii (Kellogg) Abrams, Eriogonum inflatum Torrey & Fre- mont, E. fasciculatum ssp., Acacia greggii Gray, and Fallugia par ado xa (D. Don) Endlicher. This beetle has the widest ecological and geographic range of any species described herein. It seems to be most abundant in the higher desert regions, as few specimens have been taken below 1500 feet above sea level, and has been collected at nearly 7000 feet in semi-arid desert mountain ranges. VARIATION : Specimens of A. actactospilota available for study ranged from 6.5 to 10.7 mm in length and from 2.2 to 3.8 mm in width. This species exhibits great variation in elytra! maculation. The spots are almost always scattered in a vermiculate pattern, although some specimens may have the spots arranged in a regular pattern, thus causing confusion with A. lanata. Ninety-eight per cent of the specimens studied have a basal, discal spot which is often elongate and oblique. The other spots may be variably coalesced. Rarely, a condition exists where all the median spots are connected across one or both elytra. In either case the resultant marking does not extend across the suture and often it resembles a reversed or inverted “C”. Ninety per cent of all specimens studied were found to have a humeral spot, although it was found only on one side in some instances. A specimen from Ichthyosaur State Park, Nevada, has a yellow spot on one side of the pronotum. Light or dark coppery hair was found to occur on the pronotum in 94 per cent of the speci- mens studied. In addition, the head usually bears this colored hair. Sometimes the coloration may be faint and only a few such hairs may be present, even on the pronotum. The humeral interval was almost always found to be poorly developed. Most specimens have a well-developed subapical plate. Specimens from the Colorado Desert region are, in general, more strongly marked and exhibit a greater degree of spot coalescence. In addition, the vestiture is more dense, particularly on the venter. Specimens from central Nevada average larger in size and are less densely clothed with hairs. They present a slightly different facies. DIAGNOSIS AND COMPARISON : Acmaeodera atactospilota is most apt to be confused with some forms of the sympatric A. lanata (Fig. 5) and will key out to such in Fall ( 1899). In his discussion of the latter. Fall refers to “confusedly maculate” forms. Van Dyke ( 1919) refers to a phase of A. pubiventris having the markings numerous, small, and irregular. It seems likely that both authors had specimens of A. atactospilota before them. Kerremans ( 1906) figured a specimen which he labeled A. lanata, but which proves to be A. atactospilota. This species can usually be separated from 1971 Five New Species of Acmaeodera 11 A. lanata by its confused maculation. Specimens of the latter with the vittae broken into spots rarely have additional spots between vittae, although those of the discal row are often displaced laterally. In most all cases the presence of coppery hair on the pronotum, coupled with the submarginal row of broken spots on the elytra of A. atactospilota, will serve to separate the two species. Any doubt remaining would be quickly dispelled upon comparison of the overall shape and length of the female genitalia (Figs. 12, 13). Al- though A. atactospilota shows close affinity with A. lanata, the presence of coppery hair and the structure of the ovipositor give evidence of a closer relationship to A. verity i. The two, however, are readily separated by dif- ferences in elytral maculation. A few specimens of A. atactospilota examined from the Panamint Range, California, possess dark hairs near the apices of the elytra, which is characteristic of A. mojavei, but only one specimen was seen which had elytral markings similar to that species. Acmaeodera atacto- spilota can be readily separated from A. mojavei by the denser vestiture of the former. The name for this species was derived from the Greek “ataktos,” mean- ing not arranged or irregular, and the Greek “spilotos,” meaning spotted. Acmaeodera verityi, new species Figures 7, 8, 14 FEMALE: Medium-sized, robust. Head and pronotum black with slightly metallic reflections. Each elytron black, shining, with occasional slight purplish or bluish reflections, with a discal and submarginal row of four paired yellow spots beginning at basal fourth and extending to apical fifth, the basal discal spot farthest from its submarginal counterpart, the median discal spot displaced laterally and closely approximating its submarginal counterpart, humeral spot absent. Head densely clothed with long, erect to suberect, silky white hair, a few dark hairs present on upper fourth punctures coarse, densely placed. Pronotum slightly narrower than elytra, 1.65 times as wide as long, widest on basal half; surface with slight trace of median depression in front of base, densely clothed with long, erect to subrecumbent, silky white hair except for an apical portion of disc which possesses mostly coppery brown hair, punctures dense, medium-coarse at middle of disc, becoming denser and much coarser toward sides; lateral margins arcuate, entire, much more promi- nent anteriorly, not visible from above. Elytra widest just behind base; sides subparallel for basal three-fifths then moderately converging to apex; lateral margins serrate on apical two-fifths; surface moderately clothed with long, erect to recumbent, silky white hair, a few dark hairs along sides on apical fourth, outer discal striae closer together and with coarser, deeper punctures than inner striae, punctures elongate at base, interstrial punctures medium-sized, shallow, confused basally; humeral interval moderately developed, coarsely punctured, not attaining level of front margin. 12 Contributions in Science No. 209 Ventral surface with prosternal margin trisinuate, with a moderately developed, rounded prominence on either side of middle; abdomen sparsely clothed with long, suberect to recumbent, silky white hair at middle of first visible sternite, becoming densely clothed at sides; punctures medium-sized, becoming coarser and denser toward sides, much coarser anterolaterally on first visible sternite, second visible sternite distinctly more densely punctate at middle than first, last visible sternite with a small, acute subapical plate. Female genitalia as in Fig. 14. Length: 8.8 mm. Width: 3.1 mm. TYPE MATERIAL AND LOCATION : Holotype female, 5 miles northwest of Palm Springs, Riverside County, California, May 10, 1958 (G. H. Nelson), deposited in the California Academy of Sciences. Two hundred forty-three paratypes from the following localities in California: San Bernardino County: Morongo Valley, V-26-1941 (ECV), V-4, VI-22-1957 (RLW), V-18-1958 (GHN), V-3-1959, VI-5-1960 (DSV). Riverside County: Whitewater, IV- 17-1937 (AEM), VI-28-1937 (ATM), IV-20-1951 (RCB); 5 mi W Desert Hot Springs, VI-22-1957 (DSV); 5 mi E Cabazon, IV-2-1961 (DSV); Snow Creek, IV-27 & 28, V-4 & 5, VI-1-1957 (DLM, DSV, RLW), V-1-1960 (RLW); Palm Springs Station, VI-27- 1952 (ATM); 6 mi NW Palm Springs, IV-15-1961 (DSV); 5 mi NW Palm Springs, VI-18-1954, V-4-14, VI-22- 1958, IV-23-1959, VI-12-1960 (GHN), VI-22 & 23-1957, V-10 & 11, VII- 6-1958, IV-9-1961 (DSV, RLW); Palm Springs, “April” (AF), III- 1929 (FCC), VII-1-1941 (BEW) ; Mission Creek, V-1-1952. Imperial County: Chocolate Mts., IV-9-1939 (RPA). Paratypes in the collections of the American Museum of Natural History, California Academy of Sciences, Los Angeles County Museum of Natural History, Nevada State Department of Agriculture, United States National Museum, University of California at Davis, W. F. Barr, J. M. Davidson, D. L. Mays, G. H. Nelson, F. H. Parker, D. S. Verity, and the author. Specimens from the following localities in California are not included as paratypes : Inyo County: 13 mi S Death Valley Jet, V-20-1965 (ASM) ; Death Valley, IV-27-1950 (DWD). Ventura County: 5 mi N Simi, V-4-1959 (DLM). Riverside County : 3.3 mi E Nightingale, V-1-1960 (RLW) ; Hwy. 74, Santa Rosa Mts., 3500', V-14-1958 (GHN) ; slope below Pinyon Flat, 2000', IV-10-1949 (ESR); Deep Canyon, V-3-1963 (EIS). San Diego County: 4 mi E Jacumba, VI-7-1959, V-20-1961 (DSV). DISTRIBUTION : California: Colorado Desert and outlying areas. BIONOMICS: Adults are most commonly taken resting on stems of Ephedra californica, which most likely is the larval host plant. They have also been taken on Dalea californica Watson where these species grow in close associa- tion. One specimen was taken on a flower of Viguiera deltoidea Gray. They have been taken on other yellow-flowered Compositae. Acmaeodera verityi has been found primarily in the upper Colorado Desert area from Morongo Valley to Palm Springs, California. Four speci- 1971 Five New Species of Acmaeodera 13 mens labeled “five miles north of Simi, Ventura County, California” present an ecological and distributional problem. This locality is in a Coastal Sage Scrub-Chaparral zone miles away from any desert region and well over 100 miles from any other known locality where A. verity i occurs. Also collected at this site were two specimens of A. lanata. Ephedra, which is a known host of the latter and a possible host of the former, was sought at this locality but to no avail. The significance of this entire situation can only await further col- lecting and study. The possibility of mislabeling exists. Two specimens from four miles east of Jacumba, San Diego County show a somewhat different facies than Colorado Desert material, and their status remains in doubt. A female specimen from south of Death Valley Junction exhibits the general facies of A. mojavei, but examination of the genitalia showed it to be con- specific with A. verity i. Another specimen, a male, from Death Valley is assigned tentatively to this species. These localities are far removed from the known range of this beetle and further collecting is needed to correctly interpret its range and variation. VARIATION: Specimens of A. verityi examined ranged from 6.3 to 10.0 mm in length and from 2.1 to 3.8 mm in width. This species exhibits extensive variation in elytral maculation. In the series examined many specimens exhibit more or less than the number of spots present on the holotype. Very rarely a discal spot at the base and/or a humeral spot occurs. The discal spot at the basal fourth is variable in size but always roundish in shape. It is the one most commonly absent, occurring in only 63 per cent of the specimens studied. More rarely the median discal spot is absent and when this is the case (common in specimens from Morongo Valley) the preceding condition occurs concomitantly. The median discal and submarginal spots of each elytron were found to be coalesced in 70 per cent of the specimens studied. Sometimes the discal markings are present only in the form of one or two spots near the apex of each elytron (Fig. 8). The extreme in reduced maculation was seen on one specimen which had a total of three spots on one elytron and two on the other. Five specimens were seen with pronotal spots either on one or both sides. The occurrence of dark hair on the dorsal surface varies. This is almost always present on the pronotum and in most cases can be found near the elytral apices where it is often inconspicuous. Occasionally this hair is present on the basal portions of the elytra. DIAGNOSIS AND COMPARISON: Acmaeodera verityi has been placed under A. pubiventris (Fig. 4) in most collections, and will key out next to this species in Fall (1899). The elytral markings and ground color and their completely allopatric distribution readily separates the two species. Difficulty may arise, however, in separating this species from A. mojavei. In general appearance A. verityi is more robust and the markings are less numerous. Acmaeodera mojavei usually has the pronotum distinctly narrower than the elytra, while in A. verityi this character is not usually so distinct. Acmaeodera 14 Contributions in Science No. 209 verityi is more densely pubescent on the venter, and the dark hairs near the apices of the elytra are usually less discernible. These hairs are rarely as abundant as in A. mojavei. The front margin of the prosternum is always trisinuate in the former and often emarginate in the latter. In A. mojavei the second visible sternite usually is much less densely punctate at the middle in relation to the middle of the first visible sternite. The subapical plate of A. mojavei is less prominent. The two species are readily separated on the basis of the structure of the female genitalia, which in A. verityi have the styli placed closer together. In addition there is a marked difference in sclerotization of the apical portions of the dorsal baculi, it being much broader and lighter in A . mojavei. I take great pleasure in naming this species after Mr. D. S. Verity, who has collected a large number of the specimens used in this study and who has contributed greatly to my interest in this genus. Acmaeodera mojavei, new species Figures 9, 15 FEMALE: Medium-sized, moderately elongate. Head black with coppery luster. Pronotum bluish black with coppery reflections along extreme front and at sides. Each elytron black, shining, with yellow spots as follows : A discal row of five, beginning at base and extending to apical fifth, the median and fourth spot located slightly laterad to the others, and a submarginal row of four paired with those of disc and beginning just behind umbone (apical spot missing on right side), humeral spot absent. Head densely clothed with long, erect to suberect, silky white hair which is interspersed with dark hair on upper portion, coarsely, densely punctate, reticulate. Pronotum narrower than elytra, 1.82 times as wide as long, widest be- hind middle, wider at base than at apex; surface with a slight depression at middle in front of base and in front of basal pits, disc moderately densely clothed with long, erect to subrecumbent, silky white hair on basal three- fourths, apical fourth with coppery brown hairs, front margin and sides clothed with shorter silky white hair, punctures coarse and dense on disc, becoming deeper towards sides; lateral margins arcuate, entire, flanged, very prominent, not visible from above. Elytra widest behind base; sides subparallel for basal two-thirds then moderately converging to apex; lateral margins serrate on apical two-fifths; surface moderately clothed with long, mostly erect, silky white hair infused with dark hair on apical fourth, strial punctures medium to coarse, coarser, deeper, and denser toward sides, intestrial punctures medium, shallow; hu- meral interval feebly developed, heavily punctured. Ventral surface with prosternal margin shallowly, arcuately emarginate; abdomen moderately clothed (more densely at sides) with semierect to re- cumbent, silky white hair, punctures small, moderately spaced, becoming 1971 Five New Species of Acmaeodera 15 coarser and dense towards sides of first visible sternite, second visible sternite slightly more densely punctate at middle than first, last visible sternite with a feebly developed, obtusely rounded, subapical plate. Female genitalia as in Fig. 15. Length: 8.2 mm. Width: 2.8 mm. TYPE MATERIAL AND LOCATION: Holotype female, Panamint Moun- tains south of Emigrant Junction, Inyo County, California, elevation 4500- 5000 feet. May 24, 1959 (G. H. Nelson), deposited in the California Academy of Sciences. One hundred twenty-nine paratypes from the following localities in California: Inyo County: same locality as holotype, V-11, V-24- 1959 (GHN). Kern County: Mojave, V-1-1905 (AF), VI-1917 (ECV); 33 mi N Mojave, VI-10-1962 (GHN); Searles Sta., IV-28-1950 (DWD); Boron, V-10-1955 (WRM). San Bernardino County: 6-9 mi. NE Red Mtn., V-1 1-1959 (GHN), 7, 11 and 17 mi N Red Mtn., V-24-1970 (RLW); 5 mi SW Victorville, V-1 0-1 959 (GHN) ; Barstow, V-23-1947 (GPM); Mitchell’s Cavern, VI-16-1962 (DSV); Helendale, V-18-1955 (WRR); Apple Valley, V-6-1955 (WRM); Saddleback Butte, IV-20-1959 (JCH); 5 mi S Twenty- nine Palms, VI-5-1960 (DSV). Los Angeles County: Palmdale, V-26-1957 (DSV); Littlerock, V-30-1957 (WES), V-10-1959 (DSV); Lancaster, V-2-1905 (AF); Valyermo, IV-21-1940 (RMB). Paratypes in the collections of the American Museum of Natural History, California Academy of Sci- ences, California Department of Agriculture, Canada Department of Agri- culture, Los Angeles County Museum of Natural History, United States National Museum, University of California at Davis and Riverside, G. H. Nelson, D. S. Verity, W. F. Barr and the author. Thirty-six specimens from the following localities are not included as paratypes : UTAH: “Utah” (MAC). NEVADA: Esmeralda County: 15 mi S Goldfield, VI-2- 1961 (DSV). ARIZONA: Mohave County: Wickieup, IV- 17- 1962 (GEB). Pinal County: 3-13 mi N Florence, IV-17-1964 (WFB), IV-13 and 16-1966 (WFB, MAC, JMD, JHD); Florence Jet, IV-5-1954 (TRH). CALIFORNIA: Inyo County: Lone Pine, V-2, V-24-1937 (ECV); Lone Pine Campground, V-28-1960 (DSV). Riverside County : Desert Center, IV-14-1964 (AEL). San Diego County: San Felipe Valley, V-5-1962 (DSV). DISTRIBUTION : Southwestern Utah and southern Nevada to southern Ari- zona and California. BIONOMICS: The majority of specimens were taken near Red Mtn., San Bernardino Co., California, visiting the flowers of Cassia armata Watson on which I observed them feeding. This beetle also has been taken visiting the flowers of Encelia farinosa, Sphaeralcea rusbyi eremicola, and C. covesii Gray. It is the only species discussed herein that has not been associated with Ephedra. VARIATION : Specimens of A. mojavei available for study from California ranged in length from 6.4 to 1 1.5 mm and in width from 2.3 to 4.0 mm. This 16 Contributions in Science No. 209 species exhibits wide variation in the form of the prosternal margin, which ranges from emarginate to strongly trisinuate. Only 25 per cent of the speci- mens studied were found to have this margin trisinuate and most of them only feebly so. Almost all specimens exhibiting a strongly trisinuate pros- ternal margin were of larger size. The arrangement of elytral spots is fairly consistent, but variation in their size and number is evident. A basal discal spot, occasionally on one side only, is present in 91 per cent of the specimens studied and 35 per cent of the specimens have the median spots in each row coalesced to form an oblique fascia on each elytron. One specimen was seen with the elytral maculation coalesced and reduced to a short vitta on the apical portion of each elytron, with a tiny, median, lateral spot on one elytron. This condition has also been observed in A. lanata. The humeral spot was found to be present in 5 per cent of the specimens. Occasionally a specimen will have one or two pronotal spots. One specimen has two spots on one side and one on the other. The great majority of specimens have dark hairs on the head and pronotum, and all possess them on the apical portions of the elytra. The subapical plate is usually poorly developed and rarely does it give the effect of a double margin. Specimens from Lone Pine, California, have the plate most strongly developed. A series of 25 specimens from southern Arizona differs slightly from the California forms. The elytral spots are generally smaller and there seems to be much less of a tendency for the median spots to coalesce. Only one specimen was found to possess a humeral spot. None of these specimens exhibits a strongly trisinuate prosternal margin. Although the California and Arizona forms appear somewhat different to the eye, subspecific separation seems unwarranted at this time. DIAGNOSIS AND COMPARISON : The affinities of A. mojavei are diffi- cult to ascertain. In Fall’s (1899) key it may run to A. lanata (Fig. 5) and A. pubiventris (Fig. 4) , but a comparison of elytral maculation will serve to readily separate it from those species. It may also key to A. fenyesi, the elytral markings of which are mere specks, or A. recticollis, which differs at once in having apical, red elytral markings. Van Dyke (1919) confused specimens, which proved to be this species, from Mojave, California, with A. pubiventris (Fig. 4) . One would tend to place specimens with an emargi- nate prosternum in the group “Emarginatae,” as defined by Horn (1878), but the female genitalia are definitely not of the type found in that group. The structure of the female genitalia is, in fact, quite unlike that of the other species discussed herein and may suggest that A. mojavei should stand by itself. Based on this study A. mojavei appears most closely related to A. verityi with which it is most apt to be confused. Some specimens may also resemble A. panamintensis. Differences are discussed in the sections dealing with those species. Two specimens from San Felipe Valley, California, present an enigma. They are far removed from any known locality of A. mojavei, however they 1971 Five New Species of Acmaeodera 17 agree well with that species in general appearance and the female genitalia corroborate this. Perhaps this gives evidence that A. mojavei was once wide- spread over southern California, but could not adapt to the hotter, drier conditions of the lower desert regions. Acmaeodera panamintensis, new species Figures 3,11 FEMALE: Large, moderately robust. Head and pronotum black with feeble coppery reflections. Each elytron black, shining, with orangish yellow mark- ings as follows: a median discal spot at base, paired discal and submarginal spots at basal fourth, a median fascia reaching second discal stria, paired discal and submarginal spots near apical third, a transverse spot midway to another pair near apical fifth, right elytron with an additional small spot near apex, humeral spot absent. Head densely clothed with silky white hair on apical half, with long fuscous hair on basal half, coarsely, reticulately punctured. Pronotum as wide as elytra, 1.87 times as wide as long, wider on basal half, widest in front of base; surface with a broad, shallow, median basal depression, and with a shallow, oblique impression in front of lateral pits, densely clothed with long, suberect, fuscous hair on apical two-fifths, except on extreme front and sides which are clothed with silky white hair, basal three-fifths with silky white hair interspersed with a few fuscous hairs, disc coarsely, densely punctured, punctures becoming coarser and denser toward sides; lateral margins arcuate, entire, flanged on apical two-thirds, part of flange visible from above. Elytra widest on basal ninth; sides sinuate behind umbone, gradually rounded from middle to apex; lateral margins serrate on apical half; surface moderately clothed with long, erect to suberect, fuscous hair interspersed with silky white hair, except on basal two-thirds at sides where there are long, recumbent to subrecumbent, silky white hairs, inner striae separated by about one and one-half to two times the width of punctures, outer striae much closer, with punctures coarser and deeper, interstrial punctures medium- sized, shallow; humeral interval well developed, elevated basally, heavily punctured posteriorly. Ventral surface with prosternal margin trisinuate, with a well developed, rounded prominence on either side of middle; abdomen sparsely to moder- ately clothed at middle with long, recumbent to subrecumbent, silky white hair which becomes dense at sides, with medium-sized punctures moderately spaced at middle, becoming dense at sides, anterolateral punctures of first visible sternite coarser, second visible sternite more finely, densely punctate at middle than first, last visible sternite with a well-developed, broadly rounded, subapical plate. Female genitalia as in Fig. 11. Length: 10.6 mm. Width: 3.9 mm. 18 Contributions in Science No. 209 TYPE MATERIAL AND LOCATION: Holotype female, Wildrose Canyon, Panamint Mountains (Range), Inyo County, California, elevation 7000 feet, June 15, 1961 (G. H. Nelson), on Cowania stansburiana, genitalia slide RLW 314, deposited in the California Academy of Sciences. The following para- types from the same locality: Eleven, 7000 feet, June 15, 1961; four, 8100 feet, June 17, 1961 (GHN); 7400 feet, April 22, 1964 (RLW), dug from stem of Ephedra viridis; 7200 feet, April 21, 1965 (RLW, WEB, ARG), reared from Ephedra viridis. Twelve paratypes, Lee Canyon, Clark County, Nevada, July 1, 1966 (RLW), June 2, 1970 (JMD). Paratypes deposited in the collections of Los Angeles County Museum of Natural History, W. F. Barr, J. M. Davidson, G. H. Nelson, and the author. The following specimens are placed under this species, but are not included in the type series : Two, Benton, Mono County, California, May 27, 1954 (MAC) ; one. May 17, 1942 (WMP) ; one, Mt. Montgomery, Nevada, 7000 feet, June 21, 1942 (RMB). DISTRIBUTION : Southeastern California and southwestern Nevada: Pana- mint, White, and Spring Mts. BIONOMICS: Adults have been taken on the flowers of Cowania mexicana var. stansburiana (Torrey) Jepson and Fallugia paradoxa. Field and labora- tory studies during 1964 and 1965 proved that the larvae bore in Ephedra viridis, which is usually the only species present in the Pinyon- Juniper Wood- land plant community (Munz, 1963) where A. panamintensis occurs. VARIATION : Specimens of A. panamintensis examined ranged from 8.9 to 11.1 mm in length and from 3.3 to 4.3 mm in width. The type series shows variation in the size of the elytral spots, but the arrangement is quite uniform. Their color ranges from light yellow to orangish yellow; the latter color may be due to a slight discoloration often seen in dead specimens of some species of this genus. The basal discal spots are absent in most specimens, but when present are always small. Four specimens have humeral yellow spots and three of these also show lateral yellow or brownish pronotal spots. Several of the specimens examined possess spots which almost reach the apices of the elytra. The hairs of the elytral disc are predominantly whitish in some speci- mens from Lee Canyon, Nevada. The ground color of some individuals exhibits slight bluish reflections. Three bluish specimens from Benton, which is about 130 miles northwest of the type locality and at an elevation of only 5600 feet, can most likely be assigned to this species. DIAGNOSIS AND COMPARISON : This species will key to A. pubiventris in Fall (1899). However the relationship of A. panamintensis to other species is not clear. Superficially it appears to be closely related to A. pubiventris or A . mojavei and in general appearance it may resemble either of those species. However, A. panamintensis is less elongate and less convex. Based upon the structure of the female genitalia it is probably closer to A. atactospilota or A. verityi. Acmaeodera panamintensis is readily distinguished from any of the above by having dark hairs throughout the surface of the elytral disc. 1971 Five New Species of Acmaeodera 19 The occurrence of this species at such high elevations is an interesting facet in the distribution of the group studied herein. The association of this species with Ephedra correlates well with other species under consideration, but this cannot be used alone to establish relationships. A specimen of an entirely unrelated group was also reared from that plant. Additional material and distributional data are needed to more fully interpret the taxonomic position of this beetle. Literature Cited Fall, H. C. 1899. Synopsis of the species of Acmaeodera of America north of Mexico. J. New York Entomol. Soc. 7:1-37. Horn, G. H. 1878. Revision of the species of Acmaeodera of the United States. Trans. Amer. Entomol. Soc. 7:2-27. Kerremans, C. 1906-07. Monographic des Buprestides. Vol. II. Bruxelles. 623 p. Munz, P. a. 1963. A California flora. Univ. Calif. Press, Berkeley and Los Angeles. 1681 p. Tanner, V. M. 1927. A preliminary study of the genitalia of female Coleoptera. Trans. Amer. Entomol. Soc. 53:5-50. Van Dyke, E. C. 1919. New species of Buprestidae (Col.) from the western United States, with supplementary notes concerning others. Entomol. News. 30:186- 190. White, B. E. 1939. A new species of Acmaeodera (Coleoptera: Buprestidae) with biological notes on others from Santa Barbara County, California. Pan-Pacific Entomol. 15:69-75. Accepted for publication November 2, 1970 ■.^M M 1*^ ff.-/'. -t Printed in Los Angeles, LOS ANGELES j COUNTY I; MUSEUM CONTRIBUTIONS in science lUMBER 210 February 17, 1971 OBSERVATIONS ON THE BIOLOGY OF LAND CRABS AND THEIR BURROW ASSOCIATES ON THE KENYA COAST By Charles L. Hogue and Donald B. Bright I I Los Angeles County Museum of Natural History • Exposition Park I Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 OBSERVATIONS ON THE BIOLOGY OF LAND CRABS AND THEIR BURROW ASSOCIATES ON THE KENYA COAST By Charles L. Hogue^ and Donald B. Bright^ Abstract: Brief notes concerning the identification and burrowing habits of the two most common species of land crabs on the south Kenya Coast, Cardisoma carnifex and Sesarma meinerti, are given, as a result of observations made in the dry season of 1968. Records of two insect associates, the flies Aedes pembaensis (Culicidae> and Psychoda martini (Psychodidae), are reported along with pertinent data from a small collection of crabs of the above and others made in connection with the field studies. Introductory Remarks and Acknowledgment Through the generosity of Dr. Purvis L. Martin of San Diego, California, sponsor of a general zoological expedition to Kenya, East Africa, January- March, 1968, Hogue had the opportunity to collect material and data per- taining to four species of burrowing land crabs and certain of their insect associates. A summary of the more significant field observations is presented to help fill the large gaps presently existing in the ecological knowledge of these animals. This report is one in a series evolving from our general study of tropical land crabs and their burrow associates (LCBA, see Hogue and Bright, 1969). Study Sites Observations were made and specimens taken at two stations near the town of Kilifi on the shores of the large bay (Kilifi Creek) flooding the mouth of the Voi River. A map and description of the general area is found in Mog- gridge (1950:301-304). The first station is on the south shore, immediately west of the ferry building (Figs. 1, 2, 3a). It is a gently sloping, coral sand beach, well overgrown with moderate-sized mangroves (Rhizophora mucro- nata and Avicennia marina), bounded shoreward by a banana grove. High steep cliffs close the beach on either side. The second station is situated on the north shore of Kilifi Creek, some two miles west of the ferry landing (Fig. 3b). The terrain here is similar to the first but only a few Avicennia marina trees provide cover. The soil here is a sandy loam with a gravelly surface. Additional collections of crabs were made further north at Mida Creek, near Cede. A map and description of this area is given by Lumsden (1955). ^Senior Curator of Entomology, Los Angeles County Museum of Natural History, Los Angeles, California 9000*7. ^Research Associate, Los Angeles County Museum of Natural History; and Associ- ate Professor of Biology, California State College at Fullerton, Fullerton, Cali- fornia 92631. 1 2 Contributions in Science No. 210 Crabs Observed at Kilifi Creek The most abundant supralittoral burrowing land crab at Kilifi Creek is Sesarma meinerti (Grapsidae) although the less common Car disoma carni- fex (Gecarcinidae) may often attract more attention because of its large size and striking violaceous color. The former measures up to 39 mm across the squarish carapace and displays bright red chelipeds. The rounded, heart- shaped carapace of the latter reaches breadths up to 87 mm and is overall dark purplish with the exception of the lighter purple chelipeds. Just below the high tide zone (predominantly in the mid-littoral zone) two additional burrowing crabs are often in evidence but were not investi- gated. These are the fiddler crab, Uca annulipes (Ocypodidae), with a black- mottled white carapace and a pinkish major cheliped, and the smaller Sesarma ortmanni (Grapsidae) which is similar to S. meinerti in shape but is smaller and has orange chelipeds. Cardisoma appears to be adapted to drier conditions than Sesarma. At both study sites on Kilifi Creek the former occupies a zone landward of the latter but overlapping seaward to a considerable extent. Apparently the highest tide line generally defines the lower boundary of Cardisoma. Sesarma, although occupying a lower zone, is probably flooded only during the highest tides. This is supported further by the mutual occurrence of Avicennia and Sesarma, the plant being normally adapted for occasional flooding. The soil where Cardisoma burrows are found may become very dry during the dry season. All observations (February 2-4, 1968) were made during the later part of the dry season (November to March), and some burrows, well above the highest high tide area, were being actively maintained in essentially dry soil. Yet, in all occupied burrows excavated there was some free water in the bottom due to the crab’s digging to depths sufficient to reach the water table. The surface appearance of the burrows of Cardisoma and Sesarma is quite different (Fig. 2a). Sesarma, being the smaller, constructs narrow- mouthed burrows. The average diameter of the mouths of 25 burrows taken at random was 41 mm. The burrows of juvenile Cardisoma sometimes are confused with those of Sesarma, but frequently the latter are distinguishable by their tendency to be clumped and from the small (13 mm) round feeding casts found about the entrances. Cardisoma burrows can often be distinguished by the appearance of fecal pellets being rod shaped, and terminally truncate. Cardisoma burrows also often have a hood or cap of dried mud which is not noted for Sesarma. These hoods are always relatively small (Fig. 2a) ; the extensive “castles” reported by Silas and Sankarankutty ( 1960) were not seen. Groups of Sesarma burrows characteristically occur under the protection of small Avicennia trees and thus are shaded during most of the day (Fig. lb). The burrows of Cardisoma are situated more at random but many are associ- ated with the trunks and roots of larger trees. 1971 The Biology of Land Crabs on the Kenya Coast 3 Figure 1. Kilifi Creek, a, Station 1. Avicennia thicket, habitat of Cardisoma carni- fex and Sesarma meinerti. b, Burrows of Sesarma meinerti under cover of small Avicennia (Site 1). 4 Contributions in Science No. 210 b Figure 2. a, Large hooded burrow of Cardisoma surrounded by smaller Sesarma burrows. Note spherical feeding casts by the latter (Site 1); b, Active non-hooded burrow of Cardisoma (Site 2). Note fresh mud mound at left (length of forceps 29 cm). 1971 The Biology of Land Crabs on the Kenya Coast 5 Figure 3. a, Adult Cardisoma carnifex at mouth of burrow associated with the roots of a large Avicennia (Site 1); b, Station 2. Cardisoma burrows indicated by arrows. Highest, high water line approximated by dashed line. 6 Contributions in Science No. 210 Six burrows of Car disoma were excavated to determine their under- ground configuration. The results of these excavations are diagrammed in Figure 4. The burrows are simple, that is, with one mouth opening and no interconnections with other burrows or auxiliary tunnels or chambers. In some cases, a small appendix (usually just at ground level) may be present. This is in agreement with our findings for the burrows of other Cardisoma species in the New World and the observations by Bruce-Chwatt and Fitz- Figure 4. Diagrams representing configuration of six burrows of Cardisoma carni- fex excavated on north shore of Kilifi Creek at Station 2. Wavy lines represent level of water table (not available for f); dark tone approximates position of decomposing material when present; figure of crab indicates position where en- countered during excavation. 1971 The Biology of Land Crabs on the Kenya Coast 7 John (1951) for C. armata on the west African coast. Occasionally the upper portion of the burrow throat will have numerous small side tunnels but these have always been found to represent the burrowing activities of other crabs, e.g., Uca and Sesarma (Fig. 4d). Generally, the throat of the burrow slopes gradually (50 to 60 degrees) although in many cases the main axis of the shaft is practically perpendicular to the ground surface. Five of the six burrows excavated clearly possess an enlarged terminal chamber, three of which were filled at the bottom with an accumulation of silt and organic matter undergoing active decomposition as evidenced by its black color and sulphide odor. This feature is similar to other Cardisoma we have studied. Only one Sesarma burrow was excavated. It was a simple vertical canal, approximately 1.3 meters deep, with a slightly enlarged terminal chamber. Cursory examination of many additional Sesarma burrows with a flashlight and probe indicated that the above is typical. Associates Two species of flies were found consistently dwelling in the burrows of both of these crabs : Aedes (Skusea) pembaensis Theobald, 1901 (Culi- cidae) and Psychoda martini Hogue, 1970 (Psychodidae). The mosquito Aedes {Skusea) pembaensis thrives in large numbers in association with both crabs. Formerly only the burrows of Sesarma were recorded as hosting this species (Heisch et al. 1956; Brooke Worth et al. 1961 for S. meinerti; Lumsden 1955:171 for S. eulimene—misidentihcation ? = ortmanni). Adult mosquitoes commonly rest on the walls of the burrow and emerge readily to bite. Larvae and pupae often occur in dense numbers in the water which fills the terminal chambers of all burrows. The eggs of this mosquito have been found on the legs of S. meinerti by Goiny et al. (1957) ; however, no specimens of this crab collected during the study carried mosquito eggs. On the other hand almost all of the C. carnifex were positive for eggs, the latter being found or concentrated in large numbers along certain portions of the anterior parts of the carapace and on the mouth parts. Specifically the following surfaces are utilized by the mosquitoes as oviposition sites (Fig. 5): 1) upper one-half of eye socket; 2) outer surface of exopodite of eye and rarely on eye stalk itself; 3) triangular area at the base of the orbital region; 4) epistomal region; 5) dorso-lateral surface of buccal cavity; 6) outer and inner surfaces of palps and meri of 3rd maxillipeds. This species of mosquito has been incriminated in the transmission of filariasis and may be involved in the dissemination of pathogenic viruses (Heisch et al., 1957). Mosquito surveys for public health purposes should definitely take into account its existence in crabholes and this breeding site should not be overlooked as an important source of large numbers of this potentially dangerous mosquito. 8 Contributions in Science No. 210 Figure 5. Anterior view of carapace of Cardisoma carnifex showing eggs of Aedes pembaensis in areas typically used as oviposition sites by the females. Eyes omitted. A lengthy series of a moth fly (family Psychodidae) of the genus Psychoda was also collected from the burrows of both crabs. It has proven to be an undiscovered species and has been named and described by Hogue, 1970, Psychoda martini. Only adult flies were in evidence at the collection site resting and moving over the burrow walls. No early stages were found in the numerous water samples. Land Crab Records The following general accounts are given for the four species of crabs collected during this study to establish their occurrence and facilitate their recognition by future workers. Cardisoma carnifex (Herbst, 1794) DISTRIBUTION: Mauritius, Madagascar, Andaman Islands, east coast of Africa, and whole of Indo-Pacific. For much of their range they occur in sympatry with C. hirtipes Dana (see Miyake, 1939, for a key to these two species). LOCALITY: Kilifi Creek. HABITAT: Common inhabitants of muddy shores, mangrove swamps and of saline lowland soils near the coast. ECOLOGY: Constructs well-defined burrows in soft soils where there is available ground water during the dry season. Females repair to the sea to introduce the prezoea to the required sea water environment. COLOR: Carapace dark purple; chelipeds light purple to dark cream. REFERENCES: Barnard, 1950; Macnae, 1963; and Miyake, 1939. 1971 The Biology of Land Crabs on the Kenya Coast 9 Sesarma (Sesarma) meinerti de Man, 1887 DISTRIBUTION : Andamans and Madras, Mozambique, Mauritius, Madagascar; east coast of Africa (south to Port St. John’s), Australia, and Indo-Pacific. LOCALITIES: Kilifi Creek, Mida Creek. HABITAT : Sandy-clay areas and muddy banks from upper mid-littoral to the lower limits of the supralittoral. Generally associated with mangroves, e.g., Avicennia, but in upper dryer areas of mangroves also associated with Juncus kraussii and Fimbristylis abbreviata. ECOLOGY : Burrows most common in areas with hard, baked mud. A few burrows have been observed with hoods. Crabs are generally nocturnal, remaining well within the burrow during the day. COLOR: Carapace black to grey; chelipeds bright red. REFERENCES: Chace, 1942; Day, Millard and Broekhuysen, 1954; and Macnae and Kalk, 1969. Uca annuli pes (Latreille, 1820) DISTRIBUTION : East coast of Africa to Indo-Pacific (as far east as the Mariana and Caroline Islands). LOCALITIES: Kilifi Creek, Mida Creek. HABITAT : Mud flats and clay-sandy areas in and adjacent to tropical and subtropical estuaries and mangroves. ECOLOGY : Constructs colonial, intersecting burrows among Avicennia roots, and on open areas between the mangroves. Not reported living within the mangroves proper. Known to migrate long distances (several hundred feet) from the burrows to feed. Females are very inconspicuous and seldom seen out during the day. COLOR: Carapace grayish, chelipeds salmon pink. REFERENCES: Alcock, 1900; Chace, 1942; Macnae and Kalk, 1969; and Miyake, 1939. Note: This is still often listed as Gelasimus annulipes. Sesarma (Holometopus) ortmanni Crosnier, 1965 DISTRIBUTION : East coast of Africa, Madagascar. LOCALITIES: Mida Creek, Kilifi Creek. HABITAT : Muddy soils in mangroves. ECOLOGY : Constructs shallow burrows among exposed pneumato- phores of Avicennia. COLOR: Carapace greenish brown; claws dull orange. REFERENCES : Crosnier, 1965; and Macnae and Kalk, 1 969. Literature Cited Alcock, A. 1900. Materials for a carcinological fauna of India. No. 6. The Brachyura Catometopa, or Grapsoidea. J. Asiatic Soc. Bengal, 69:279-486. Barnard, K. H. 1950. Descriptive catalogue of South African decapod crustaceans. Ann. S. Afr. Mus. 38:1-837. 10 Contributions in Science No. 210 Brooke Worth, C., J. De Sousa and M. P. Weinbren. 1961. Studies on the life history of Aedes (Skusea) pembaensis (Theobald) (Diptera, Culicidae). B. Ent. Res. 52:257-261. Bruce-Chwatt, L. J. and R. A. Fitz-John. 1951. Mosquitoes in crab-burrows on the coast of West Africa and their control. J. Trop. Med. Hyg. 54:116-121. Chace, F. a. 1942. Scientific results of a fourth expedition to forested areas in eastern Africa. III. Decapod Crustacea. Bull. Mus. Comp. Zool. 91: 185-233. Crosnier, a. 1965. Cructaces decapodes, Grapsidae et Ocypodidae. Faune de Madagascar 18:1-143, pi. 1-11. Day, j. H., N. A. H. Millard and G. J. Broekhuysen. 1954. The ecology of South African estuaries. Part IV: The St. Lucia System. Trans. Roy. Soc. S. Afr. 34:129-156. Goiny, H., E. C. C. Van Someren and R. B. Heisch. 1957. The eggs of Aedes {Skusea) pembaensis Theobald discovered on crabs. E. Afr. Med. J. 34: 1-2. Heisch, R. B., H. A. Goiny and M. Ikata. 1956. A new vector of filariasis in East Africa. Trans. Roy. Soc. Trop. Med. Hyg. 50:421-422. Hogue, C. L. 1970. A new moth fly of the genus Psychoda from crabholes on the Kenya Coast (Diptera: Psychodidae). Los Angeles Co. Mus., Contrib. Sci. 204:1-5. , AND D. B. Bright. 1969. Study of the biologies of land crabs and their burrow associates. Assoc. Trop. Biol. Newsletter 18:9-11. Lumsden, W. H. R. 1955. Entomological studies, relating to yellow fever epi- demiology, at Gede and Taveta, Kenya. B. Ent. Res. 46: 149-183. Macnae, W. 1963. Mangrove swamps in South Africa. J. Ecol. 51:1-25. , AND M. Kalk (Ed.). 1969. A natural history of Inhaca Island, Mozam- bique. Witwatersrand Univ. Press, Johannesburg. 163 p. + 11 pi. Miyake, S. 1939. Notes on Crustacea Brachyura collected by Professor Teiso Esaki’s Micronesia expeditions 1937-1938 together with a checklist of micro- nesian Brachyura. Rec. Oceanogr. Works in Japan 10: 168-247. Moggridge, j. Y. 1950. The relations of the coastal tsetse of Kenya to the plant communities. B. Ent. Res. 41:301-315, pi. X. Silas, E. G., and C. Sankarankutty. 1960. On the castle building habit of the crab Cardisoma carnifex (Herbst) (Family Gecarcinidae), of the Andaman Islands. J. Mar. Biol. Assoc. India 2:237-240. Accepted for publication October 23, 1970 LOS ANGELES COUNTY MUSEUM CONTRIBUTIONS Vim IN SCIENCE 'mber 211 April 7, 1971 THE BIRDS OF THE LOWLANDS OF BWAMBA, ! TORO PROVINCE, UGANDA ! 1 1 ! i 1 1 1 |l By Herbert Friedmann and John G. Williams I j ] i Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 BIRDS OF THE LOWLANDS OF BWAMBA, TORO PROVINCE, UGANDA By Herbert Friedmann^ and John G. Williams^ Abstract: The forested areas of the lowlands of Bwamba, extreme western Uganda, are, in effect, the eastern outliers of the great Ituri Forest of the Congo, and it is in Bwamba that many Congo birds reach their eastern limits. The Bwamba Forest was one of the west Uganda forests covered by the four year survey of the Los Angeles County Museum of Natural His- tory under National Science Foundation Grants GB-5107 and GB-7787. The Bwamba collections in this Museum comprise about 2200 specimens of 216 species, which, altogether with a few additions previously recorded, add some 81 species to the Bwamba avifauna as compiled in 1949 by the van Somerens, a total of 380 species in all. In the case of a good number of species these Bwamba records are the first, and only, indications of their occurrence in Uganda. The present report is a complete list of the Bwamba lowland avifauna, but it avoids useless repetition by giving details only where they add to previous knowledge. Our west Uganda survey concentrated on the forest fauna, as it is the wooded sections, and not the surrounding grassy areas, that are in danger of being altered by man, and as a result most of the additional species are sylvan dwellers. The sad fact remains that at the rate at which these forests are disappearing these woodland species may also disappear from the Uganda extremi- ties of their respective ranges. Bwamba County, Toro District, Uganda, lies along the Uganda-Congo border, 0° 40' N. to 0° 54' N., 29° 47' E. to 30° 10' E. It is bounded on the north by the Semliki river, on the east and southeast by the Ruwenzori moun- tains, and on the southwest by the Lamia river. It is isolated from the rest of Uganda by the Ruwenzori range and the Lake Albert escarpment, which effectively demarcate Bwamba from the high plateau of Toro. Although the eastern boundaries of Bwamba County follow the high watershed of the northern part of Ruwenzori, most of the county lies at a much lower altitude in the trough of the Albertine rift. The rich, wet forests of the Bwamba lowlands, now only a fragment of what they were not too many years ago, are essentially Congolese in their fauna. Inasmuch as it is in these forests that many central African birds reach their easternmost limits, it is biologically desirable to know as accurately as possible the longitudinal and latitudinal, as well as the ecological, terminal points for each species. While it is recognized that faunas have limits to which their included species adhere, the precise factors that control or that limit their distribution and the extent to which each of them are effective are frequently ^Research Associate in Ornithology, Los Angeles County Museum of Natural History. ^Research Associate in Zoology, Los Angeles County Museum of Natural History. 1 2 Contributions in Science No. 211 different for different species. The accurate mapping of its range is part of the descriptive knowledge of any given kind of bird (or other animal). There is an urgency about gathering such information and in preserving for future study adequate specimen material of each, as every year sees more and more of the original forest cleared for general agriculture, due to dense population pressure, with all the local changes in biotic distribution that such alterations of the habitat are bound to produce. Although it is undeniable that the lowland forests of Bwamba (2200 to 3500 feet) are no longer the dense, high canopy, tropical habitat they once were, they still are sufficiently like their original condition in some places to have retained their vertebrate fauna. Our knowledge of this fauna has grown greatly in the last few years. This is especially true of the birds, the part of the total fauna dealt with in this paper. This new information is due chiefly to very extensive collecting surveys made in the Bwamba forests by field parties under the direction of the Los Angeles County Museum of Natural History, first in 1963 by the Knudsen-Machris Expedition, and later, in 1967 and 1968-1969, with the financial sponsorship of National Science Founda- tion grants GB-5107 and GB-7787. In all three surveys [the collecting team consisting of A. L. Archer (1963 and 1967) and R. Glen and A. Williams (1968-1969] the field work was concentrated in the dense forest itself, and only small and incidental collections were made in the adjacent grasslands and savannas. A relatively short time was spent in the Ruwenzori highlands as well, but the collections made there, while highly desirable for the Museum, added little to previous knowledge, as the montane fauna is relatively limited and had, apparently, been well explored by earlier parties. Accordingly, in this report, we are leaving out of our discussion the Ruwenzori montane material. The comprehensive report by the van Somerens, issued in 1949, offers a firm base from which to extend our knowledge of Bwamba ornithology. Be- tween that publication and the present one only a small number of papers have appeared that deal wholly or in part with the birds of Bwamba. These, all listed in the bibliography of this report, are as follows : Eggeling 1954; Friedmann 1966, 1968a, 1968b, 1969a, 1969b; Friedmann and Williams 1968; Keith 1968; Keith and Twomey 1968; Ridley, Percy and Percy 1953; Weekes 1949; Williams 1951, 1957. In addition, it may be mentioned that Smart made a number of observations in Bwamba; his sight records were turned over to Keith, who included them in his papers. The van Somerens’ report dealt with the birds of the whole of Bwamba County, which ranges ecologically from the lowland forest, the area of special interest in the present paper, to the alpine highlands of Ruwenzori, where vegetation extends up to about 12,000 feet. The low country, forming Zone 4 of the van Somerens’ paper, is the great stretch of undulating forested terrain, ranging from 2200 to 3500 feet in elevation, watered by numerous transecting streams flowing across to the Semliki. Each of these streams is in a broad and fairly deep valley, originally, and still, in part, covered by dense, high-canopy forest, or, in some instances, with grassy swamps. The areas between these 1971 The Birds of the Lowlands of Bwamba 3 valleys are largely clothed with stands of tall elephant grass. The dense, ever- green forests, which extend in many places up to the Semliki, are merely extensions of the great Ituri forest on the other side of that river, and form the eastern limit of what Chapin (1932: 90) termed the Lower Guinea Forest District of the Congo. The Bwamba forests are characterized by their high canopy, averaging 80 feet, but with trees of lower height as well, and have thick, luxuriant undergrowth. The humidity and temperature are high in these lowlands. According to the van Somerens the temperature varies little and the humidity is usually near the saturation point, with an annual rainfall of about 60 inches a year spread throughout the year, but with the precipitation less in January and February than in other months. At Lesse, on the Congo side of the Semliki valley, Chapin (1932: 48) noted that rain fell almost daily throughout a great part of the year. As Chapin (1932: 81) has pointed out, temperature alone appears to exert no marked influence on the distribution of birds (and other animals and plants as well) in central Africa, at elevations below 5000 feet. Inasmuch as the Bwamba forest is well below this elevation we need not be concerned with temperature, particularly because it is so stable. Although the quantity and the seasonal distribution of rain in the lowlands does affect the biota, there are no differences in this regard in the various parts of the Bwamba forest, which is, in a geographic sense, a relatively small and uniform area. In this whole area the highest monthly mean temperatures occur during the first quarter of the year, usually in March and April, while the lowest minimum temperatures occur in July and August, but the difference between the ex- tremes is only about 10° Centigrade or 18° Fahrenheit. In their discussion of the distribution of birds in this area the van Somerens stressed the fact that the local aspects of the occurrence of the component species showed the Bwamba forest to be more or less stratified— the high canopy; the intermediate trees and tall shrubs; and the dense under- growth—and that each of these strata has its characteristic birds, insects, etc. They gave further information about these forests, which are still applicable even in their present reduced condition, and from which the following facts have been extracted. Some parts of the forest consist of relatively pure stands of ironwood (Cyanometra alexanderi) and in these areas the intermediate stratum is largely absent. While the undergrowth is relatively sparse, it still is sufficient to afford good ground cover habitats for the birds of the forest floor, which have thereby managed to persist in spite of extensive deforestation. The van Somerens noted the heavy, tangled, almost impenetrable under- growth in the mixed forest areas, denser in some places than in others, but all with very dense vegetation along their margins, while in the wetter, soggy portions the forest floor was carpeted with a tall growth of Zingiberaceae (the ginger family) and Marantaceae (arrowroot family). The van Somerens listed among the larger trees ironwood, already mentioned, which is the domi- nant tree at Ntandi, one of our collecting localities, mvule (Chlorophora 4 Contributions in Science No. 211 excelsa), Uganda mahogany (Khaya anthotheca) , mbande (Afzelia bella), and two species of Cordia {millenii and abyssinica) . At Mongiro, another locality where our collectors worked, there are also extensive stands of oil palms (Elaeis guineensis). In places there are many large fig trees (Ficus) whose fruits attract many pigeons, touracos, and other frugivorous birds. Other trees mentioned by the van Somerens include munyamaizi (Mitragyna stipulosa), kesuba (Antiaris toxicaria), nsambya (Markhamia platycalyx) and three others for which they give no local native names, A Istonia congensis, Mildbraediodendron excelsum, and Spathodea campanulata, the flame tree. The material on which the present report is based is housed in the perma- nent collections of the Los Angeles County Museum of Natural History, and comprises 2281 specimens of 216 species. Further collections, containing 98 specimens of 37 species, in the Western Foundation of Vertebrate Zoology, have kindly been made available as requested in connection with this report. The combined collections were made in almost all months of the year, the greatest part in April, May, June, July, October, November, and December. Aside from small number of European migrants, such as some of the sand- pipers and a very few of the wagtails and flycatchers, the rest of the species are resident, completely non-migratory birds. It has not been possible for us to search for unpublished Bwamba material in other museums. To present a clear picture of the great increase in our knowledge of Bwamba ornithology, we may eliminate all the Ruwenzori highland birds from the van Somerens’ list, as we have done from our present collection. The van Somerens reported 344 species in all, but of these 45 were from above 5000 feet elevation, leaving a total of 299 kinds of birds from the Bwamba lowlands, including both the forests and the immediately adjacent open country. To this number we may now add 81 more, bringing the total up to 380 species, a very large list for an area of that size. These 81 additional birds include a number of species already reported in print since the van Somerens’ basic paper, but these are included here because the records were published a few at a time in widely scattered papers and journals, difficult to bring to- gether. It may be noted that our field parties made no special efforts to collect birds away from the forested areas. Consequently the list for the Bwamba lowlands is incomplete for the birds of the open country, but it is, we think, now reasonably comprehensive for the birds of the evergreen forests. Of the birds now added to the lowland Bwamba bird list, nearly two-thirds are forest species. It must be remembered that of the non-sylvan species, the majority were taken at the very edges of, and probably intrude short distances into, the forest. Localities on their labels are identical with those of some strictly woodland birds. In the following list of additions to the Bwamba avifauna, the non-forest birds are marked with an asterisk after their names. Most of the species that were listed by the van Somerens, but not obtained by our collectors, are species of the open country, not of the forest. Moreau (1966: 288) took the van Somerens’ 1949 list, to which he added data from Williams (1951) and unpublished notes supplied him by 1971 The Birds of the Lowlands of Bwamba 5 Keith, and from this total he reduced the list of what he considered the true forest birds of the Bwamba lowlands to 128 species. This number was arrived at by eliminating many species that we consider properly designated as forest dwellers. It is obvious that no two compilers will agree on all cases, but this, in itself is relatively minor, as it is often a matter of judgment rather than of data. The important thing that Moreau brought out, and which stresses the great richness of the Bwamba avifauna, is the following. The area of about twenty square miles of Bwamba lowland forest collected in by the van Somerens and their assistants, by Williams and by Keith, was found to be “. . . inhabited by nearly half as many species of birds as the entire Congo, and nearly two-thirds as many as the 70,000 square miles of the whole Upper Guinea forest . . .” Our present additions to the known Bwamba Forest ornis increase its relative richness still more. The lowland forests of Bwamba must possess one of the greatest concentrations of bird species of any area in the entire African continent. The additions to the known avifauna of the Bwamba lowlands since the van Somerens’ basic report are : Podiceps ruficollis capensis* Nycticorax leuconotus* Butorides striatus atricapillus^ Necrosyrtes monachus^ Accipiter badius sphenurus* Accipiter castanilius Buteo buteo vulpinus* Hieraaetus dubius Falco subbuteo^ Francolinus latharni Sarothrura elegans Himantornis haematopus haematopus Tringa glareola* Tringa ochropus* Tringa hypoleucos* Calidris minuta^ Larus fuscus fuscus"^ Columba malherbii iriditorques Streptopelia capicola tropica'^' Crinifer zonurus* Cercococcyx mechowi Cercococcyx olivinus Glaucidium tephronotum medje Glaucidium castaneum Caprimulgus tristigma tristigma* Caprimidgus batesi Chaetura ussheri sharpei* Alcedo leucogaster leopoldi Merops apiaster* M crops superciliosus superciliosus* Upupa epops epops* Pogoniulus scolopaceus Pogoniulus subsulphureus flavimentum Pogoniulus atro-flavus Trachyphonus purpureus elgonensis Indicator variegatus* Indicator minor riggenbachi Indicator conirostris conirostris Indicator exilis pachyrhynchus Indicator pumilio Melichneutes robustus Melignomon zenkeri V erreauxia africana Dendropicos gabonensis gabonensis Smithornis rufolateralis budongoensis Hirundo daurica emini* Phyllanthus atripennis bohndorfi Chlorocichla simplex Phyllastrephus lorenzi Phyllastrephus icterinus tricolor Bleda eximia ugandae Criniger barbatus weileri Muscicapa striata striata* Trochocercus albicauda Trochocercus albiventris toroensis 6 Contributions in Science No. 211 Erythropygia leucosticta collsi Erithacus erythrothorax mabirae Erithacus cyornithopsis lopezi Cossypha natalensis intensa Zoothera oberlaenderi Campephaga phoenicea phoenicea'^ Sphenoeacus mentalis mentalis* Cisticola natalensis strangeV^ Cisticola brachyptera brachyptera* Malaconotus cruentus adolfi-friederici Oriolus oriolus oriolus Onychognathus fulgidus hartlaubii Ptilostomus afer^ Anthreptes longuemariae haussarum Anthreptes collaris somereni Nectarinia rubescens rubescens Nectarinia afra stuhlmanni Nectarinia bouvieri Ploceus pelzelni pelzelni Ploceus melanocephalus fischeri Ploceus albinucha holomelas Malimbus malimbicus malimbicus Euplectes hordeaceus craspedopterus^ Nigrita luteifrons luteifrons Spermophaga poliogenys Parmoptila woodhousei jamesoni In the present state of our knowledge of African birds no useful purpose would be served by a fully annotated catalogue of all the Bwamba bird speci- mens in the Los Angeles County Museum of Natural History, as in many cases they merely repeat or corroborate the earlier report of the van Somerens. However, there is an advantage in presenting a complete list of the known Bwamba avifauna. To simplify this, where we have no new specimen data or observations, we merely list the species with a reference to the appropriate page in the van Somerens’ 1949 report. Even in the case of species where we do have new data the details are given only where they add to previous knowledge. The nomenclature used is that of White’s lists, except where more recent reviews have been published, or where we have reason to disagree with his treatment. In order to reduce the bulk of the following catalogue the references to the van Somerens’ report are abbreviated to v. S. 1949; followed by the page number. Bwamba Collecting Localities Bubukwanga (16) Bumaga; one mile northeast of Ntandi (9) Bundibugyo ( 1 ) Bundikeke (17) Bundimusuba (2) Buranga Pass (3) Bweramule (not on map; about 6 miles northeast of map limits) Hakitengya (4) Humia River (5) Kanyamwerima (15) Kirimia River (14) Kisuba(18) Korongo (6) Mongiro (7) Nkarara (8) Ntandi (9) Ntoroka; open country near Ntandi (9) Ntotoro (10) Nyankiro (11) Rwansenge (12) Sempaya (13) Tokwe River; close to Humia (5) 1971 The Birds of the Lowlands of Bwamba 7 Figure 1. Bwamba Collecting Localities Family Podicepitidae Podiceps ruficollis capensis Salvadori, Ridley, Percy, and Percy (1951: 161) noted a few dabchicks on the forest-edged Lake Nyangasa. Family Pelecanidae Pelecanus rufescens Gmelin. v. S. 1949: 14. Family Phalacrocoracidae Phalacrocorax africanus africanus (Gmelin). v. S. 1949: 14. Family Anhihgidae Anhinga rufa rufa (Daudin). v. S. 1949: 14. 8 Contributions in Science No. 211 Family Ardeidae Nycticorax nycticorax nycticorax (Linnaeus), v. S. 1949: 15. Nycticorax leuconotus (Wagler). One female, Ntandi, 2300 feet, December 11, 1968; ovary not enlarged; weight 436 grams; bill horn black, the basal two-thirds of the mandible green- ish yellow; iris orange-brown; eye lids blackish edged with yellow; feet pale greenish yellow. The white-backed night heron appears to be a rare, or, at least, seldom collected, bird in Uganda. Ardeola ibis ibis (Linnaeus), v. S. 1949: 15. Butorides striatus atricapillus (Afzelius) . Ridley, Percy, and Percy (1951 : 161 ) reported this heron from the Semliki river, below Sempaya, and another “almost certainly of this species” from the hot swamp near Mbuga Hot Springs. Egretta alba melanorhynchos (Wagler). v. S. 1949: 14. Egretta garzetta garzetta (Linnaeus), v. S. 1949: 15. Ardea cinerea cinerea (Linnaeus), v. S. 1949: 14. Ardea melanocephala Vigors and Children, v. S. 1949: 14. Ardea goliath Cretzschmer. v. S. 1949: 14. Family Scopidae Scopus umbretta umbretta Gmelin. v. S. 1949: 15. Family Ciconiidae Ciconia ciconia ciconia (Linnaeus), v. S. 1949: 15. Ciconia abdimii Lichtenstein, v. S. 1949: 15. Anastomus lamelligerus lamelligerus Temminck. v. S. 1949: 15. Leptoptilus crumeniferus (Lesson), v. S. 1949: 16. Ridley, Percy, and Percy (1953: 164) saw a few of these storks at Bweramule. Ibis ibis (Linnaeus), v. S. 1949: 16. Family Threskiornithidae Threskiornis aethiopicus aethiopicus (Latham), v. S. 1949: 16. Bostrychia hagedash brevirostris (Reichenow). v. S. 1949: 16. Platalea alba Scopoli. v. S. 1949: 16. 1971 The Birds of the Lowlands of Bwamba 9 Family Anatidae Dendrocygna viduata (Linnaeus), v. S. 1949: 16. Alopochen aegyptiaca (Linnaeus), v. S. 1949: 17. Plectropterus gambensis (Linnaeus), v. S. 1949: 17. Sarkidiornis melanota melanota (Pennant), v. S. 1949: 17. Anas sparsaleucostigma {KuppeW) 1949: 16. Anas undulata undulata Dubois, v. S. 1949: 16. Netta erythrophthalma brunnea Eyton. v. S. 1949: 16. Family Accipitridae Aegypius monachus (Linnaeus), v. S. 1949: 17. Gyps bengalensis africanus Salvador!, v. S. 1949: 17. Necrosyrtes monachus (Temminck). Ridley, Percy, and Percy (1953: 164) noted hooded vultures at Bwera- mule. Gypohierax angolensis (Gmelin). One adult male, Ntandi, 2300 feet, June 27, 1967, testes not enlarged; weight 1575 grams; bill whitish gray; iris yellow; feet pale yellowish brown; stomach contents palm nut husks and fibre. Also recorded from Bwamba by the van Somerens (1949: 19). Polyboroides radiatus typus Smith. One immature female in worn, brown plumage, Nkarara, 2200 feet, July 2, 1967; ovary not enlarged; weight 652 grams; cere yellow; bill black, grayish at base; iris brown; feet yellow. Also recorded from Bwamba by the van Somerens (1949: 21). Our specimen was raiding the nests of palm swifts when collected. Dryotriorchis spectabilis batesi Sharpe. Added to the Bwamba list by Ridley, Percy, and Percy (1953: 162) on the basis of a sight record, “almost certainly of this species,” near Hakitengya, September 30. Terathopius ecaudatus (Daudin). v. S. 1949: 19. Accipiter tachiro sparsimfasciatus (Reichenow). Six specimens, 1 5,5 $ , Ntandi, 2300 feet, June to December; weights 235 grams 5 ; 300 to 493.5 grams $ ; stomach contents small birds and a rodent. Also recorded from Bwamba by the van Somerens (1949: 20-21). 10 Contributions in Science No. 211 Accipiter badius sphenurus (Riippell) . One adult male, Ntandi, 2300 feet, October 26, 1968; testes slightly enlarged; weight 108 grams; bill black, gray at base of mandible; iris orange; feet bright yellow; eye skin orange-yellow. The specimen is somewhat inter- mediate between sphenurus and polyzonoides, but is well placed in the former race. Accipiter castanilius Bonaparte. This rare hawk is represented by a single male, taken April 1, 1963; testes not enlarged; bill black, yellowish at base; feet yellow. As reported earlier (Friedmann, 1966: 15) this is the first record for the species in Uganda, and the easternmost example thus far reported. Accipiter erythropus zenkeri Reichenow. The fact that three examples of this hawk were taken in as many localities in Bwamba indicates that it must be fairly common, although like most forest raptors it is not numerous in any one spot. The specimens are : 1 9 , Rwan- senge, 2200 feet, July 13, ovary not enlarged; 1 $ , Ntandi, 2300 feet, Decem- ber 8, testes slightly enlarged; 1 $ , Nyankiro, 3000 feet, November 24, ovary not enlarged; weights recorded for the male 94 grams, and 170 grams for one of the females. Bill black, grayish black on the lower mandible; yellowish at the base in one 9 ; cere and gape chrome yellow; iris red to bright crimson; feet chrome yellow to lemon-yellow; stomach contents a small bird in one case, and a large caterpillar in another. The specimen collected at Ntandi was seen to fly into the upper foliage of a tall tree, almost smashing its way among the leaves. It appeared to be attempting to catch some small birds, making short but rapid flights among the branches. It did this four times before it was collected. The Nyankiro bird was one of a pair in courtship display. It flew over the -other individual in circles and then landed on a branch, when it began to swing its tail back and forth for a few seconds. The fact that the specimen is a female raises the question of the activities of the sexes in the courtship performance. Also recorded from Bwamba by the van Somerens, 1949: 20. Melierax metabates mechowi Cabanis. v. S. 1949: 21. Melierax gabar (Daudin). v. S. 1949: 21. Urotriorchis macrourus (Hartlaub). A male, taken at Ntandi, 2300 feet, November 27, 1968, is the first actual specimen record from Uganda. The van Somerens (1949: 21) saw the long-tailed hawk at Ntandi in August 1946, and reported that Haddow had seen the species in Bwamba on several occasions. While there was little reason to question the accuracy of these sight records, it is good to have them reinforced by a specimen. The bird was not in breeding state. Our specimen is in nearly fully adult plumage, but still has some of the brown feathers of 1971 The Birds of the Lowlands of Bwamba 11 the immature stage among the newer blue-gray ones of the upperparts, and also has some white showing among the chestnut bay of the underparts; bill black, grayish at the base; cere and gape yellow; iris orange-brown; eye ring pale grayish yellow; feet yellow; weight 491.5 grams. The specimen was col- lected in the upper undergrowth of mixed forest. When first seen it was low down, and when disturbed, it flew through the forest at the same low level, finally landing higher up, about 30 feet from the ground. Its stomach con- tained one small rodent. On two other occasions the collectors noted examples of this hawk, but were unable to obtain them, an adult at Nyankiro, November 24, and a young bird at Ntandi, November 29. Kaupifalco monogrammicus (Temminck). Lizard buzzards were collected on two occasions, a male at Ntandi, 2300 feet, December 11, and a female at Nyankiro, 3000 feet, November 25; both were in non-breeding condition; bill black; cere and gape orange; iris dark chestnut brown; eye ring dull orange; feet pale orange; stomach of one speci- men contained a single large beetle, the other was empty; one bird weighed 309 grams. Buteo rufofuscus augur (Riappell). v. S. 1949: 20. Buteo buteo vulpinus (Gloger). Two males of the steppe buzzard were collected at Ntandi, 2300 feet, one on November 3 and the other on December 10; both with small testes, bill black, grayish basally; cere yellow; iris cream brown; feet yellow. The Novem- ber example was seen to fly up from the ground with a rodent in its claws, but this was lost when the bird was shot. Its stomach contained some rodent fur; the other bird had some snake scales in its stomach; one bird weighed 427 grams. This hawk is, of course, a winter visitor from the Palaearctic, and, as such, is only seasonally a member of the Bwamba ornis. Lophoaetus occipitalis (Daudin). One adult, unsexed, crested hawk eagle was taken at Ntandi, 2300 feet, June 13; bill gray becoming black terminally; iris and eye ring yellow; feet pale yellow; stomach contents remains of a rodent, Lophuromys. Also reported from Bwamba by the van Somerens (1949: 18). Stephanoaetus coronatus (Linnaeus). One female, Ntandi, 2300 feet, June 25; bill black, cere and gape yellow; iris cream yellow; feet yellow. The van Somerens (1949: 18) listed numerous Bwamba localities for the crowned hawk-eagle, showing its general presence in the forests there. Polemaetus bellicosus (Daudin). v. S. 1949: 18. 12 Contributions in Science No. 211 Hieraaetus dubius (Smith). The very rare Ayres’ hawk-eagle is represented by an adult male, testes slightly enlarged, taken at Ntandi, 2300 feet, June 25; bill black, grayish basally; cere and iris yellow; feet pale yellow; weight 685 grams; remains of a small bird in stomach. This specimen, together with another from the Kibale forest, have been discussed by the present authors previously (1968: 12). Aquila wahlbergi Sundevall, v. S. 1949: 18. Haliaeetus vocifer (Daudin). v. S. 1949: 18. Milvus migrans parasitus (Daudin). v. S. 1949: 17. Ridley, Percy, and Percy (1953: 164) saw this kite at Bweramule. Elanus caeruleus caeruleus (Desfontaines). v. S. 1949: 17. Macheirhamphus alcinus anderssoni (Gurney), v. S. 1949: 18. Family Pandionidae Pandion haliaetus haliaetus (Linnaeus), v. S. 1949: 22. Family Falconidae Falco cuvieri Smith, v. S. 1949: 17. Falco subbuteo Linnaeus. The hobby was added to the Bwamba list by a single example in worn plumage, a male with small gonads, taken at Ntandi, 2300 feet, at the edge of the forest, November 29; bill pale yellowish gray darkening to black at the tip; cere greenish yellow; iris dark brown; eye skin yellowish; feet pale yellow; weight 125 grams; one beetle in stomach. Falco ardosiaceus Bonnaterre and Vieillot. v. S. 1949: 17. Family Phasianidae Francolinus lathami Hartlaub. A Latham’s francolin was caught in a trap on the ground near a stream at Ntandi, 2300 feet, November 30. The bird is a fully adult male; testes not enlarged; bill black; iris dark brown; feet dull lemon yellow; weight 254 grams; stomach contents small seeds, grit, and unidentifiable insect remains. This species reaches its eastern limits in western Uganda, where it has been collected only a few times. Francolinus squamatus schuetti Cabanis. Two specimens: 1 5 , testes not enlarged, Bwamba Valley, April 30; 1 5 , testes not enlarged, Bundimusuba, July 11; bill orange; iris brown; feet orange; weight 552 grams; stomach contents seeds and grit. Also recorded from Bwamba by the van Somerens (1949: 22). Coturnix delagorguei delagorguei Delagorgue. v. S. 1949: 22. 1971 The Birds of the Lowlands of Bwamba 13 Family Numididae Numida meleagris toruensis Neumann. One downy chick, $ ?, Ntoroka, 2800 feet, June 23; bill pale yellow- brown; iris grayish brown; feet dull orange-brown. Also recorded from the Bwamba lowlands by the van Somerens (1949: 23). Ntoroka is not a forest locality, and the helmeted guineafowl is, of course, a bush country bird. Guttera edouardi sethsmithi Neumann. One 5 , testes not enlarged; Ntandi, 2300 feet, July 2; bill pale greenish horn, the lower mandible darker, especially basally; iris dark brown; feet blackish; weight 1026 grams; stomach contents grit, vegetable and insect remains. Recorded by van Someren ( 1949: 22) from other Bwamba localities as well as from Ntandi. Family Rallidae Limnocorax flavirostra (Swainson) v. S. 1949: 23. Sarothrura elegans (Smith) The little buff-spotted crake may be added to the Bwamba avifauna on the basis of an adult male, testes not enlarged, taken at Ntandi, 2300 feet, June 29; bill blackish horn, paler on the mandible; iris dark brown; feet gray- brown; weight 41 grams; stomach content mainly snails. Sarothrura pulchra centralis Neumann. Common in the Bwamba forest; six specimens from Ntandi, 2300 feet, and Bundimusuba, 3000 feet; weights 43 to 49 grams; all in non-breeding condition (June, July, November); stomach contents grit, snails, and insect remains. These rails respond to an imitation of their call, a series of five notes given in quick succession. One that was called in by Glen came so close, almost on his shoe, that it was too close to shoot. Williams then had to call it toward him and was able to collect it. Himantornis haematopus haematopus Hartlaub. This large forest rail is here reported for the first time from Uganda, and is the easternmost known specimen record. An adult $ , testes not enlarged, was collected at Ntandi, 2300 feet, December 2, 1968; bill black, pale green- gray at the base of the mandible; gape black; iris orange-brown; eye skin blackish; feet pinkish red; weight 438 grams; stomach contents large bits of grit and some seeds. This specimen was trapped in a noose set on an animal path in the forest. Another individual, possibly its mate, was also present but retreated when the bird was removed from the snare by the collector. Gallinula chloropiis meridionalis (Brehm). v. S. 1949: 23. Family Burhinidae Burhinus vermiculatus vermiculatus (Cabanis). v. S. 1949: 24. 14 Contributions in Science No. 211 Family Charadriidae Charadrius hiaticula tundrae (Lowe), v. S. 1949: 23. Charadrius tricollaris tricollaris Vieillot. In the open country at the edge of the forest, at Ntandi, 2300 feet, one male, testes not enlarged, was collected July 19; weight 32 grams. Also recorded by the van Somerens ( 1 949 : 23) in all months of the year. Family Scolopacidae Tringa nebularia (Gunnerus). Ridley, Percy, and Percy (1953: 162) reported one at Mongiro, 2300 feet, October 16. T ringa stagnatilis (Bechstein) One unsexed marsh sandpiper was collected at Mongiro, November 7. An earlier record from the same place was listed by Ridley, Percy, and Percy (1953: 162). Tringa gl areola (Linnaeus). The wood sandpiper was found to be common at Mongiro by Ridley, Percy, and Percy ( 1953: 162) ; our collectors did not find it. Tringa ochropus (Linnaeus). Reported from Bwamba only by Ridley, Percy, and Percy ( 1953: 162) at Mongiro. T ringa hypoleucos Linnaeus. Same as for preceding species. Gallinago media (Latham), v. S. 1 949 : 24. Crocethia alba (Pallas), v. S., 1949: 24. Calidris minuta (Leisler). Known from Bwamba only on the basis of a sight record by Williams (1951: 110). Philomachus pugnax (Linnaeus), v. S. 1949: 24. Himantopus himantopus himantopus (Linnaeus), v. S. 1949: 24. Family Laridae Larus fuscus fuscus (Linnaeus). Known from Bwamba on the basis of one example reported by Ridley, Percy, and Percy ( 1953: 162), and another captured at Bundibugyo, July 2, recorded by Eggeling (1954: 198). The latter was a bird originally banded in Sweden. 1971 The Birds of the Lowlands of Bwamba 15 Family Columbidae Columba arquatrix arquatrix Temminck. Three specimens of this common pigeon were collected: 1 male, 1 female, Bundimusuba, 3000 feet, November 20, 27; 1 male. Mongiro, 2300 feet, November 6; all in non-breeding plumage; bill yellow; iris yellow-brown; bare skin around eye lemon yellow; feet yellow; weights 309 ( $ ), 408, 429 grams (5s). The female has the occiput and nape duskier than do the males. This species was also recorded from Bwamba by the van Somerens (1949: 24) and by Ridley, Percy, and Percy (1953: 162). Large flocks visit the hot springs at Mongiro to drink, usually in the late afternoon. A few Columba albinucha sometimes associate with these flocks. Columba albinucha Sassi. Ridley, Percy, and Percy (1953: 162) collected a young white-naped pigeon near Mongiro, October 7. Also reported earlier by the van Somerens (1949: 25). Columba malherbii iriditorques (Cassin). Keith (1968: 5) suggested that a pigeon whose voice he tape-recorded in Bwamba was of this species. He did not see or collect the bird, and at the time thought he was listening to C. delegorguei, a pigeon with which he was familiar. It was later, on realizing that delegorguei does not occur in Uganda or the adjacent part of the Congo, that he reidentified the recording as iriditorques, a species known to live in the lowland forest just across the Semliki river, and known to have a call very similar to that of delegorguei. The place of iriditorques in the Bwamba fauna is highly likely, but is yet to be corroborated by a specimen record. Recently we have reported (1970: 16) the first Uganda specimen of this pigeon from the lower level (5300 feet) of the Impenetrable Forest, extreme southwestern Uganda. Streptopelia semitorquata (Ruppell). This common pigeon was collected at three Bwamba localities: Ntandi, 2300 feet, December 6, one male, testes not enlarged; Rwansenge, 2200 feet, July 14, one male, testes enlarged; Bwamba Valley, April 24, one unsexed bird; feet dark maroon; weights 193 to 200 grams; stomach contents seeds. Also recorded by the van Somerens ( 1949: 25). Streptopelia capicola tropica (Reichenow). Ridley, Percy, and Percy (1953: 162) reported this ring-necked dove only east of the Buranga Pass, but not on the Bwamba side of Ruwenzori. Turtur tympanistria (Temminck). A long series of 16 specimens is at hand from Mongiro, Nkarara, Ntandi, and Bundimusuba; weights 52 to 58 grams ( $ ); 69 to 79 grams ( 5 ). Also recorded by the van Somerens (1949: 26). 16 Contributions in Science No. 211 Turtur afer (Linnaeus), v. S. 1949: 26. Aplopelia larvata jacksoni (Sharpe), v. S. 1949: 26. Treron australis uellensis (Reichenow). The Uelle green pigeon is represented by ten specimens from Ntandi, 2300 feet; Bundimusuba, 3000 feet; and Bwamba Valley. One of the birds, taken on July 6, had an enlarged ovary; the others were not in breeding con- dition; bill whitish gray, the soft parts pinkish red; iris bright metallic blue; bare skin around eye gray; feet lemon yellow in five specimens, red in the other five; weights 130 (2), 182 to 216 grams ($). Also reported by the van Somerens (1949: 26-27) and Ridley, Percy, and Percy (1953: 162). If uellensis and gibberifrons differ in the color of their feet, half of our Bwamba birds are of the former race, with yellow feet, and half are gibberifrons, with red feet. Family Psittacidae Poicephalus meyeri saturatus (Sharpe), v. S. 1949: 31. Psittacus erithacus erithacus (Linnaeus). One male, Ntandi, 2300 feet, December 8; bill black; iris gray; feet gray; weight 402 grams. Also recorded by the van Somerens (1949: 31). Psittacula krameri krameri (Scopoli). v. S. 1949: 31. Agapornis pullaria ugandae Neumann, v. S. 1949: 31. Agapornis swinderniana emini Neumann. Two males, Ntandi, 2300 feet, November 4; testes not, or only slightly, enlarged; two males, Nyankiro, 3000 feet, November 24; testes not enlarged; maxilla blackish gray, darker at the tip; mandible pale yellowish gray, darker on the tomium; iris Naples yellow; feet olivaceous gray; weights 39 to 41 grams. This relatively rarely collected love bird was seen at the edge of an isolated patch of forest, where as many as four birds were seen at once perch- ing on the top of a tall tree. Noisy screeching sounds were heard from them. Previously recorded from Bwamba by the van Somerens (1949: 32) as well. Theirs was the first announcement of this species in Uganda; the present specimens are the only others taken in that country. The species feeds largely on the fruits of a small fruited fig tree, and has also been seen visiting the large red blossoms of the flame tree, Spathodea. Family Musophagidae Tauraco schutti emini (Reichenow). Emin’s turaco is a common bird in the Bwamba forests; four specimens collected: Ntandi, 2300 feet, two males, June 18 and November 1; Mongiro, 2300 feet, one male, November 1; Bwamba Forest, one unsexed bird, April 24; bill black, reddish at base of mandible; iris dark brown; bare eye skin 1971 The Birds of the Lowlands of Bwamba 17 red; feet black; weight 215, 229, 235 grams; stomach contents fruits. Also reported from several Bwamba sites by the van Somerens (1949: 31). Musophaga rossae Gould. Common in Bwamba; seven specimens collected: Bwamba Forest, one unsexed bird, April 19; Bundimusuba, 3000 feet, two males, one female, one unsexed bird, July 6, November 20, one of the July birds with large testes, the other and the November example with small gonads; Korongo, Semliki River, 2200 feet, one male, July 14, testes small; Mongiro, 2300 feet, one female, November 19, ovary slightly enlarged; maxilla yellow, with grayish tomium; cere orange; mandible orange-brown becoming yellow at the tip; iris dark brown; eye skin yellow; feet black; weight 398 grams ( $ ), 415 grams {$)’, stomach contents fruits and seeds. The van Somerens (1949: 30) also reported this turaco in the Bwamba forests. Crinifer zonurus (Riippell). Not in the forest, but in the adjacent open country; one female, ovary small, Bwamba Valley, April 24; bill greenish yellow; feet black. Corytheola cristata (Vieillot). The giant turaco is common in the Bwamba forests, as was first reported by the van Somerens ( 1949: 30). Our collectors noted it on several occasions and collected one male at Ntandi, 2300 feet, June 30; bill deep yellow on basal half, orange-red on the terminal half, the culminal ridge and tip black- ish; iris dark brown; feet brownish black; weight 857 grams. Family Cuculidae Clamator glandarius (Linnaeus), v. S. 1949: 28. Clamator jacobinus pica (Hemprich and Ehrenberg). v. S. 1949: 28. Clamator levaillantii (Swainson). One male, Ntandi, 2300 feet, June 18; testes not enlarged; bill black; iris dark brown; feet dull gray; weight 115 grams; stomach contents beetle larvae, small and large. Also reported by Fladdow {in van Somerens, 1949: 28). This cuckoo is not a sylvan bird, and must have been found just outside the true forest. It probably is not abundant as only one specimen was collected and as it was not seen years ago by the van Somerens in Bwamba. Cuculus solitarius solitarius Stephens. The red-chested cuckoo is represented by two examples, an adult male, testes slightly enlarged, Nkarara, 2200 feet, July 3, and an unsexed, juvenal bird, Ntandi, 2300 feet, November 1. In the adult the soft parts were recorded as follows : bill black, the mandible yellowish gray and basally yellow in one, merely somewhat grayish in the other; iris dark brown; eye-skin blackish; feet pale brownish yellow; stomach contents hairy caterpillars and other insect remains. 18 Contributions in Science No. 211 The adult bird is solitarius and not clamosus gabonensis, but it approaches the latter species in that it has the dark abdominal bars slightly darker and broader than in any of a series of solitarius from Kenya. Indeed, it agrees in this respect, as well as in the creamy white background of its abdomen and under tail coverts, with our Bwamba example of gabonensis, which, in turn, is unusually pale below for its species. It may be recalled that Ogilvie-Grant (1910; 424) recorded a similar solitarius specimen from the Ruwenzori. The range of plumage variations in solitarius, as outlined long ago by Ogilvie-Grant and others, and in clamosus gabonensis, as described by Bates (1911: 500-502) and by Bannerman (1921: 89-95) still need further study with more material with more accurate and more detailed documentation, plus some facts, as opposed to guesses, as to the inheritance pattern of these characters. There can be no serious doubt about the close relationship of the two taxa, and it may be that the presently puzzling variations in the birds from areas of sympatry, such as Bwamba, may be the result of original mixing of the two before they had attained specific distinctness. The calls of the two species are, of course, quite distinct, but we have no way of knowing how or when, their vocal differentiation took place. Cuculus clamosus gabonensis Lafresnaye. One adult male, in non-breeding state, was taken at Ntandi, 2300 feet, December 12; bill black, base of mandible and sides more grayish; iris dark brown; eye ring black; feet pale brownish yellow; weight 89 grams. This speci- men is considered to be gabonensis and not solitarius because of its darker, more glossy blue-black upper parts, the reduction of white marks on its rec- trices, and because the reddish color of the breast extends across the throat and chin. However, the posterior lower parts are much paler than any other gabonensis examined, and agree, in fact, in the coloration of these areas, with the adult solitarius from Nkarara, mentioned under the preceding species account. The specimen would thus seem to come within the “rnabirae” group of gabonensis. The collector noted that this bird was heard calling a mournful whistle of three notes in an ascending scale— fwoo, twoo, twoo. This is certainly in agreement with the known notes of gabonensis and not of solitarius and corroborates our identification of this unusually pale-bellied specimen. The van Somerens (1949: 28) also heard gabonensis in Bwamba. Williams (1951: 107) obtained a specimen there also. That this species and solitarius are sympatric in that area is not surprising; the two have been recorded together in a good number of places in the eastern Congo. Cercococcyx mechowi Cabanis. One female, Bwamba Forest, April 23, ovary small; one male, testes not enlarged, Bundimusuba, 3000 feet, July 10; bill blackish, the mandible green- ish horn for its basal two-thirds, the tip black; iris dark brown; eye lids yellow; feet pale yellow with an orange tint; weight 52 grams; stomach contents insect 1971 The Birds of the Lowlands of Bwamba 19 fragments. The April 23 specimen is the one mentioned in an earlier paper (Friedmann, 1966: 18). Cercococcyx olivinus Sassi. Our collectors did not meet with this cuckoo, but Keith (1968: 6) ob- tained one in a net in the Bwamba Forest, July 14, 1963. This is the only Bwamba record to date. Chrysococcyx ftavigularis Shelley. This cuckoo would appear to be commoner in the Bwamba Forest than previous records suggest it is in the Congo Forest on the other side of the Semliki. Between November 3 and December 10, eight specimens, seven males and one female, were collected at Mongiro, 2300 feet, and Ntandi, 2300 feet; bill yellow-green, the tip and tomium yellow; iris cream white to cream yellow; eye-ring yellowish green; feet greenish yellow; weight 27.5 to 31 grams; stomach contents caterpillars. This remarkable series of a rarely collected bird has already been reported in detail (Friedmann 1969). The yellow-throated cuckoo had been reported previously from Bwamba by the van Somerens (1949: 29) and by Ridley, Percy, and Percy (1953: 163). Bwamba is the easternmost area, and the only one in Uganda, from which this species has been collected. In the senior author’s 1968 review of the glossy cuckoos it was pointed out that C. ftavigularis was more closely related to C. klaas than to any other species of the genus Chrysococcyx, and that these two, together with C. cupreus, formed a group apart from their congeners. At the time that study was completed the only critical information on the anatomical similarities and differences in the various species of glossy cuckoos was what Berger (1955) had been able to record. When he made his dissections, no anatomi- cal material of ftavigularis was available to him, as none existed in any museum. Since then we have obtained the specimens mentioned above, and the bodies of these birds were preserved as well as their skins. In an attempt to add to his earlier data, the body of one of our Bwamba specimens was sent to Berger, who very kindly agreed to study it. Although such a carcass, without the head, wings, and legs, and largely lacking the appendicular bones and muscles, presents only limited characters for exami- nation, it is of interest to report that Berger found nothing to contradict or to minimize the close relationship of ftavigularis and klaas, and he did find two anatomical features that further support this presumed relationship. The following data were generously sent to us, for our use, by Dr. Berger. The structure of the iliotibialis muscle in ftavigularis is like that in klaas and in cupreus and not like that in caprius. Differences in this muscle were looked upon, in Berger’s 1955 report, as among the most important charac- ters separating caprius from klaas and cupreus. Therefore the fact that its structure in ftavigularis agrees with that in klaas and cupreus does support the close affinity existing between these species. 20 Contributions in Science No. 211 Furthermore, flavigularis has a single pair of intrinsic syringeal muscles, as do the other glossy cuckoos Berger had previously dissected, but here again flavigularis agrees with klaas and cupreus in having these muscles inserted on the fourth bronchial semiring, whereas in caprius and lucidus they are inserted on the third one. Through the kind cooperation of Dr. Anton De Roo, of Tervuren, we have been able to study some young males of flavigularis, which enable us to add to the 1968 account of its plumages (loc. cit.: 114). The first, juvenal, plumage has the upperparts of the head, body, and wings dark, slightly glossy green (as in the adult) but the feathers from the crown to the upper tail coverts with narrow, broadly spaced, bars of pale tawny; the underparts finely barred as in the female. The second stage shows the feathers of the head and upper back, but not those of the rump and wing and tail coverts, without the tawny bars; under- parts fairly similar to the first stage but with a slight yellowish tinge on the chin and throat; underparts of the body somewhat less ochraceous and paler, and with the dusky bars slightly paler than in the first stage. In November 1969 our collectors, Robert M. Glen and Andrew Wil- liams, made a quick visit to the east end of the Maramagambo Forest and heard Chrysococcyx flavigularis calling, but were unable to collect a speci- men for the record. There is no reason to question this “sound” record, as the two men became familiar with the bird during their work in Bwamba. This observation extends the Uganda range of this cuckoo to a second locality about 85 miles south of Bwamba. It must be realized that the intervening country is not suitable for this bird, and that its known range in Uganda is merely the two forested areas, Bwamba, and Maramagambo. Chrysococcyx klaas (Stephens). Common in the more open parts of the forest; two specimens were collected at Ntandi, 2300 feet, a male, testes not enlarged, December 8, and a female, July 2, also not in breeding state; bill dull greenish gray, the tip and tomium dusky; iris brown, flecked with gray; feet olive; weight 27 to 30 grams; stomach contents caterpillars and other insects. This cuckoo was previously listed from Bwamba by the van Somerens (1949: 29). A surprising note by the van Somerens informs us that two males showed a number of pale yellowish, freshly grown feathers on the throat. While it is now known that klaas and flavigularis are sympatric in Bwamba, and while it is considered that they are more closely related to each other than to any other species of the genus Chrysococcyx, this observation cannot be looked upon as suggestive of hybridization between them. It is of definite interest, however. It would be advisable for collectors to make a particular effort to note the coloration of new throat feathers in klaas away from possible sympatry with flavigularis, as it may be that some of these feathers are pale yellowish at first but quickly fade to white, as Williams has found in Kenya birds collected while in moult. 1971 The Birds of the Lowlands of Bwamba 21 We know that in museum skins even the very bright yellow ventral feathers of males of C. cupreus may fade to white. Chrysococcyx cupreus cupreus (Shaw). The emerald cuckoo is common in the Bwamba forests. At Ntandi, 2300 feet, three females were collected, July 8, 17, and December 10; all in non- breeding state; bill blue, greenish at base of mandible; iris dark brown; eye skin apple green; feet bluish greenish gray; weight 27, 38, and 45.5 grams (a surprising range); stomach contents caterpillars. The van Somerens (1949: 29) had previously recorded the emerald cuckoo from Bwamba, also. Chrysococcyx caprius (Boddaert). v. S. 1949: 29. Ceuthmochares aereus aereus (Vieillot). Five examples of this species, collected at N. Humia, 2300 feet, Decem- ber 9; Ntandi, 2300 feet, June 28, November 15, 16; and Rwansenge, July 15, attest to the fact that it is fairly numerous in the Bwamba forest, especially at the periphery of the denser parts of the woods. The specimens were found feeding and skulking among the dense foliage well up in the trees in second- ary forest growth; stomach contents caterpillars, beetles, and other insects; bill yellow, black at the base of the culmen; iris dark red; feet black; weights 58 to 63 grams. The June 28 bird showed slight gonadal enlargement; the others all had small gonads. The yellow-billed green cuckoo was also reported from the Bwamba by the van Somerens (1949: 30). Centropus monachus fischeri Reichenow. One female was taken at Ntandi, 2300 feet, July 10; ovary small; bill black; iris crimson; feet black; weight 222 grams; stomach contents insect fragments and slugs. The van Somerens (1949: 29) found this coucal to be common around the swamps in the Bwamba. Centropus superciliosus loandae C. Grant. Not a forest bird, but found in the adjacent open country. One female was taken in the “Semliki Valley,” April 20; it was in non-breeding state. The van Somerens (1949: 30) noted it in the areas at the margin of the forest, chiefly in the bush and scrub country of the Bwamba lowlands. Family Strigidae Bubo africanus africanus (Temminck). v. S. 1949: 40. Bubo lacteus (Temminck). One female Verreaux’s eagle owl was collected at Ntandi, 2300 feet, October 27; ovary not enlarged; bill whitish gray, more bluish gray on the mandible; iris dark brown; eye skin blackish; feet whitish gray. The bird was seen at dusk, sitting on the top of a dead tree flapping its wings and bowing 22 Contributions in Science No. 211 its head up and down, and simultaneously making a retching sound, as if trying to cough up a pellet of indigestible remains of its last meal. Also recorded from Bwamba by the van Somerens (1949: 40). Glaucidium tephronotum medje Chapin. One male, collected at Bundimusuba, 3000 feet, July 11; testes not enlarged. This example was discussed in an earlier paper (Friedmann and Williams, 1968: 14-15), and it is the only specimen record from Bwamba. This owl has also been taken in three other western Uganda localities— the Kibale, Kalinzu, and Impenetrable forests. The specimen had the bill yellow, tinged with greenish on the mandible; iris light yellow; eyelids yellowish; feet orange -yellow; weight 95 grams; stomach contents insect remains and small bones. Glaucidium castaneum Neumann. This rarely collected owl was added to the Bwamba avifaunal list, and, indeed, to that of Uganda, when an adult female was taken at Ntandi, 2300 feet, December 8, 1968; ovary not enlarged; bill pale yellowish horn; cere yellow; iris bright chrome yellow; eyelids grayish; feet pale yellow; weight 1 19 grams; stomach contents entirely beetle fragments. The bird was first noted perching in an ironwood tree as it was being mobbed by small birds. It flew off and was followed and obtained by the collector. Ciccaba woodfordii nuchal is (Sharpe). The wood owl was found at Ntandi, 2300 feet, where three specimens were obtained: one adult female with small ovary and one ju venal male on December 10, and one adult male with slight testicular enlargement on November 16; bill dull lemon yellow, the cere with a brownish wash in the female, not in the male; iris very dark brown; feet yellow; weight 269 grams (male), 285 grams (female), 265 grams (young bird); stomach contents grass- hoppers and caterpillars. The female is much darker, less rufescent, than the male, especially on the upper parts. The young bird, still with much of the natal down clinging to the new feathers, is very pale. This owl was also noted in Bwamba by the van Somerens (1949: 40). Asio capensis capensis (Smith). The African marsh owl does not occur in the true forest, but in the open moist country outside it. The van Somerens ( 1949: 40) reported one at Bundi- bugyo, and others along the Bwamba road. Family Caprimulgidae Caprimulgus pectoralis nigriscapularis Reichenow. Williams (1951: 108) collected an adult male in breeding condition and a nestling of this nightjar at Ntotoro, March 13, 1948. Earlier still the van Somerens (1949: 41) found the species at Ntandi. 1971 The Birds of the Lowlands of Bwamba 23 Caprimulgus tristigma tristigma Riippell. The freckled nightjar was added to the Bwamba avifauna by Ridley, Percy, and Percy (1953: 163), who collected one on the road near Mongiro. Caprimulgus bate si Sharpe. Two specimens, the first to be taken in Uganda, were collected in Bwamba, one at Ntandi, 2300 feet, June 22, and one at Nkarara, 2200 feet, July 3; both were females in non-breeding condition. These two specimens were discussed in an earlier paper (Friedmann and Williams, 1968: 15). Still earlier, Williams (1951: 111) saw what he thought was this nightjar at Ntotoro, March 18, 1948. Caprimulgus fossii fossii Hartlaub. v. S. 1949: 41. Macrodypteryx vexillarius (Gould), v. S. 1949: 41. Family Apodidae Apus melba maximus (O. -Grant), v. S. 1949: 41. Apus aequatorialis aequatorialis (Von Muller), v. S. 1949: 41. Cypsiurus parvus myochrous (Reichenow). Palm swifts were found at Nkarara, 2200 feet, where seven specimens, all in non-breeding state, were taken on June 27 and 28, and at Ntandi, 2300 feet, where one male with large testes was collected December 6; bill black; iris dark brown; feet dark grayish brown; weight 12 to 15 grams; stomach contents small flying ants and other insect fragments. Also recorded from Bwamba by the van Somerens (1949: 41), and by Ridley, Percy, and Percy (1953: 163). Chaetura sabini Gray. Three examples, one male, two females, were taken at Ntandi, 2300 feet, October 26 and December 4; all in non-breeding state; bill black; iris dark brown; feet grayish washed with violet; weights 15.5 to 17 grams. The van Somerens (1949: 42) had only sight records by Dr. Haddow. Our present specimens have wing lengths of 122 and 123 mm, and thereby support White’s conclusion (1965: 215) that the supposedly short-winged race ogowensis cannot be maintained. Recently, in discussing a west Uganda specimen from the Kibale forest, we (1968: 16) did find it to have a very short wing, which seemed to support the recognition of ogowensis. This conclusion is certainly upset by the present Bwamba specimens, even though examination again of the Kibale bird confirms its short wing length. Keith (1968: 7) saw this swift quite frequently in the Bwamba forest in July, 1963. Chaetura ussheri sharpei Neumann. The mottled-throated spine-tail is reported as ranging east to Bwamba by White (1965: 215), probably on the basis of the records by Williams 24 Contributions in Science No. 211 (1951: 108), the only ones to date. We have (1968: 16) reported on examples from the Kibale forest, the only other west Ugandan area from which this swift was then known. Family Coliidae Colius striatus kiwuensis Reichenow. This coly does not occur in the forest, but is common on its fringes. We have specimens from Bundimusuba, 3000 feet, November 22; Ntandi, 2300 feet, July 16; and Bwamba valley, April 23. Birds taken in all three months showed enlarged gonads. Also reported from Bwamba by the van Somerens (1949: 42). Family Trogonidae Apaloderma narina brachyurum Chapin. Two examples of the narina trogon were taken at Ntandi, 2300 feet, December 4 and 9, respectively; both were males in non-breeding state; bill pale yellowish gray, pale yellow on the basal half; iris dark brown; bare skin yellowish apple green with a pale blue wash around the eyes and the gape; feet dull pinkish dark brown; weight 70.5 and 72.5 grams. The van Somerens (1949: 42) found this bird to be scarce (“not much in evidence”) although they did collect one at Sara, and noted that Haddow had reported it at Mongiro. Family Alcedinidae Ceryle maxima (Pallas), v. S. 1949: 33. Ceryle rudis rudis (Linnaeus), v. S. 1949: 33. Alcedo quadribrachys guentheri (Sharpe). Although the van Somerens (1949: 33) did not meet with this kingfisher along any of the Bwamba streams, our collectors found it to be quite numer- ous at Ntandi, 2300 feet, where ten specimens were collected in June, July, November, and December, and at Rwansenge, 2200 feet, where two others were taken in July. Some of the June and July birds had enlarged gonads; the November and December ones did not; bill black, whitish at tip; iris blackish brown; feet pale flesh color; weights 25.5 to 39 grams. One of these shining blue kingfishers was seen at Ntandi, perched on a thin dead root sticking out of the mud bank of a small forest stream; it was quite fearless and permitted the collector to approach within eight yards. Alcedo leucogaster leopoldi (Dubois) The use of mist nets has caused a complete change in our estimate of the status of the white-bellied kingfisher. The work of the van Somerens was done without these nets and it is not surprising that they failed to obtain any specimens. The species was first recorded from Bwamba by Williams (1957: 159-160) and later by Friedmann (1966: 22-23) on a single example apiece; 1971 The Birds of the Lowlands of Bwamba 25 and by Keith (1968: 8). We now have 13 additional specimens from Bwamba, which show that it is common there: Ntandi, 2300 feet, June, July, Novem- ber and December; Nkarara, 2200 feet, July; mostly in non-breeding state; weights 12 to 16 grams. The bulk of this series has been discussed in an earlier paper by the senior author (1969: 1-6). Ceyx picta picta (Boddaert). Common in Bwamba; we have 13 specimens from Nkarara, 2200 feet, June and July; Ntandi, 2300 feet, July, November, December; Rwansenge, 2200 feet, July; mostly in non-breeding gonadal state; weights 11 to 12.5 grams. The pigmy kingfisher had been recorded from Bwamba by the van Somerens (1949: 33) earlier. Ceyx lecontei (Cassin). The red-headed pigmy kingfisher is common along the Bwamba forested streams; we have 14 examples from Ntandi, 2300 feet, July, October, Novem- ber, December; Mongiro, November; Nkarara, July; and Rwansenge, July; all with no or only little sign of gonadal enlargement; weights 11 to 12 grams. Previously reported from Bwamba by the van Somerens (1949: 33) as well. Halcyon senegalensis (Linnaeus). The red and black-billed kingfisher is common in Bwamba, as was pre- viously noted by the van Somerens (1949: 34). We have 11 specimens from Ntandi, 2300 feet, June, July, November, December; Mongiro, 2300 feet, November; Nyankiro, 3000 feet, November; Nkarara, 2200 feet, July; and “Bwamba Forest”, April. Only the April specimens showed enlarged gonads; their weights ranged from 41 to 62 grams. Halcyon malimbica malimbica (Shaw). The van Somerens (1949: 34) considered this species less common in the Bwamba than H. senegalensis, but the experience of our collectors was otherwise. They obtained 15 specimens at Nkarara, 2200 feet, June, July; Ntandi, 2300 feet, July, October, December; Mongiro, 2300 feet, November; and Bundimusuba, 3000 feet, November. One June and one November speci- men had enlarged gonads; the rest of the series did not; weights 77 to 88 grams. Halcyon badia Verreaux. Two specimens of the chestnut-backed kingfisher were collected, an adult male, in non-breeding state, at Ntotoro River, west of Mongiro, 2250 feet, November 5, and an unsexed example on the N. Humia River, 2300 feet, December 9; bill brownish blood red; iris dark brown; eye ring red; feet dark reddish brown; weights 61.5 to 64 grams. The van Somerens (1949: 34) con- sidered it a scarce bird in Bwamba, and only saw it once. The bird collected west of Mongiro was catching insects on the wing about 20 feet in the air. 26 Contributions in Science No. 211 and was noted as making a purring sound with its wings; the other was about 12 feet up in a tree when seen. Halcyon chelicuti chelicuti (Stanley), v. S. 1949: 35. Halcyon leucocephala leucocephala (Muller). The gray-headed kingfisher was found and collected, one specimen at Mongiro, 2300 feet, November 12, and one at Ntandi, 2300 feet, December 6; weights 41, 42 grams, respectively. Neither was in breeding state. These two suggest that the species is more widely distributed in Bwamba than the van Somerens (1949: 35) suggested when they reported it as restricted to the northern end of Bwamba, toward Sempaya. As they noted, Haddow had recorded it throughout the year, in the open scrubby growth outside the forest. Family Meropidae Merops apiaster Linnaeus. Recorded from Bwamba by Ridley, Percy, and Percy (1953: 163), who noted flocks of 60 or more of these birds in and around the native village at Hakitengya; not in the forest. This bee-eater is, of course, only a winter visitor in the Bwamba area. Merops superciliosus superciliosus Linnaeus. Williams (1951: 107) saw several blue-checked bee-eaters and obtained one at Bundibugo, in March 1948. One male, taken at Korongo, Semliki River, by our collectors, July 15, is the only other Bwamba record known to us. Merops albicollis Vieillot. Very common in the open country outside the forest in Bwamba. Our collectors obtained specimens at Ntandi, 2300 feet, November 4 to 18; Mon- giro, 2300 feet, November 5; Bwamba valley, April 22 to 23. Also reported by the van Somerens (1949: 35). Our examples had weights of from 22 to 25 grams. Merops pusillus meridionalis (Sharpe). The van Somerens (1949: 35) considered this little bee-eater as not very common in the open country of Bwamba. Our collectors met with it fre- quently, and obtained specimens in the Bwamba valley, April 23; Rwansenge, 2200 feet, July 6 to 13; Ntandi, 2300 feet, July 12. Merops lafresnayii oreobates (Sharpe), v. S. 1945: 35. Merops gularis australis (Reichenow). This central African bee-eater is numerous in Bwamba. We have six specimens from Ntandi, 2300 feet, June 21 to 24 and November 10 to 12; all in non-breeding state; bill black; iris red-brown; feet black; weight 29 to 34 grams; stomach contents beetles. Also reported from Bwamba by the van Somerens (1949: 35), and by Williams (1951: 108), and by Keith (1968: 9). 1971 The Birds of the Lowlands of Bwamba 27 Family Coraciidae Eurystomus glaucurus afer (Latham). First recorded from Bwamba by the van Somerens (1949: 32), this broad- billed roller has been found to be common and widely distributed there. We have specimens from Nkarara, 2200 feet, June 27; Humia, 2300 feet, July 11; Rwansenge, 2200 feet, July 15; and Nyankiro, 3000 feet, November 25; all our examples were in non-breeding state when collected; weights 116 to 119 grams. Eurystomus gularis neglectus Neumann. The blue-throated roller seems to be less numerous in Bwamba than the preceding species. Our collectors met with it only at Ntandi, 2300 feet, where an adult female in non-breeding condition was obtained on June 13. The van Somerens (1949: 32) also found this bird in Bwamba, as did Williams (1951: 107). Family Upupidae Upupa epops epops Linnaeus. The European hoopoe was added to the Bwamba list by Ridley, Percy, and Percy (1953: 163). The bird is, of course, not in the forest, and is only a winter visitor from Europe. Family Phoeniculidae Phoeniculus bollei jacksoni (Sharpe). Specimens were obtained at Nyankiro, 3000 feet, November 24, and at Bundimusuba, 3200 feet, November 24 and 27; all with small gonads; bill dull pinkish crimson, horn white at the tip; iris dark brown; bare skin dull red; feet red; weight 48 to 59 grams. The white-headed kakelaar was also reported from Bwamba by the van Somerens (1949: 39-40). Phoeniculus castaneiceps brunneiceps (Sharpe). The distribution of the chestnut-headed kakelaar seems to be somewhat local and unpredictable in the forests of western Uganda. Thus, the van Somerens (1949: 40) and Williams (1951: 108), found it in Bwamba while our collectors did not, but obtained a good series in the Bugoma Forest, at only a little greater elevation. Phoeniculus cyanomelas schalowi (Neumann), v. S. 1949: 40. Family Bucerotidae Tockus camurus camurus Cassin. The red-billed dwarf hornbill is fairly common in the Bwamba forest. Our collectors obtained specimens there on April 22 and 23; at Ntandi, 2300 feet, June, October, November, and December; and at Nkarara, 2200 feet, July 1. None of the specimens were in breeding condition; bill red with black tips; iris cream; feet brownish black; weight 96 to 104 grams for the females, 28 Contributions in Science No. 211 122 grams for one male. Parties of three birds were seen feeding in the tree tops, and the collectors noted the birds gave a soft peau note repeated several times. The van Somerens (1949: 38-39) considered the call of this hornbill as similar to the whimpering of a very young puppy. Tockus hartlaubi granti (Hartert). The Congo dwarf hornbill was first reported from Bwamba by the van Somerens (1949: 38). Our collectors obtained further specimens at Bundi- keke, 3000 feet, July 11— a male with enlarged testes; at Ntandi, 2300 feet, November 2; and at Kisuba, 2500 feet, November 2. Bwamba is the only Uganda locality for this hornbill. Tockus fasciatus fasciatus (Shaw). A common hornbill in Bwamba, especially in the open portions of the forest and in the clearings adjacent to it. Our collectors obtained examples in Bwamba, April 22 and 23; at Ntandi, 2300 feet, in July and November; at Nkarara, 2200 feet, in June; and at Bundimusuba, 3000 feet, in November; all in non-breeding gonadal condition; weights 227 to 270 grams. The van Somerens (1949: 39) had previously found this bird to be very common in the Bwamba area. Tockus alboterminalis geloensis (Neumann), v. S. 1949: 39. (Not in the true Bwamba lowlands; records only from the Sempaya hills above the hot springs.) Tropicranus albocristatus cassini (Finsch). Listed from Bwamba by White (1965: 249-250), probably on the basis of the van Somerens’ 1949 record (p. 39). Ceratogymna atrata (Temminck). The black-wattled hornbill is fairly common in the Bwamba Forest. We have specimens from “Bwamba”, April 23; Mongiro, 2300 feet, November 1; Nkarara, 2200 feet, July 2; and Bundimusuba, 3000 feet, July 5. The April specimens have been discussed by the senior author in an earlier paper (1966: 25); the July 2 bird from Nkarara had enlarged testes; the other examples were in non-breeding state; bill dark horn gray, the lower basal half washed with brown, and the tip of the casque pale brown; iris reddish mauve; bare skin pale blue around the eye, the throat black with the edges of the wattle pale blue; feet black; weight 1600 grams. The species was seen in palm swamp forest, and occasionally in small parties flying overhead. The birds were heard giving a repeated croaking wawoo sound. This hornbill had been reported from Bwamba by the van Somerens (1949: 38). Bycanistes fistulator duboisi Sclater. First recorded from Bwamba by the van Somerens (1949: 36) who found it not uncommon there. Our collectors procured further specimens in 1971 The Birds of the Lowlands of Bwamba 29 “Bwamba”, April 23, and Ntandi, 2300 feet, and at Nkarara, 2200 feet, in late June; all specimens in non-breeding state; bill ivory becoming grayer at the tip; iris dark brown; feet gray, the scales black; weights 413 to 485 grams; stomach contents large seeds and fruit remains. Keith (1968: 9) writes that this hornbill is common in Bwamba. Bycanistes cylindricus albotobialis (Cabanis and Reichenow). At Bundimusuba, 3000 feet, July 8, one unsexed example of the white- thighed hornbill was collected. Previously the van Somerens (1949: 37) had recorded it, with extensive field notes, from Bwamba, the first Uganda locality for the species. Since then it has been found there by Ridley, Percy, and Percy (1953: 163) as well, and also in the Budongo Forest (Friedmann, 1966: 25). Bycanistes subcylindricus subquadratus Cabanis. The black and white casqued hornbill is common in the Bwamba Forest. We have specimens from Ntandi, 2300 feet, June 22, and from Bundimusuba, 3000 feet, July 8. The van Somerens (1949: 36-37) were the first to record the species from Bwamba. Family Capitonidae Lybius bidentatus aequatorialis (Sharpe). The double-toothed barbet must be quite local and sporadic in its dis- tribution in Bwamba as the 1963 field party of the Knudsen-Machris expedi- tion obtained three examples there on April 23 and 24, but neither our 1967 nor our 1968-69 collectors met with it at all. The van Somerens (1949: 42) apparently knew of it in Bwamba only through Dr. Haddow’s observations and specimens. Lybius hirsutus ansorgii (Shelley). The Uganda hairy-breasted barbet was collected at “Bwamba” in April; Ntandi, 2300 feet, in June, July, November, and December; at Nkarara, 2200 feet, in July; and at Mongiro, 2300 feet, in November. All the specimens were in non-breeding state; weights 51 to 55 grams. The van Somerens (1949: 42) found this species at other Bwamba sites as well. Gymnobucco bonapartei cinereiceps Sharpe. This brown barbet was met with by our field parties only at Bundimu- suba, 3000 feet, in July 1967, and again in November 1968. A total of nine specimens, including six females and three males, were collected, only one of which, a male taken on July 11, was found to show gonadal enlargement; weights varied from 50 to 65 grams. The van Somerens (1949: 42) reported the species from other Bwamba localities as well. 30 Contributions in Science No. 211 Pogoniulus scolopaceus aloysii (Salvadori). One female was taken at Mongiro, 2300 feet, November 3; a bird in non- breeding state; weight 14 grams. The Uganda speckled tinker-bird would seem to be uncommon in Bwamba as this was the only time our collectors met with it on three separate trips, and as the van Somerens did not include it in their list. Pogoniulus bilineatus leucolaema (Verreaux). The yellow-rumped tinker-bird is common in the Bwamba forest, as was first reported by the van Somerens (1949: 43). Our collectors obtained addi- tional specimens at Ntandi, 2300 feet, July 10 to 17, and October 31 to November 20; and at Bundimusuba, 3000 feet, November 22 to 29. A few of the July and of the November examples had enlarged gonads; the majority did not; weights ranged from 10 to 14 grams. Pogoniulus subsulphureus fiavimentum (Verreaux). The fact that our collectors obtained a series of 16 examples of this tinker-bird at two localities in Bwamba suggests that it must be a common bird there. It is, therefore, surprising that the large collections amassed by the van Somerens years ago failed to include this species. Our present examples are from Ntandi, 2300 feet, July, October, and November; and from Bundi- musuba, 3000 feet, November; all were in non-breeding state; their weights varied from 9 to 12 grams. Pogoniulus atro-flavus (Sparrman). The red-rumped tinker-bird was added to the Bwamba avifaunal list, and, indeed to that of Uganda, when two specimens were taken at Ntandi, 2300 feet, July 3 and 12, 1967, as reported by us in an earlier paper (1968: 16-17), where further details were given concerning them, and also a sight record by R. W. Smart (Keith, 1968: 9). Trachyphonus purpuratus elgonensis (Sharpe). The Elgon yellow-billed barbet was found in a number of places in the Bwamba; at “Bwamba Forest”, April 21; at Ntandi, 2300 feet, June 28 and October 26 to November 17; and at Bundimusuba, 3000 feet, July 6 to 12. None of our specimens showed gonadal enlargement; their weights varied from 71 to 85 grams; bill yellow; iris red; bare skin yellow, feet pale olive yellow with gray-black scales. This barbet had also been recorded from Bwamba by Williams (1951; 108) and by Ridley, Percy, and Percy (1953: 163). Family Indicatoridae Indicator maculatus stictithorax Reichenow. Bwamba is the easternmost area and the only place in Uganda where the spotted honey-guide is known to occur. The van Somerens were the first to report it from there (1949: 43) thereby adding it to the Uganda avifauna. 1971 The Birds of the Lowlands of Bwamba 31 Williams (1951: 108) collected a male in full breeding condition in thick forest near Mongiro on March 22, 1948, and the Knudsen-Machris expedi- tion obtained a further example in the Bwamba forest on April 23, 1963. These three records of a single bird each made it seem that this honey-guide was probably uncommon there, but our field teams in 1967 and 1968-1969 collected a long series at Ntandi, 2300 feet, in June, July, October, November, and December; Mongiro, 2300 feet, in November; and Bundimusuba, 3000 feet, in November. Most of the specimens showed little or no gonadal enlarge- ment, but females with swollen ovaries were taken in July and December. The weights of our specimens varied from 39.5 to 51 grams; bill blackish, browner at base of mandible; iris dark brown; bare eye skin olive gray; feet gray with olive scales. In our surveys in 1967 and 1968-1969, special attention was paid to the honey-guides with results that clearly demonstrate the misleading picture that may easily result from general, more or less random, collecting. This is demonstrated in the present species, for which, as mentioned above, there were only three single records from 1944 to 1963, while in our two surveys no fewer than 24 specimens were obtained. In some of the other honey- guides— icator minor riggenbachi, 1. conirostris conirostris, and 7. exilis pachyrhynchus— the increase in specimen data was even more extensive, and there is no reason to think that any of these species have become more numer- ous in the Bwamba area since 1963. The van Somerens’ 1949 basic report listed two species of honey-guides from Bwamba; we now list nine. The great increase in specimen and species records is due in part to the use of mist nets placed near opened wild bees’ nests. It was necessary to procure ample mate- rial of some of the species because of the difficulty of distinguishing them in the field, and because of the need to elucidate the status of such sibling species as 1. minor and /. conirostris, and to determine whether I. exilis was really all one species or included odd specimens of pumilio and possibly of will cock si and meliphilus. Indicator variegatus Lesson. The scaly-throated honey-guide is sympatric with I. maculatus in Bwamba; specimens of both were taken in the same localities and at the same time. The data on variegatus are as follows: Ntandi, 2300 feet, June 24 and October 26, one on each date; Mongiro, 2300 feet, November 4 to 7, three specimens; all with small gonads; weights ranging from 47 to 55 grams. Indicator minor riggenbachi Zedlitz. This race of the lesser honey-guide and the thick-billed honey-guide, 7. conirostris, are sympatric in Bwamba, where a most informative series of both were collected. These birds, 25 specimens of riggenbachi and 15 of conirostris, have been discussed in detail in another paper by the senior author (in press, Proc. 3rd Pan-African Ornithological Congress), and all that need 32 Contributions in Science No. 211 be stated here is that the Bwamba population of /. minor is of the subspecies riggenbachi, known earlier from the Ituri forest on the other side of the Semliki, but not reported in print from Uganda; Ntandi, 2300 feet, October to December; Mongiro, 2300 feet, November; and Bundimusuba, 3000 feet, November; most of the specimens had small gonads, but three had enlarged ones; weights ranged from 25 to 31 grams. Indicator conirostris conirostris (Cassin). Specimens of the thick-billed honey-guide were taken at Bundimusuba, 3000 feet, July 1, and November 25 and 26; at Ntandi, 2300 feet, October to December; and at Mongiro, 2300 feet, November 8 to 13. Two females taken in late November at Bundimusuba had enlarged ovaries; all the other speci- mens were inmon-breeding plumage. These birds showed a greater range in weight than the preceding species, 25.5 to 38 grams, as compared with 25 to 31 grams in a longer series of /. minor riggenbachi. Like the latter group, the present series are discussed in detail in the paper referred to in the account of that taxon. Suffice it to say that the bulky evidence of sympatry of the two forms in Bwamba and their distinctness there as contrasted with the inter- gradations found in populations in parts of western Africa, leads to the con- clusion that conirostris first diverged from minor in western Africa and that the perplexing population still found in places there is a result of their ancient mixture prior to the achievement of specific status, while in Bwamba coni- rostris and minor came together again after their divergence had attained specific rank with the result that there they act as sympatric, but distinct, species. Indicator exilis pachyrhynchus (Heuglin). This race of the least gray-breasted honey-guide is very common in the Bwamba forest. A very long series of 68 specimens were collected at Ntandi, 2300 feet, in July, October, November, and December; at Bundimusuba, 3000 feet, in July and November; at Mongiro, 2300 feet, November 5 to 15; and at Humia, 2400 feet, undated. The reason for collecting so many speci- mens was that there was always a possibility that one of them might turn out to be one of the sibling species closely resembling exilis. Of the 68 examples, 47 are adults with white loreal streaks and whitish chins and black gular streaks, while 21 are immature and lack these characters, and have less dusky on the tips of the rectrices and have these feathers a little more pointed at the tip. These young birds vary more than the adults in their ventral coloration, the extremes being almost as different in degree of darkness as are /. coni- rostris and I. minor riggenbachi. Most of the specimens had weights of from 15 to 19.5 grams, but one immature female weighed only 12 grams, which is less than some /. pumilio, a still smaller bird! The soft parts of this series were noted as follows: bill black, the base of the mandible pinkish gray; iris dark brown; bare skin dark gray; feet olive gray, stomach contents insect frag- 1971 The Birds of the Lowlands of Bwamba 33 ments and flakes of beeswax. Bwamba specimens average slightly more green- ish, less grayish below than a series from the Impenetrable forest, Kigezi, but the difference is small and is not completely constant. Lest it seem shocking that so long a series of 7. exilis was collected, especially to readers with only a casual acquaintance with this species in life, whose limited field experience might suggest to them that the species is far from abundant, we may point out an experience reported by one of our col- lectors. He wrote us that within a matter of six hours no less than 1 8 honey- guides were seen (12 collected) on a single tree. Of the 12 birds collected, eight were /. exilis. Furthermore, it may be noted that much of the forest where this series was collected is now (February 1970) felled and given over to agriculture, and exilis is absent now, although common only a short time ago. Indicator pumilio Chapin. As we reported in another paper (1968: 18-19), one example of this smallest of the Indicators was taken at Ntandi, 2300 feet, July 13, 1967. A reexamination of the specimen does not cause us to question the identifica- tion, even though the occurrence of the species at so low an elevation is still surprising. Melichneutes robustus (Bates). The lyre-tailed honey-guide may be added to the Bwamba avifauna on the basis of the fact that its distinctive notes have been heard there by Haddow, whose identification was accepted by Chapin. The bird has not been collected in Bwamba as yet, but inasmuch as it does occur in the Iturbi Forest, its presence east of the Semliki in Bwamba is not surprising. (Friedmann, 1955: 237). Melignomon zenkeri Reichenow. Only one Bwamba record; Bundimusuba, 3000 feet, July 9, 1967, adult breeding female with enlarged ovary. This specimen has already been dis- cussed by us (1968: 19-20), and also by the senior author (1968) in greater detail. Prodotiscus insignis insignis (Cassin). One adult male, in non-breeding state, was collected at the forest edge at Ntandi, 2300 feet, November 4, 1968; bill black; iris dark brown; feet gray; weight 11 grams. The van Somerens (1949: 43) had noted this small honey-guide at Ntandi as well as at other Bwamba localities. It apparently is not uncommon around the perimeter of the forested areas. Family Picidae V erreauxia africana (Verreaux). The green piculet is common in Bwamba, which is its easternmost known locus of occurrence. The first two specimens, taken on April 20 and 21, 1963, 34 Contributions in Science No. 211 were included in an earlier paper (Friedmann, 1966: 28), but since then a notable series of twelve more specimens have been collected at Ntandi, 2300 feet, June 16 to July 6, and November 2; and at Nkarara, 2200 feet, June 28; bill dark horn gray; iris crimson; bare skin around eye dull maroon; feet dull maroon; weights 7.5 to 9.5 grams. Three of the June examples from Ntandi had enlarged gonads; the other specimens had small ones. Keith (1968: 9) has also noted this bird in Bwamba, on the basis of Smart’s sight records. Campethera cailliautii permista (Reichenow). When the van Somerens (1949: 44) recorded this green-backed wood- pecker from Bwamba it was a new bird to the Uganda list as well. Our collec- tors obtained additional specimens in the Bwamba forest, April 24, one female in breeding condition; Ntandi, 2300 feet, July 16, one non-breeding female, December 8, one non-breeding male; bill horn gray, somewhat grayer on the mandible; iris dark brown; feet olive; weights 49 to 52 grams; stomach con- tents small beetle larvae. Campethera caroli (Malherbe) . The brown-eared woodpecker is common in Bwamba; we have 30 specimens from Ntandi, 2300 feet, June, July, October, November and December; Nkarara, 2200 feet, June and July; Bundimusuba, 3000 feet, November; Mongiro, 2300 feet, November; and “Bwamba”, April. All our birds had small gonads; bill dull gray; iris dark brown; feet olive; weights 55 to 66 grams (two specimens were marked 44 grams) ; stomach contents ants, mantis, insect fragments. The van Somerens ( 1949: 44) also noted this wood- pecker in several places in Bwamba; as did Williams ( 1951 : 108). Campethera nivosa herberti (Alexander). This woodpecker was found to be not uncommon, and specimens were obtained at Ntandi, 2300 feet, in July, November, and December; at Bundi- musuba, 3000 feet, in July; and at Rwansenge in July. All of our nine exam- ples were in non-breeding condition; bill dark horn gray, paler below; iris red-brown; feet olive; weights 31 to 37 grams; stomach contents ants and other insect fragments. This species was not represented in the van Somerens’ collection, but was reported from Bwamba by Ridley, Percy, and Percy (1953: 163), and earlier by Williams ( 1951 : 109). This woodpecker often feeds low down in the forest undergrowth; several were taken in mist nets set at ground level. Dendropicos fuscescens lepidus (Cabanis and Heine), v. S. 1949: 45. Dendropicos poecilolaemus (Reichenow). The Uganda spotted woodpecker is a bird of the open, but wooded clearings, rather than of the dense forest. Our collectors obtained one speci- men at Bundimusuba, 3000 feet, July 8, and another at Ntandi, 2300 feet, November 3; both in non-breeding state; bill dull silvery gray on maxilla, paler 1971 The Birds of the Lowlands of Bwamba 35 on mandible; iris dark red-brown; feet olive gray; weights 25 to 29 grams. This species was also noted from Bwamba by the van Somerens ( 1 949 : 45 ) . Dendropicos gabonensis gabonensis (Verreaux). Two specimens taken at Ntandi, June 15 and 23, are the only specimen records for Bwamba, and for Uganda. They were discussed in our earlier paper (1968: 20). Mesopicos goertae centralis Reichenow. A bird of the open woodlands, not of the dense forest, the Uganda gray woodpecker was met with by our collectors but a single time, at Ntandi, 2300 feet, November 20, when they collected one male, in non-breeding state; bill blackish gray, paler at the base of the mandible; iris dark brown; feet dark gray; weight 51 grams; stomach contents small beetles. Also obtained in Bwamba by the van Somerens (1949: 44). Mesopicos xantholophus (Hargitt). The golden-crowned woodpecker is common and widely distributed in Bwamba: specimens were taken at Ntandi, 2300 feet, June, July, and Decem- ber; Bundimusuba, 3000 feet, July and November; and Nyankiro, 3000 feet, in November; all in non-breeding state; bill gray, paler below; iris dark red; feet olive gray; weights 59 to 66 grams; stomach contents insect fragments and, in one case, one spider. Also noted from Bwamba by the van Somerens (1949: 44). Family Eurylaemidae Smithornis rufolateralis budongoensis van Someren. The red-sided broadbill is fairly common in Bwamba; specimens were taken at Ntandi, 2300 feet, December 7; at Nkarara, 2200 feet, June 29 and July 2; Bundimusuba, 3000 feet, July 16; and Rwansenge, 2200 feet, July 14; two of the July birds having enlarged gonads, the rest small ones; maxilla blackish horn, mandible whitish; iris dark horn; feet pale brown tinged with olive, the toes greener; weight 19.5 to 29 grams; stomach contents ants, small beetles, and other insect remains. At Nkarara and at Bundimusuba individuals were noted flying in small circles above their perches, calling loudly as they did so. The call is a loud brrr note. As early as 1951 Williams (112) pub- lished a sight record of this broadbill from Bwamba. Family Pittidae Pitta reichenowi Madarasz. v. S. 1949: 45. Family Hirundinidae Riparia riparia riparis (Linnaeus), v. S. 1949: 79. Ridley, Percy, and Percy (1953: 164) also noted sand martins in Bwamba. 36 Contributions in Science No. 211 Riparia cincta cincta (Boddaert). v. S. 1949: 80. Hirundo rustica riistica Linnaeus. Ridley, Percy, and Percy (1953: 164) reported very large flocks of barn swallows over the Mongiro swamp in October. The van Somerens (1949: 79) had earlier recorded it from the Bwamba area on the strength of Haddow’s observations. Hirundo angolensis Bocage. One male, in non-breeding state, was collected at Ntandi, 2300 feet, July 19; weight 17 grams. This was the only time our collectors met with this swallow. The van Somerens (1949: 79) found the species breeding in Bwamba from April to August. Hirundo senegalensis saturatior Bannerman. The mosque swallow was met with at Ntandi, 2300 feet, where two males, in non-breeding condition, were collected June 2; weights 48 to 50 grams. The van Somerens (1949: 79) had earlier found this species com- monly throughout the more open areas of Bwamba. Hirundo abyssinica unitatis Sclater and Praed. The lesser striped swallow was found to be common in Bwamba by the van Somerens (1949: 79). Our field party obtained three examples, two of them with enlarged gonads, June 15 to July 9, at Ntandi, 2300 feet; weights 16.5 to 19 grams. Cecropis daurica emini (Reichenow). The red-rumped swallow was reported from Bwamba by Ridley, Percy, and Percy (1953: 164), at Hakitengya. Cecropis semirufa gordoni (Jardine). Our collectors obtained two examples at Korongo-Semliki, 2200 feet, July 14; weights 20 to 22 mm.; the Knudsen-Machris expedition collected one in Bwamba, April 23, 1963. Delichon urbica urbica (Linnaeus), v. S., 1949: 80. Psalidoprocne pristoptera ruwenzori Chapin, v. S., 1949: 80. Psalidoprocne albiceps albiceps Sclater. Three examples were taken at Ntandi, 2300 feet, July 7 to 19, and November 30; one of the July birds was a male with large testes, the other two were non-breeding females; weights 1 1 to 12 grams. The van Somerens (1949: 80) found the white-headed rough-winged swallow to be common in the open country in Bwamba. Family Motacillidae Motacilla flava Linnaeus, v. S. 1949: 46. 1971 The Birds of the Lowlands of Bwamba 37 Motacilla clam torrentium Ticehurst. v. S. 1949: 45. Motacilla alba vidua Sundevall. Regular in the open country of Bwamba, whence reported by the van Somerens (1949: 45); Ridley, Percy, and Percy (1953: 163); and by our collectors who obtained several males at Ntandi, 2300 feet, June 24 to July 14, and November 16 to December 2; and one at Mongiro, 2300 feet, on November 8. Anthus trivialis trivialis (Linnaeus). One example of this European migrant was taken at Nyankiro, 3000 feet, November 24; obviously not in the forested country. Family Timaliidae Trichastoma fulvescens ugandae (van Someren). Very common in Bwamba, from where we have a long series of speci- mens, taken in many places during April, June, July, October, November, and December. The van Somerens (1949: 46) reported this thrush babbler as “not uncommon in the undergrowth of the forest but . . . hard to detect . . .”, so it would seem that our series must be attributed to the use of mist nets. Although most of our specimens were in non-breeding state, one June bird had large gonads; our examples weighed from 26 to 37 grams (one male was said to weigh only 17 grams, but this may have been an error in writing on the label). Trichastoma rufipenne rufipenne Sharpe. This species seems to be uncommon in Bwamba. The van Somerens (1949: 47) encountered it but seldom; Williams (1951: 109) obtained one at Ntotoro, March 19, 1948. In a report on the Knudsen-Machris collection, the senior author (1966: 29) reported three Bwamba examples, but these we now have reidentified as T. albipectus. Our collectors in 1967 and in 1968-1969, obtained many examples of that species, but none that may be considered rufipenne. Trichastoma albipectus (Reichenow). Common in Bwamba; we have many specimens from Ntandi, 2300 feet; Nkarara, 2200 feet; and Bundimusuba, 3000 feet; weights 28 to 34 grams. The fact that this species was not obtained by the van Somerens can only be assumed to have been due to the absence at that time of mist nets as col- lecting aids. Some of our July birds had enlarged gonads; others did not; none of the October or November specimens showed any gonadal swelling. Phyllanthus atripennis bohndorffi (Sharpe). The chestnut babbler was collected in the “Bwamba Forest” April 23; at Bundimusuba, 3000 feet, in July and November; and at Ntandi, 2300 feet, October 28 to November 5; one specimen, October 30, was in breeding con- 38 Contributions in Science No. 211 dition; the ten others had small gonads; bill pale yellow horn; iris deep red brown; lower eye lid recorded as blackish in one case, yellow in another!; feet pale greenish gray, the claws whitish; weights 67 to 99 grams; stomach con- tents beetles and other insect remains. The three April specimens are referred to in an earlier paper by the senior author (1966: 29). Bwamba appears to be the only area in Uganda where this bird is known to occur. It must be a lowland forest species entirely as it is not present in extensive collections from the Kibale, Bugoma, or Budongo areas, all of which are at higher elevations. Family Pycnonotidae Pycnonotus barbatus minor Heuglin. Very common in Bwamba; our collectors obtained specimens at Ntandi, 2300 feet, July, October, November, December; Nkarara, 2200 feet, July. The van Somerens ( 1949: 48) also found it numerous in the more open for- ests and in the areas adjacent to the forest cultivated by the natives. None of our specimens had enlarged gonads, although the van Somerens reported eggs for most months of the year. Our specimens had body weights of 39 to 41 grams. In our accounts of this species, and of all the bulbuls, we follow the nomenclature of Rand ( 1960, in Peters, 9: 221-300). Pycnonotus virens holochlorus van Someren. This green bulbul is very numerous in Bwamba, where the experience of our field teams agreed with that of the van Somerens (1949: 53) who found it in the taller undergrowth throughout the forested area. Because of the difficulty of distinguishing this species from some of the other bulbuls in life many specimens were taken, as follows : Bwamba Forest, April 20 to 23; Ntandi, 2300 feet, June, July, October, November, and December; Bundi- musuba, 3000 feet, July and November; Mongiro, 2300 feet, November. Birds with enlarged gonads were taken in July and November, but others taken at the same time showed no gonadal swelling; weights 18 to 26 grams. Pycnonotus gracilis ugandae (van Someren). Another common bulbul in Bwamba; we have a long series of specimens from Ntandi, 2300 feet, June, July, November, and December; Bundimusuba, 3000 feet, July and November; and Bundikeke, 3000 feet, November. Only a single specimen, taken July 18, had enlarged gonads; weights of our 22 specimens ranged from 18 to 26 grams; bill black; iris dark brown; feet olive to olive gray; stomach contents insect remains and small fruits and green berries. Pycnonotus curvirostris curvirostris (Cassin). The sombre bulbul must be somewhat erratic, although common, in Bwamba, as our collectors obtained ten specimens at three localities : Ntandi, 2300 feet, Nkarara, 2200 feet, and Bundimusuba, 3000 feet, in 1967, but 1971 The Birds of the Lowlands of Bwamba 39 failed to meet with the species in 1968-1969. Two of our July birds, from Ntandi, had enlarged gonads, agreeing with the van Somerens’ (1949; 52) July breeding record from Ntotoro. The Knudsen-Machris 1963 expedition collected this bulbul in Bwamba as well. Our specimens were noted as having the bill black in the males, dark grayish brown horn with paler tomial edges in the females; iris red-brown to dark brown; feet pale olive to olive gray; weights 19 to 29 grams. This species and P. gracilis are so very similar in appearance that one wonders what isolating mechanism keeps them apart. P. curvirostris is usually, but not always, larger (wings 74 to 84 mm.) than P. gracilis (65 to 74 mm.), and usually the former lacks pale grayish feathering on the upper and lower eyelids found, as a rule, in the latter, and also curvirostris averages less yellowish, more dusky olive, on the abdomen, but all these characters are not completely trenchant. Chapin (1953: 105, 107) did not know any way to distinguish the voices of the two. In the Congo forest Chapin found them together, “. . . and probably in equal numbers. The habits are the same . . .” Pycnonotus latirostris eugenius (Reichenow). The Uganda yellow-whiskered bulbul was found by our collectors to be common in the Bwamba lowland forests. It is therefore somewhat surprising that the van Somerens (1949: 53) found it only in the mountain forests of the Ruwenzori slopes at altitudes of from 6000 to 8500 feet. Our 24 speci- mens were taken as follows: “Bwamba Forest,” April 24; Ntandi, 2300 feet, October 28 to December 3, June 16 to July 12; Bundimusuba, 3000 feet, November 21, 22, July 6; and Nkarara, 2200 feet, July 1; weights 19 to 30 grams. One of our November birds had enlarged gonads; all the others showed no enlargement. The van Somerens reported breeding birds in April, July, and August. The soft parts of our series were noted as follows: bill dark brown horn to blackish with tomium yellowish to orange-brown; iris dark brown; feet yellowish horn color. Pycnonotus gracilirostris chagwensis (van Someren). The slender-billed greenbul was common and widespread in the higher branches of the trees in Bwamba, where our collectors obtained a dozen specimens at Ntandi, 2300 feet, November 8 to 16, July 13 to 18; Bundi- musuba, 3000 feet, July 20 and November 20; and Rwansenge, 2200 feet, July 14; weights 29 to 31 grams; bill black; iris red; feet black; stomach contents insect remains and small fruits. One of the November 20 birds had nest building material in its bill. Baeopogon indicator indicator (Verreaux). The white-tailed greenbul was collected at Ntandi, 2300 feet, November 10 to December 10, and June 17 to July 10; at Bundimusuba, 3000 feet, November 20 and July 12; and at Mongiro, 2300 feet, November 8; weights 37.5 to 52 grams. The fact that our collectors were able to obtain a dozen 40 Contributions in Science No. 211 examples at these places suggests that the species must be fairly common, but the van Somerens (1949: 52) noted very few, and, indeed they had no records for two years, 1944 and 1945. Our birds show a surprising variation in the width of the bill, the narrowest ones apparently not due to the collectors having tied them tightly; the narrowest one is only 3.5 mm broad at the base of the nares, while the broadest one is 7.5 mm or more than twice as wide. The collectors’ field notes on our specimens inform us that the bill is black; the iris varying from white to grayish white to pale brownish gray in males, from dark gray to dark brown in females; feet silvery gray to dark gray; stomach contents large black ants, beetles, and small fruits. Chlorocichia simplex (Hartlaub). This bulbul is less common than the preceding several species, and was not even recorded in the 1949 list of the two van Somerens. Our collectors obtained two examples at Ntandi, 2300 feet, one on November 24 and one on July 3, and two others at Nkarara, 2200 feet, June 28 and 29; weights 43 grams; bill black, the tomial edges grayish; iris dark brown; feet gray. Chlorocichia flavicollis pallidigula (Sharpe). This is another relatively uncommon species in Bwamba, and is an addition to the 1949 list of the two van Somerens. The Knudsen-Machris 1963 expedition obtained two males with enlarged gonads in the Bwamba Forest, April 20 and 23; our 1967 field team collected one at Rwansenge, July 6, while in 1968-1969 the species was not observed. Our specimens had the bill black; iris whitish to cream with a brownish or a pinkish tone; feet gray-brown to blackish; weights 37 to 54 grams; stomach contents insect fragments. Thescelocichla leucopleiira (Cassin). When the van Somerens (1949: 49) first collected the swamp palm bulbul in Bwamba it was a new bird for Uganda. As indicated in another paper (Friedmann, 1966: 31), the Knudsen-Machris expedition obtained four examples there April 20 to 23. In 1967 our field team collected one at Rwansenge, 2200 feet, July 6, and in 1968-1969 the species was not found at all. It would appear to be somewhat erratic and uncommon in the Bwamba area; weights of our birds ranged from 60 to 65 grams; bill black; iris whitish to cream with a pale pinkish tinge; feet gray-brown; stomach contents insect fragments. Phyllastrcphus scandens orientalis (Hartlaub). Not listed by the van Somerens, but noted from Bwamba by Rand (1958: 194; 1960: 264). The Knudsen-Machris 1963 expedition obtained several specimens on April 22 to 24 (Friedmann, 1966: 31); our subsequent field teams obtained a series of 18 others at Ntandi, 2300 feet, October 30 to December 9, and June 26 to July 15; at Mongiro 2300 feet, November 8; at 1971 The Birds of the Lowlands of Bwamba 41 Bumaga, 2300 feet, June 27; and at Nkarara, 2200 feet, on June 29. One of our July Ntandi birds had enlarged gonads, the others were in non-breeding state; weights 30.5 to 44 grams; bill blackish to pale gray horn, the mandible paler than the maxilla; iris creamy brown to deep sepia; feet pale gray; stomach contents insect fragments. The fact that our collectors Vvcre able to procure good numbers of this bird and that earlier Rand (1958: 194) had eight Bwamba specimens is evidence that the species is common there. Phyllastrephus hypochloris (Jackson). Ntandi, 2300 feet, December 1, 3, two specimens; Nkarara, 2200 feet, July 1, one; Bundimusuba, 3000 feet, November 20 to 24, five examples; weights 19 to 28 grams. We have elsewhere (1969: 26) commented on a series of this olive greenbul from Malabigambo forest, Sango Bay, in which account most of our Bwamba specimens also figured. The collectors noted the soft parts of our birds as follows: bill black to blackish gray becoming whitish along the tomium, and paler on the mandible than on the maxilla; iris brown; feet gray; stomach contents insect remains and a small fruit seed. A pair of the Bundimusuba birds were building a nest in a creeper-covered tree when collected on November 22. Phyllastrephus lorenzi Sassi. One specimen of this very rarely collected bulbul was taken at Ntandi, 2300 feet, June 17, 1967, constituting the first record for the species not only in Bwamba, but in Uganda, as pointed out by us earlier (1968: 23). Our 1968-1969 party failed to obtain a second example. Phyllastrephus fischeri sucosus Reichenow. Although the van Somerens (1949: 50) collected specimens of this species only above 5000 feet, they wrote that it also occurs at lower elevations. For this reason we include it in the present list of the birds of the Bwamba lowlands. Rand (1958: 210) reports specimen data from “Bwamba Pass”. Phyllastrephus icterinus tricolor (Cassin). In Bwamba our collectors met with this bulbul only at Ntandi, 2300 feet, where they obtained three specimens June 21 to 27, and one on November 16. A full discussion of these birds as well as of its sympatric sibling, P. xavieri, has been given elsewhere by the senior author (1968: 110-112). Phyllastrephus xavieri xavieri (Oustalet). This bird is much commoner in Bwamba than is P. icterinus. It was collected there by the van Somerens (1949: 50), by the Knudsen-Machris 1963 expedition, and also by our 1967 and 1968-1969 field parties, who took 33 specimens at the following localities: Ntandi, 2300 feet, October 3 to December 9 and June 25 to July 7; at Bundikeke, 3000 feet, November 23; at Mongiro, 2300 feet, November 3; at Nkarara, 2200 feet, June 30; and at Rwansenge, 2200 feet, July 14; weight 18 to 29 grams. Of all these individuals 42 Contributions in Science No. 211 only two, taken on June 30 at Nkarara and on July at Ntandi, showed gonadal enlargement. A fuller discussion of this species is given in the paper referred to under P. icterinus. Bleda syndactyla woosnami Ogilvie-Grant. The bristle-billed bulbul is common in the Bwamba forest, as indicated by 21 specimens taken at Ntandi, 2300 feet, October 13 to November 17 and June 15 to July 3; at Mongiro, 2300 feet, November 5; at Bundimusuba, 3000 feet, November 20 and July 6 to 9; at Rwansenge, 2200 feet, July 19; at Nikara, 2200 feet, June 29; and “Bwamba Forest”, April 21 to 24. It was reported earlier from Bwamba by the van Somerens (1949: 48) who con- sidered it rather scarce although they noted it at several places. Our series had weights of from 29.5 to 51 grams. The minimum weight given may be an error for 39.5, as a series from the Malabigambo forest (Friedmann and Williams, 1969: 27) ranged in weight from 40 to 47.5 grams; bill dull dark gray, paler on the mandible; iris dark red to dark red-brown; feet pale grayish with a pinkish or a brownish tinge; stomach contents caterpillars, beetles, and other insect fragments. Bleda eximia ugandae van Someren. The yellow-tailed bristle-bill is common and sympatric with B. syndac- tyla in Bwamba, although it apparently eluded the collectors working for the van Somerens. We have 15 specimens from the following localities: Ntandi, 2300 feet, November 12 to 18 and July 6 to 12; Nkarara, 2200 feet, June 29 and 30; Bundimusuba, 3000 feet, July 6; and Rwansenge, 2200 feet, July 16 to 19; weight 35 to 41 grams; bill dark gray to blackish, paler below and on the tomium; iris cream yellow to lemon yellow; feet gray; stomach contents grasshoppers, beetles, and other insect fragments. Nicator chloris (Valenciennes). This species is common in Bwamba. The van Somerens ( 1949: 81) were the first to record it there, and the Knudsen-Machris 1963 expedition obtained five examples with enlarged gonads April 21 to 23. Our 1967 and 1968-1969 field teams obtained 24 others as follows: Ntandi, 2300 feet, October 30 to December 6, and June 16 to July 13; Mongiro, 2300 feet, November 1 to 11; Bundimusuba, 3000 feet, November 20; Nkarara, 2200 feet, June 27; and Rwansenge, 2200 feet, July 6; weights 36 to 52 grams; bill black to blackish horn, the gape pale yellow; iris dark brown to deep grayish buff; feet bluish gray; stomach contents caterpillars, beetles, and other insect fragments. Nicator vireo Cabanis. This smaller species of Nicator exists sympatrically with its larger rela- tive, N. chloris, throughout Bwamba. The Bwamba area appears to be the eastern limit of the range of N. vireo, while N. chloris extends eastward at least to the Uganda-Kenya border. Rand (1958: 217) had two Bwamba 1971 The Birds of the Lowlands of Bwamba 43 specimens of N. vireo when writing his paper. The Knudsen-Machris 1963 expedition collected two April 20 and 23; our later field parties took two more specimens, one at Bundimusuba, 3000 feet, November 22, and one at Rwansenge, 2200 feet, July 15; weight 26 grams. We also have one taken at Ntotoro, 2250 feet, in March. Soft parts: bill grayish brown, basally paler; iris brown to grayish brown; feet gray; stomach contents orthoptera, beetles, and other insect fragments. Criniger barbatus weileri Gyldenstolpe. Robert Glen collected an adult female of this “bearded” bulbul in Bwamba Forest, 3500 feet, January 20, 1966; the specimen is now in the collection of the Western Foundation of Vertebrate Zoology (no. 18352); bill brown; iris brown; feet gray; stomach contents insect and fruit remains. This appears to be the first record for this species in Uganda. That it should occur in Bwamba is not surprising, although it must occur only in the few unexploited portions of the forest. Chapin (1953: 131) wrote that in the Congo in, “. . . the Ituri and Semliki forests it is very particular as to haunts. Only in wide stretches of unspoiled forest may one hope to see or hear the bird . . .” Our identification of this specimen to the subspecies weileri is based on the fact that Chapin included the Semliki Forest in the range of that race; we have seen no comparative material. The specimen is in fairly fresh plumage. Criniger c alums emini Chapin. This white-throated bulbul is very common in Bwamba, as attested by a long series of specimens from “Bwamba,” 2600 feet, April; Ntandi, 2300 feet, June, July, November; Rwansenge, 2200 feet, July; and Bundimusuba, 3000 feet, November. In studying these birds we have had the advantage of a series of both C. calurus and C. ndussumensis, identified by Rand, loaned to us by the Field Museum. The two are said to be almost identical but ndiissu- mensis supposedly has the bill somewhat more slender, the loreal area grayish or grayish white, the flanks somewhat more dusky green, and the undertail coverts more buffy yellow. These characters do not hold with complete con- stancy even in the two series identified by Rand, but all our Bwamba birds seem to be C. calurus emini. Of the 35 specimens, 4 have fairly grayish loreal areas but these do not have particularly narrow bills; one that has a bill as narrow as any of the ndussumensis has no grayish on lores. All of our speci- mens have the rectrices dusky russet edged with greenish; in fresh plumage the edges are more extensive, giving the feathers a more greenish, less russet look; all have the undertail coverts greenish yellow, not buffy yellow. It should be noted that our April “Bwamba” birds were published as ndussumensis (Friedmann, 1966: 32, following van Someren, 1949: 48). All of our speci- mens were in non-breeding state; bill blackish on maxilla, bluish gray on mandible, with whitish tomial edges; iris chocolate brown to reddish; feet 44 Contributions in Science No. 211 bluish gray; weight 23 to 31 grams; stomach contents small insects, fruits, and seeds. In spite of the fact that both calurus and ndussumensis are both supposed to occur in the forests of the Semliki valley, we know of no indubita- ble ndussumensis from Bwamba. Chapin (1953: 132-136) called both taxa races of the same species and treated all the Semliki birds, with which the Bwamba population should be in agreement, as ndussumensis. We cannot see any difference between the two taxa as far as their rectrices and their upper coverts are concerned. Family Campephagidae Campephaga phoenicea (Latham). The situation in Bwamba of this cuckoo-shrike is highly peculiar and puzzling. The van Somerens (1949: 80) reported only flava; we have three examples that are clearly flava, but we also have one adult male phoenicea with bright red shoulder patches! If this were to be interpreted as sympatry (none of the birds, however, were in breeding condition when collected), it would raise the question as to the specific distinctness of phoenicea and fiava, a question that cannot be decided on such scanty material. The only other interpretation would be to assume the male phoenicea was a wanderer from northern Uganda, where that form is known to occur. This is the interpreta- tion we are following here. The male C. p. phoenicea was taken at Mongiro, 2300 feet, on November 12; the specimens of C. p. flava, one male, one female, one probable female, were taken at Bundimusuba, 3000 feet, July 8 to 11. Aside from the fact that we have here two races of the same species, it is of interest to note that the Bundimusuba birds are definitely flava and not petiti, the race that occurs in the Kibale Forest. The male has the inner webs of the remiges clearly tinged with olive green as in several flava from Kenya, not blackish like the rest of the feathers as in petitv, the two females have the underparts white, marked with wavy transverse black bars as in flava from the Samburu district, Kenya, and not yellow as in petiti. One of them has some yellowish areas on the breast and upper abdomen, but is still predomi- nantly white below. The male C. p. phoenicea weighed 29 grams, the three flava 30 to 34 grams; in the males of both the bill was black (blackish horn in the females of flava) ; iris dark brown; feet blackish. The red shouldered male was watched when it was catching insects on the wing as it moved through the undergrowth. Family Muscicapidae Subfamily Muscicapinae Muscicapa striata striata (Pallas). The spotted flycatcher of Europe is only a winter visitor to the open country of Bwamba, where it was first recorded by Ridley, Percy and Percy 1971 The Birds of the Lowlands of Bwamba 45 (1953: 164). Our collectors obtained another specimen at Bundimusuba, 3000 feet, November 22. Muscicapa aquatica infulata Hartlaub. v. S., 1949: 54. Muscicapa caerulescens brevicauda Ogilvie-Grant. Fairly common in Bwamba, whence we have ten specimens from Ntandi, 2300 feet; Nyankiro, 3000 feet; Nkarara, 2200 feet; and Bundimusuba, 3000 feet; taken in June, July, November, and December; bill black, the basal part of the mandible paler bluish gray; iris dark brown; feet brownish gray to gray; weights 15 to 19 grams; stomach contents insect fragments. At Nkarara, June 28, a nearly fully grown young bird in juvenal plumage was collected together with an adult female, probably a recent fledgling with its mother. The van Somerens (1949: 54) noted this flycatcher in Bwamba and found that, while predominantly insectivorous, it does eat berries as well. Muscicapa griseigularis Jackson. The gray-throated flycatcher was first reported from Bwamba by the van Somerens (1949: 54), who found it was not numerous there. Our collec- tors obtained four speeimens at Ntandi, 2300 feet, November 17, and at Bundimusuba, 3000 feet, July 7, and November 26; bill black paling to gray- ish horn on the mandible; iris dark brown; feet dark blue-gray; weights 11.5 to 14 grams; stomach contents beetle larvae and other insect fragments. Muscicapa comitata (Cassin). Seven specimens; Bundimusuba, 3000 feet, November 22, and Ntandi, 2300 feet, July 15 to 22, and November 14 to 28; bill black; iris dark brown; feet gray to black; weights 13 to 15 grams; stomach contents insect remains. This flycatcher was found along a forest road, not deep in the forest according to the experience of the van Somerens (1949: 54-55), but it is a true forest species nearly always found along streams, where it perches on stones sticking up out of the water, or on roots, and dead branches hanging over the water. Muscicapa infuscata (Cassin). v. S., 1949: 55 (as Artomyias fuliginosa) . Myopornis plumbeus plumbeus (Hartlaub). v. S., 1949: 60. Me gaby as flammulata aequatorialis Jackson. Specimens of this flycatcher were obtained at Ntandi, 2300 feet, and at Nkarara, in June, July, and November; all in non-breeding state; bill black; iris red to brown; feet pale maroon to dull maroon; weight 26 to 30 grams; stomach contents insect fragments. All of our six specimens are females. Bias musicus musicus (Vieillot). In the experience of our collectors this black and white flycatcher was uncommon and local; it was found only at Ntandi, 2300 feet, where two examples were obtained October 29 and December 7. In contrast, the van 46 Contributions in Science No. 211 Somerens (1949: 56) found it to be common in the more open parts of the forest. Our birds had the bill black; iris yellow; feet pale yellow in one and pale olive brown in the other; weight 21 grams; stomach contents insect fragments. Platysteira cyanea nyansae Neumann. The absence of this species in our collection is surprising in view of its abundance in Bwamba, according to the van Somerens’ experience 1949: 56). Platysteira castanea castanea Fraser. The chestnut wattle-eye is very common in the Bwamba forest. A long series of specimens was collected at Ntandi, 2300 feet, June, July, October, November, and December; at Bundimusuba, 3000 feet, in July and Novem- ber; and at Rwansenge, 2200 feet, in July; bill black; iris dark brown; wattles dull maroon in males, deep purple to mauve maroon in females; feet dull maroon to deep purple; weight 1 1 to 16 grams; stomach contents insects. According to the van Somerens (1949: 56-57) this wattle-eye frequents chiefly the lower trees and the tall undergrowth in the forest, an observation agreeing with what our collectors found as well. Platysteira blissetti jamesoni (Sharpe). The van Somerens ( 1949: 57) considered Jameson’s wattle-eye less numerous in Bwamba than the preceding species, but our collectors found it to be very common and obtained many specimens at “Bwamba Forest,” April; Ntandi, 2300 feet, June, July, October, November, December; Bundi- musuba, 3000 feet, July and November; Rwansenge, 2200 feet, July; Bumaga, 2300 feet, June. Two of our specimens, taken on June 28, had much enlarged gonads; the others were definitely not in breeding condition; bill black; wattles turquoise blue; iris maroon to dark brown; feet maroon to purplish maroon; weights 9 to 12 grams; stomach contents insects. Platysteira concreta graueri (Hartert). v. S., 1949: 57. Trochocercus longicauda teresitus (Antinori). The van Somerens ( 1949: 58) found the blue fantail flycatcher common in cultivated areas adjacent to the forest. Carcasson collected a male in the Bwamba lowlands on May 23, 1956; the specimen is now in the collection of the Western Foundation of Vertebrate Zoology. T rochocercus nitens nitens Cassin, The experience of our collectors showed the blue-headed crested fly- catcher to be common and wide-spread in Bwamba, in contrast to that of the van Somerens (1949: 58) who considered it rather scarce. We have some 15 specimens from the following localities: Ntandi, 2300 feet, June, October, November; Mongiro, 2300 feet, November; Bundimusuba, 3000 feet, July, November; Bundikeke, 3000 feet, July; and Rwansenge, 2200 feet, July; bill 1971 The Birds of the Lowlands of Bwamba 47 grayish blue with blackish tip; iris dark brown; feet gray blue; weights 10 to 12.5 grams; stomach contents insect fragments. Birds with enlarged gonads were taken on July 8 (a female with a well-developed, but still unshelled, egg) , November 16 and 22. The van Somerens reported a nest and eggs in April, so it would seem that the breeding season is prolonged. Trochocercus nigromitratus (Reichenow). Not as common as the last species, this flycatcher is represented in our collections by seven specimens: “Bwamba Forest”, April 21, 22; Ntandi, 2300 feet, June 24, July 3, November 16; Bundimusuba, 3000 feet, July 9; bill black; iris dark brown; feet deep blue-gray to blackish gray; weights 9 to 10.5 grams; stomach contents insects. Terpsiphone rufiv enter somereni Chapin. The red-bellied paradise flycatcher is very common in the fairly open forests of Bwamba. The van Somerens (1949: 59) found it outnumbered the other species of paradise flycatchers there, as did our collectors as well. Ridley, Percy, and Percy (1953: 164) also noted it there. We have a long series of specimens from “Bwamba Forest,” 2600 feet, April; Ntandi, 2300 feet, June, July, October, November, and December; Nkarara, 2200 feet, June and July; Bumaga, 2300 feet, July; Mongiro, 2300 feet, November; Bundimusuba, 3000 feet, November; and Rwansenge, 2200 feet, July; bill cobalt blue, the tip and the tomium black; iris dark brown; feet dark blue-gray; weights 13 to 19 grams; stomach contents insects. Terpsiphone viridis ferreti (Guerin). This paradise flycatcher, with its great variety of plumage colors, is widely distributed in the Bwamba lowlands. One of our present series, from Nyankiro, has the back and long tail feathers white; the others are in the brown or so-called “normal” plumage. We are in agreement with Chapin’s (1963) conclusions about the nature of the perplexing color phases of this bird, and no further comment is called for at this point. We have a long series of specimens from: “Bwamba Forest,” April; Rwansenge, 2200 feet, July; Korongo, Semliki river, 2200 feet, July; Ntandi, 2300 feet, November and December; Bundimusuba, 3000 feet, November; and Nyankiro, 3000 feet, November; bill pale blue with the tip and tomium blackish; iris dark brown; feet blue-gray; weight 11 to 16 grams; stomach contents insect remains. Terpsiphone batesi Chapin, v. S., 1949: 59. Hyliota slat ini Sassi. v. S., 1949: 53. Subfamily Turdinae Erythropygia hartlaubi Reichenow. The brown-backed scrub-robin is found in the cultivated areas just out- 48 Contributions in Science No. 211 side the Bwamba forests. One example was taken there April 22. The van Somerens (1949: 66) found it numerous there many years earlier. Erythropygia leucosticta collsi Alexander. One record, a subadult male taken in Bwamba Forest, July 23, 1960, by Twomey (Keith and Twomey, 1968: 545). Erithacus cyornithopsis lopezi (Alexander) . Our collectors found this species only at Ntandi, 2300 feet, November 1 1 to 20, where they obtained four specimens, all in nonbreeding condition; bill black, the base of the mandible yellowish; iris dark brown; feet silvery gray; weight 17 to 19 grams; stomach contents insects and small snails. This is an extremely illusive denizen of the dense, virgin forest, and would be very difficult to collect without the use of mist nets. This may account for its absence in the van Somerens’ list. Erithacus erythrothorax mabirae (Jackson). The little forest robin was very common in the forests of Bwamba; a long series of specimens was taken at “Bwamba” April 19 to 25; Ntandi, 2300 feet, October, November, December, June, and July; Mongiro, 2300 feet, November 3 to 17; Bundimusuba, 3000 feet, November 21 to 29; and Nkarara, 2200 feet, June 29, July 1. Three of the July birds had enlarged gonads; one of the April specimens was a partly grown young bird. In an earlier paper (Friedmann, 1966: 36) it was pointed out that this young bird, hardly more than a week out of the nest, had the chin, throat, and much of the abdomen pale yellow, not “creamy white” as described in the literature. This difference must be due to post-mortem fading, as the same specimen is now less yellowish, more whitish than it was five years ago. Our series of adults ranged in weight from 11 to 19 grams; bill black; iris dark brown; feet grayish violet to grayish pink; stomach contents insect remains. The van Somerens did not include this species in their report. Its absence there and its abundance in our more recent collections is an indication of the great advantage of mist netting in making faunal surveys, especially as far as the birds of the undergrowth are concerned. Our collectors repeatedly released many individuals of the present and other species when they already had adequate samples from each locality. Although very common, this robin is seldom observed because of its excessive shyness. Cossypha natalensis intensa Mearns. Common at Mongiro, 2300 feet, November 3 to 8, when eight specimens were collected, four of them with enlarged gonads; bill black; iris dark brown; feet pinkish gray; weight 29 to 37 grams; stomach contents insects. Cossypha cyanocampter bartteloti Shelley. The blue-shouldered robin chat was found to be common and wide- spread in the lowland forest, our collectors having the same experience with it 1971 The Birds of the Lowlands of Bwamba 49 as did the van Somerens ( 1949: 63). We have specimens from Bwamba, April 24; Ntandi, 2300 feet, June, July, October, and November; Bundimusuba, 3000 feet, July 6 to 10, November 22 to 29; and Rwansenge, 2200 feet, July 15. Some of the July birds had much gonadal enlargement, the others did not; bill black; iris dark brown; feet brownish black; weight 23.5 to 30 grams; stomach contents chiefly small insect remains, one small snail in one instance. Cossypha niveicapilla (Lafresnaye) . The snowy-headed robin chat was obtained in the Bwamba Forest on April 21 by the Knudsen-Machris expedition of 1963, but was not found again by our field parties in 1967 or in 1968-1969. This is surprising in view of the fact that the van Somerens ( 1949: 63) encountered it in most parts of the forested areas. Alethe diademata woosnami Ogilvie-Grant. Our collectors found the fire-crested alethe to be very numerous, and obtained a long series of specimens from “Bwamba,” April 20 to 21; Ntandi, 2300 feet, June, July, November, and December; Bundimusuba, 3000 feet, November; Mongiro, 2300 feet, November; Bumaga, 2300 feet, June 27; Nkarara, 2200 feet, June 28; Ntotoro, 2250 feet, March 6. The experience of the van Somerens (1949: 64) was quite otherwise, as they met with this bird in only one place in Bwamba. Williams (1951: 109) obtained this thrush in Bwamba as well. None of our specimens were in breeding condition; bill black; iris dark brown; feet gray; weight 28 to 34 grams; stomach contents largely beetles, some other insect remains, and in one case, a small snail. Alethe poliocephala carruthersi Ogilvie-Grant. As in the preceding species, the van Somerens (1949: 64) found this thrush scarce while our field parties met with it frequently and obtained a long series of specimens at “Bwamba,” April 29 to May 5; Ntandi, 2300 feet, June, July, October and November; Mongiro, 2300 feet, November 5 to 7; Bundimu- suba, 3000 feet, July 6, November 22 to 24; Nkarara, 2200 feet, June 28 to July 1; and Rwansengi, 2200 feet, July 14 to 16. One of the late June birds from Ntandi had enlarged gonads; all the other examples were in non-breeding condition; bill black; iris reddish brown to dark brown; feet pale pinkish flesh; weight 21.9 to 31 grams; stomach contents insect remains. Stizorhina fraseri vulpina Reichenow. Both our collectors and the van Somerens agreed in finding the rufous flycatcher-thrush to be common and widespread in the Bwamba Forest. We have many specimens from Ntandi, 2300 feet, June, July, October and November; Bundimusuba, 3000 feet, November 22; Mongiro, 2300 feet, November 1 1 to 16; Nkarara, 2200 feet, June 27 to July 1; birds with enlarged gonads were taken in June and in November; bill black; iris dark brown; feet brown; weight 27 to 37 grams; stomach contents insects. 50 Contributions in Science No. 211 Neocossyphus rufus gahonensis Neumann. Not as numerous as Stizorhina, or as the next species, but not uncom- mon in the Bwamba. Our collectors obtained eight examples at Ntandi, 2300 feet, June 29, November 6 to 19; at Mongiro, 2300 feet, November 4 to 12; and at Nkarara, 2200 feet, July 1. Two of them, taken June 29 and mid November, had enlarged gonads; the others were in non-breeding condition; bill black; iris dark brown; feet pale pinkish gray; weight 55 to 72 grams; stomach contents beetles and other insects. The van Somerens (1949: 65) also found this bird to be relatively scarce. Neocossyphus poensis praepectoralis Jackson. In contrast with the experience of the van Somerens ( 1949: 65) who met with this ant thrush only twice, our collectors obtained a long series of specimens as follows : Bwamba Forest, April 21 to 25; Ntandi, 2300 feet, June, July, November; Mongiro, 2300 feet, November 4 to 8; Bundimusuba, 3000 feet, July 6, November 2; Nkarara, June 30; birds with enlarged gonads were taken June 30 and July 5; the others were in non-breeding state; bill black; iris dark brown; feet pinkish white; weight 42 to 51 grams; stomach contents largely beetles but also some other insect fragments. SaxicoJa rubetra (Linnaeus), v. S., 1949: 67. Saxicola torquata axillaris (Shelley), v. S., 1949: 67. Thamnolaea cinnamomeiventris subrufipennis Reichenow. v. S., 1949: 67. Zoothera oberlaenderi (Sassi). We have a single example of this rare thrush, an adult female, with an enlarged ovary, taken at Ntandi, 2300 feet, on July 17, as discussed by us in an earlier paper ( 1968: 27). Keith ( 1968: 10-11), and Keith and Twomey (1968: 544) have reported on the other few Bwamba records of this species, which have also been discussed by Prigogine ( 1965: 242). T Urdus olivaceus centralis Reichenow. This thrush is common in the cultivated clearings at the edges of the forested areas. In addition to three birds with enlarged gonads, taken April 21 to 23, by the 1963 Knudsen-Machris expedition (Friedmann, 1966: 38), we have six other specimens taken at Ntandi, 2300 feet, June 15 to 21, and November 1; and Mongiro, 2300 feet, November 8. The van Somerens (1949: 61) also reported this bird from the Bwamba lowlands. Our specimens had weights of 65 to 66 grams. Subfamily Sylviinae Acrocephalus natalensis niloticus (Neumann), v. S., 1949: 67. Chloropeta natalensis massaica Fischer and Reichenow. v. S., 1949: 60. 1971 The Birds of the Lowlands of Bwamba 51 Sphenoeacus mentalis mentalis (Fraser). Four specimens of this large warbler were obtained: Nkarara, 2200 feet, June 27, July 2; Korongo, Semliki River, 2200 feet, July 14; and Rwansenge, 2200 feet, July 15; bill black with the tomial edges whitish gray; iris pale orange to Naples yellow; feet blue-gray to brownish gray; weight 29 to 40 grams; stomach contents small insect fragments. We are following White in considering Melocichla the same as Sphenoeacus, but with less than complete assurance that they are congeneric. Sylvia borin (Boddaert). At Ntandi, 2300 feet, November 11 to 16, this European winter visitor was found in the open country adjacent to the forest and three examples were collected; weight 19 to 25 grams. Phylloscopus trochilus trochilus (Linnaeus). The common willow warbler of Europe winters in most parts of Africa, and the cultivated areas of the Bwamba lowlands are no exception. At Ntandi, 2300 feet, November 1 to December 3, four specimens were taken; weight 8.5 to 10 grams. Cisticola erythrops sylvia Reichenow. v. S., 1949: 67. Cisticola galactotes amphilecta Reichenow. This grass warbler was found in the grass and scrub outside the forest at Nkarara, 2200 feet, June 27, and Rwansenge, 2200 feet, July 6. The Nkarara bird was in breeding condition; the other had small gonads; bill black on the maxilla, pale gray on the mandible; iris light brown; feet pale brown; weight 17 and 18 grams, respectively; stomach contents insects. Also reported as common in Bwamba by the van Somerens (1949: 68). Cisticola carruthersi Ogilvie-Grant. v. S., 1949: 68. Cisticola natalensis strangei (Fraser). Two examples of this grass warbler were taken at Nkarara, 2200 feet, June 27 and 29; bill blackish, the mandible lighter— more grayish; iris buff brown; feet pale whitish flesh to pale flesh brown; weight 21.5 to 34 grams (a surprising range for two specimens!); stomach contents insects, including a beetle and a grasshopper. Cisticola brachyptera brachyptera (Sharpe). One of this species was taken in the same place and date as one of the preceding one, Nkarara, 2200 feet, June 27; bill blackish on the maxilla, pale gray on the mandible; iris buff brown; feet pale flesh horn; weight 8.5 grams; stomach contents insects. Prinia subfiava melanorhyncha (Jardine and Fraser). This long-tailed warbler was met with only at Nkarara, 2200 feet, June 52 Contributions in Science No. 211 27, when two examples were obtained, both with small gonads; bill black, gray at the base; iris light brown; feet pale flesh brown; weight 8 to 8.5 grams; stomach contents insects. The van Somerens (1949: 68) found it to be a common bird in the cultivated clearings in the Bwamba lowlands. Prinia leucopogon reichenowi (Hartlaub). v. S., 1949: 68. Prinia bairdii obscura (Neumann). The banded long-tailed warbler appears to be uncommon, or at least local, in Bwamba. The van Somerens (1949: 68) had only a single specimen; our collectors obtained two at Ntandi, 2300 feet, one on June 15 and the other on November 14; bill black; iris yellow ochre to light brown; feet silvery gray; weight 14 to 16 grams; stomach contents small beetles. Apalis rufogularis nigrescens (Jackson) . The van Somerens (1949: 68-69) refer to the red-throated forest warbler as the, “. . . common Apalis of the lowland forests . . .”, but it was the only member of its genus that they or our field teams found there, the other nearby species of the genus being montane birds of the Ruwenzori slopes. We have seven specimens, six from Ntandi, 2300 feet, June 19 to July 19, and December 4; and one from Nkarara, 2200 feet, July 1. All were in non-breeding condition; bill black; iris brown; feet orange-brown to flesh gray; weight 7.5 to 9 grams; stomach contents small insects. All the males the van Somerens collected in July (some at Ntandi) had enlarged gonads, unlike our birds. Eminia lepida Hartlaub. v. S., 1949: 72. Bathmocercus cerviniventris vulpinus Reichenow. Common in the Bwamba Forest. Our collectors took specimens at Ntandi, 2300 feet, June 18 to 24; Nkarara, 2200 feet, June 28 and 29; Tokwe River, 2200 feet, June 28; Bundimusuba, 3000 feet, July 6 and November 23, while the Knudsen-Machris expedition collected three examples in the Bwamba Forest, March 4 to April 23, 1963. A bird taken on July 6 at Bundimusuba, had enlarged gonads; the others were in non-breeding condi- tion; bill black; iris brown; feet gray; weight 14 to 21 grams; stomach con- tents insects. Camaroptera superciliaris (Fraser). Fairly common; we have two specimens from each of three localities: Bundimusuba, 3000 feet, July 2; Korongo, Semliki River, 2200 feet, July 14; and Ntandi, 2300 feet, November 27 and December 10. One of the July 14 birds was in breeding condition; the other specimens showed little or no gonadal enlargement; bill black; iris dark brown; feet grayish brown to dark brown, the toes paler; weight 9 to 1 1 grams; stomach contents insects. 1971 The Birds of the Lowlands of Bwamba 53 Camaroptera chloronota toroensis (Jackson). A very common bird along the forest edges. We have a long series: Ntandi, 2300 feet, June 16 to July 17, and October 31 to November 20; Bundimusuba, 3000 feet, July 7 to 12, and November 20 to 25; Mongiro, 2300 feet, November 4; Bumaga, 2300 feet, July 6 and 7; Rwansenge, 2200 feet, July 13 and 14; and Bwamba Forest, April 22 to 24; bill black; iris dark brown; feet black; weight 9 to 12 grams (one bird marked 18 grams) ; stomach contents insects. The van Somerens ( 1 949 : 71) also found this species fre- quently at the margins of the Bwamba woodlands. None of our birds showed any seasonal gonadal swelling. Camaroptera brachyura tincta (Cassin). This species was also common, but was more in the cultivated areas just outside the forest than the preceding one. Specimens were obtained at Nkarara, 2200 feet, June 28 to July 7; Ntandi, 2300 feet, July 2, November 16 to 19; Rwansenge, 2200 feet, July 13; Mongiro, 2300 feet, November 5; and Bundimusuba, 3000 feet, November 21; of these birds, four, taken in early July and mid November, had swollen gonads; the others were in non- breeding condition; bill black; iris light brown; feet flesh color; weight 10 to 13 grams; stomach contents small insects. Eremomela badiceps badiceps (Fraser). The rufous-crowned eremomela was met with by our collectors only once, at Ntandi, 2300 feet, November 11, when a single example in breeding state was obtained. This is quite different from the experience of the van Somerens (1949: 69) who had specimens from at least four Bwamba locali- ties and saw up to a dozen birds on one occasion. Our specimen had the bill black; iris dark brown; feet flesh brown; weight 10 grams; stomach contents small caterpillars. Sylvietta virens baraka Sharpe. Our field teams found this little crombec to be common and widely dis- tributed in the lowland forest. Specimens were taken at Ntandi, 2300 feet, June 15 to July 11, and December 8; Nkarara, 2200 feet, June 27; Bumaga, 2300 feet, July 2; and Bundimusuba, 3000 feet, November 20; bill dark to pale horn brown, paler on the mandible; iris light brown to buff brown; feet pale buff brown; weight 8 to 10 grams (one marked as only 5 grams) ; stomach contents insects. This species was found by the van Somerens (1949: 70) to be common chiefly in the undergrowth and among creepers. Macrosphenus flavicans hypochondriacus (Reichenow) . This race of the yellow longbill is common in the Bwamba woodlands. We have some ten specimens: Ntandi, 2300 feet, June 27, July 17, October 30 to December 1; Nkarara, 2200 feet, July 1; Bundikeke, 3000 feet, November 23; Bundimusuba, 3000 feet, July 7, November 20 and 21; bill black with the 54 Contributions in Science No. 211 tomium and the tip and under side whitish; iris yellow; feet blue-gray; weight 12.5 to 15.5 grams; stomach contents seeds. Hylia prasina prasina (Cassin). The green hylia was found to be very common, and a long series of specimens was taken: “Bwamba,” April 20 to 23; Ntandi, 2300 feet, June 15 to July 16 and October 28 to December 4; Bundimusuba, 3000 feet, July 6, and November 21 to 29; Rwansenge, 2200 feet, July 14; and Mongiro, 2300 feet, November 3 and 4. Five of the June-July birds from Ntandi had enlarged gonads; the others were in non-breeding state; bill black, the tomium of the mandible basally pale yellow; iris dark brown; feet olive; weight 1 1 to 15 grams; stomach contents small insects. The van Somerens (1949: 73) wrote that the stomachs of their series contained only small black scale insects and scale wax, very much as in the sharp-billed honey-guide, Pro- dotiscus insignis. They also noted that the behavior of these two unrelated birds was quite similar, searching about for scale insects. Many years ago, in Cameroon, Bates also noted the preference for scale insects shown by the green hylia. Family Dicruridae Dicrurus adsimilis coracinus J. and E. Verreaux. This drongo is common in the Bwamba Forest; our collectors obtained a series of specimens at Bwamba, April 20; Ntandi, 2300 feet, June 13 to 24 and November 1 to December 9; and at Bundimusuba, 3000 feet, July 11. The April birds, and some of those taken in other months, had enlarged gonads; the birds ranged in weight from 45 to 53 grams; stomach contents insect remains, chiefly beetles. This species was noted by the van Somerens (1949: 80, 81) under two specific headings— D. modestus { — D. adsimilis coracinus of the present report) and D. adsimilis divaricatus. According to Vaurie’s map (1949: 225), in western Uganda one might expect coracinus in the forests and nominate adsimilis in the adjacent open country. Three of our twelve specimens from Ntandi lack the deep velvety black upper back present in the other nine and have this area as glossy as the rest of the head and body, and might, therefore, be considered as examples of typical adsimilis. However, they have the inner portions of the remiges blackish, not ashy fuscous as in specimens of adsimilis from Kenya and Tanzania. Inasmuch as there is open country at Ntandi just outside the forest it is possible that these three specimens were taken outside the true forest and are nearer to adsimilis than to coracinus. They do not agree with D. atripennis, which is said to extend east to the Semliki Valley and might, therefore, be expected in Bwamba. This species has a shallower tail fork and is slightly smaller. Through the courtesy of the American Museum of Natural History, four examples of atripennis were available for direct comparison. 1971 The Birds of the Lowlands of Bwamba 55 The van Somerens noted coracinus chiefly in those parts of the Bwamba forests where there was little undergrowth. This suggests that they found the velvety-backed drongo well in the interior, and not around the margins, of the woodlands. Williams (1951: 110) also found coracinus in the forests in Bwamba. Family Laniidae Prionops caniceps mentalis (Sharpe). The red-billed helmet-shrike is another common bird of the Bwamba woodlands. Both the van Somerens and our field parties saw many of them. We have some 14 specimens from: Bwamba, April 24; Ntandi, 2300 feet, June 28, November 20 to December 9; and Ntotoro, 2600 feet, February 28; bill dark crimson, the tip more orange-red; iris dull brownish gray with the outer ring orange or yellow; eye ring whitish; feet orange-red; weight 39 to 50 grams; stomach contents caterpillars, beetles, praying mantis, and other insect fragments. Dryoscopus gambensis malzacii (Heuglin). v. S., 1949: 83. Drysocopus senegalensis (Hartlaub). When the van Somerens (1949: 83) collected their first specimen at Bwamba the red-eyed puff-back shrike was an addition to the known avifauna of Uganda. They obtained others subsequently, and our collectors also found the species at Ntandi, 2300 feet, November 20, and succeeded in collecting two individuals, one in breeding condition; bill black; iris orange-yellow to orange-red; feet bluish gray; weight 30 to 34 grams; stomach contents insects, mainly beetles. Tchagra minuta minuta (Hartlaub). v. S., 1949: 84. Tchagra australis emini (Reichenow). v. S., 1949: 84. Laniarius liihderi liihderi Reichenow. This bush shrike was met with only at Ntandi, 2300 feet, December 10, when two specimens, in non-breeding state, were collected: bill black; iris dark brown; feet blue-gray; weight 45 to 50.5 grams; stomach contents beetles and small caterpillars. Laniarius ferrugineus major (Hartlaub). v. S.„ 1949: 82. Lariarius leucorhynchus (Hartlaub). While the van Somerens (1949: 82) reported the sooty boubou as a scarce species in the Bwamba Forest, our field teams met with it often and collected 14 specimens at Ntandi, 2300 feet, June 15 to July 11, and October 6 to December 4; Rwansenge, 2200 feet, July 6; Bundimusuba, 300 feet, July 11; at “Bwamba” April 25; some of the July specimens had enlarged gonads; bill black; iris dark brown; feet black; weight 45 to 55 grams; stomach con- 56 Contributions in Science No. 211 tents insects, mainly beetles. At Ntandi this bush shrike was heard to give a five note whistle, first two, then a pause followed by three notes. It is a skulking bird of the dense creepers in secondary undergrowth where it keeps under cover as a rule. Telophorus bocagei jacksoni (Sharpe), v. S., 1949: 84. Malaconotus cruentus adolfi-friederici Reichenow. The fire-breasted bush-shrike is common in the forests of Bwamba; our collectors obtained 8 specimens: Ntandi, 2300 feet, July 5 to 7, December 4 to 13; Rwansenge, 2200 feet, July 15; and Bwamba Forest, April 24. Two of the July birds were in breeding condition; the others were not; bill black; iris gray to pale gray; fee gray; weight 70 to 75 grams; stomach contents insects, including a grasshopper, small beetles, a butterfly, and moth eggs. Williams ( 1957: 159) was the first to record this species from the Bwamba Forest. One of these birds was attracted by Robert Glen’s imitation of its call and hopped about excitedly apparently trying to find the caller. The call was a series of five or six deep-toned whistled poo poo notes given at intervals of about a second. Lanins mackinnoni Sharpe. Three specimens, Ntandi, 2300 feet, July 2 and 3; and Nyankiro, 3000 feet, November 25; bill black; iris dark brown; feet black; weight 35 grams each; stomach contents insects. The van Somerens (1949: 84) found this gray shrike common in Bwamba. Lanius minor Gmelin. Our collectors obtained an example of the lesser gray shrike at Bwamba, April 25, and another at Kisuba, 2500 feet, November 2; bill black, paling to gray basally; iris dark brown; feet black; weight 48 grams; stomach contents insects. Lanius excubitoroides excubitoroides Prevost and Des Murs. v. S., 1949: 85. Family Oriolidae Oriolus oriolus oriolus (Linnaeus). Specimens of the European oriole wintering in the Bwamba lowlands were taken at Bundimusuba, 3000 feet, November 20, and at Bundikeke, 3000 feet, November 23; weight 68 to 74 grams; stomach contents insects. Oriolus auratus notatus Peters, v. S., 1949: 85. Oriolus brachyrhynchus laetior Sharpe. The black-headed oriole was found to be common in the Bwamba wood- lands; 15 specimens were taken: Ntandi, 2300 feet, June 16 to July 19 and November 1 to December 7; Nkarara, 2200 feet, June 30; and Mongiro, 2300 feet, November 14; some of them with large, others with small, gonads; bill 1971 The Birds of the Lowlands of Bwamba 57 dull brownish pink to dill red (blackish in an immature bird); iris dark red to reddish brown; feet dark gray to bluish gray; weight 45 to 58 grams; stom- ach contents fruit seeds, and small caterpillars. Oriolus nigripennis nigripennis Verreaux. The van Somerens ( 1949: 85) were the first to record the black-winged oriole from Bwamba, and, indeed, from Uganda. Our field team found it but once, at Ntandi, 2300 feet, July 18; bill dull pinkish red; iris dark red; feet gray; weight 55 grams; stomach contents seeds, fruit, and insects. Family Corvidae Ptilostomus afer (Linnaeus). The piapiac is not a forest bird, but occurs in the open country immedi- ately adjacent to the forest. One female was collected in the Semliki Valley, Bwamba, 2300 feet, April 21; bill black; iris violet; feet black. Ridley, Percy, and Percy (1953: 164) also reported taking two examples of this species from a flock on the banks of the Semliki where it emerges from the forest. Corvus albus Muller, v. S., 1949: 86. Corvus albicollis (Latham), v. S., 1949: 86. Ridley, Percy, and Percy (1953: 164) reported several whitenecked ravens at Bweramule. Family Sturnidae Poeoptera lugubris Bonaparte. This forest starling was added to the Uganda ornis when the van Som- erens (1949: 87) reported it in considerable numbers in the Bwamba low- lands. Later, Ridley, Percy, and Percy (1953: 164) found small flocks at Bubukwanga. Our collecting teams did not meet with it at any time, which raises the possibility that the birds may wander about and stay only where shrubs or trees are fruiting. This is suggested by the experience of Williams who found them around fruiting fig trees and another unidentified forest tree with small green fruits. Our field team collected numerous specimens of this starling in other west Ugandan forests and were, therefore, familiar with it and its habits. The specimens they obtained in these other areas had fruit pulp and seeds in their stomachs. Onychognathus fulgidus hartlaubi Grey. Two specimens of this rufous-winged starling were taken, one at Ntandi, 2300 feet, December 11, and one at Nkarara, 2200 feet, June 28; both in non-breeding condition; bill black; iris red; feet black; weight recorded as 40 to 1 17 grams (but we think that 40 must be an error for 110, an easy slip in writing on a label); stomach contents fruit seeds and a large brown fruit. This starling is found in the more open parts of the forest, especially around 58 Contributions in Science No. 211 large trees left in clearings made by the natives for cultivation. Unlike some of the other starlings it does not go in flocks but is usually seen singly or in pairs. Williams (1951: 110) first reported this species from Bwamba (Bundibugyo). Lamprotornis purpureiceps (J. & E. Verreaux). Both the van Somerens (1949: 86-87) and our collectors found the purple-headed glossy starling commonly in various places in the Bwamba forests. We have a good series of specimens from Ntandi, 2300 feet, June 15 to July 10 and December 6 to 8; Bundimusuba, 3000 feet, July 8 to 10; and Bwamba Forest, April 25; birds with large and with small gonads were col- lected at the same time; bill black; iris dark brown; feet black; weight 60 to 70 grams; stomach contents fruit (?) seeds, and insect fragments. Lamprotornis splendidus splendidus (Vieillot). Our collectors found this handsome glossy starling very common in Bwamba, as did the van Somerens many years earlier (1949: 87). Ridley, Percy, and Percy (1953: 164) saw several hundred of them at a communal roost near the Kirimia River. We have half a dozen specimens: Ntandi, 2300 feet, June 22; Bundimusuba, 3000 feet, November 20 and 23; Rwansenge, 2200 feet, July 15; all in non-breeding condition; bill black; iris white; feet black; weight 110 to 142 grams; stomach contents fruit pulp, seeds, and, in one case, some insect fragments. One of the Bundimusuba birds was collected from a party of over 100 individuals feeding in the fruiting trees. Cinnyricinclus leucogaster verreauxi (Bocage). Reported as seen in large flocks at times by the van Somerens (1949: 86), our collectors found this starling in the forest clearings in smaller num- bers, only one or a few at a time, usually well up in the trees. They obtained a few specimens at Ntandi, 2300 feet, November 29, and at Bundimusuba, 3000 feet, November 22; bill black; iris bright lemon yellow in two males, brownish gray with a pale yellow outer ring in a female; feet blackish; weight 45 to 48 grams; stomach contents fruits and fruit seeds. At Bundimusuba a party of 6 or 7 of these starlings was seen feeding in a fruiting wild fig tree. Buphagus erythrorhynchus erythrorhynchus (Stanley), v. S., 1949: 88. Family Zosteropidae Zosterops senegalensis toroensis Reichenow. v. S., 1949: 88. Family Nectariniidae Anthreptes fraseri axillaris (Reichenow). The gray-headed sunbird is common in the Bwamba forests. The van Somerens (1949: 78) found it chiefly in the tree-tops; our collectors, and, earlier, the junior author, collected many specimens at Ntandi, 2300 feet. 1971 The Birds of the Lowlands of Bwamba 59 June, July, October, November, December; Nkarara, 2200 feet, June and July; Mongiro, 2300 feet, November. Some birds with enlarged gonads and others with small ones were taken during the same months; bill blackish, grayer on the mandible; iris brown to red-brown; feet gray with an olive wash; weight 9.5 to 13 grams; stomach contents insect fragments, including at least one caterpillar. Anthreptes longuemariae haussarum Neumann. There are two specimens of this white-bellied sunbird from Mongiro, 2300 feet, July 1 and 1 1, in the Williams collection of sunbirds, now in the Los Angeles County Museum of Natural History; bill dark brown; iris brown; feet black. The sunbird occurs in the open, grassy areas next to the forest, but not in the dense woodlands. Anthreptes rectirostris tephrolaema (Jardine and Fraser). Fairly common, although recorded by the van Somerens (1949: 78) as somewhat scarce, but possibly overlooked. We have specimens from Ntandi, 2300 feet, June 20, and November 8 to 25; Mongiro, 2300 feet, November 7 and 12; the June bird and one of the November ones had enlarged gonads; bill blackish horn; iris deep reddish brown; feet blackish brown; weight 6 to 10.5 grams; stomach contents small fruit seeds and insect remains. Anthreptes coUaris somereni Chapin. Our collectors found this sunbird common and obtained a series of specimens at Ntandi, 2300 feet, June, July, and November; Nkarara, 2200 feet, July; Mongiro, 2300 feet, November; and Rwansenge, 2200 feet, July. Yet neither the van Somerens nor the Knudsen-Machris expedition collected this species in Bwamba. Two of our July birds had enlarged gonads; bill black; iris dark brown; feet black; weight 7.5 to 9 grams; stomach contents small insect fragments. This sunbird is commoner around the edges of clear- ings in the forest than in the depths of the forest. Nectarinia seimundi traylori Wolters. Our collectors met with the little green sunbird only at Ntandi, 2300 feet, July 6 to 17, when they collected a number of specimens; bill dull horn gray, paling to yellowish horn on the mandible; iris brown; feet dark gray; stomach contents small spiders, fig seeds and pulp, and other small seeds. There are other Bwamba examples in the Williams sunbird collection, and the van Somerens (1949: 78) found the species to be plentiful, but easily overlooked. In the Congo, Chapin (1954: 205) also found it, “. . .one of the sunbirds most easily overlooked ...” Nectarinia olivacea cephaelis (Bates). The olive sunbird is common in the forests of Bwamba; we have a long series of specimens from: Ntandi, 2300 feet, June, July, and November; Bundimusuba, 3000 feet, November; Mongiro, 2300 feet, November; and 60 Contributions in Science No. 211 Bwamba, April. Some of the April and July birds had enlarged gonads; bill black; iris dark brown; feet dark brown; weight 9 to 13 grams; stomach contents insects. Previously recorded from Bwamba by the van Somerens (1949: 77) and by Williams (1951: 110). Nectarinia verticalis viridisplendens (Reichenow). v. S., 1949: 77. Nectarinia cyanolaema octaviae (Amadon). The blue-throated brown sunbird is common in the Bwamba Forest. We have a good series of specimens from: Ntandi, 2300 feet, June, July, Novem- ber, and December; Mongiro, 2300 feet, November; Bundimusuba, 3000 feet, November; and the north Humia River, 2300 feet, December; bill black; iris dark brown; feet black; weight 12 to 18 grams; stomach contents insect remains. Nectarinia rubescens rubescens (Vieillot). This sunbird is fairly common and widely distributed in the Bwamba Forest. The van Somerens ( 1949: 76) reported it chiefly in the more open parts of the forest and on the margins of the denser parts; our collectors met it in many places, and obtained specimens at Ntandi, 2300 feet, July, October, December; Mongiro, 2300 feet, November; and Bwamba, March; many other specimens are in the Williams sunbird collection; bill black; iris dark brown; feet black; weight 10 to 13 grams; stomach contents insects and spider fragments. Nectarinia senegalensis lamperti (Reichenow). Williams ( 1951 : 109) refers to several Bwamba specimens. Previously this sunbird had been reported from Bwamba by the van Somerens (1949: 76) as well. Nectarinia venusta igneiventris (Reichenow). v. S., 1949: 74. Nectarinia chloropygia orphogaster (Reichenow). The olive-bellied sunbird is very common in Bwamba, chiefly in the cultivated areas outside the true forest; we have many specimens : Ntandi, 2300 feet, June, July, November, and December; Bumaga, 2300 feet, July; Nkarara, 2200 feet, July; Rwansenge, 2200 feet, July; and Bwamba Forest, April. Some of the July birds had enlarged gonads; bill black; iris dark brown; feet black; weight 7 to 1 1.5 grams; stomach contents small insect remains. The van Somerens ( 1949: 74) found nesting pairs in April and July. Nectarinia minulla minuUa (Reichenow). First recorded for Bwamba by the van Somerens ( 1949: 75), many specimens were collected later by or for Williams, and are now in his sun- bird collection in Los Angeles; our present collectors obtained an additional seven specimens at Ntandi, 2300 feet, June 17 to July 10, and December 2. Most of the June- July birds were in breeding condition. This species fre- 1971 The Birds of the Lowlands of Bwamba 61 quents the second growth and the borders of the heavy forest, into the depths of which it seldom penetrates. Nectarinia bouvieri (Shelley). Williams (1951: 109) obtained two examples of this seldom collected sunbird at Ntotoro, 2250 feet, March 22, 1948, in secondary growth in the forest. No one else has noted it in Bwamba. Nectarinia cuprea cuprea (Shaw). The coppery sunbird is not found in the dense forest but at the edges of such areas, especially where there are numerous flowering trees. The van Somerens (1949: 76), Williams (1951: 109) and the Knudsen-Machris 1963 expedition (Friedmann, 1966: 47) all obtained specimens. Since then our collectors met with this species but once, at Ntandi, 2300 feet, July 12 to 18, when they obtained three examples, one in breeding condition, the other two not; bill black; iris dark brown; feet black; weight 9.5 grams; stomach contents small insects. Nectarinia purpureiventris (Reichenow). v. S., 1949: 74. Nectarinia erythrocerca Hartlaub. v. S., 1949: 74. Nectarinia superba buvuma (van Someren). Our collectors obtained specimens of this sunbird only at Ntandi, 2300 feet, June 29 to July 14, and November 7 to 11. Two of the November birds had enlarged gonads; bill black; iris dark brown; feet black; weight 17 to 19 grams; stomach contents insects. The van Somerens (1949: 76) found the superb sunbird at Bundibugyo and Kanyamwerima. This sunbird is more a bird of the borders rather than the depths of the heavy forest. It is much attracted to banana flowers in the cultivations along the forest margins. Nectarinia kilimensis kilimensis Shelley, v. S., 1949: 74. Family Ploceidae Passer griseus griseus (Vieillot). The common gray-headed sparrow is a bird of the open, cultivated areas, not of the forest; specimens were taken at Ntandi, 2300 feet, Nyankiro, 3000 feet, and “Bwamba Valley,” 2600 feet, in April, July, and November; bill black; iris light brown; feet brown; weight 28 to 32.5 grams; stomach contents seeds and grit. Amblyospiza albifrons melanota (Heuglin). Common in the grassy areas, not entering the forest. This bird has been noted by everyone who has written about Bwamba birds, and our specimens add nothing new; we have a series from several localities in Bwamba, April, July, October, and November; bill blackish horn in males, yellowish brown with blackish tip in females; iris dark brown; feet dark grayish brown; weight 62 Contributions in Science No. 211 40 to 44 grams; stomach contents seeds. A party of grossbeak weavers was seen feeding in a fruiting tree at Nyankiro, November 25. Ploceus baglafecht stuhlmanni (Reichenow). v. S., 1949: 94. Ploceus pelzelni pelzelni (Hartlaub). One specimen, with enlarged gonads, was collected at Rwansenge, 2200 feet, July 14; bill black; iris dark brown; feet gray with a browish tinge; weight 13 grams; stomach contents insects. The fact that our collectors met with this bird only once, coupled with the absence of any mention of it in the earlier Bwamba literature, suggests that it is an uncommon bird there. Ploceus ocularis crocatus (Hartlaub). v. S., 1949; 93. Ploceus nigricollis nigricollis (Vieillot). Fairly common; our field teams obtained a number of specimens at Korongo, Semliki River, 2200 feet, July 14; Ntandi, 2300 feet, November 1; and Mongiro, 2300 feet, November 8 to 12; bill black; iris dark brown; feet gray; weight 24 to 28 grams; stomach contents insects. Ploceus xanthops (Hartlaub). v. S., 1949: 94. Ploceus cucullatus bohndorffi Reichenow. A very common bird in the clearings, cultivated areas, and native villages. We have many specimens from Bwamba valley, 2600 feet, April; Humia, 2300 feet, July and December; Ntandi, 2300 feet, July and November. All earlier collectors in Bwamba have similarly found this weaver to be very numerous. Our specimens had the bill black; iris orange-red; feet pale brown; weight 38 to 45 grams; stomach contents seeds and some beetle and other insect fragments. Ploceus nigerrimus nigerrimus Vieillot. Another common weaver of the Bwamba lowlands, as noted long ago by the van Somerens (1949: 92). We have series of specimens from Ntandi, 2300 feet, June, October, and November; Mongiro, 2300 feet, November; and Bundimusuba, 3000 feet, July; bill black; iris dark red-brown to dark brown; feet pale brown to gray-brown; weight 27 to 33 grams; stomach contents seeds and insect remains. Ploceus melanocephalus fischeri Reichenow. This weaver is represented in the collection by two males taken at Rwansenge, 2200 feet, July 14 and 15; bill black; iris dark brown; feet pale brown; weight 26, 27 grams; stomach contents seeds, and insects. Ploceus tricolor interscapularis Reichenow. The van Somerens (1949: 92-93) considered this species as scarce, but admitted that it might have been overlooked in the forest. Our collectors obtained a series of 13 specimens at Ntandi, 2300 feet, June 13 to 21, Octo- 1971 The Birds of the Lowlands of Bwamba 63 her 26 to November 28; Mongiro, 2300 feet, November 7; and Bundimusuba, 3000 feet, July 12; bill black; iris dark red-brown to dark brown; feet pale brown to gray-brown; weight 27 to 33 grams; stomach contents seeds and small insects. At Ntandi, on November 24, eight or more of these weaver birds were watched as they flew into the upper branches of a tall ironwood tree, where they busily moved about searching for food among the pale green foliage at the tips of the branches. Ploceus al binucha holomelas Sassi. This weaver was first found in the Bwamba lowlands by Williams ( 1951 : 110) and seemed to be a scarce bird there. Our collectors obtained two speci- mens at Ntandi, 2300 feet, December 8; bill black; iris white; feet grayish black; weight 26 and 29.5 grams; stomach contents small caterpillars. These two individuals were shot from a large flock high up in the top- most foliage of tall trees in the ironwood forest. The birds were extremely shy, and were followed for some distance before the collectors could get within gun shot range. The birds remained wholly in the upper leafy branches and were difficult to see when perching, becoming visible chiefly when begin- ning to move. They had a constant chick chick note, apparently given by all the members of the party. Ploceus superciliosus (Shelley). This is a bird of the tall grass, not of the forest. The van Somerens ( 1949: 94) found it at Bundibugyo; our collectors obtained a few examples at Nkarara, 2200 feet, June 27 and July 2, and at Ntandi, 2300 feet, July 14. One of the Nkarara birds was in breeding condition; bill black on the maxilla, gray on the mandible; iris dark brown; feet pale brown; weight 22 to 23 grams; stomach contents seeds, but, in one case, mainly insect remains. Ploceus bicolor mentalis (Hartlaub). This weaver is not numerous, but is probably widely distributed in Bwamba. The van Somerens (1949: 94) found it in at least two localities, and we have two specimens from a third one— Bundimusuba, 3000 feet, November 20; bill blue-gray with the tomium darker; iris deep crimson; feet pale brownish flesh color; weight 32 and 34 grams; stomach contents insects. Malimbus nitens (Gray). Our collectors found this species in a number of localities, which makes it surprising that the van Somerens failed to report it. We have specimens from Nkarara, 2200 feet, July 1, 2; Rwansenge, 2200 feet, July 13; Korongo, Semliki River, 2200 feet, July 14; and Mongiro, 2300 feet, November 7 to December 10. At Mongiro three examples were obtained all of which were in breeding condition; bill pale bluish gray; iris red to red-brown; feet dark gray; weight 30 to 37 grams; stomach contents insects and small flat seeds. 64 Contributions in Science No. 211 A specimen taken at Mongiro on November 7 was watched building its nest on a hanging branch over a small stream. Another was seen feeding among oil palm fronds. The bird gave a warbling chrr-rr-rr note, as well as a sharper chrick call. This forest weaver was first reported from Bwamba, and, indeed, from Uganda, by Williams ( 1957: 159). Malimbus rubricoUis rubricollis (Swainson). Although the van Somerens (1949: 94) found this species throughout the lowland forest of Bwamba, our collectors met with it only once, at Bundi- musuba, 3000 feet, July 11 and 12, where three specimens, one in breeding state, were taken; bill black; iris dark brown; feet blackish; weight 39 to 44 grams; stomach contents ants (?), beetles, and other insects. Malimbus erythrogaster Reichenow. v. S., 1949: 95. Malimbus malimbicus malimbicus (Daudin). A numerous bird of the Bwamba forests; we have a long series from seven localities, 2600 to 3000 feet in elevation, taken in April, June, July, October, November, and December. Most were in non-breeding condition, but five out of fourteen taken from the end of October to early in December had enlarged gonads; bill black; iris dark brown; feet blackish gray; weight 26 to 43 grams; stomach contents insects, with many beetles in some cases. One female at Ntandi, December 7, was seen picking up fibers from a dead creeper, presumably for nest-building. Others were noted feeding high up in a tall tree. Quelea quelea aethiopica (Sundevall). v. S., 1949: 97. Euplectes gierowii ansorgei (Hartert). Reported as plentiful near Bundibugyo by the van Somerens (1949: 96), our collectors only once encountered this bird of the tall grasslands, at Mon- giro, 2300 feet, November 1, when they were able to collect a single specimen; bill black; iris dark brown; feet dark brown; weight 30 grams; stomach contents seeds. Euplectes hordeaceus craspedopterus (Bonaparte). Not a bird of the forest. One of these bishop birds was collected in Bwamba valley, 2600 feet, on April 19, by the Knudsen-Machris expedition; bill black; feet dark flesh color. Euplectes axillaris phoeniceus (Heuglin). v. S., 1949: 96. Euplectes ardens concolor (Cassin). v. S., 1949: 96. Family Estrildidae Parmoptila woodhousei jamesoni (Shelley). This seldom collected little bird was met with at Ntandi, 2300 feet. 1971 The Birds of the Lowlands of Bwamba 65 where four specimens were obtained, November 16 to December 4. One of them had enlarged gonads; the others did not; bill black; iris dark brown; feet whitish brown; weight 10 to 10.5 grams; stomach contents small insect remains. This little bird has a wide range of food foraging. One of them was seen hopping about in dense undergrowth about two feet from the ground, where it was with another individual, possibly its mate, apparently searching for food. Another was collected close to the forest floor, but a third one was shot about 30 feet up from the ground on a frond of an oil palm, where it was hanging upside down searching for insects. The species was noted as having a soft twee twee call note, and as being a fast flier. Keith ( 1968: 13-14) noted a Bwamba sight record by Smart. Nigrita fusconota fusconota Fraser. This species is found throughout the Bwamba forests. The van Som- erens (1949: 98) reported it from four localities, to which our collection adds a fifth. At Ntandi, 2300 feet, July 10 to 17 and again from November 12 to 16, nine specimens were obtained; six of the July birds were in breed- ing state; the November birds were not; bill black; iris dark brown; feet dark gray; weight 8 to 10 grams; stomach contents very small insects. Nigrita bicolor brunnescens Reichenow. Common in the outer fringes of the forest; our collectors obtained 13 specimens at Ntandi, 2300 feet, June 16 to July 31, while a single one was taken in the “Bwamba valley” April 23, by the Knudsen-Machris expedition. The birds taken at Ntandi varied in their gonadal state from small to much enlarged; bill black; iris dull red to dark brown; feet brown to grayish brown; weight 9 to 13 grams; stomach contents insect fragments. This species was first recorded from Bwamba by the van Somerens ( 1 949 : 99 ) , who thereby established its place in the Uganda avifauna. One of the Ntandi specimens, taken on July 5, was building a nest of dry leaves when collected. Nigrita luteifrons luteifrons J. and E. Verreaux. Apparently the least numerous species of its genus in the Bwamba low- lands, as it was not reported by the van Somerens’ collectors there. We have three specimens, all taken at Ntandi, July 12, November 11 and 12; bill black; iris yellowish white; feet brownish white; weight 11 to 12 grams; stomach contents insect fragments. This species was first reported from Bwamba by Williams (1957: 160). Keith (1968: 13) also obtained a speci- men there. Nigrita canicapilla sparsimguttata Reichenow. This is the commonest of the four species of Nigrita in the Bwamba low- lands, where it is found chiefly about the edges of the forest. It was reported by all the earlier workers on Bwamba ornithology, and our present series of 29 66 Contributions in Science No. 211 specimens adds little new. The Bwamba population is intermediate in char- acter between sparsimguttata and schistacea, but we place our birds under the former name to conform to the treatment in the last revision of the Estrildidae (Traylor, 1968: 310-389). Our specimens, taken in “Bwamba Forest,” 2600 feet; Ntandi, 2300 feet, and Bundimusuba, 3000 feet, in April, June, July, October, November, and December, had the bill blackish basally, reddish distally; iris orange-red to dull orange; feet dark brown; weight 16 to 21 grams; stomach contents fruit skins and some insect fragments. Nesocharis ansorgei ansorgei (Hartert). v. S., 1949: 101. Mandingoa nitidula schlegeli (Sharpe). The van Somerens (1949: 101) observed this little twin-spot only once, but our collectors obtained a long series of specimens at no less than seven localities, which indicates that the bird is abundant and widely dis- tributed in the Bwamba forests: Bwamba, 2600 feet, April; Bumaga, 2300 feet, June; Nkarara, 2200 feet, June and July; Ntandi, 2300 feet, June, July, October, November, December; Rwansenge, 2300 feet, July; Mongiro, 2300 feet, November; and Bundimusuba, 3000 feet, November. Birds with enlarged gonads were taken together with others in non-breeding state, in July and November; bill blackish basally, the distal half or more red; iris dark brown; feet brown; weight 10 to 12.5 grams; stomach contents small seeds. Pyrenestes ostrinus ostrinus (Vieillot). The black-bellied seed-cracker is common in the Bwamba woodlands, where our collectors obtained a dozen specimens at Ntandi, 2300 feet, June 18 to July 7, and December 10; and at Rwansenge, 2300 feet, July 13. Two of the June- July birds were in breeding state; bill metallic blue-gray, the tip darker; iris dark brown; feet brown; weight 18 to 27 grams; stomach contents crushed white seeds. Everyone who has had occasion to study specimens of this bird has been puzzled at the enormous size variability of their bills, and has been tempted to divide them into sympatric species. However, Chapin’s conclusions (1924 and 1954: 490-498) still seem the best we have, and the only ones that fit the biological requirements of the problem. As the van Somerens remarked (1949: 100) there is an almost parallel case in Amblyospiza. Spermophaga poliogenys (Ogilvie-Grant). This relatively uncommon species is a denizen of the deep forest, where it is apt to be either overlooked or confused by the observer with the next, and much commoner species. It was first reported from Bwamba by Williams (1951: 110) who thereby added it to the known avifauna of Uganda. We have a small series of specimens, all taken at Ntandi, 2300 feet, in July, November, and December; one of the July birds was in breeding state; the others had small gonads; bill metallic blue-gray, turning to orange at the tip 1971 The Birds of the Lowlands of Bwamba 67 and along the tomium; iris dark brown; feet grayish brown; weight 20 to 21 grams; stomach contents small seeds. Spermophaga ruficapilla ruficapilla (Shelley). Very common in the Bwamba forests; the van Somerens (1949: 99) noted it in many places and our collectors obtained many specimens at the following localities: “Bwamba,” April; Nkarara, 2200 feet, June and July; Ntandi, 2300 feet, June, July, October, November, and December; Bundi- musuba, 3000 feet, July and November; Mongiro, 2300 feet, November. In all these places and months birds with large gonads and others with small, resting ones, were taken together; bill basally blackish with a bright metallic blue wash, and becoming reddish or reddish purple distally; iris dark brown; feet brown with gray or olive wash, eye ring pale blue; weight 16 to 26.5 grams; stomach contents small seeds exclusively. An adult male with enlarged gonads, taken at Ntandi, November 20, was seen to fly into a dense thicket of dead scrubs with a stem of grass in its bill, presumably for nest-building. This suggests that the male takes part in the task of nest-building; hitherto nothing was recorded in print about the role of the sexes in nidification in this species, although the van Somerens collected a pair “. . . engaged in nest-building, though we did not know this till the female was shot. As we were going forward to pick her up, a male flew out from the base of an oil palm, and on searching the spot a collection of green material was found, hidden under leaves in the axils of drooping fronds. It was a rather large structure, domed and with a side entrance . . .” Lagonosticta rubricata congica Sharpe. This fire finch occurs in the grassy areas at the outside edge of the forest or in grassy clearings inside the forest. We have only a single example, a bird in breeding condition, taken at Nkarara, 2200 feet, on July 3. The van Som- erens ( 1949: 101) had reported the species from Bwamba many years earlier, but gave no indication as to its numerical status there. Estrilda melanotis kilimensis (Sharpe), v. S., 1949: 102. Estrilda paludicola paludicola Heuglin. The fawn-breasted waxbill is common in the bush and grass country close to the forests. We have a few specimens from “Bwamba,” April 23; Nkarara, 2200 feet, June 28; and Ntandi, 2300 feet, November 16; bill bright orange; iris red; feet dull gray brown; weight 5 to 8 grams; stomach contents small grass seeds. Although the limits of the ranges of typical paludicola and of the race roseicrissa are uncertain in western Uganda, we follow Chapin ( 1954: 538) and the van Somerens (1949: 101) in consid- ering the Bwamba population to be of the nominate subspecies. The Bwamba area is between the ranges of the two races as given by Traylor (1968: 339). 68 Contributions in Science No. 211 Estrilda astrild adesma Reichenow. The common waxbill is abundant in the open country of the Bwamba lowlands as it is in suitable areas in most of Africa. The only question is whether Bwamba birds are to be considered adesma or nyansae, or if the two are really separable. Chapin (1954; 547-548) treated adesma as a synonym of nyansae; we are following Traylor ( 1968: 342) in considering adesma distinct, as a matter of conforming to the latest revision, as we do not have the material on which to decide otherwise. Our collectors paid little attention to the birds outside the forest in Bwamba and obtained but a single specimen, at Ntandi, 2300 feet, December 2; bill red; iris dark brown; feet blackish; weight 7 grams; stomach contents grass seeds. Estrilda nonnula nonnula Hartlaub. The black-crowned waxbill is also a very common bird of the open cultivated areas adjacent to the forests. Our collectors obtained examples at Nkarara, 2200 feet, June 29; Ntandi, 2300 feet, June 23 and 26; and at Bundimusuba, 3000 feet, November 23; the Ntandi birds were in breeding condition; the others were not; bill black with the sides of the culmen and the base of the mandible dull red; iris dark brown; feet black; weight 7 to 8 grams; stomach contents grass seeds. Amandava subfiava subflava (Vieillot). v. S., 1949: 102. Lonchura cucullata cucullata (Swainson). The little bronze mannikin is very common in the open areas outside of the forest. We have a series of specimens from Ntandi, 2300 feet, June 15 to July 19, and October 30 to November 20; and from Mongiro, 2300 feet, November 13; bill black on maxilla, gray on mandible; iris dark brown; feet blackish; weight 8 to 1 1 grams; stomach contents grass seeds. Lonchura bicolor poensis (Fraser). As pointed out by the van Somerens ( 1949: 98) the black and white mannikin is more of a dweller in the forest margins and less in the open cul- tivated areas than is the bronze mannikin. Our collectors obtained specimens only at Ntandi, 2300 feet, June 15, and October 29 to November 29; mostly in breeding state; bill pale blue-gray; iris dark brown; feet black; weight 10 grams in every case; stomach contents grass seeds. Lonchura fringilloides (Lafresnaye). When the van Somerens ( 1 949 : 97) first recorded the magpie mannikin from the cultivated, open areas of Bwamba, it was a “new” bird to Uganda. It still is a relatively seldom reported species, but our collectors obtained a small series at the following localities : Ntandi, 2300 feet, June 19 and Octo- ber 26 to November 15; Nkarara, 2200 feet, July 13; Rwansenge, 2300 feet, July 14; and Humia, 2300 feet, December 4; some of the July and November 1971 The Birds of the Lowlands of Bwamba 69 birds had enlarged gonads; the others had small ones; bill black on the maxilla, pale blue-gray on the mandible; iris dark brown; feet dark gray; weight 17 to 20 grams; stomach contents small seeds. Vidua macroura (Pallas), v. S., 1949: 97. Family Fringillidae Serinus frontalis frontalis Reichenow. v. S., 1949: 103. Serinus mozambicus barbatus (Heuglin). v. S., 1949: 103. Serinus sulphuratus shelleyi Neumann, v. S., 1949: 103. Literature Cited Bannerman, D. a. 1921. Remarks on rare and otherwise interesting birds con- contained in collections made by Mr. G. L, Bates in southern Cameroon. Ibis, Ser. 11, 3: 81-121. Bates, G. L. 1911. Further notes on the birds of southern Cameroon. Part 1. Ibis, Ser. 9, 5: 479-545. Berger, A. J. 1955. On the anatomy and relationship of Glossy Cuckoos of the genera Clirysococcyx, Lampromorpha and Chaleites. Proc. U.S. Nat. Mus. 103; 585-597. Chapin, J. P. 1924. Size variation in Pyrenestes, a genus of weaverfinches. Amer. Mus. Nat. Hist. Bull. 49: 415-441. 1953. The birds of the Belgian Congo. Part III. Amer. Mus. Nat. Hist, Bull. 75A: 1-821. 1954. The Birds of the Belgian Congo. Part IV. Amer. Mus. Nat. Hist. Bull. 75B: 1-846. 1963. The supposed “gray mutants” of Terpsiphone viridis. Ibis 105: 198-202. Eggling, W. j. 1954. The birds of Bwamba. Uganda Journ. 18: 198-199. Friedmann, H. 1955. The Honey-guides. U.S. Nat. Mus. Bull. 208. 292 p. — 1966, A contribution to the ornithology of Uganda. Bull. Los Angeles County Mus. Nat. Hist., Science 3: 1-55. 1968a. Zenker’s honey-guide. Journal f. Omith. 109: 276-283, — 1968b. Range and variation of the icterine bulbul in Uganda. Bull, Brit. Ornith. Club 88: 110-112. 1969a. Additions to knowledge of the yellow-throated glossy cuckoo, Chrysococcyx flavigularis. Journ. f. Ornith. 110: 176-180. 1969b. The status and distribution in Uganda of the white-bellied king- fisher, Alcedo leucogaster leopoldi. Los Angeles County Mus., Contrib. Sci. 158: 1-6. in press. Phenotypic potential and speciation in Indicator and Prodotiscus. Proceedings Third Pan-African Ornithological Congress. Friedmann, H., and J. G. Williams. 1968. Notable records of rare or little-known birds from western Uganda. Rev, Zool. Bot. Afr. 77, nos. 1-2: 11-36. 1969. The birds of the Sango Bay Forests, Buddu County, Masaka District, Uganda. Los Angeles County Mus., Contrib. Sci. 162: 1-48. 70 Contributions in Science No. 211 1970. Additions to the known avifauna of the Bugoma, Kibale, and Im- penetrable Forests, west Uganda. Los Angeles County Mus., Contrib. Sci. 198: 1-20. Keith, G. S. 1968. Notes on birds of East Africa, including additions to the avifauna. Amer. Mus. Novitates 2321. Keith, G. S., and A. Twomey. 1968. New distributional records of some east Afri- can birds. Ibis 110: 537-548. Moreau, R. E. 1966. The bird faunas of Africa and its islands. Academic Press, New York, London. 424 p. Ogilvie-Grant, W. R. 1910. Ruwenzori expedition reports. 16. Aves, Trans. Zool. Soc. London 19: 253-480. Prigogine, a. 1965. Notes sur quelques Geokichla de la Republique du Congo. Rev. Zool. Bot. Afr. 7 1 : 3-4. Rand, A. L. 1958. Notes on African bulbuls. Fieldiana, Zool. 85, no. 6: 145-220. 1960. African Pycnonotidae. in Peters, Check List of birds of the world, 9: 221-300. Ridley, M. W., H. E. Percy, and R. C. Percy. 1953. The birds of Bwamba. Further additions. Uganda Journ. 17: 161-165. Traylor, M. A. 1968. Estrildidae (African), in Peters, Check List of birds of the world, 14: 306-397. VAN SoMEREN, V. G. L., and G. R. C. van Someren. 1949. The birds of Bwamba. Uganda Journ. 13 (Special Suppl. ) : 1-111. Vaurie, C. 1949. A revision of the bird family Dicruridae. Amer. Mus. Nat. Hist. Bull 93, art. 4: 203-342. Weekes, J. T. 1949. The birds of Ruwenzori. Uganda Journ. 13, no. 2: 130-144. White, C. M. N. 1965. A revised check list of African non-passerine birds. Lusaka, Govt. Printer. 299 p. Williams, J. G. 1951. The birds of Bwamba: some additions. Uganda Journ. 15: 107-111. 1957. Four new bird records for eastern Africa. Bull. Brit. Omith. Club 77: 159-160. Accepted for publication October 20, 1970 Printed in Los Angeles, California by Continental Grap i ' LOS ANGELES ) COUNTY i! 1MUSEUM 1 i 1 CONTRIBUTIONS IVii’S IN SCIENCE J viBER 212 March 31, 1971 tl; NOTES ON THE BAT FLIES OF COSTA RICA ! (Diptera: Nycteribiidae) l! ji I By B. V. Peterson ji i| 1 ;l l| i! j! i' 'I !| Lo^ Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 NOTES ON THE BAT FLIES OF COSTA RICA (Diptera: Nycteribiidae) By B. V. Peterson^ Abstract: Five species of Nycteribiidae are now known from Costa Rica of which Basilia ortizi, B. rondanii, plus one undetermined species, Basilia sp. (c), are recorded for the first time. Basilia myotis is synonymized with B. ferrisi, and a lecto- type is designated for the latter species. A key to the females and males of the Costa Rican species of Basilia is also provided. The family Nycteribiidae is represented in the New World by about 45 species including three new species now in press. The latest treatment of the American genera and species is found in Theodor’s (1967) comprehen- sive treatment of this family. Only two species of Basilia have been reported from Costa Rica, viz. B. costaricensis Guimaraes and B. ferrisi Schuurmans Stekhoven Jr. The study of eight recently collected specimens shows that at least five species occur there. The purpose of this paper is to report the results of the study of these eight specimens plus the eight specimens of the type series of Basilia myotis Curran as well as 17 of the type series of 21 specimens of B. ferrisi Schuurmans Stekhoven Jr., and to provide a key to the species of bat flies known from Costa Rica. I extend my thanks to the following individuals who provided the speci- mens used in this study: Dr. C. L. Hogue, Los Angeles County Museum of Natural History, Los Angeles, California; Dr. A. Stone, Systematic Ento- mology Laboratory, Agricultural Research Service, U. S. Department of Agriculture, Washington, D. C.; Dr. P. W. Wygodzinsky, Department of Entomology, American Museum of Natural History, New York, New York; and Dr. J. R. Anderson, Division of Parasitology, University of California, Berkeley, California. This paper is, in part, a result of research supported by U. S. Army Medical Research and Development Command Grant DA-MD- 49-193-63-G94, E. S. Truxel and C. A. McLaughlin, Principal Investigators, Los Angeles County Museum of Natural History. Key to the Costa Rican Species of Basilia Females 1. Posterior margin of mesonotum without a dorsal, upright, digitiform pro- cess. Tergal plate I elongate, somewhat triangular, its hind margin narrow and sides straight or slightly concave; long setae of hind margin reaching ^Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario. 1 2 Contributions in Science No. 212 or crossing the hind margin of tergal plate II. Posterior margin of tergal plate II broadly rounded, not produced into lobes. ferrisi Posterior margin of mesonotum with a variable sized, dorsal, upright, digitiform process. Tergal plate I shorter, more rectangular or broadly rounded along hind margin; long setae of hind margin shorter, at most, reaching about middle of tergal plate II. Posterior margin of tergal plate II produced into at least short lobes whose posterolateral margins are some- what concave 2 2. Sternite VI longitudinally divided on the midline. Digitiform process of posterior margin of mesonotum short, triangular. Tergal plate I moderately long, hind margin broadly rounded and entirely fringed with long setae, this fringe with only a narrow gap in the middle. Posterior margin of tergal plate II with two short, broad, distinct lobes which are separated by a broad notch costaricensis Sternite VI entire, not longitudinally divided on the midline. Digitiform process of posterior margin of mesonotum elongate. Tergal plate I short, its hind margin broadly rounded and with fringe of short setae laterally, or hind margin produced as two very short lobes with long setae only on these lobes. Posterior margin of tergal plate II produced into long, or short and narrowly separated lobes 3 3. Posterior margin of tergal plate I produced into two very short lobes, each bearing about 7-9 long setae. Tergal plate II produced posteriorly into two moderately long, slender lobes ortizi Posterior margin of tergal plate I broadly rounded, and fringed with short setae, this fringe with a wide gap in the middle. Tergal plate II produced posteriorly into two short, moderately broad, rounded lobes which are separated by a small, shallow notch rondanii Males! 1. Posterior margin of synsternite V + VI with about 30-38 short spines in two or three rows. Aedeagus parallel-sided, with dorsal serrations near base; apex triangular, with its angle ventral in position rondanii Posterior margin of synsternite V + VI with about 13-22 short spines in two rows. Apex of aedeagus not as above 2 2. Sternites III-IV with numerous discal setae in addition to those on their hind margins. Aedeagus without dorsal or ventral serrations 3 Sternite III usually with only a few scattered discal setae; sternite IV with- out discal setae, setae present only along its hind margin. Aedeagus with serrations or at least an indication of serrations (not known for ortizi) 4 !This key to the males must be considered tentative until more specimens are avail- able for study. 1971 Notes on the Bat Flies of Costa Rica 3 3. Posterior margin of synsternite V + VI with about 20 short spines. Only tergite II with discal setae; tergites III-VI without discal setae. Aedeagus broad, parallel-sided, tapering apically to a rounded end which curves ventrally costaricensis Posterior margin of synsternite V + VI with 13 short spines. Both ter- gites II-III with discal setae, those of tergite III sparse centrally, more dense laterally; tergites IV-VI without discal setae. Aedeagus slender, parallel-sided, apex broadly rounded with a short but distinct ventral projection Basilia sp. (c) 4. Tergites II-VI without discal setae except for a few setae at sides of tergite II. Aedeagus broad, parallel-sided or widening distally, with at least indica- tions of serrations dorsally and ventrally, apex broadly rounded. . .ferrisi Tergite II with more numerous, scattered discal setae; tergites III-VI with- out discal setae. Form of aedeagus not known ortizi Basilia costaricensis Guimaraes and D’Andretta Basilia costaricensis Guimaraes and D’Andretta, 1956, Arq. Zool. Est. 5. Paulo 10:106; figs. 146, 164, 169-174, 178 (female, male). This species, known only from Costa Rica, was described from one male and two females collected at Bebedero, near Las Canas, Guanacaste Province, 12-VI-1930, by Reimoser, from an undetermined bat. The holotype female and allotype male are deposited in the collection of the Naturhis- torisches Museum, Vienna, Austria, and the female paratype apparently is in the British Museum (Natural History) (Theodor, 1967). To my knowledge, additional specimens have not been collected. Basilia ferrisi Schuurmans Stekhoven Jr. Basilia ferrisi Schuurmans Stekhoven Jr., 1931, Zeitschr. Parasitenk. 3:217 (new name for B. speiseri of Ferris, 1924, not Miranda Ribeiro). Scott, 1936, J. Linn. Soc. Lond., Zool. 39:502 (new name for B. speiseri of Ferris, 1924, not Miranda Ribeiro). Scott, 1939, Allan Hancock Pacific Exped. 2:168 (citation). Scott, 1940, Proc. R. Ent. Soc. Lond. (B) 9:61 (correction of author of the species). Guimaraes, 1940, Papeis Avulsos Depto. Zool. 1:5-7. (correction of author of the species). Schuurmans Stek- hoven Jr., 1942, Beit. Fauna Perus 3:100 (citation). Del Ponte, 1944, An. Inst. Med. Reg., Tucuman 1:118 (citation and key). Guimaraes, 1946, Arq. Zool. Est. S. Paulo 5:16 (citation and key). Guimaraes and D’Andretta, 1956, Arq. Zool. Est. S. Paulo 10:75 (distribution and comments). Maa, 1965, J. Med. Ent. 1:380 (citation, in subgenus Pseudelytromyia) . Gui- maraes, 1966, In Ectoparasites of Panama, p. 399 (citation). Theodor, 1967, Brit. Mus. (Nat. Hist.) Publ. 655:278; fig. 429 (citation and key). Gui- maraes, 1968, In A Catalogue of the Diptera of the Americas south of the United States 101:2 (citation). 4 Contributions in Science No. 212 Basilia speiseri, Ferris, not Miranda Ribeiro, 1924, Ent. News 35:198; figs. A-D (female, male). Stiles and Nolan, 1931, Bull. Nat. Inst. Health 155:648 (part) (citation). Curran, 1935, Amer. Mus. Novit. 765 :4 (part) (citation). Basilia myotis Curran, 1935, Amer. Mus. Novit. 765:3; figs. 3-5 (female, male) New Synonymy. Scott, 1936, J. Linn. Soc. Lond., Zool. 39:497 (cita- tion). Bequaert, 1942, Bol. Ent. Venezolana 1:84 (new distribution). Del Ponte, 1944, An. Inst. Med. Reg., Tucuman 1:118 (citation and key). Gui- maraes, 1946, Arq. Zool. Est. S. Paulo 5: 16 (citation and key). Guimaraes and D’Andretta, 1956, Arq. Zool. Est. S. Paulo 10:76; figs. 85, 106-1 11, 124, 129, 145 (female, male; distribution and comments). Peterson, 1960, Proc. ent. Soc. Ont. 90 (1959) :31; figs. 7-8 (female; key). Maa, 1965, J. Med. Ent. 1:380 (citation, in subgenus Pseudelytromyia) . Guimaraes, 1966. In Ectoparasites of Panama, p. 398 (comments and key). Theodor, 1967, Brit. Mus. (Nat. Hist.) Publ. 655:281; figs. 422, 441, 480-481 (female, male) . Guimaraes, 1968. In A Catalogue of the Diptera of the Americas south of the United States 101 : 3 (citation). Basilia bellardii, Schuurmans Stekhoven Jr., not Rondani, 1931, Zeitschr. Parasitenk. 3:207; figs. 1-6 (female, male). Hase, 1931, Zeitschr. Parasitenk. 3:220; figs. 1-17 (habits). Hase, 1931, Zeitschr. Parasitenk. 3:836 (correct name of host). Scott, 1936, J. Linn. Soc. Lond., Zool. 39:497 (part) (cita- tion). Bequaert, 1942, Bol. Ent. Venezolana 1:83 (new distribution). Gui- maraes, 1946, Arq. Zool. Est. S. Paulo 5:62; figs. 73-80 (female, male) (part). Karaman, 1948, Rad, Acad. Yougosl. 273: 123; figs. 4a-b (female, male). Guimaraesia bellardii, Schuurmans Stekhoven Jr., 1951, Acta Zool. Lilloana 12: 112; fig. 4a-b (female, male; transfer to new genus, and rede- scription). There has been much confusion in the past concerning the identity of this species as is evident from the above reasonably complete list of synonymy. Ferris, in 1924, redescribed and illustrated both sexes of a species he identified as Basilia speiseri (Miranda Ribeiro). He based his notes on eleven females and ten males from Myotis nigricans, collected in Sipurio, Costa Rica, and indicated these specimens were from the collection of the U.S. National Museum. (The name Pseudelytromyia speiseri Rib. is written on the labels of the slides on which these specimens are mounted, while in his paper Ferris used Basilia for the generic name. ) Schuurmans Stekhoven Jr. (1931) in comparing B. bellardii (considered by later authors to be B. myotis) with B. speiseri noted that B. speiseri of Ferris and B. speiseri as described by Miranda Ribeiro himself, were probably different species. Accordingly, Schuurmans Stekhoven Jr. named the species of Ferris B. ferrisi. In his discussion of B. speiseri, Curran (1935), apparently unaware of Schuurmans Stekhoven Jr.’s paper, also pointed out that Ferris’ specimens were probably of another species. Curran described and illustrated B. myotis in the same 1971 Notes on the Bat Flies of Costa Rica 5 paper and stated that the female differed markedly from B. speiseri as figured by Ferris. After examining Ferris’ specimens, Scott (1936) agreed with Cur- ran that this species was distinct from B. speiseri, and proposed for it the name B. ferrisi. Scott (1939) accredited this name to himself, but in April 1940, he published a note recognizing Schuurmans Stekhoven Jr.’s earlier proposal of the name. In November of 1940, Guimaraes published the priority of B. ferrisi, accredited the name to Schuurmans Stekhoven Jr., and listed the appropriate synonymy. Guimaraes and D’Andretta (1956) suggested that B. myotis Curran might be a synonym of B. ferrisi. Although both Scott (1936) and Guimaraes (Guimaraes and D’Andretta, 1956) examined syntypes of B. ferrisi, neither selected a lectotype for the species, and Guimaraes was not certain of the possible synonymy of B. myotis with B. ferrisi. I have examined 17 of the original 21 syntypes (10 11$$) of this species. The female in the best state of preservation, mounted on a slide and bearing the data “Sipurio, Costa Rica, from Myotis nigricans,” has been labelled and is here designated the Lectotype of B. ferrisi. The lectotype is in the collection of the United States National Museum, Washington, D.C. Of the remaining 20 syntypes, 18 have been labelled Paralectotypes and are distributed as follows : 3 $ $, 2 $ $ , U.S. National Museum; 5 $ $ , 6 $ $ , California Insect Survey, Berkeley; 1 1 $ , British Museum (Natural History), London. The location of the two remaining paralectotypes is unknown. Individually the syntypes are in too poor condition to enable the reso- lution of the suggested synonymy of B. myotis with B. ferrisi. However, when examined as a group the critical characters are evident and have enabled the comparison of the species with the type specimens of B. myotis from the American Museum of Natural History. All of Ferris’ specimens were mounted on slides and it was necessary to remove a male (USNM) to examine the genitalia. The slight differences in the male genitalia fall within the normal range of variation (Figs. 1-2) (see also Figs. 422, 429 of Theodor, 1967), and no other significant differences could be found in either sex. It is evident that B. myotis Curran is the same as B. ferrisi Schuurmans Stekhoven Jr. Additional specimens of B. ferrisi examined include one male and one female from Playvelas (? Playavelas), Rio Frio, Alajuela Province, 23-V- 1964, F. S. Truxal et al., from Myotis species, and one male labelled “Costa Rica, Zent, H. S. Blair, Aug. 1915”. Basilia ortizi Machado-Allison Basilia ortizi Machado-Allison, 1963, Acta Biol. Ven. 3:455; figs. 1-6 (female, male). This species, from the bat Eptesicus melanopterus, was known only from the holotype female and one male paratype collected in Serrania de Nuvia, Bolivar State, Venezuela, 5-VIII-1962, by J. Ojasti. I know of no additional published records for this species. However, Maa (1965) included 6 Contributions in Science No. 212 Figures 1 and 2. Basilia ferrisi, male genitalia showing intraspecific variation. 1, from paratype of B. myotis { — ferrisi); 2, from paralectotype of B. ferrisi. 1971 Notes on the Bat Flies of Costa Rica 7 it in his interim world list of species, and in 1968 he mentioned it in compari- son with two species he described as new. Machado-Allison (1967) men- tions it in connection with “la Biota Amazonica”, and Guimaraes (1968) also cited it in his catalog of the species occurring south of the United States. One female of B. ortizi was taken from Eptesicus gaumeri collected at Boca de Barranca, Puntarenas Province, 30-VII-1962, F. S. Truxal. Dr. Machado-Allison kindly compared this specimen with the type female of B. ortizi and stated (personal communication), “The female does not show characteristics that would enable its separation from B. ortizi, ... I have no doubts ... it belongs to B. ortizi". Basilia rondanii Guimaraes and D’Andretta Basilia rondanii Guimaraes and D’Andretta, 1956, Arq. Zool. Est. S. Paulo 10:50; figs. 39-40, 42, 44, 47-51, 53, 55, 79 (female, male). Three specimens were available for study: one male from Volcan Poas, Alajuela Province, 15-VIII-1963, from Myotis nigricans, A. Starrett; and one male and one female, 13.5 mi S Liberia, Guanacaste Province, 3-VIII-1964, C. L. Hogue et al., from Myotis species. Basilia rondanii has been recorded from southwestern United States, Mexico, Guatemala and Honduras. Basilia sp. (c) Guimaraes and D’Andretta Basilia sp. (c) Guimaraes and D’Andretta, 1956, Arq. Zool. Est. S. Paulo 10:125; fig. 211 (male). A single male from Eptesicus species, collected near Atenas, Alajuela Province, 13-VIII-1963, by A. Starrett, appears to be the same as the male specimen that Guimaraes and D’Andretta (1956) listed as Basilia sp. (c), from Centurio senax from Guatemala. The two specimens seem to be iden- tical in chaetotaxy, in the shape of the aedeagus and paramere, and both have 12 spines on the hind margin of synsternite V + VI. Both specimens, however, were taken from hosts of different families, but Centurio senax and several species of Eptesicus are known to occur in the same habitat. Basilia sp. (c) keys to B. silvae (from Chile) in Theodor’s 1967 key, but as mentioned by Guimaraes and D’Andretta (1956) the aedeagus of B. silvae is longer and more pointed. The species is also close to B. tiptoni Guimaraes (1966) but only has eight notopleural setae instead of 12, and has 65 spines in the ctenidium of synsternite I + II as opposed to 45 spines in B. tiptoni. Other minor differences in the chaetotaxy of the two species exist as well. Guimaraes and D’Andretta suggested the possibility that Basilia sp. (c) might be the undescribed male of their B. anomala, a species known from Guatemala and Mexico from the bat Rhogeessa tumida. An association of the two sexes will be necessary before this possibility can be resolved. 8 Contributions in Science No. 212 Resumen En Costa Rica se conocen actualmente cinco especies, de las cuales la Basilia ortizi, la B. rondanii y una especie no determinada, la Basilia sp. (c) han sido registradas por primera vez. A la Basilia myotis se le ha dado el sinonimo de B. ferrisi, y se designa un tipolectivo para ultima especie. Tambien se proporciona una clave de machos y hembras de las especies costarricenses. Literature Cited Curran, C. H. 1935. New species of Nycteribiidae and Streblidae (Diptera). Amer. Mus. Novit. 765:1-15. Ferris, G. F. 1924. The New World Nycteribiidae (Diptera Pupipara). Ent. News 35:191-199. Guimaraes, L. R. 1940. Sobre a prioridade de Basilia ferrisi Schuurmans Stek- hoven, 1931. Papeis Avulsos Depto. Zool. 1:5-7. — . 1966. Nycteribiid batflies from Panama (Diptera: Nycteribiidae), p. 393-404. In R. L. Wenzel et al. (eds.). Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago. 1968. Family Nycteribiidae, 101:1-7. In N. Papavero (ed.). A catalogue of the Diptera of the Americas south of the United States, Depto. Zool., Sect, Agr. Est, S. Paulo. Guimaraes, L. R., and M. A. V. D’Andretta. 1956. Sinopse dos Nycteribiidae (Diptera) do Novo Mundo. Arq. Zool. Est. S. Paulo 10: 1-184. Maa, T. C. 1965. An interim world list of batflies (Diptera: Nycteribiidae and Streblidae). J. Med. Ent. 1:377-386. 1968. New Basilia species from Thailand, Mexico and Brazil (Diptera: Nycteribiidae). Pacif. Ins. 10:25-32. Machado-Allison, C. E. 1967. Sobre algunos ectoparasitos de mamiferos de la biota amazonica, p. 365-372. In H. Lent (ed.). Atas do simposio sobre a biota amazonica Vol. 5 (Zoologia). Schuurmans Stekhoven Jr., J. H. 1931. Eine Seltene, Ungeniigend Beschriebene Basilia- Art (Diptera Pupipara) aus Venezuela. Zeitschr. Parasitenk. 3:205-219. Scott, H. 1936. Descriptions and records of Nycteribiidae (Diptera Pupipara), with a discussion of the genus Basilia. J. Linn. Soc. Lond., Zool. 39:479-505. 1939. A new species of Nycteribiidae (Diptera Pupipara) from islands in the Gulf of California. Allan Hancock Pacific Exped. 2:167-170. 1940. Nycteribiidae from southern Brazil (Diptera). Proc. R. Ent. Soc. Lond. (B) 9:57-61. Theodor, O. 1967. An illustrated catalogue of the Rothschild collection of Nycte- ribiidae (Diptera) in the British Museum (Natural History). Brit. Mus. (Nat, Hist.) Publ. 655:1-506. Accepted for publication November 10, 1970 r j v/V -f J-'- Printed in Los Angeles, California by Continental Or. LOS ' ANGELES I COUNTY MUSEUM i' jUMBER 213 CONTRIBUTIONS cI'Lui IN SCIENCE May 27, 1971 TYPE SPECIMENS OF RECENT MOLLUSKS IN THE ‘ LOS ANGELES COUNTY MUSEUM OF NATURAL HISTORY I |. j By Gale G. Sphon i ! Il i 1 ! Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 TYPE SPECIMENS OF RECENT ' MOLLUSKS IN THE LOS ANGELES COUNTY MUSEUM OF NATURAL HISTORY By Gale G. Sphon^ Abstract: Type specimens of recent mollusks in the Los Angeles County Museum of Natural History are listed by the original published names. The original reference, the museum type number, and the locality are given. A figure with five previ- ously unfigured holotypes is included. Introduction This list is published in compliance with Recommendation 72D of the International Commission on Zoological Nomenclature ( 1964:75). It includes a total of 338 entries: 127 holotypes, 18 syntypes, and 193 paratypes of recent mollusks in the Los Angeles County Museum of Natural History. The entries marked by an asterisk (*) are part of the Allan Hancock Foundation (Univer- sity of Southern California) collection of gastropods on loan to the Los Angeles County Museum of Natural History. The list is arranged by class and alphabetically by the genus in which the species or subspecies was originally described. Each entry contains the author, date and bibliographic reference for the species or subspecies, the type category and number, and the locality. We assign each holotype a sepa- rate number. Paratypes are given a number for the entire lot unless they are from different localities. Definitions of type categories by Schenk, McMasters, Keen and Muller (1956:5-9) and the International Commission on Zoological Nomenclature (1964) are followed in this list. Pelecypoda Aligena redondoensis T. Burch T. BURCH, 1941:50 2 Paratypes 1010; 75 Fms off Redondo Beach, Los Angeles County, California Astarte vernicosa Dali DALL, 1903:948 3 Paratypes 1204; 10-16 Fms, Nazan Bay, Atka Is., Alaska Benthocardiella pusilla Powell POWELL, 1930:533 2 Paratypes 1361; 10-12 Fms, Wanganui, New Zealand ^Curatorial Assistant, Invertebrate Zoology, Los Angeles County Museum of Natural History. 1 2 Contributions in Science No. 213 Callocardia catharia Dali BALL, 1902:402 7 Paratypes 1205; 20 Fms, Guaymas, Sonora, Mexico Cardita umnaka Willett WILLETT, 1932c: 87 Holotype 1034; Umnak Is., Alaska Cardita ventricosa montereyensis Smith and Gordon SMITH and GORDON, 1948:212 2 Paratypes 1002; 63 Fms, NW of Point Pinos, Monterey Bay, Monte- rey County, California Cardita ventricosa redondoensis T. Burch T. BURCH, 1944:14 3 Paratypes 1299; 75 Fms, Redondo Beach, Los Angeles County, Cali- fornia Corbula luteola rosea Williamson WILLIAMSON, 1905b: 120 Syntypes 1421 (2 valves) ; Terminal Is, Los Angeles County, California Ensis myrae Berry BERRY, 1953a: 398 Paratype 1 104; Terminal Is, San Pedro, Los Angeles County, Cali- fornia Glycimeris keenae Willett WILLETT, 1943:114 Holotype 1065; 30 Fms, Forrester Is, Alaska Heterodonax bimaculatus purpureus Williamson WILLIAMSON, 1892:187 Syntypes 1176 (9 pair and 2 valves); San Pedro, Los Angeles County, California Heterodonax bimaculatus salmoneus Williamson WILLIAMSON, 1892:187 Syntypes 1 177 (7 pair and 1 1 valves); San Pedro, Los Angeles County, California Lasaea nipponica Keen KEEN, 1938:26 Paratype 1079; Watanoha, Matsusima, Japan Legrandina aucklandica Powell POWELL, 1933:233 Paratype 1356; Faith Harbour, Auckland Is, New Zealand Limopsis diegensis Dali BALL, 1908:203 5 Paratypes 1166; 80 Fms, off San Diego, San Diego County, California 1971 Type Specimens of Recent Mollusca 3 S) ^ . Q u ^ O O 43 3 a £•§ S 5 33 0\ s' «5 P-Si e c 43 •S3 ^ Q I/-J iT ' ^ S 43 *J zi m - O 43 Os •S os o so Pu "4 Willett, 1935a, holotype; Diam. 16.8mm, Alt. 10.4mm, AV2 whorls. Figs. 10-12, Micrarionta chocolata Willett, 1935a, holotype; Diam. 16.7mm, Alt. 10.3mm, AVi whorls. Figs. 13-15, Micrarionta granitensis Willett, 1935b, holotype; Diam. 14.8mm, Alt. 8.5mm, AVi whorls. Figs. 16-18, Micrarionta brunnea Willett, 1935a, holotype; Diam. 13.5mm, Alt. 7.9mm, 41/3 whorls. Figs. 19-21, Micrarionta mccoiana Willett, 1935b, holotype; Diam. 12.8mm, Alt. 7mm, 4% whorls. Photographs by Armando Solis. 4 Contributions in Science No. 213 Lithophaga (Labis) attenuata rogersi Berry BERRY, 1957:76 6 Paratypes 1 1 17; NW side of Cholla Cove, Sonora, Mexico Macoma inflatula Dali DALE, 1897:11 2 Paratypes 1328; 12 Fms, Captain’s Bay, Unalaska, Alaska Musculus phenax Dali DALE, 1915:138 17 Paratypes 1077; St. George Is, Pribiloff Group, Alaska Nucula pigafettae Dali DAEE, 1908:203 Paratypes 1167 (4 valves); 677 Fms, off Valdivia, Chile Nuculana burchi Willett WIEEETT, 1944:71 Holotype 1066; 50 Fms, Redondo Beach, Eos Angeles County, Cali- fornia Nuculana redondoensis J. Q. Burch J. Q. BURCH, 1944:9 Paratypes 1316 (5 pairs and 5 valves); Redondo Beach, Eos Angeles County, California Pecten (Cyclopecten) catalinensis Willett WIEEETT, 193 lb: 65 Holotype 1020; 100 Fms, off White’s Eanding, Santa Catalina Is, Eos Angeles County, California Penitella fitchi Turner TURNER, 1955:71 2 Paratypes 1444; North side of Bahia San Bartolome, Baja California, Mexico Pseudochama clarionensis Willett WIEEETT, 1938b:48 Holotype and paratype 1058; 30 Fms, SW side of Clarion Is, Revilla- gigedo Is, Colima, Mexico Yoldia ensifera Dali DAEE, 1897:9 3 Paratypes 1327; 37 Fms, Monterey Bay, Monterey County, California Yoldia montereyensis Dali DAEE, 1893:29 4 Paratypes 1125; 382 Fms, Monterey Bay, Monterey County, Cali- fornia 1971 Type Specimens of Recent Mollusca 5 Gastropoda Acanthina tyrianthina Berry BERRY, 1957:78 Paratype 1007; Man of War Cove, Magdalena Bay, Baja California, Mexico Acmaea (Collisella 1) apicina Dali DALE, 1879:341 2 Paratypes 1425; East side of Simeohoff Is, Shumagin Is, Alaska “Acmaea” stanfordiana Berry BERRY, 1957:76 3 Paratypes 1012; Punta Penasco, Sonora, Mexico Acteocina culcitella intermedia Willett WILLETT, 1928:38 Holotype 1015; 30 Fms; off Santa Catalina Is, Los Angeles County, California Acteon punctocoelata vancouverensis I. Oldroyd I. OLDROYD, 1927:25 Paratype 1078; 10-15 Fms, off Brandon Is, Departure Bay, British Columbia, Canada Agaronia murrha Berry BERRY, 1953b:417 Paratype 1105; Corinto, Nicaragua A lei t hoe arabica jaculoides Powell POWELL, 1924:108 Paratype 1320; 40-50 Fms, off Whakatane, New Zealand A lectrion limacina Dali DALL, 1917:577 2 Paratypes 1441; 18 Fms, near La Paz, Baja California, Mexico Allogona lombardi A. G. Smith A. G. SMITH, 1943:545 2 Paratypes 1221; Along Meadow Creek, 1.5 mi S of Selway Falls, Idaho County, Idaho *Amaea (s.l.) context a DuShane DuSHANE, 1970a: 5 Holotype 1 199; 5-10 Fms, Petatlan Bay, Guerrero, Mexico *Amaea (Scalina) deroyae DuShane DuSHANE, 1970b:330 Holotype 1236, 4 paratypes 1239; 30 Fms, Tagus Cove, Albemarle Is, Galapagos Is, Ecuador 6 Contributions in Science No. 213 Amaea (Scalina) tehuanarum DuShane and McLean DuSHANEand McLEAN, 1968:4 Holotype 1162, paratype 1163; 59-68 meters, Gulf of Tehuantepec, Mexico Anachis berryi Shasky SHASKY, 1970:190 Holotype 1413, 2 paratypes 1414; El Pulmo Reef, Baja California, Mexico Anachis (Glyptanachis) hilli Pilsbry and Lowe PILSBRY and LOWE, 1932:73 Paratype 1009; Mazatlan, Sinaloa, Mexico Ancilla matthewsi Burch and Burch BURCH and BURCH, 1967:81 Paratype 1248; 16 Fms, Fortaleza, Ceara, Brazil Ancilla osculata Sowerby SOWERBY, 1900:3 2 Syntypes 1325; The Kowie, South Africa Annularia (Bermudezia) capestanyi Torre and Bartsch TORRE and BARTSCH, 1941:358 3 Paratypes 1353; Palenque de Taguayabon, near Remedios, Santa Clara, Cuba Annularia (Bermudezia) obliterata Torre and Bartsch TORRE and BARTSCH, 1941:358 Paratype 1354; Cerro de Guajabana, Santa Clara, Cuba Aplysia vaccaria Winkler WINKLER, 1955:5 2 Paratypes 1231 ; Point Fermin, Los Angeles County, California Arena adust a McLean McLEAN, 1970d:123 Holotype 1391, 20 paratypes 1392; Intertidal to 6 feet, Espiritu Santo Is, Baja California, Mexico Arene echinata McLean McLEAN, 1970a: 311 2 Paratypes 1277; Fernandina Is, Galapagos Is, Ecuador ^ Arene ferruginosa McLean McLEAN, 1970d:123 Holotype 1393; S of White Friars, Guerrero, Mexico Arene guttata McLean McLEAN, 1970a: 3 10 Holotype 1275, 103 paratypes 1276; Academy Bay, Santa Cruz Is, Galapagos Is, Ecuador 1971 Type Specimens of Recent Mollusca 7 Arene stellata McLean McLEAN, 1970d:123 Holotype 1394, 4 paratypes 1395; Partida Bay, Espiritu Santo Is, Baja California, Mexico Arene winslowae Pilsbry and Lowe PILSBRY and LOWE, 1932:86 3 Paratypes 1262; San Juan del Sur, Nicaragua Ashmunella lenticula Gregg GREGG, 1953:74 Paratype 1091; North fork of Horseshoe Canyon, Cochise County, Arizona Ashmunella thomsoniana porterae Pilsbry and Cockerell PILSBRY and COCKERELL, 1899:49 2 Syntypes 1229; Beulah, New Mexico Aspella (1 Dermomurex) myrakeenae Emerson and D’Attilio EMERSON and D’ATTILIO, 1970:88 Paratype 1237; Banderas Bay, Nayarit, Mexico Balds delmontensis Smith and Gordon SMITH and GORDON, 1948:219 Paratype 1005; 10 Fms, Monterey Bay, Monterey County, California Bela sculpturata Dali DALL, 1887:299 2 Paratypes 1442; Chiachi Is, Canada Bellaspira acdivicosta McLean and Poorman McLEAN and POORMAN, 1970:6 Holotype 1193; 15-20 Fms, Bahia San Carlos, Sonora, Mexico *Bellaspira darionensis McLean and Poorman McLEAN and POORMAN, 1970:9 Holotype 1366, paratype 1367; 28-45 Fms, Sulphur Bay, Clarion Is, Revillagigedo Is, Mexico * Bellaspira margaritensis McLean and Poorman McLEAN and POORMAN, 1970:5 Holotype 1337, 2 paratypes 1138; 17-21 Fms, N of Margarita Is, Vene- zuela Borsonia (Borsonella) diegensis Dali DALL, 1908:275 Paratype 1443; 822 Fms, off San Diego, San Diego County, California Bucdnum pemphigus major Dali DALL, 1919a:325 2 Paratypes 1206; 100 Fms, SE of Kamchatka, Alaska 8 Contributions in Science No. 213 Calliostoma gordanum McLean McLEAN, 1970c:422 Paratype 1268; 70 Fms, Gorda Bank, Baja California, Mexico Calliostoma jacquelinae McLean McLEAN, 1970c: 425 Paratype 1274; 82 Fms, Academy Bay, Santa Cruz Is, Galapagos Is, Ecuador ^'Calliostoma keenae McLean McLEAN, 1970c: 425 Holotype 1272; 58-60 Fms, 12 mi off Laguna Beach, Orange County, California Calliostoma sanjaimense McLean McLEAN, 1970c:423 Holotype 1269, 2 paratypes 1 270; 75 Fms, off Cape San Lucas, Baja California, Mexico ^Calliostoma veleroae McLean McLEAN, 1970c: 423 Holotype 1271; 54 Fms, 3 mi S of Isla Ladrones, Panama Cant hams (Gemophos) berryi McLean McLEAN, 1970a:314 Holotype 1286, 120 paratypes 1287; 10-15 Fms, La Cruz, Banderas Bay, Jalisco, Mexico Cerion (Strophiops) fernandina Clench CLENCH, 1937:21 2 Paratypes 1 304; 8 mi SE of Simms, Long Is, Bahama Is Cerion hughe si Clench CLENCH, 1952:107 4 Paratypes 1295; Sandy Point, Savannah Sound, Eleuthera Is, Bahama Is Cerion (Strophiops) malonei Clench CLENCH, 1937:20 2 Paratypes 1303; 3.5 mi SEof Simms, Long Is, Bahama Is Cerion marielinum Torre and Pilsbry TORRE and PILSBRY, 1927:74 6 Paratypes 1296; Mariel, Pinar del Rio, Cuba Cerion (Strophiops) mcleani Clench CLENCH, 1937:22 3 Paratypes 1305; 1 mi E of O’Neills, Long Is, Bahama Is Cerion salvatori Torre and Pilsbry TORRE and PILSBRY, 1927:74 6 Paratypes 1 297 ; Playa de Santa Fe, Habana Province, Cuba 1971 Type Specimens of Recent Mollusca 9 Chilina oldroydae Marshall MARSHALL, 1924:4 Paratype 1076; Lake Fetalafquen, Chubut, Argentina Chondrothyretes affinis Torre and Bartsch TORRE and BARTSCH, 1938:303 7 Paratypes 1354; Dos Hermanos, Vinales, Pinar del Rio, Cuba Chondrothyretes barbouri barbouri Torre and Bartsch TORRE and BARTSCH, 1938:299 5 Paratypes 1349; Luis Lazo, Pinar del Rio, Cuba Chondrothyretes cerina cerina Torre and Bartsch TORRE and BARTSCH, 1938:298 6 Syntypes 1350; Sitio del Infierno, Sierro del Infierno, Vinales, Cuba Chrysodomus pribiloffensis Dali DALE, 1919a:323 Paratype 1289; 62 Fms, W of Pribiloff Is, Alaska Cinclidotyphis myrae DuShane DuSHANE, 1969:343 Holotype 1 194; Tenacatita, Jalisco, Mexico Cingula eyerdami Willett WILLETT, 1934c: 103 Holotype 1037; Elrington Is, Alaska Cingula forresterensis Willett WILLETT, 1934c: 103 Holotype 1038; Forrester Is, Alaska Clathrodrillia (Carinodrillia) bicarinata Shasky SHASKY, 1961:21 Paratype 1013; 45-90 Fms, off Espirtu Santo Is, Baja California, Mexico Columbella socorroensis Shasky SHASKY, 1970:194 Holotype 1418; 2 meters, Braithwaite Bay, Socorro Is, Revillagigedo Is, Colima, Mexico Conus poormani Berry BERRY, 1968:156 2 Paratypes 1261; 24-26 Fms, off Morro Colorado, Sonora, Mexico Coralliophila macleani Shasky SHASKY, 1970:189 Holotype 141 1, paratype 1412; N of Saladita Bay, Guaymas, Sonora, Mexico Crepidula coei Berry BERRY, 1950:35 Paratype 1008; Near Sunset Beach, Orange County, California 10 Contributions in Science No. 213 Crepidula rugosa naticarum Williamson WILLIAMSON, 1905a: 50 2 Syntypes 1026; San Pedro Bay, Los Angeles County, California Crepidula rugosa norrisiarum Williamson WILLIAMSON, 1905a: 50 4 Syntypes 1027; San Pedro, Los Angeles County, California Cryptogemma quentinensis Dali BALL, 1919b:32 2 Paratypes 1423; 822 Fms, off San Diego, San Diego County, Cali- fornia Cytharella electra Dali DALL, 191 9b: 77 2 Paratypes 1429; Head of Concepcion Bay, Baja California, Mexico Diodor a arnoldi McLean McLEAN, 1966:5 Holotype 1 143; 9 paratypes 1 144; 3 mi S of Carmel, Monterey County, California * Diodor a punctifissa McLean McLEAN, 1970b: 365 Holotype 1311, 5 paratypes 1312; 100-150 Fms, Wenman Is, Gala- pagos Is, Ecuador Duplicaria (Duplicaria) crakei R. D. Burch R. D. BURCH, 1965:245 Paratype 1212; Broome, West Australia Elaeocyma ricaudae Berry BERRY, 1969:164 2 Paratypes 1445; 24-26 Fms, near Boca Soledad, Baja California, Mexico Emarginula angusta McLean McLEAN, 1970b: 363 Holotype 1308; 170-160 meters off San Felix Is, Chile Emarginula dictya McLean McLEAN, 1970b: 3 64 Holotype 1309, paratype 1310; 170-160 meters off San Felix Is, Chile Epiphragmophora californiensis miwoka Bartsch BARTSCH, 1919:248 4 Paratypes 1334; Point Reyes, Marin County, California Epitonium (Acirsa) cerralvoensis DuShane DuSHANE, 1970b: 3 Holotype 1200, paratype 1201 ; 7 Fms, Cerralvo Is, Baja California, Mexico 1971 Type Specimens of Recent Mollusca 11 *Epitonium (Nitidiscala) hancocki DuShane DuSHANE, 1970a: 332 Holotype 1235, paratype 1238; Galapagos Is, Ecuador Epitonium (Asperiscala) huffmani DuShane and McLean DuSHANE and McLEAN, 1968:1 Holotype 1159; Cholla Bay, Sonora, Mexico Epitonium (Asperiscala) longinosanum DuShane DuSHANE, 1970b: 1 Holotype 1 196; Rancho Inocentes, Baja California, Mexico Epitonium (Asperiscala) macleani DuShane DuSHANE, 1970b: 3 Holotype 1197, 8 paratypes 1 198; 15-20 Fms, off Punta Ventana, Baja California, Mexico * Epitonium (Acirsa) murrha DuShane DuSHANE, 1970b: 4 Holotype 1232; 30 Fms, 10 mi SW of Secas Is, Panama Epitonium pedroanum Willett WILLETT, 1932c:88 Holotype 1036; 20 Fms, San Pedro, Los Angeles County, California Epitonium (Epitonium) shyorum DuShane and McLean DuSHANE and McLEAN, 1968:2 Holotype 1 160; 12-13 Fms, Manzanillo, Colima, Mexico Epitonium tinctum bormanni Strong STRONG, 1941:47 Holotype 1064; Mission Bay, San Diego, San Diego County, California Euchelus guttarosea Dali DALE, 1889:382 Paratype 1427; 116 Fms, Samana Bay, Santo Domingo Eulima randolphi Vanatta VANATTA, 1899:256 4 Paratypes 1336; Dutch Harbor, Unalaska Bay, Unalaska Is, Alaska Fissurella (Cremides) barbouri Farfante FARFANTE, 1943:9 Paratype 1218; 1 mi W of Nassau, Bahama Is Fissurella (Cremides) decemcostata McLean McLEAN, 1970b: 365 Holotype 1313, paratype 1314; Playa Caleta, Acapulco, Guerrero, Mexico 12 Contributions in Science No. 213 Fissurella (Cremides) deroyae McLean McLEAN, 1970d:118 Holotype 1375, 6 paratypes 1376; Academy Bay, Santa Cruz Is, Galapagos Is, Ecuador ^Fissurella (Clypidella) morrisoni McLean McLEAN, 1970d:117 Holotype 1374; Bahia Honda, Panama '^Fusinus allyni McLean McLEAN, 1970a: 314 Holotype 1290, 3 paratypes 1291; 70-80 Fms, Daphne Minor Is, Galapagos Is, Ecuador Gibbula harrisi Baker and Spicer BAKER and SPICER, 1930: 177 2 Paratypes 1014; 4 Fms, Basilan Channel, off Zamboanga, Mindanao, Philippine Is Haliotis corrugata oweni Talmadge TALMADGE, 1966:1 Holotype 1 147, 5 paratypes 1 148; 20 ft. West Anchorage, Guadalupe Is, Baja California, Mexico Haliotis cracherodii splendidula Williamson WILLIAMSON 1892:198 Holotype 1035; Point Vincente, Los Angeles County, California Haliotis fulgens guadalupensis Talmadge TALMADGE, 1964:375 Paratype 1210; South tip, Morro Sur, Guadalupe Is, Baja California, Mexico Haliotis (Padollus) roberti McLean, 1970 McLEAN, 1970d:115 Holotype 1368, 7 paratypes 1369; 40-47 Fms, Chatham Bay, Cocos Is, Costa Rica Hastula (Punctoterebra) betsyae R. D. Burch R. D. BURCH, 1965:243 Paratype 1214; Honokohau, Maui, Hawaii Helicostyla leucophthalma tagolandensis Pilsbry PILSBRY, 1908:47 Syntype 1294; Tagolandung Is, Celebes Group Helminthogly pta arrosa mattolensis A. G. Smith A. G. SMITH, 1938:83 4 Paratypes 1222; Between Cape Mendocino and the mouth of the Mattole River, Humboldt County, California 1971 Type Specimens of Recent Mollusca 13 Helminthoglypta arrosa williamsi A. G. Smith A. G. SMITH, 1938:79 3 Paratypes 1220; Hog Is, Tomales Bay, Marin County, California Helminthoglypta benitoensis H. N. Lowe H. N. LOWE, 1930:43 Holotype 1021; Pinnacles National Monument, San Benito County, California Helminthoglypta caruthersi Willett WILLETT, 1934b: 57 Holotype 1039; Morris Canyon, Kern County, California (Figs. 1-3). Helminthoglypta cuyama Hanna and Smith HANNA and SMITH, 1937:15 6 Paratypes 1224; Cuyama River Valley, E of Santa Maria, Santa Barbara County, California Helminthoglypta cuyamacensis piutensis Willett WILLETT, 1938a: 53 Holotype 1056; Piute Mountains, Kern County, California Helminthoglypta fontiphila Gregg GREGG, 1931:49 Holotype 1032; Little Rock Creek, Los Angeles County, California Helminthoglypta greggi Willett WILLETT, 1931a: 124 Holotype 1031; 3.5 mi S of Mohave, Kern County, California Helminthoglypta reediana Willett WILLETT, 1932b: 134 Holotype 1030; Lowe Canyon, Monterey County, California Helminthoglypta similans Hanna and Smith HANNA and SMITH, 1937:13 2 Paratypes 1223; .75 mi SE of Oil City, Fresno County, California Helminthoglypta stageri Willett WILLETT, 1938a: 52 Holotype 1055; Erskine Creek, Piccte Mountains, Kern County, Cali- fornia Helminthoglypta traski pacoimensis Gregg GREGG, 1931:47 Holotype 1033; Pacoima Canyon, Los Angeles County, California H elminthoglypta tudiculata rex Church and Smith CHURCH and SMITH, 1938:119 Paratype 1227; Middle Fork, Tule River, 2 mi above Springville, Tulare County, California 14 Contributions in Science No. 213 Kerguelenella stewartiana Powell POWELL, 1939:237 4 Paratypes 1215; Aker’s Point, Stewart Is, New Zealand Lamellaria koto Schwengel SCHWENGEL, 1944:17 Paratype 1216; Near Braidenton Beach, Florida Lamellaria sharoni Willett WILLETT, 1939:123 Holotype 1059, 2 paratypes 1092; Anaheim Bay, Orange County, Cali- fornia Lapsigyrus myriosirissa Shasky SHASKY, 1970:189 Holotype 1410; Point Tiburon, Mazatlan, Sinaloa, Mexico *Latiaxis (Babelomurex) santacruzensis Emerson and D’Attilio EMERSON and D’ATTILIO, 1970:272 Paratype 1234; 48-73 Fms off Barrington Is, Galapagos Is, Ecuador Leptothyra engbergi Willett WILLETT, 1929a: 27 Holotype 1017; Olga, Washington *Leurolepas roseola McLean McLEAN, 1970b: 365 Holotype 1315; 13 Fms, Magdalena Is, Tres Marias Is, Nayarit, Mexico Liotia miniata Dali DALE, 1889:390 3 Paratypes 1428; Nassau, Bahama Is Lirularia bicostata McLean McLEAN, 1964:129 4 Paratypes 1134; 45 feet, off Middle Coronado Is, Baja California, Mexico Littorina albicarinata McLean McLEAN, 1970d:127 Holotype 1399, 50 paratypes 1400; El Requeson, Concepcion Bay, Baja California, Mexico Littorina sitkana atkana Dali , DALE, 1886:211 3 Paratypes 1203; Kyska Is, Alaska Lucapinella eleanorae McLean McLEAN, 1967:350 Holotype 1153; 10 Fms, off La Cruz, Banderas Bay, Jalisco, Mexico 1971 Type Specimens of Recent Mollusca 15 Macrarene coronadensis Stohler STOHLER, 1959:439 Paratype 1110; “Rockpile,” SE of South Coronado Is, Baja California, Mexico The original designation as hypotype no. 11 has no nomenclatural standing and this specimen, currently in our collection, is considered a paratype. ■^Macrarene lepidotera McLean McLEAN, 1970d:125 Holotype 1396; 70-75 Fms, off Braithwaite Bay, Socorro Is, Revilla- gigedo Is, Mexico Macrarene spectabilospina Shasky SHASKY, 1970:188 Holotype 1409; Gulf of Tehuantepec, Chiapas, Mexico Mangelia arteaga Dali and Bartsch DALE and BARTSCH, 1910 Paratype 1424; 8-34 Fms, Barkeley Sound, Vancouver Is, British Columbia, Canada Mangelia (Kurtziella) cyrene Dali DALE, 1919b: 32 3 Paratypes 1440; 26 Fms, off La Paz, Baja California, Mexico Melanella rosa Willett WILLETT, 1944:72 Holotype 1075; off Redondo Beach, Los Angeles County, California Micrarionta brunnea Willett WILLETT, 1935a: 15 Holotype 1042, 4 paratypes 1357; Chuckwalla Spring, Riverside County, California (Figs. 16-18). Micrarionta (? Eremarionta) chacei Willett WILLETT, 1940a: 80 Holotype 1062, paratype 1093; 9 mi N of Ensenada, Baja California, Mexico Micrarionta chocolata Willett WILLETT, 1935a: 15 Holotype 1041; Beal’s Well, Chocolate Mountains, Imperial County, California (Figs. 10-12). Micrarionta chuckwallana Willett WILLETT, 1935a: 15 Holotype 1040; 1 mi S of Corn Springs, Chuckwalla Mountains, River- side County, California (Figs. 7-9). 16 Contributions in Science No. 213 Micrarionta granitensis Willett WILLETT, 1935b: 1 Holotype 1043; NW end of the Granite Mountains, Riverside County, California (Figs. 13-15). Micrarionta harperi carrizoensis Willett WILLETT, 1937b: 123 Holotype 1049; Above Painted Gorge, Carrizo Mountain, Imperial County, California Micrarionta hutsoni amboiana Willett WILLETT, 1931a: 123 Holotype 1029; 6 mi NW of Amboy, San Bernardino County, California Micrarionta hutsoni hilli Willett WILLETT, 1930a: 6 Holotype 1024; Sheep Hole Mountains, San Bernardino County, Cali- fornia Micrarionta hutsoni unifasciata Willett WILLETT, 1930a:6 Holotype 1023; Newberry Springs, San Bernardino County, California Micrarionta immaculata Willett WILLETT, 1937a: 6 Holotype 1051; 7 mi S of Vidal, San Bernardino County, California Micrarionta indioensis cathedralis Willett WILLETT, 1930b: 115 Holotype 1022; Cathedral Canyon, Riverside County, California Micrarionta indioensis remota Willett WILLETT, 1937b: 124 Holotype 1050; Borego Mountain, San Diego County, California Micrarionta mccoiana Willett WILLETT, 1935b: 2 Holotype 1044; McCoy Well, McCoy Mountains, Riverside County, California (Figs. 19-21). Micrarionta ora Willett WILLETT, 1929b: 17 Holotype 1018; North end of the Fish Mountains, Imperial County, California (Figs. 4-6). Micrarionta orocopia Willett WILLETT, 1934a: 15 Holotype 1060; 2.5 mi N of Dos Palmas, Riverside County, California Microdochus floridanus Rheder RHEDER, 1943:193 12 Paratypes 1317; Missouri Key, Florida 1971 Type Specimens of Recent Mollusca 17 Mirachelus galapagensis McLean McLEAN, 1970d:119 Holotype 1377, 2 paratypes 1378; 75-100 meters off Canal Bolivar, near Tagus Cove, Albemarle Is, Galapagos Is, Ecuador Monadenia fidelis beryllica E. P. and E. M. Chace E. P. and E. M. CHACE, 1935:48 5 Paratypes 1225; 8 mi N of Brookings, Oregon Monadenia fidelis pronotis Berry BERRY, 1931:122 6 Paratypes 1226; Point St. George, 3 mi N of Crescent City, Del Norte County, California Monadenia (Corynadenia) hillebrandi mariposa A. G. Smith A. G. SMITH, 1957:24 2 Paratypes 1112; McLean Cave, Mariposa County, California '^Morum (Cancellomorum) veleroae Emerson EMERSON, 1968:53 Holotype 1170, 2 figured paratypes 1171, 1 unfigured paratype 1172; 31-50 Fms, Cocos Is, Costa Rica galapaganus Emerson and D’Attilio EMERSON and D’ATTILIO, 1970:271 Paratype 1233; 50-100 Fms, N of Hood Is, Galapagos Is, Ecuador Murexiella (Murexiella) keenae E. H. Yokes E. H. YOKES, 1970a: 328 Holotype 1259; Yenado Beach, Canal Zone, Panama Murexiella (Murexiella) laurae E. H. Yokes E. H. YOKES, 1970a: 328 Holotype 1260; 17 Fms, Punta de Juluapan, Manzanillo, Colima, Mexico Nassarina (Zanassarina) anitae Campbell CAMPBELL, 1961:26 Paratype 1096; 10-25 Fms, Cabo Haro, Guaymas, Sonora, Mexico ^'Nassarius shaskyi McLean McLEAN, 1970d:128 Holotype 1405, 2 paratypes 1406; 20 Fms, off Gorgona Is, Colombia Natica brunneolinea McLean McLEAN, 1970a:313 Holotype 1282, figured paratype 1283; 50-100 Fms, Academy Bay, Santa Cruz Is, Galapagos Is, Ecuador Natica sigillata McLean McLEAN, 1970a: 313 Holotype 1284, paratype 1285; Galapagos Is, Ecuador 18 Contributions in Science No. 213 Neptunea stile si A. G. Smith A. G. SMITH, 1968:117 Paratype 1 152; 88-85 Fms, Queen Charlotte Sound, British Columbia, Canada. Paratype 1211; 100-125 Fms, between La Perouse Bank, Washington and Cape Scott, British Columbia, Canada *Nesta galapagensis McLean McLEAN, 1970b: 363 Holotype 1307; 80-100 Fms, Tagus Cove, Albemarle Is, Galapagos Is, Ecuador *Niso emersoni McLean McLEAN, 1970d:128 Holotype 1403, figured paratype 1404; 7-1 1 Fms, off San Jose Point, Guatemala Ocenebra keenae Bormann BORMANN, 1946:40 2 Paratypes 1001; White’s Point, Los Angeles County, California Oliva reject a J. Q. and R. L. Burch J. Q. and R. L. BURCH, 1962: 166 Paratype 1006; La Paz, Baja California, Mexico Olivella inusta Sowerby SOWERBY, 1915:165 2 Syntypes 1335; Florida Olivella sphoni Burch and Campbell BURCH and CAMPBELL, 1963:124 Paratype 1124; 12 meters, off Dos Hermanos Is, Mazatlan, Sinaloa, Mexico Opalia chacei Strong STRONG, 1937:5 Holotype 1045; Crescent City, Del Norte County, California Parapholyx effusa nevadensis Henderson HENDERSON, 1934:90 3 Paratypes 1080; Winnemucca Lake, Nevada Pectinodonta arcuata Dali DALL, 1882:409 Paratype 1426; 226 Fms, off Santa Lucia, Lesser Antilles Persicula bandera Coan and Roth COAN and ROTH, 1965:67 Holotype 1141, 18 paratypes 1 142; Banderas Bay, Jalisco, Mexico Philine alba Mattox MATTOX, 1958:98 Paratype 1118; 35 Fms, 1.8 mi SE of Long Point, Santa Catalina Is, Los Angeles County, California 1971 Type Specimens of Recent Mollusca 19 Philine calif ornica Willett WILLETT, 1944:71 Holotype 1074; 50 Fms, off Redondo Beach, Los Angeles County, California *Poirieria (Panamurex) velero E. H. Yokes E. H. YOKES, 1970b: 47 Holotype 1407, 7 paratypes 1408; 10-22 Fms, 1-2 mi SW of Cabo de Yela, Guajira Peninsula, Colombia Pseudomelatoma moesta maculata Williamson WILLIAMSON, 1905b: 122 Holotype 1052; Point Fermin, Los Angeles County, California Pseudomelatoma semiinflata redondoensis T. Burch T. BURCH, 1938:21 3 Paratypes 1011; 25 Fms, Redondo Beach, Los Angeles County, California Pterotyphis (Tripterotyphis) arcana DuShane DuSHANE, 1969:344 Holotype 1195; Mazatlan, Sinaloa, Mexico Pterotyphis (Tripterotyphis) fayae Keen and Campbell KEEN and CAMPBELL, 1964:54 Paratype 1 133; Barra de Navidad, Jalisco, Mexico Pterynotus (Purpurellus) macleani Emerson and D’Attilo EMERSON and D’ATTILO, 1969:146 Holotype 1230, 25 Fms, Loreto Channel, Baja California, Mexico. * Figured paratype 1273; 12 Fms, Secas Is, Panama Puncturella (Cranopsis) decorata Cowan and McLean COWAN and McLEAN, 1968:105 2 Paratypes 1175; 106 Fms, Queen Charlotte Is, British Columbia, Canada Pupa sterkiana Pilsbry PILSBRY, 1889:412 45 Syntypes 1333; San Quintin Bay, Baja California, Mexico Pyrene aureola Howard HOWARD, 1963:1 6 Paratypes 1209; Norse Beach, Punta Penasco, Sonora, Mexico Radwinia tehuantepecensis Shasky SHASKY, 1970:191 Holotype 1415; Between San Simeon and Puerto Madero, Gulf of Tehuantepec, Chiapas, Mexico 20 Contributions in Science No. 213 Rissoella hertleini Smith and Gordon SMITH and GORDON, 1948:224 Paratype 1003; 10 Fms, off Cabrillo Point, Monterey Bay, Monterey County, California Rissoina hannai Smith and Gordon SMITH and GORDON, 1948:219 2 Paratypes 1004; 25 Fms, Carmel Bay, Monterey County, California *Ruthia ecuadoriana Shasky SHASKY, 1970:193 Holotype 1417; 4 meters off Cape San Francisco, Ecuador Ruthia mazatlanica Shasky SHASKY, 1970:192 Paratype 1416; Chivos Is, Mazatlan, Sinaloa, Mexico Scalina (Ferminoscala) billeeana DuShane and Bratcher DuSHANE and BRATCHER, 1965:160 Paratype 1095; 8-10 feet, SW end of Ceralbo Is, Baja, California, Mexico *Scissurella (Anatoma) epicharis McLean McLEAN, 1970d:117 Holotype 1370, paratype 1371; 12 Fms, Cartago Bay, Albemarle Is, Galapagos Is, Ecuador *Scissurella (Anatoma) keenae McLean McLEAN, 1970d:117 Holotype 1372, 6 paratypes 1373; 46 Fms, Angel de la Guarda Is, Baja California, Mexico Scrobs ovata Powell POWELL, 1927a: 547 2 Paratypes 1362; 12 Fms, Ranganu Bay, off Awanui, New Zealand Solariella lewisii Willett WILLETT, 1946:32 Holotype 1084; 15 Fms, Spiridon Bay, Kodiak Is, Alaska Sonorelix (Herpeteros) angelus Gregg GREGG, 1948:100 Paratype 1085; 2 mi E of Solemint, Los Angeles County, California Sonorella allynsmithi Gregg and Miller GREGG and MILLER, 1969:90 9 Paratypes 1246; Phoenix Mountains, Maricopa County, Arizona Sonorella baboquivariensis cossi Miller MILLER, 1966:46 2 Paratypes 1244; Arch Canyon, Organ Pipe Cactus National Monu- ment, Pima County, Arizona 1971 Type Specimens of Recent Mollusca 21 Sonorella neglecta Gregg GREGG, 1951:156 Paratype 1087; 3 mi W of Portal, Chiricahua Mountains, Cochise County, Arizona Sonorella simmonsi Miller MILLER, 1966:46 3 Paratypes 1245; Picacho Mountains, Pinal County, Arizona Sonorella waltoni Miller MILLER, 1968:61 3 Paratypes 1247; West Doubtful Canyon, Peloncillo Mountains, Co- chise County, Arizona Speleodiscoides spirellum A. G. Smith A. G. SMITH, 1957:34 3 Paratypes 1111; Violin Cave, Amador County, California Steeriana (Steeriana) cajamarcana Weyrauch and Zilch WEYRAUCH and ZILCH, 1954:73 5 Paratypes 1329; Hacienda Cochambul, near Cajamarca, Peru Steeriana (Steeriana) celendinensis Weyrauch and Zilch WEYRAUCH and ZILCH, 1954:70 5 Paratypes 1330; 3 km S of Celendin, Peru Steeriana celendinensis isidroensis Weyrauch and Zilch WEYRAUCH and ZILCH, 1954:72 5 Paratypes 1331; Mount San Isidro, near Celendin, Peru Strombina (Cotonopsis) mendozana Shasky SHASKY, 1970:194 Holotype 1419, paratype 1420; 33-73 meters. Gulf of Fonseca, El Salvador Strombus raninus nanus Bales BALES, 1942:18 Paratype 1358; Lake Worth, Palm Beach County, Florida ^Tegula (Agathistoma) bergeroni McLean McLEAN, 1970d:119 Holotype 1379, paratype 1380; Cabita Bay, near Cabo Corrientes, Colombia Tegula (Agathistoma) corteziana McLean McLEAN, 1970d:119 Holotype 1381, 71 paratypes 1382; South side of Cabo Tepoca, Sonora, Mexico Tegula (Agathistoma) felipensis McLean McLEAN, 1970d:121 Holotype 1383, 16 paratypes 1384; Punta San Felpe, Baja California, Mexico 22 Contributions in Science No. 213 Tegula (Agathistoma) mendella McLean McLEAN, 1964:131 Paratype 1135; Mission Bay, San Diego, San Diego County, California * Tegula (Agathistoma) picta McLean McLEAN, 1970d:121 Holotype 1385, 14 paratypes 1386; W of Manta, Ecuador Tegula (Agathistoma) verdispira McLean McLEAN, 1970d:122 Holotype 1387, paratype 1388; South Anchorage, Maria Magdalena Is, Tres Marias Is, Nayarit, Mexico Tegula (Agathistoma) verrucosa McLean McLEAN, 1970d:122 Holotype 1389, 5 paratypes 1390; Palo Seco, Canal Zone, Panama Temesa (Temesa) omissa Weyrauch WEYRAUCH, 1957:18 5 Paratypes 1332; Tarmatabo, near Tarma, Peru Terebra allyni Bratcher and Burch BRATCHER and BURCH, 1970a:298 Paratype 1254; 5-10 Fms, Maria Madre Is, Tres Marias Is, Nayarit, Mexico Terebra apicitincta Sowerby SOWERBY, 1900:5 2 Syntypes 1324; The Kowie, South Africa ^Terebra brandi Bratcher and Burch BRATCHER and BURCH, 1970a: 296 Holotype 1252, paratype 1253; 5-10 Fms, Petatlan Bay, Oaxaca, Mexico * Terebra dorothyae Bratcher and Burch BRATCHER and BURCH, 1970a: 297 Holotype 1250, figured paratype 1251; 7-11 Fms, San Jose Point, Guatemala Terebra (Strioterebrum) dushanae Campbell CAMPBELL, 1964:135 4 Paratypes 1208; Puertecitos, Baja California, Mexico "^Terebra hancocki Bratcher and Burch BRATCHER and BURCH, 1970a:299 Holotype 1255, 2 paratypes 1256; 8-10 Fms, La Libertad, Santa Elena Bay, Ecuador * T erebra hertleini Bratcher and Burch BRATCHER and BURCH, 1970b:l Paratype 1288; 15-25 Fms, Academy Bay, Santa Cruz Is, Galapagos Is, Ecuador 1971 Type Specimens of Recent Mollusca 23 *Terebra jacquelinae Bratcher and Burch BRATCHER and BURCH, 1970b: 2 26 Paratypes 1179; 10-25 Fms, Academy Bay, Santa Cruz Is ,Galapagos Is, Ecuador Terebra (Strioterebrum) ninfae Campbell CAMPBELL, 1961:27 Paratype 1106; Trawled between Puerto Madero and the Guatemalan border, Chiapas, Mexico '^Terebra purdyae Bratcher and Burch BRATCHER and BURCH, 1970b: 5 Holotype 1182, paratype 1183; 70-80 Fms, N of Charles Is, Galapa- gos Is, Ecuador Terebra shyana Bratcher and Burch BRATCHER and BURCH, 1970a:295 Holotype 1249; 17-40 Fms, Manzanillo, Colima, Mexico * Terebra stabler i Bratcher and Burch BRATCHER and BURCH, 1970b: 5 Holotype 1180, paratype 1181; 4-20 Fms, Socorro Is, Revillagigedo Is, Colima, Mexico Thais floridana haysae Clench CLENCH, 1927:6 Paratype 1219; Grand Bayau, Mississippi Delta, Louisiana ^Thala jeancateae Sphon SPHON, 1969:85 Holotype 1202; 50-60 Fms, Tagus Cove, Albemarle Is, Galapagos Is, Ecuador Thoristella chathamensis cookiana Powell POWELL, 1934:154 Paratype 1360; Island Bay, Wellington, New Zealand Thyca (Bessomia) callista Berry BERRY, 1959:110 Holotype 1120; 3-4 Fms, San Carlos Bay, Sonora, Mexico Tindaria brunnea Dali DALE, 1916:401 1 1 Paratypes 1161; 1401 Fms, Midway between St. Paul and Umnak Is, Alaska Trichosirius octocarinatus Powell POWELL, 1931:376 Paratype 1359; 50 Fms, Bounty Is, New Zealand *Tricolia diantha McLean McLEAN, 1970d:125 Holotype 1397, 37 paratypes 1398; 60 Fms, off Albemarle Is, Galapa- gos Is, Ecuador 24 Contributions in Science No. 213 Trigonostoma milleri J. Q. Burch J. Q. BURCH, 1949:3 2 Paratypes 1086; Tambor, Puntarenas, Costa Rica Tritonidea insculpta Sowerby SOWERBY, 1900:2 2 Syntypes 1323; The Kowie, South Africa Trivia ovulata rubra Shaw SHAW, 1909:312 Syntype 1321; Jeffreys Bay, South Africa Trophon (Boreotrophon) albospinosus Willett WILLETT, 193 lb: 66 Holotype 1028; 25 Fms, off the north shore of Santa Catalina Is, Los Angeles County, California Trophon kowiensis Sowerby SOWERBY, 1901:213 2 Syntypes 1326; The Kowie, South Africa T ugali chilensis McLean McLEAN, 1970b: 362 Holotype 1306; Carvajal Bay, south end of Isla Mas a Tierra, Juan Fernandez Is, Chile Turbonilla (Pyrgiscus) burchi Gordon GORDON, 1938:49 Holotype 1057; 25 Fms, 1 mi W of Redondo Beach, Los Angeles County, California Turbonilla (Strioturbonilla) cayucosensis Willett WILLETT, 1929a: 26 Holotype 1016, figured paratype 1083; Cayucos, San Luis Obispo County, California Turbonilla (Pyrgolampros) strongi Willett WILLETT, 193 lb: 67 Holotype 1019; 25 Fms, off the north shore of Santa Catalina Is, Los Angeles County, California T urcicula bairdii Dali DALL, 1889:376 5 Paratypes 1207; 414 Fms, off San Clemente Is, Los Angeles County, California Turritella (Zeacolpus) ahiparana Powell POWELL, 1927b: 297 18 Paratypes 1319; 23 Fms, Ahipara, New Zealand Turritella anactor Berry BERRY, 1957:78 Paratype 1447; Playa de Almejas, north of San Felipe, Baja California, Mexico 1971 Type Specimens of Recent Mollusca 25 Turritella kowiensis Sowerby SOWERBY, 1900:6 2 Syntypes 1322; The Kowie, South Africa Turritella parkeri McLean McLEAN, 1970d:127 Holotype 1401, 27 paratypes 1402; 45-65 Fms, W of Espiritu Santo Is, Baja California, Mexico Turritella willetti McLean McLEAN, 1970a: 3 12 Holotype 1280, 2 paratypes 1281; 7-12 Fms, Santiago Bay, Manzanillo, Colima, Mexico Urocoptis alleni Torre^ TORRE, 1929a: plate 4, fig. 7 11 Syntypes 1298; Pena Blanca, Sierra Anafe, Pinar del Rio, Cuba Urocoptis cuestai Torre TORRE, 1929b: 445 3 Syntypes 1351; 14 km Mogote on the road from Pinar del Rio to Vinales, Cuba Urocoptis (Gongylostoma) fortiuscula Torre TORRE, 1912:57 3 Paratypes 1302; “El Mamey,” Sagua la Grande, Santa Clara, Cuba Urocoptis livida barbouri Torre and Clench TORRE and CLENCH, 1930:15 4 Paratypes 1300; La Portuguesa, Central Soledad, Cienfuegos, Cuba Urocoptis mellita Torre TORRE, 1932:88 10 Paratypes 1217; “Finca El Ingles,” Sierra del Grillo, Madruga, Habana Province, Cuba Urocoptis nodulifera Torre TORRE, 1929:444 4 Paratypes 1352; Mogote Dos Hermanos, Vinales, Cuba Urocoptis sinistra Torre^ TORRE, 1928: plate l,fig. 8 8 Paratypes 1301; Near Marianao Beach, Habana, Cuba ^The taxon, Urocoptis alleni, has never been described. The name was validated by the publication of a drawing of the species in 1929 with the caption of “U. alleni, Torre, Antilla, Caleta de Ovando, Oriente.” Our specimens were obtained from M. L. Jaume, an associate of Torre. While they are from a different locality (Pena Blanca, Sierra Anafe, Pinar del Rio, Cuba), Jaume’s original label was marked “Co-Typus” and I consider them as syntypes. 2The taxon, Urocoptis sinistra Torre, was validated in 1928 by the publication of a photograph of the holotype with the caption, “Urocoptis sinistra Torre, n. sp. Havana.” Our specimens are from an associate of Torre (M. L. Jaume) and are from the type locality. 26 Contributions in Science No. 213 V ermicularia frisbeyae McLean McLEAN, 1970a:311 Holotype 1278, paratype 1279; 30-40 Fms, off the lighthouse, Manza- nillo, Colima, Mexico V espericola karokomm Talmadge TALMADGE, 1962:28 2 Paratypes 1228; Sawmill Gulch, 2 mi E of Orleans Ranger Station, Humboldt County, California Volvulella tenuissima Willett WILLETT, 1944:71 Holotype 1073; 75 Fms, Redondo Beach, Los Angeles County, Cali- fornia Xenophalium harrisonae Powell POWELL, 1928:640 Paratype 1318; Mason’s Bay, Stewart Is, New Zealand Amphineura Callistochiton connellyi Willett WILLETT, 1937c: 423 Holotype 1271; 54 Fms, 3 mi S of Isla Ladrones, Panama Chaetopleura (Pallochiton) euryplax Berry BERRY, 1945:491 Paratype 1263; Bahia de Adair, Sonora, Mexico Dendrochiton p suites Berry BERRY, 1963:135 Paratype 1448; Near entrance to Mission Bay, San Diego, San Diego County, California Ischnochiton (Lepidozona) catalinae Willett WILLETT, 1940b: 185 Holotype 1063; 2 paratypes 1000; 30 Fms, White’s Landing, Santa Catalina Is, Los Angeles County, California Lepidozona pella Berry BERRY, 1963:137 2 Paratypes 1449; San Felipe, Baja California, Mexico Mopalia pedroana Willett WILLETT, 1932a: 101 Holotype 1025; 12 Fms, off San Pedro, Los Angeles County, California 1971 Type Specimens of Recent Mollusca 27 Index acclivicosta, Bellaspira, p. 7 adusta, Arene, p. 6 affinis, Chondrothyretes, p. 9 aphiparana, Turritella (Zeacolpus), p. 24 alba, Philine, p. 18 albicarinata, Littorina, p. 14 albospinosus, Trophon (Boreotrophon), p. 24 alleni, Urocoptis, p. 25 allyni, Fusinus, p. 12 allyni, Terebra, p. 22 allynsmithi, Sonorella, p. 20 amboiana, Micrarionta hutsoni, p. 16 anactor, Turritella, p. 24 angelus, Sonorelix (Herpeteros), p. 20 angusta, Emarginula, p. 10 anitae, Nassarina (Zanassarina), p. 17 apicina, Acmaea, p. 5 apicitincta, Terebra, p. 22 arabica jaculoides, Alcithoe, p. 5 arcana, Pterotyphis (Tripterotyphis), p. 19 arcuata, Pectinodonta, p. 18 amoldi, Diodora, p. 10 arrosa mattolensis, Helminthoglypta, p. 12 arrosa williamsi, Helminthoglypta, p. 13 arteaga, Mangelia, p. 15 atkana, Littorina sitkana, p. 14 attenuata rogersi, Lithophaga (Labis), p. 4 aucklandica, Legrandina, p. 2 aureola. Pyrene, p. 19 baboquivariensis cossi, Sonorella, p. 20 bairdii, Turcicula, p. 24 bandera, Persicula, p. 18 barbouri, Chondrothyretes, p. 9 barbouri, Fissurella (Cremides), p. 11 barbouri, Urocoptis livida, p. 25 benitoensis, Helminthoglypta, p. 13 bergeroni, Tegula (Agathistoma), p. 21 beryllica, Monadenia fidelis, p. 17 berryi, Anachis, p. 6 berryi, Cantharus (Gemophos), p. 8 betsyae, Hastula (Punctoterebra), p. 12 bicarinata, Clathrodrillia (Carinodrillia), p. 9 bicostata, Lirularia, p. 14 billeeana, Scalina (Ferminoscala), p. 20 bimaculatus purpureus, Heterodonax, p.2 bimaculatus salmoneus, Heterodonax, p. 2 bormanni, Epitonium tine turn, p. 11 brandi, Terebra, p. 22 brunnea, Micrarionta, p. 15 brunnea, Tindaria, p. 23 brunneolinea, Natica, p. 17 burchi, Nuculana, p. 4 burchi, Turbonilla (Pyrgiscus), p. 24 cajamarcana, Steeriana (Steeriana), p.21 californica, Philine, p. 19 californiensis miwoka, Epiphragmorpha, p. 10 callista, Thyca (Bessomia), p. 23 capestanyi, Annularia (Bermudezia), p. 6 carrizoensis, Micrarionta harperi, p. 16 caruthersi, Helminthoglypta, p. 13 catalinae, Ischnochiton (Lepidozona), p. 26 catalinensis, Pecten (Cyclopecten), p. 4 catharia, Callocardia, p. 2 cathedralis, Micrarionta indioensis, p. 16 cayucosensis, Turbonilla (Strioturbonilla), p. 24 celendinensis, Steeriana (Steeriana), p.21 celendinensis isidroensis, Steeriana, p.21 cerina, Chondrothyretes, p. 9 cerralvoensis, Epitonium (Acirsa), p. 10 chacei, Micrarionta (? Eremarionta), P. 15 chacei, Opalia, p. 18 chathamensis cookiana, Thoristella, p. 23 chilensis, Tugali, p. 24 chocolata, Micrarionta, p. 15 chuckwallana, Micracionta, p. 15 clarionensis, Bellaspira, p. 7 clarionensis, Pseudochama, p. 4 coei, Crepidula, p. 9 connellyi, Callistochiton, p. 26 cookiana, Thoristella chathamensis, p. 23 contexta, Amaea (s.l.), p. 5 coronadensis, Macr arene, p. 15 corrugata oweni, Haliotis, p. 12 28 Contributions in Science No. 213 corteziana, Tegula (Agathistoma), p.21 cossi, Sonorella baboquivariensis, p. 20 cracherodii splendidula, Haliotis, p. 12 crakei, Duplicaria (Duplicaria), p. 10 cuestai, Urocoptis, p. 25 culcitella intermedia, Acteocina, p. 5 cuyama, Helminthoglypta, p. 13 cuyamacensis piutensis, Helminthoglypta, p. 13 cyrene, Mangelia (Kurtziella), p. 15 decemcostata, Fissurella, p. 11 decorata, Puncturella (Cranopsis), p. 19 delmontensis. Balds, p. 7 deroyae, Amaea (Scalina), p. 5 deroyae, Fissurella (Cremides), p. 12 diantha, Tricolia, p. 23 dictya, Emarginula, p. 10 diegensis, Borsonia (Borsonella), p. 7 digensis, Limopsis, p. 2 dorothyae, Terebra, p. 22 dushanae, Terebra (Strioterebrum), p. 22 echinata, Arene, p. 6 ecuadoriana, Ruthia, p. 20 effusa nevadensis, Parapholyx, p. 18 eleanorae, Lucapinella, p. 14 electra, Cytharella, p. 10 emersoni, Niso, p. 18 enbergi, Leptothyra, p. 14 ensifera, Yoldia, p. 4 epicharis, Scissurella (Anatoma), p. 20 euryplax, Chaetopleura (Pallochiton), p. 26 eyerdami. Cingula, p. 9 fayae, Pterotyphis (Tripterotyphis), p. 19 felipensis, Tegula (Agathistoma), p. 21 fernandina, Cerion (Strophiops), p. 8 ferruginosa, Arene, p. 6 fidelis beryllica, Monadenia, p. 17 fidelis pronotis, Monadenia, p. 17 fitchi, Penitella, p. 4 floridana haysae, Thais, p. 23 floridanus, Microdochus, p. 16 fontiphila, Helminthoglypta, p. 13 forresterensis. Cingula, p. 9 fortiuscula, Urocoptis (Gongylostoma), p. 25 frisbeyae, Vermicularia, p, 26 fulgens guadalupensis, Haliotis, p. 12 galapaganus, “Murex”, p, 17 galapagensis, Mirachelus, p. 17 galapagensis, Nesta, p. 18 gordanum, Calliostoma, p. 8 granitensis, Micrarionta, p. 16 greggi, Helminthoglypta, p. 1 3 guadalupensis, Haliotis fulgens, p. 12 guttarosea, Euchelus, p. 11 guttata, Arene, p. 6 hancocki, Terebra, p. 22 hannai, Rissoina, p. 20 harperi carrizoensis, Micrarionta, p. 16 harrisi, Gibbula, p. 12 harrisonae, Xenophalium, p. 26 haysae, Thais floridana, p. 23 hertleini, Rissoella, p. 20 hertleini, Terebra, p. 22 hillebrandi mariposa, Monadenia, p. 17 hilli, Anachis, p. 6 hilli, Micrarionta hutsoni, p. 16 huffmani, Epitonium (Asperiscala), p. 11 hughesi, Cerion, p. 8 hutsoni amboiana, Micrarionta, p. 16 hutsoni hilli, Micrarionta, p. 16 hutsoni unifasciata, Micrarionta, p. 16 immaculata, Micrarionta, p. 16 indioensis cathedralis, Micrarionta, p. 16 indioensis remota, Micrarionta, p. 16 inflatula, Macoma, p. 4 insculpta, Tritonidea, p. 24 intermedia, Acteocina culcitella, p. 5 inusta, Olivella, p, 18 isidroensis, Steeriana celendinensis, p.21 jacquelinae, Calliostoma, p. 8 jacquelinae, Terebra, p. 23 jaculoides, Alcithoe arabica, p. 5 jeancateae, Thala, p. 23 karokorum, Vespericola, p. 26 keenae, Calliostoma, p. 8 keenae, Glycimeris, p. 2 keenae, Murexiella (Murexiella), p. 17 keenae, Ocenebra, p. 1 8 keenae, Scissurella (Anatoma), p. 20 koto, Lamellaria, p. 14 kowiensis, Trophon, p. 24 kowiensis, Turritella, p. 25 laurae, Murexiella (Murexiella), p. 17 lenticula, Ashmunella, p. 7 lepidotera, Macarene, p. 15 leucophthalma tagolandensis, Helicostyla, p. 12 1971 Type Specimens of Recent Mollusca 29 lewisii, Solariella, p. 20 limacina, Alectrion, p. 5 iivida barbouri, Urocoptis, p. 25 lombardii, Allogona, p. 5 longinosanum, Epitonium (Asperiscala), p. 11 luteola rosea, Corbula, p. 2 macleani, Coralliophila, p. 9 macleani, Epitonium (Asperiscala), p. 11 macleani, Pterynotus (Purpurellus), p. 19 maculata, Pseudomelatoma moesta, p. 19 major, Buccinum pemphigus, p. 7 malonei, Cerion (Strophiops), p. 8 margaritensis, Bellaspira, p. 7 marielinum, Cerion, p. 8 mariposa, Monadenia hillebrandi, p. 17 matthewsi, Ancilla, p. 6 mattolensis, Helminthoglypta arrosa, p. 12 mazatlanica, Ruthia, p. 20 mccoiana, Micrarionta, p. 16 mcleani, Cerion (Strophiops), p. 8 mellita, Urocoptis, p. 25 mendella, Tegula (Agathistoma), p. 22 mendozana, Strombina (Cotonopsis), p.21 milleri, Trigonostoma, p. 24 miniata, Liotia, p. 14 miwoka, Epiphragmorpha califomiensis, p. 10 moesta maculata, Pseudomelatoma, p. 19 montereyensis, Cardita ventricosa, p. 2 montereyensis, Yoldia, p. 4 morrisoni, Fissurella (Clypidella), p. 12 murrha, Agaronia, p. 5 murrha, Epitonium (Acirsa), p. 11 myrae, Cinclidotyphis, p. 9 myrae, Ensis, p. 2 myrakeenae, Aspella ( ? Dermomurex), p.7 myriosirissa, Lapsigyrus, p. 14 nanus, Strombus raninus, p. 21 naticarum, Crepidula rugosa, p. 10 neglecta, Sonorella, p. 21 nevadensis, Parapholyx effusa, p. 18 ninfae, Terebra (Strioterebrum), p. 23 nipponica, Lasaea, p. 2 nodulifera, Urocoptis, p. 25 norrisiarum, Crepidula rugosa, p. 10 obliterata, Annularia (Bermudezia), p. 6 octocarinatus, Trichosirius, p. 23 oldroydae, Chilina, p. 9 omissa, Temesa (Temesa), p. 22 ora, Micrarionta, p. 16 orocopia, Micrarionta, p. 16 osculata, Ancilla, p. 6 ovata, Scrobs, p. 20 ovulata rubra. Trivia, p. 24 oweni, Haliotis corrugata, p. 12 pacoimensis, Helminthoglypta traski, P. 13 parkeri, Turritella, p. 25 pedroana, Mopalia, p. 26 pedroanum, Epitonium, p. 11 pella, Lepidozona, p. 26 pemphigus major, Buccinum, p. 7 phenax, Musculus, p. 4 picta, Tegula (Agathistoma), p. 22 pigafettae, Nucula, p. 4 piutensis, Helminthoglypta cuyamacensis, p. 13 poormani. Conus, p. 9 porterae, Ashmunella thomsoniana, p. 7 pribiloffensis, Chrysodomus, p. 9 pronotis, Monadenia fidelis, p. 17 psaltes, Dendrochiton, p. 26 punctifissa, Diodora, p. 10 punctocoelata vancouverensis, Acteon, p.5 purdyae, Terebra, p. 23 purpureus, Heterodonax bimaculatus, p.2 pusilla, Benthocardiella, p. 1 quentinensis. Cryptogemma, p. 10 randolphi, Eulima, p. 11 raninus nanus, Strombus, p. 21 redondoensis, Aligena, p. 1 redondoensis, Cardita ventricosa, p. 2 redondonensis, Nuculana, p. 4 redondoensis, Pseudomelatoma semiinflata, p. 19 reediana, Helminthoglypta, p. 13 rejecta, Oliva, p. 18 remota, Micrarionta indioensis, p. 16 rex, Helminthoglypta tudiculata, p. 13 ricaudae, Elaeocyma, p. 10 roberti, Haliotis (Padollus), p. 12 rogersi, Lithophaga (Labis) attenuata, p. 4 rosa, Melanella, p. 15 rosea, Corbula, p. 2 30 Contributions in Science No. 213 roseola, Leurolepas, p. 14 rubra, Trivia ovulata, p. 24 rugosa naticanim, Crepidula, p. 10 rugosa norrisiarum, Crepidula, p. 10 salmoneus, Heterodonax bimaculatus, p.2 salvatori, Cerion, p. 8 sanjaimense, Calliostoma, p. 8 santacruzensis, Latiaxis (Babelomurex), p. 14 sculpturata, Bela, p. 7 semiinflata redondoensis, Pseudomelatoma, p. 19 sharoni, Lamellaria, p. 14 shaskyi, Nassarius, p. 17 shyana, Terebra, p. 23 shyorum, Epitonium (Epitonium), p. 11 sigillata, Natica, p. 17 similans, Helminthoglypta, p. 13 simmonsi, Sonorella, p. 21 sinistra, Urocoptis, p. 25 sitkana atkana, Littorina, p. 14 socorroensis, Columbella, p. 9 spectabilospina, Macarene, p. 15 sphoni, Olivella (Olivella), p. 18 splendidula, Haliotis cracherodii, p. 12 spirellum, Speleodiscoides, p. 21 stageri, Helminthoglypta, p. 13 stanfordiana, “Acmaea”, p. 5 stellata, Arene, p. 7 sterkiana, Pupa, p. 19 stewartiana, Kerguelenella, p. 14 stilesi, Neptunea, p. 18 stohleri, Terebra, p. 23 strong!, Turbonilla (Pyrgolampros), p. 24 tagolandensis, Helicostyla leucophthalma, p. 12 tehuanarum, Amaea (Scalina), p. 6 tehuantepecensis, Radwinia, p. 19 tenuissima, Volvulella, p. 26 thomsoniana porterae, Ashmunella, p. 7 tinctum bormanni, Epitonium, p. 11 traski pacoimensis, Helminthoglypta, p. 13 tudiculata rex, Helminthoglypta, p, 13 tyrianthina, Acanthina, p. 5 umnaka, Cardita, p. 2 unifasciata, Micrarionta hutsoni, p. 16 vaccaria, Aplysia, p. 6 vancouverensis, Acteon punctocoelata, p. 5 velero, Poirieria (Panamurex), p. 19 veleroae, Calliostoma, p. 8 veleroae, Morum ( Cancellomorum ) , p. 17 ventricosa montereyensis, Cardita, p. 2 ventricosa redondoensis, Cardita, p. 2 verdispira, Tegula (Agathistoma), p. 22 vernicosa, Astarte, p. 1 verrucosa, Tegula (Agathistoma), p. 22 waltoni, Sonorella, p. 21 willetti, Turritella, p. 25 williamsi, Helminthoglypta arrosa, p. 13 winslowae, Arene, p. 7 Literature Cited Baker, F. C. and V. D. P. Spicer. 1930. New species of mollusks. Trans. San Diego Soc. Nat. Hist. 6(6) : 177-178. Bales, B. R. 1942. A new subspecies of S trombus rainus Gmelin. Nautilus 56(1 ) : 18-19. Bartsch, P. 1919. Two new landshells from California. Proc. Biol. Soc. Wash. 32:248. Berry, S. S. 1931. A new California race of Monadenia. Nautilus 44(4) : 122-123. 1945. Two new chitons from the Gulf of California. Amer. Midi. Nat. 34(2):491-92. 1950. A partial review of some west American species of Crepidula. Leaf- lets in Malacoloty 1(8) :35-39. 1953a. West American razor-clams of the genus Ensis. Trans. San Diego Soc. Nat. Hist. 11 (15): 198-400. 1953b. Notices of new west American marine Mollusca. Trans. San Diego Soc. Nat. Hist. 11(16) :417-19. 1957. Notices of new eastern Pacific Mollusca. — I. Leaflets in Malacology l(14):76-79. 1971 Type Specimens of Recent Mollusca 31 1959. Notices of new eastern Pacific Mollusca. — III. Leaflets in Malacol- ogy 1(18):110-11. 1963. Diagnoses of new eastern Pacific chitons. — II. Leaflets in Mala- cology 1(22): 135-38. - 1968. Notices of new eastern Pacific Mollusca. — VII. Leaflets in Mala- cology l(25):156-57. ■ 1969. Notices of new eastern Pacific Mollusca. — VIII. Leaflets in Mala- cology 1(26): 159-66. Bormann, M. 1946. A survey of some west American Ocenebras, with description of a new species. Nautilus 60(2) : 37-43. Bratcher, T. L. and R. D. Burch. 1970a. Five new species of Terebra from the eastern Pacific. The Veliger 12(3) :295-300. 1970b. Four new Terebrid gastropods from eastern Pacific islands. Los Angeles Co. Mus., Contrib. Sci. 188: 1-6. Burch, J. Q. 1944. West North American marine Mollusca from San Diego, Calif, to Alaska. Min. Conch. Club So. Calif. 33:7-14. 1949. A new Trigonostoma from central America. Min. Conch. Club So. Calif. 94:3-4. Burch, J. Q. and R. L. Burch. 1962. New species of Oliva from west Mexico. Nautilus 75(4) : 166. 1967. Anew Ancilla from Brazil. Nautilus 80(3): 81-82. Burch, J. Q. and G. B. Campbell. 1963. Four new Olivella from Gulf of Cali- fornia. Nautilus 76(4) : 120-26. Burch, R. D. 1965. New Terebrid species from the Indo-Pacific Ocean and from the Gulf of Mexico, with new locality records and provisional lists of species collected in western Australia and at Sabah, Malaysia (Mollusca: Gastropoda). The Veliger 7(4): 241-54. Burch, T. A. 1938. A new Pseudomelatoma from California. Nautilus 52(1 ) : 21-22. 1941. A survey of the west American Aligenas with a description of a new species. Nautilus 55(2) :50-51. 1944. West North American Mollusca from San Diego, California to Alaska. Min. Conch. Club So. Calif. 39:5-26. Campbell, G. B. 1961. Four new Panamic gastropods. The Veliger 4(1) :25-28. 1964. New Terebrid species from the eastern Pacific. The Veliger 6(3) : 132-38. Chace, E. P. and E. M. Chace. 1935. A new variety of Monadenia fidelis from Curry County, Oregon. Nautilus 49(2) :48-49. Clench, W. J. 1927. A new subspecies of Thais from Louisiana. Nautilus 41(1): 6-8. 1937. Three new species of Cerions from Long Island, Bahamas. Nautilus 51(l):20-23. 1952. Land and freshwater mollusks of Eleuthera Island, Bahama Islands. Revista de la Sociedad Malacologica “Carlos de la Torre” 8(3) : 107. CoAN, E. V. AND B. Roth. 1965. A new species of Persicula from west Mexico. The Veliger 8(2) :67-69. Cowan, I. McT. and J. H. McLean. 1968. A new species of Puncturella (Cranopsis) from the northeastern Pacific. The Veliger 11(2): 105-08. Church, C. C. and A. G. Smith. 1938. A giant race of Helminthoglypta from Tulare Co., California. Nautilus 51(4) : 1 19-22. 32 Contributions in Science No. 213 Dale, W. H. 1879. Report on the limpets and chitons of the Alaskan and arctic regions, with descriptions of genera and species believed to be new. Proc. U.S. Nat. Mus. 1:281-344. 1882. On certain limpets and chitons from the deep waters off the eastern coast of the United States. Proc. U.S. Nat. Mus. 4(246) : 400- 14. 1886. Contributions to the natural history of the Commander Islands. No. 6 — -report on Bering Island Mollusca collected by Mr. Nicholas Grebnitzki. Proc. U.S. Nat. Mus. 9:211-12. 1887. Supplementary notes on some species of mollusks of the Bering Sea and vicinity. Proc. U.S. Nat. Mus. 9(571) :297-309. 1889. Reports on the results of dredging, under the supervision of Alex- ander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U.S. Coast Survey steamer “Blake,” Lieut.-Commander C. D. Sigsbee, U.S.N., and Commander J. R. Bartlett, U.S.N. commanding. XXIX. — Report on the Mollusca. Bull. Mus. Comp. Zool. 18:1-492. 1893. On a new species of Yoldia from California. Nautilus 7(3) : 29-30. - 1897. Notice of some new or interesting species of shells from British Columbia, and the adjacent region. Nat. Hist. Soc. Brit. Columbia Bull. 2:9. 1902. Synopsis of the family Veneridae and of the North American recent species. Proc. U.S. Nat. Mus. 26(1312) :402. 1903. Synopsis of the family Astartidae with a review of the American species. Proc. U.S. Nat. Mus. 26:945-49. 1908. Reports on the dredging operations off the west coast of central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Com- mission steamer “Albatross,” during 1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding. XXXVII. Reports on the scientific results of the expedi- tion to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission steamer “Albatross,” from October, 1904, to March, 1905, Lieut. L. M. Garret, U.S.N. commanding. XIV. The Mollusca and the Brachi- poda. Bull. Mus. Comp. Zool. 43:203-487. 1915. A new species of Modiolaria from Bering Sea. Nautilus 28(12) :138. 1916. Diagnoses of new species of marine bivalve mollusks from the north- west coast of America in the collection of the United States National Museum. Proc. U.S. Nat. Mus. 52(2183) :401. 1917. Summary of the mollusks of the family Alectrionidae of the west coast of America. Proc. U.S. Nat. Mus. 51 (2166): 575-79. 1919a. 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) :323-26. 1919b. Descriptions of new species of mollusks of the family Turritidae from the west coast of America and adjacent regions. Proc. U.S. Nat. Mus. 52(2288): 1-86. Dale, W. H. and P. Bartsch. 1910. New species of shells collected by Mr. John Macoun at Barkley Sound, Vancouver Island, British Columbia. Mem. Canad- ian Dept. Mines 14-N, pp. 1-22. DuShane, H. 1969. A new genus and two new species of Typhinae from the Pa- namic province ( Gastropoda : Muricidae ) . The Veliger 1 1(4) :343-45. 1970a. Two new Epitoniidae from the Galapagos Islands (Mollusca: Gastropoda). The Veliger 12(3) : 330-32. 1970b. Five new Epitoniid gastropods from the west coast of the Amer- icas. Los Angeles Co. Mus., Contrib. Sci. 185:1-6. 1971 Type Specimens of Recent Mollusca 33 DuShane, H. and T. L. Bratcher. 1965. A new Scalina from the Gulf of Cali- fornia. The Veliger 8(2): 160-61. DuShane, H. and J. H. McLean. 1968. Three new Epitoniid gastropods from the Panamic province. Los Angeles Co. Mus., Contrib. Sci. 145:1-6. Emerson, W. K. 1968. A new species of the gastropod genus Morum from the eastern Pacific Ocean. J. de Conchyl. 107(1) :53-57. Emerson, W. K. and A. D’Attilio. 1969. Remarks on the taxonomic placement of Purpurellus Jousseaume, 1880, with the description of a new species. The Veliger 12(2) : 146-48. 1970. Aspella myrakeenae, new species from western Mexico. Nautilus 83(3):88-95. Fare ANTE, I. P. 1943. The genera Fissurella, Lucapina and Lucapinella in the west- ern Atlantic. Johnsonia 1(10): 1-20. Gordon, M. 1938. A new Turbonilla from Redondo Beach, California. Nautilus 52(2):49-51. Gregg, W. O. 1931. Two new helicoids from Los Angeles County, California. Nautilus 45(2): 49-51. 1948. A new and unusual helcoid snail from Los Angeles County, Cali- fornia. Bull. So. Calif. Acad. Sci. 47(3) : 100-02. 1951. A new Sonorella from the Chirichaua Mountains, Arizona. Bull. So. Calif. Acad. Sci. 50(3) : 156-58. 1953. Two new land snails from Arizona. Bull. So. Calif. Acad. Sci. 52(2):74-75. Gregg, W. O. and W. Miller. 1969. A new Sonorella from Phoenix, Arizona. Nautilus 82(3):90-93. Hanna, G. D. and A. G. Smith. 1937. Some notes on an old race of California land snail with descriptions of three new forms. Nautilus 51(1) : 10-17. Henderson, J. 1934. Some western fresh water mollusks. Nautilus 47(3) : 86-91. Howard, F. B. 1963. Description of a new Pyrene from Mexico ( Mollusca-Gastro- poda). Occ. Pap. Santa Barbara Mus. Nat. Hist. 7:1-12. International Commission on Zoological Nomenclature. 1964. International code of zoological nomenclature, adopted by the XV International Congress of Zoology. Inst. Trust Zool. Nomenclature, London. 176 p. Keen, A. M. 1938. New pelecypod species of the genera Lasaea and Crassinella. Proc. Malac. Soc. London. 23(1) :26-28. Keen, A. M. and G. B. Campbell. 1964. Ten new species of Typhinae (Gastro- poda: Muricidae). The Veliger 7(1) :46-57. Lowe, H. N. 1930. A new land shell from San Benito County, California. Nautilus 44(2) :43. Marshall, B. 1924. New species of mollusks of the genus Chi Una. Proc. U.S. Nat. Mus. 66(13): 1-5. Mattox, N. T. 1958. Studies on the Opisthobranchiata : II. A new tectibranch of the genus Philine. Bull. So. Calif. Acad. Sci. 57(2) : 98- 104. McLean, J. H. 1964. New species of recent and fossil west American Aspidobranch gastropods. The Veliger 7(2) : 129-133. — — _ — 1966. A new genus of Fissurellidae and a new name for a misunderstood species of west American Diodora. Los Angeles Co. Mus., Contrib. Sci. 100:1-8. 1967. West American species of Lucapinella. The Veliger 9(3) :349-52. 34 Contributions in Science No. 213 1970a. New species of Panamic marine gastropods. The Veliger 12(3) : 310-15. 1970b. Descriptions of a new genus and eight new species of eastern Pacific Fissurellidae, with notes on other species. The Veliger 12(3) :362-67. 1970c. Notes on the deep water Calliostomas of the Panamic province, with descriptions of six new species. The Veliger 12 (4): 42 1-26. 1970d. New species of tropical eastern Pacific Gastropoda. Malacological Review 2:115-30. McLean, J. H. and L. H. Poorman. 1970. Reinstatement of the turrid genus Bella- spira Conrad, 1868 ( Mollusca : Gastropoda ) with a review of the known spe- cies. Los Angeles Co. Mus., Contrib. Sci. 189:1-11. Miller, W. 1966. Three new Sonorella from southwest Arizona. Nautilus 80(2) : 46-48. 1968. New Sonorella from Arizona. Nautilus 82(2) :61-63. Oldroyd, I. S. 1927. The marine shells of the west coast of North America. Stan- ford Univ. Pub. Geol. Sci. 2(1): 1-297. PiLSBRY, H. A. 1889. New and little-known American mollusks, no. 2. Proc. Acad. Nat. Sci. Philadelphia 41 :412. 1908. Land shells of Tangulandang (Tangolanda). Nautilus 22(4-5) : 47. PiLSBRY, H. A. AND T. D. A. CocKERELL. 1899. Another new Ashmunella. Nautilus 13(5):49-50. PiLSBRY, H. A. AND H. N. LowE. 1932. West Mexican and central American mol- lucks collected by H. N. Lowe, 1929-31. Proc. Acad. Nat. Sci. Philadelphia 84:33-144. Powell, A. W. B. 1924. Description of a new subspecies of Alcithoe arabica Mar- tyn, from New Zealand. Proc. Malac. Soc. London 16(3) : 108-09. 1927a. The genetic relationships of Australasian rissoids. Trans. New Zealand Instit. 57:547. 1927b. Mollusca from twenty-three fathoms off Ahipara, N. Z. Trans. and Proc. New Zealand Instit. 58(3) :297. 1928. The recent and tertiary cassids of New Zealand and a study in hybridization. Trans, and Proc. New Zealand Instit. 59(3) :640. 1930. New species of New Zealand Mollusca from shallow-water dredg- ings. Trans. New Zealand Instit. 60:533-34. 1931. Descriptions of some new species of recent Mollusca. Records of the Canterbury Mus. 3:376. 1933. New species of marine mollusks from the subantarctic islands of New Zealand. Proc. Malac. Soc. London. 20:233. 1934. Gastropods new to the New Zealand fauna. Trans. Royal Soc. New Zealand. 64:154-55. 1939. The Mollusca of Stewart Island. Records of the Auckland Instit. and Mus. 2:237-38. Rheder, H. 1943. New marine mollusks from the Antillean region. Proc. U.S. Nat. Mus. 93(3161): 194-94. Schenk, E. T., H. H. McMasters, A. M. Keen, and S. W. Muller. 1956. Pro- cedure in Taxonomy. Stanford Univ. Press, Stanford. 149 p. SCHWENGEL, J. S. 1944. A new Floridian Lamellaria. Nautilus 58(1 ) : 17-18. Shasky, D. R. 1961. New deep water mollusks from the Gulf of California. The Veliger 4(1): 18-21. 1970. New gastropod taxa from tropical western America. The Veliger 13(2):188-95. i f » I I 1 IL I 1971 Type Specimens of Recent Mollusca 35 Shaw, H. O. N. 1909. Notes on the genera Cypraea and Trivia. Proc. Malac. Soc. London 8(5):312. Smith, A. G. 1938. Three new subspecies of Helminthoglypta arrosa (Gould). Nautilus 51(3): 83-85. 1943. Mollusks of the Clearwater Mountains, Idaho. Proc. Cal. Acad. Sci. 4th Series, 23(36): 545-47. 1957. Snails from California caves. Proc. Cal. Acad. Sci. 4th Series, 29(2):24-26. 1968. A new Neptunea from the Pacific northwest. The Veliger 11(2): 117-120. Smith, A. G. and M. Gordon. 1948. The marine mollusks and brachiopods of Monterey Bay, California, and vicinity. Proc. Cal. Acad. Sci. 4th Series, 26(8): 147-245. SowERBY, G. B. 1900. On some marine shells from Pondoland and The Kowie, with descriptions of seventeen new species. Proc. Malac. Soc. London 4(1) :3. 1901. On seven new species of marine Mollusca collected by Dr. H. Becker at “The Kowie,” South Africa. Proc. Malac. Soc. London 4(5):213. 1915. Descriptions of new species of Mollusca from various localities. Ann. and Mag. Nat. Hist. 8th Series, 16:165. Sphon, G. G. 1969. Notes on the Mitridae of the eastern Pacific II — The genus Thala, with the description of a new species. The Veliger 12(l):84-88. Stohler, R. 1959. Two new species of west North American marine gastropods. Proc. Cal. Acad. Sci. (4th Series) 29(1 1 ) :439-444. Strong, A. M. 1937. Notes on west coast Epitoniidae. Nautilus 51(1) :5-6. 1941. Notes on Epitonium (Nitidoscala) tinctum (Carpenter). Nautilus 55(2) :47. Talmadge, R. R. 1962. A new land snail from the Klamath Mountains, California ( Mollusca :Pulmonata: Poly gyridae). The Veliger 5(l):28-29. 1964. The races of Haliotis fulgens Philippi: (Mollusca: Gastropoda). Trans. San Diego Soc. Nat. Hist. 13(18): 375. 1966. A new Haliotid from Guadalupe Island, Mexico (Mollusca: Gastro- poda). Los Angeles Co. Mus., Contrib. Sci. 109:1-4. Torre, C. de la. 1912. New Cuban Urocoptis of the U. cinerea group. Nautilus 26(4):57. 1928. No description, figure only. Nautilus 42(l):plate 1, fig. 8. 1929a. No description, figure only. Nautilus 42(3):plate 4, fig. 7. — -. 1929b New Cuban Urocoptidae. Proc. Acad. Nat. Sci. Philadelphia 81:445. — 1932. Some new Cuban Urocoptis. Nautilus 45(3):88-89. Torre, C. de la and P. Bartsch. 1938. The Cuban operculate land shells of the subfamily Chondropominae. Proc. U.S. Nat. Mus. 85(3039) : 193-403. 1941. The Cuban operculate land mollusks of the family Annulariidae, exclusive of the subfamily Chrondropominae. Proc. U.S. Nat. Mus. 89(3096): 131-385. Torre, C. de la and W. J. Clench. 1930. West Indian mollusks. no. 1: — Two new varieties of Urocoptis livida Torre. Nautilus 44(1): 15-16. Torre, C. de la and H. A. Pilsbry. 1927. Cuban land shells of the Lowe-Hand- Emery expedition. Nautilus 40(3): 74-75. Turner, R. 1955. The family Pholadidae in the western Atlantic and eastern Pacific Part II — Martesiinae, Jouannetiinae and Xylophaginae. Johnsonia 3(34): 65-160. 36 Contributions in Science No. 213 Vanatta, E. G. 1899. West American Eulimidae. Proc. Acad. Nat. Sci. Philadel- phia for 1889. p. 256. Yokes, E. H. 1970a. The west American species of Murexiella, (Gastropoda: Muricidae) including two new species. The Veliger 12(3) :325-29. 1970b. Cenozoic Muricidae of the western Atlantic region Part V — Pterynotus and Poirieria. Tulane Studies in Geol. and Paleon. 8(1 & 2): 1-50. Weyrauch, W. 1957. Sieben neue Clausiliiden aus Peru. Archiv Fiir Mollusken- kunde 86(1-3) : 18-21. Weyrauch, W. and A. Zilch. 1954. Landschnecken aus Peru, 2. Archiv Fur Mol- luskenkunde 83 ( 1-3) : 70-73. Williamson, M. B. 1892. An annotated list of the shells of San Pedro Bay and vicinity. Proc. U.S. Nat. Mus. 15(898) : 187. 1905a. New varieties of Crepidula rugosa Nutt, found on Natica and on Norrisia. Nautilus 19(5) :50-51. 1905b. Some west American shells — including a new variety of Corbula luteola Cpr. and two new varieties of gastropods. Bull. So. Calif. Acad. Sci. 4(8):118-29. Willett, G. 1928. Notes on some Pacific coast Acteocinas, wtih description of one new subspecies. Nautilus 42(2): 37-38. 1929a. Description of two new species of molluscs from the west coast of North America. Nautilus 43(1) : 26-28. 1929b. Description of two new land shells from Southern California. Bull. So. Calif. Acad. Sci. 28(2): 16-18. 1930a. Desert helicoids of the Micrarionta hutsoni group. Nautilus 44(1):6. 1930b. The Micrariontas of the indioensis group, with the description of a new subspecies. Nautilus 43(4) : 115-16. 1931a. Two new helicoids from the Mohave Desert, California. Nautilus 44(4): 123. 1931b. Three new marine mollusks from Catalina Island, California. Nautilus 45(2) :65-67. 1932a. A new chiton from Southern California. Nautilus 45(3) : 101-02. 1932b. A new Helminthoglypta from Monterey County, California. Nau- tilus 45(4) : 134-35. 1932c. A new Cardita from the Aleutian Islands and a new Epitonium from Southern California. Trans. San Diego Soc. Nat. Hist. 7(9) :85-90. — 1934a. Micrariontas of desert ranges bordering the east side of Coachella Valley and Salton Sink, California. Bull. So. Calif. Acad. Sci. 38(1) : 15. — 1934b. A new Helminthoglypta from the east slope of the Sierra Nevada, California. Bull. So. Calif. Acad. Sci. 33(2) :57. 1934c. Two new Cingulas from Alaska. Nautilus 47(3) : 103-04. 1935a. Three new Micrariontas from the central Colorado Desert, Cali- fornia. Nautilus 49(1 ) : 15. 1935b. Further notes on the desert snails of Riverside County, California. Bull. So. Calif. Acad. Sci. 34(1): 1-2. 1937a. A new land shell from the Riverside Mountains, Colorado Desert. Bull. So. Calif. Acad. Sci. 36(l):6-7. 1937b. Micrariontas of the southwestern Colorado Desert. Nautilus 50(4): 123-24. 1937c. A new Callistochiton from Lower California. Nautilus 51(1): 25-26. 1971 Type Specimens of Recent Mollusca 37 1938a. Two new land shells from Kern County, California. Bull. So. Calif. Acad. Sci. 37(2):53-54. 1938b. A new Pseudochama from Clarion Island, Mexico. Nautilus 52(2) : 48-49. 1939. Description of a new mollusk from California. Nautilus 52(4): 123-24 . 1940a. A new land shell from Lower California. Bull. So. Calif. Acad. Sci. 39(2):80-82. 1940b. Contributions from the Los Angeles Museum-Channel Islands Biological Survey 13. A new Ischnochiton from Catalina Island, California. Bull. So. Calif. Acad. Sci. 39(3) : 185-86. 1943. Northwest American species of Glycimeris. Bull. So. Calif. Acad. Sci. 42(3):114. 1944. New species of mollusks from Redondo, California. Bull. So. Calif. Acad. Sci. 43(2):71-72. 1946. A new mollusk of the genus Solariella from Alaska. Bull. So. Calif. Acad. Sci. 45(1) :32. Winkler, L. R. 1955. A new species of Aplysia on the Southern California coast. Bull. So. Calif. Acad. Sci. 45(1): 5-7. Accepted for publication November 19, 1970. i .■■A..''''-; '■ ./.’A-v; ..•rr«“-}: -A'' - ‘tf'A ‘ W 4-M - ■t A' ••=. ' ■■ ' »»»TkV ■vit».'«.- .... ,.;•’.... A.^-.AAAAAf T' ""' "" .' ■' •,.* i':..i'‘y ■ ', '= '-■ ■ " V.' i' J .>a,.:AAj^A;aa^^ '"■‘■ AAA;t:;y^^ vA',;;;* .^VA ' -.A-:" A::j^ Cvfclf J .' •. • ^ h V- 0- ■:.i|:,;..'-P.,. . ■'"A^ :-A 4^' ■ r'. %: .^’is A'-' . a'A , 54a "’ V. t- . ^ ' -Aeiv - • i Printed in Los Angeles, California by Continental Graphics LOS ANGELES COUNTY " MUSEUM CONTRIBUTIONS IN SCIENCE jjMBER 214 May 28, 1971 STUDIES ON CALIFORNIA ANTS. 6. THREE NEW SPECIES OF MYRMECOCYSTUS (HYMENOPTERA: FORMICIDAE) :i 1 j| i I By Roy R. Snelling i i Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 STUDIES ON CALIFORNIA ANTS. 6. THREE NEW SPECIES OF MYRMECOCYSTUS (Hymenoptera: Formicidae) ‘ ' -Ry Roy R. Smelling^ Abstract: Three new species of honey ants are herein described from southern California: M. ewarti from Shaver’s Well, Riverside Co. {mexicanus group) ; M. creightoni from Pear- blossom, Los Angeles Co. (lugubris group); M. wheeled from Los Angeles Co. {wheeled group). All castes of the three species are described and illustrated. The following three new species are described in advance of my forth- coming revision of Myrmecocystus to provide names for a study to be pub- lished by G. C. Wheeler. Detailed distribution data, other than citation of type material are not given. These data will be presented in the revision. In the interests of conserving space in the description, some features of the head and thorax are indicated by code letters: Cl, Cephalic Index — (HW/ HL) (100); EL, Eye Length— With the head in full face view, the maximum length of the compound eye; HL, Head Length— With the head in full face view, the anterior clypeal margin and occipital margin on the same plane, the maximum distance between these points along the midline; HW, Head Width— With the head in full face view, the greatest measurable distance between head margins below the eyes (worker and female) or at the level of the upper eye margin (males) ; lOD, Interocellar Distance— With head in full face view, the maximum distance between the inner margins of the posterior (or lateral) ocelli (Sexual castes only) ; OD, Ocellar Diameter— The transverse diameter of the anterior (or middle) ocellus, with the head in full face view (Sexual castes only) ; OMD, Oculo-Mandibular Distance — With the head in full face view, the distance between the lower margin of the eyes and mandibular bases; OOD, Ocellar-Ocular Distance— With the head in full face view, the minimum hodzontal distance between the posterior ocelli and the inner eye margin (Sexual castes only) ; PW, Pronotal Width— In dorsal aspect, the greatest width of the pronotum (workers) immediately anterior to the wing bases (sexual castes) ; SI, Scape Index — (HW/SL) ( 100) ; SL, Scape Length— Maximum measurable scape length, excluding basal condyle; WL, Weber’s Length— Diagonal length in lateral aspect of thorax, from anterior pronotal margin to posterior margin of propodeum ( = epinotum). Measurements are in millimeters (mm). Each description begins with a series of measurements, indicating the range in the material studied. Each range is followed by a number in parentheses, indicating the measurement for the holotype, thus: HL 0.93-1.23 (1.20) reads [HL = 0.93-1.23 mm (holo- type, 1.20 mm)]. The term “propodeum” is used for the posterior portion ^Entomology Section, Los Angeles County Museum of Natural History, Los Ange- les, Calif. 90007. 1 2 Contributions in Science No. 214 of the alitrunk in preference to “epinotum,” more widely used by myrmecolo- gists. The usage here is harmonious with that of other studies on hymenopter- ous insects (Wilson, 1955; Cole, 1968). Holotypes of the three species described below are in the collection of the Los Angeles County Museum of Natural History. Paratypes will be distributed to the American Museum of Natural History (AMNH), Museum of Comparative Zoology (MCZ), Museum of Natural History, Geneva (MNHG), United States National Museum of Natural History (USNM) and the private collections of Professors A. C. Cole, Jr., W. S. Creighton, and G. C. Wheeler. Figures for this paper were prepared by Ruth A. DeNicola under Grant No. 4494, the Penrose Fund, of the American Philosophical Society. Myrmecocystus ewarti, new species Figure 1 Diagnosis. — A member of the mexicanus group closely related to M. pyramicus M. Smith: propodeum produced upward at junction of anterior and posterior faces; petiolar scale strongly compressed in profile; pronotum with two or more fully erect white hairs; disc of first gastric tergite with scat- tered erect white hairs; hind tibiae with few or no erect hairs. Measurement. -m. 0.76-1.30 (1.23); HW 0.70-1.30 (1.23); SL 0.93-1.36 (1.36); WL 1.10-1.80 (1.70); PW 0.46-0.83 (0.83). Head shape varying from longer than broad in most workers to slightly broader than long in largest workers. Cl 88-104 (100), a little shorter than scape; SI 102-136 (110). In frontal view head broadest at lower margin of eyes, sides slightly convex to straight, narrowed toward mandibular insertions. Occiput, in frontal view, somewhat flattened in middle, sides convex, not at all angulate. Eye large, 1.5 x length of first flagellar segment, EL 0.90-1.15 (0.93) X OMD. Mandible with seven distinct teeth, often with a small intercalary denticle between the penultimate and basal teeth. Thorax slender to moderately robust, PW 0.37-0.50 (0.48) x WL. Basal face of propodeum pyramidally produced upward at juncture with posterior face, about half as long as posterior face. Petiole compressed when viewed in profile, crest thin, weakly angularly excised in middle; in dorsal view twice as wide as long. Vestiture.-lB.iQct hairs sparse on head, confined to clypeus, frontal lobes and occipital areas. Erect pronotal pilosity sparse, but with at least a pair of fine, fully erect hairs which are about as long as apical width of scape; mesonotum with 3-6 erect hairs; propodeum without erect hairs at summit of declivity, or with one or two which are less than half as long as those of mesonotum. Petiolar scale with a few very short, inconspicuous erect hairs on crest. Disc of first tergite with scattered, short, fully erect blunt hairs; second and succeeding tergites with progressively longer discal hairs. Tibiae with very sparse, fine, decumbent to subdecumbent hairs on outer surfaces. 1971 Studies on California Ants 3 these shorter and finer than the row of gradated bristles on the inner surface. Appressed pubescence very fine, sparse on head, a little more abundant on occiput, especially behind eyes; thoracic pubescence denser than cephalic, but not obscuring surface, denser on sides of propodeum than elsewhere. First three gastric tergites with dense, fine pubescence which does not obscure surface, fourth and fifth with very sparse pubescence. Integument. — T>u\\, everywhere very finely coriaceous, with conspicuous round sparse punctures on clypeus; cheeks more densely coriaceous, with scattered elongate punctures; frontal lobes with scattered micropunctures. Co/or. — Light brownish yellow, legs and gaster more yellowish; mandibu- lar margins somewhat ferruginous. FEMALE. Measurements: HL 1.47; HW 1.63; SL 1.42; EL 0.57; OMD 0.52; WL 3.6; PW 1.89. Head broader than long. Cl = 90, longer than scape; SI = 87. In frontal view head broadest behind eyes, sides converging slightly toward mandibular insertions. Occiput, in frontal view flat, with well-rounded corners. Eye large, 1.8 X length of first flagellar segment; removed from mandibular insertion by a little less than its length (11:10). Lateral ocelli separated by three times diameter of anterior ocellus. Thorax robust, PW 0.52 x WL. In profile posterior two-thirds of meso- scutum and anterior half of scutellum on same plane; me^anotum not protruding. Petiole, in profile, compressed, crest sharp; distinctly notched; from above, about three times wider than long (not clearly visible in only available specimen). Vestiture. — ^XQci cephalic pilosity as described for worker; pronotum with a few erect hairs, along anterior margin; mesoscutum and scutellum with scattered long erect yellowish hairs arising from coarse punctures; pleurae with about a dozen long, erect yellowish hairs; propodeum without con- spicuous erect hairs; first tergite with scattered subdecumbent to erect yellow- ish hairs on disc; second and following tergites with fully erect yellowish hairs longer, a little more abundant; inner face of fore femora without conspicuous fully erect hairs, though 15 + are present on ventral surface; middle and hind tibiae with numerous decumbent, fine yellowish hairs. Pubescence long, yellowish, appressed to decumbent on head, thorax and appendages; fully appressed and abundant on first three tergites, con- spicuously sparser on fourth. MAl.'E.-Measurements: HL 0.66-0.76; HW 0.63; SL 0.73-0.76; EL 0.33; OMD 0.16; WL 1.33-1.43; PW 0.83-1.00. Margins of head slightly convergent toward mandibular insertions; head a little longer than broad (Cl 90-95), a little shorter than scape; SI 115-121; OMD 0.50 X EL; anterior ocellus little smaller than lateral ocelli; lOD 3.0-3. 5 X OD; OOD 1.0- 1.5 x OD. Mandible with preapical tooth or preapical notch 4 Contributions in Science No. 214 1971 Studies on California Ants 5 before apical tooth. Clypeus with or without obscure preapical transverse depression. Petiole, in profile, distinctly higher than long, sharply cuneate; in frontal view, sides convergent toward narrow, flat, medially notched crest; in dorsal view, about twice as wide as long. Terminalia: Fig. ICc-e. V e St it ure. — Erect hairs yellowish, sparse on head and thorax, longest on scutellum, where they are about equal to minimum diameter of eye; propodeum without conspicuous erect hairs; tibiae and scapes without erect hairs. First two tergites with sparse, short erect hairs, third and following segments with conspicuous long, scattered hairs. Pubescence sparse and inconspicuous on head and thorax, conspicuously denser on propodeum above and on first two tergites, forewings with conspicuous fringe hairs from stigma to apex, around apical margin to vein Cu-A; apical and hind margins of hind wing fringed. Integument. — ModevditeXy shiny, with piligerous micropunctures, a few scattered coarse punctures on scutum and mesopleura. Co/or. — Uniformly brownish, appendages yellowish to yellowish brown. Wings whitish hyaline, stigma and veins pale yellowish. Type material.— EioXotype worker, allotype male, 19, 16^^, 223 9? paratypes, Shaver’s Well, 3 mi W, Riverside Co., California, 1 March 1964 (R. R. Snelling). Holotype, allotype and most paratypes in LACM; nine paratypes in each of the following: AMNH, MCZ, MNHG, USNM and pri- vate collections of A. C. Cole, Jr., W. S. Creighton, and G. C. Wheeler. Etymology. — Th\s> species is named for Dr. William Ewart of the Uni- versity of California, Riverside, evidently the first person to collect this species. Dr. Ewart’s series was sent to me by Professor Creighton who had recognized the species as new and suggested the name. Present data indicate that this species replaces the similar M. pyramicus in southern California on the Mojave and Colorado Deserts. It is probably to be found also in adjacent portions of Arizona, Sonora, and Baja California. Discussion.— The workers of M. ewarti may be separated from those of M. pyramicus by the presence of a number of erect hairs on the pronotum and first tergite. A pair of pronotal hairs, which seems always to be present in M. ewarti, is as long as or longer than the apical breadth of the scape. Although occasional specimens of M. pyramicus may have one or two erect pronotal hairs, they are always much shorter. Erect hairs are present on the hind tibiae of M. pyramicus but absent in M. ewarti. The median area of the clypeus of Figure 1. Myrmecocystus ewarti, new species. A, female, lateral view; Aa, female head, frontal view; Ab, female mesoscutum. B, worker major, lateral view; Ba, worker major, head, frontal view; Bb, worker minor, head, frontal view; Be, worker petiole, profile and posterior views. C, male, lateral view; Ca, male head, frontal view; Cb, male mandible; Cc, male ninth sternite; Cd, male volsella, inner view; Ce, male aedeagus, lateral view. 6 Contributions in Science No. 214 M. ewarti has four or more long, erect hairs; in M. pyramicus clypeal hairs are confined to the margins. The single female available of M. ewarti is very similar to those of M. pyramicus but has a number of erect hairs on the clypeal disc, the antennal scapes possess numerous fine suberect hairs, long erect hairs are abundant on the outer face of the fore femora and there are erect hairs on the hind tibiae. The cheeks of M. ewarti are less sharply coriaceous, the punctures below the eyes are larger and are round (clearly elongate in M. pyramicus) . The best character to separate the females of these species seems to be that of mesocutal punctation. The area between the parapisides in M. ewarti is rather uniformly finely, sparsely piligerously punctate. There are, in addition, a number of much coarser, setigerous punctures scattered over the disc. The piligerous punctures, laterad of the parapsides, are little coarser than those of the median area, and are mostly separated by two or more times a puncture diameter. In M. pyra- micus, the center of the mesoscutum is virtually impunctate, the setigerous punctures are fewer and less conspicuous, and the punctures laterad of the parapsides are much coarser than those of the median area, and are mostly separated by a puncture diameter or less. The males are very similar and, until more specimens of M. pyramicus are available, the differences noted here must be considered provisional. In size, M. pyramicus males are conspicuously longer; head length of males of this species exceeds 0.90 mm, while that of M. ewarti is less than 0.80 mm. However, Myrmecocystus males vary greatly in size within a single colony, so the size difference must be considered with this variability in mind. The lower margins of the fore femora of M. ewarti have a number of long, erect hairs as well as many extremely fine, short ones. The few males seen of M. pyramicus possess, in addition, about as many long hairs and an equal, or greater, number of hairs about half as long as the longer. The most conspicu- ous difference is the presence of a discoidal cell in M. ewarti males and its lack in those of M. pyramicus. Finally, M. ewarti males have a well developed fringe of hairs on the apical margin of the fore wing and apical and posterior margins of the hind wing. There is no fringe on the forewing of M. pyramicus and on the hind wing it is extremely sparse, most of the hairs separated by much more than their own lengths. Myrmecocystus creightoni, new species Figure 2 Diagnosis.— A member of the lugubris group identifiable by the presence of abundant reclinate hairs on the antennal scapes and virtual lack of erect hairs on the thoracic dorsum of the workers; female mesoscutum finely, uni- formly punctate and with marginal fringes on the wings; male without erect hairs on scapes and tibiae, wings with marginal fringe. 1971 Studies on California Ants 7 WORKER. Measurements.-m.0.1?>-\.02> (0.90) ; HW 0.60-0.93 (0.80); SL 0.80-1.13 (1.00); WL 0.96-1.50 (1.30); PW 0.40-0.63 (0.56). Sides of head straight (slightly convex in largest workers), converging slightly toward mandibular insertions, longer than broad in all sizes. Cl 78-93 (88), somewhat shorter than scape; SI 108-138 (125). Occiput broadly flat- tened in frontal view, with poorly indicated lateral corners. Eye small, about as long as first flagellar segment, EL 0.50-0.72 x OMD (0.63). Mandible with seven teeth on cutting margin, rarely with a minute intercalary denticle be- tween basal and penultimate teeth. Thorax slender to moderately robust, PW 0.36-0.50 x WL (0.41). In larger workers rear of mesonotum dropping sharply to metanotum. Basal face of propodeum as long as posterior face or slightly shorter, juncture of two faces distinctly rounded. Petiole erect, in profile about twice higher than thick, not at all cuneate, summit rounded; crest flat in frontal view, not, or barely, impressed in middle; in dorsal view about twice as wide as thick. — Pubescence scattered on cheeks, frons and occiput, nowhere concealing surface, longer than an ocellar diameter; longer and denser on thorax, especially on sides of propodeum, longer and denser on gaster, but usually not concealing surface, sparse at sides of tergites. Hairs of scapes abundant, reclinate (rarely fully erect); occipital hairs shorter than maximum thickness of hind femora; thoracic dorsum usually lacking erect hairs, but three or four inconspicuous ones may be present toward sides; petiolar crest without conspicuous erect hairs; first gastric tergite with discal erect hairs shorter than minimum thickness of hind femora; inner face of fore femora without erect hairs; outer face of middle and hind tibiae with few or no erect hairs. Integument. — VoXishQd on clypeus, cheeks and frons, duller on occiput, with scattered fine piligerous punctures on frons, clypeus and cheeks, those on cheeks often coarse and elongate. Thorax shiny, but no less so than head, densely coriaceous, especially on sides of propodeum; gaster moderately shiny, densely and finely coriaceous. Co/or. — Uniformly medium to dark brownish, legs lighter; mandibles and lateral clypeal lobes lighter, often yellowish. FEMALE. Measurements. -m. 1.40; HW 1.43-1.46; SL 1.33; EL 0.40- 0.43; OMD 0.50; WL 2.7-2.9; PW 1.8-1. 9. Head in full face view with margins straight, converging slightly toward mandibular insertions, broad. Cl 102-104; a little longer than scape; SI 90-93. Occiput, in frontal view, broad, flat, with rounded lateral angles. Eye small, barely longer than first flagellar segment; EL 0.80-0.86 x OMD. Posterior ocelli separated by about 3 x diameter of anterior ocellus, removed from eye margin by about 3.5 x diameter of anterior ocellus. Mandible with seven teeth. Thorax robust, PW 0.65-0.66 x WL. In profile, posterior two-thirds of 8 Contributions in Science No. 214 1971 Studies on California Ants 9 mesoscutum slightly convex, more flattened caudad, posterior margin below anterior margin of scutellum; scutellum, in profile, broadly convex; metanotum not protruding. Petiole, in profile, compressed, about twice as high as thick at level of spiracle, crest thin; in frontal view, deeply notched; from above about 3.5 x wider than thick. Vestiture. — CQ^hdilic pubescence yellowish, as long or longer than an ocellar diameter, much of it reclinate rather than appressed; scattered, not obscuring surface except on cheeks above mandibular insertions. Thoracic and gastric pubescence white, appressed to reclinate; long, not concealing surface, but quite dense on gaster. Scape hairs reclinate; cheeks with a few long erect hairs in frontal view; thoracic dorsum with numerous erect yellowish hairs, longest on scutellum; erect hairs on disc of first gastric tergite no longer than basal thickness of hind tibiae; petiolar scale with a few short erect hairs on crest; fore femora without erect hairs on inner face; middle and hind tibiae with abundant suberect hairs on outer face. Wings without marginal fringe, but membrane with abundant short, erect hairs. Integument.— Head shiny, with coarse setigerous punctures on clypeus and upper half of cheeks; frons and occiput with fine well-separated piligerous punctures; frontal lobes with dense, fine, punctures. Pronotum finely and densely punctate; mesopleurae moderately shiny, densely coriaceous, with fine punctures peripherally and scattered coarse, setigerous punctures; meta- pleurae duller, densely tesselate, abundantly punctate, punctures finer than those in middle of mesopleura; mesoscutum densely, finely punctate, except for sparsely punctate area along midline, punctures laterad of parapsidal lines sparser than those of disc; mesocutellar punctures equal to those of mesoscu- tum, sparser in middle; metanotum dull, densely coriaceous and micropunc- tate; propodeum slightly shiny, roughened and densely coriaceous. Gaster moderately shiny, with fine, dense piligerous punctures. Color. — Uniformly light brownish except for yellowish mandible, lateral clypeal lobes and lower half, or less, of genae. Wings hyaline, veins and stigma light brownish. MALE. Measurements. -UL 0.63-0.71; HW 0.60-0.63; SL 0.70-0.73; EL 0.23-0.25; OMD 0.13-0.15; WL 1.53-1.66; PW 0.93-1.00. In full face view, sides of head strongly convergent toward mandibular insertions, margins straight; head a little longer than broad. Cl 90-94; slightly shorter than scape; SI 1 14-115; OMD 0.56-0.60 x EL; ocelli subequal to one Figure 2. Myrmecocystus creightoni, new species. A, female, lateral view; Aa, female head, frontal view; Ab, female mesoscutum. B, worker major, lateral view; Ba, worker major, head, frontal view; Bb, worker minor, head, frontal view; Be, worker petiole, profile and posterior views. C, male, lateral view; Ca, male head, frontal view; Cb, male mandible; Cc, male ninth sternite; Cd, male volsella, inner view, Ce, male aedeagus, lateral view. 10 Contributions in Science No. 214 another; lOD 3. 5-4.0 x OD; OOD 2.5-3. 0 x OD. Mandibular margin simple, with preapical notch and two small denticulae basad, apical tooth as broad at base as long. Clypeus usually without preapical transverse depression. Petiolar scale in profile, higher than long, evenly thick from base to near summit where it is narrowly rounded to slightly angulate; crest, seen from front, angulate at sides, broadly and rather deeply emarginate; in dorsal view, about 2.25 x wider than long. Terminalia: Fig. 2 Cc-e. Vestiture.—V^hiiQ, appressed pubescence long, sparse on head and tho- racic dorsum; more abundant on thoracic pleurae, particularly on propodeum; longest on gaster, but nowhere concealing surface. Erect hairs sparse, short on head and thorax, yellowish. Middle of first tergite, in profile, with abundant very short, suberect whitish hairs; tergites with scattered long to very long yellowish hairs, especialy caudad and ventrally. Fore and hind wings fringed along apical and hind margins. Integument. — Head shiny, with scattered fine punctures. Mesoscutum shiny with sparse fine punctures; propodeum, meso- and metapleurae duller, densely coriaceous, with scattered fine punctures; scutellum with a few very fine punctures. Gaster shiny, finely piligerously punctate. Co/or.— Uniformly light to very dark brownish, appendages and mandi- bles lighter. Wings hyaline, veins and stigma light brownish. Type material.— HoXoiypo, worker, allotype male, 18 $$,35^^, 305 ? paratypes. Pearblossom, 2 mi S, 3500 ft, Los Angeles Co., Calif., 12 February 1967 (R. R. Snelling). Additional paratypes: 19,322 ? ?, same locality, 20 March 1966 (R.R. Snelling). Holotype, allotype and most paratypes inLACM; nine paratypes in each of the following: AMNH, MCZ, MNHG, USNM and private collections of A. C. Cole, Jr., W. S. Creighton, and G. C. Wheeler. Etymology. — It is my pleasure to dedicate this species to Professor W. S. Creighton for his outstanding contributions to the understanding of United States ants, with my thanks for his generosity and enthusiastic encouragement during this study. Habitat.— ThQ type series was taken in coarse, sandy soil above a dry wash on the edge of the Mojave Desert, in a mixed creosote bush-juniper- Joshua tree ecotone. Additional samples are from Kern and Riverside Counties, in areas of mixed creosote bush and juniper. The known elevational amplitude is from 3000-4300 feet. Discussion. — In Creighton’s key (1950) this ant will key to M. yuma Wheeler. Creighton (1956) synomymized M. yuma under M. lugubris, but I believe that he was in error and recognize M. yuma as a valid species. The absence of conspicuous erect hairs on the antennal scapes and propodeum will readily separate workers of M. creightoni from M. yuma and M. hammettensis 1971 Studies on California Ants 11 Cole. Conspicuous erect hairs are present on the pronotum and mesonotum of M. lugubris but lacking in M. cr eight oni. The female of M. cr eight oni has the mesoscutum finely and rather uni- formly closely punctate except for a small area of sparser punctation medially; those of M. yuma and M. hammettensis have the mesoscutum almost im- punctate between the parapsides with scattered coarse, setigerous punctures. The female of M. lugubris is unknown, but by analogy with other, underscribed species, should have a mesoscutum with mixed coarse and fine punctures and a broad, median impunctate area. The males of M. lugubris and M. hammettensis lack fringe hairs on the wings, present in M. creightoni. The male of M. yuma is unknown, and I do not care to speculate on its characteristics, since those of its nearest relatives also are unknown. Myrmecocystus wheeleri, new species Figure 3 Myrmecocystus melliger semirufus var. testaceus, Wheeler, 1908. Bull. Amer. Mus. Nat. Hist. 24:355-356. (in part, misidentification ) Diagnosis.— Not closely related to any known species but most closely resembles M. kennedyi Cole (= M. semirufus of authors, not Emery, 1893). Workers are readily separated from all other species by the concolorous orange ferruginous head, thorax and gaster. From variant populations of M. kennedyi with partially concolorous gaster, M. wheeleri may be separated by the presence of abundant, close appressed white hairs on the third tergite. Males are most easily recognized by the shape of the aedeagus and unusually large teeth of that structure (Fig. 3 Ce). WORKER. Measurements.-liLQ.92>-\.26 ( 1.26) ; HW 0.83-1.10 (1.10); SL 1.16-1.60 (1.60); WL 1. 6-2.0 (2.0); PW 0.60-0.83 (0.83). Head a little longer than broad, Cl = 81-96 (86), distinctly shorter than scape; SI = 130-153 ( 145) . In frontal view, head broadest at, or a little below, the eyes, sides straight or slightly, evenly convex, narrowed slightly toward mandibular insertions. Occiput in frontal view broadly rounded laterally, summit slightly convex or flattened. Eye small, barely longer than first flagellar segment; removed from mandibular insertion by 1.5-2. 1 (2.1) times its own length. Mandibles usually with seven, rarely eight, teeth. Thorax slender, PW 0.36-0.43 (0.41) x WL. Basal face of propodeum broadly rounded into posterior face. Petiole, in profile, about as thick as high, narrowed toward evenly rounded apex; crest evenly convex in frontal view; in dorsal view, scale 1.4-1. 5 x wider than long. Vestiture. — ¥.voct hairs sparse on head, confined mainly to clypeus, frontal area and occiput, those of occiput distinctly longer than eye length, of frons and clypeus variable, but mostly shorter than eye length; cheeks with scattered 12 Contributions in Science No. 214 erect hairs less than half as long as eye length. Pronotum with 12-18 erect hairs of irregular length, all shorter than eye length, a variable number of much shorter hairs on neck; mesonotum with about a dozen erect hairs, less than half eye length long; metanotum usually without erect hairs; propodeum with 12 or more erect hairs of variable length, the longest equal to half, or a little more, maximum eye length. Petiolar scale with 6-10 fine, erect short hairs on crest. Discs of gastric segments with sparse erect hairs, about equal to half of maximum eye length, hairs of tergal margins only slightly shorter than eye length. Scapes with abundant fine, short suberect hairs on inner and lower faces. Inner face of fore femora without erect hairs except along lower margin, these about as long as those of outer face. Middle and hind tibiae with abundant fine, reclinate hairs on all surfaces, these a little shorter than minimum thickness of the tibiae. Appressed pubescence sparse, short, on head, denser on occiput and vertex; distinctly longer and denser on thorax, coxae and femora, petiole and first three gastric tergites. Pubescence very sparse or absent from fourth and fifth tergites. Color. — Orange ferruginous, often with lower half of face more yellowish; fourth and fifth gastric segments often infuscate. Rarely most of alitrunk, gaster and legs infuscate in some minors. FEMALE. Measurernents.-m. 1.8; HW 1.8; SL 1.8; EL 0.4; OMD 0.7; WL 4.0; PW 2.5. Head as broad as long. Cl = 100, as long as scape; SI = 100. In frontal view, head parallel-sided, as broad at mandibular insertions as at lower eye level. Occiput rounded laterally, without evident corners, slightly convex in middle. Eye small, 1.3 times length of first funicular segment, removed from mandibular insertion by 1.6 times its length. Lateral ocelli separated by 2.5 times diameter of anterior ocellus, separated from eye margin by 3.5 times diameter of anterior ocellus. Penultimate segment of maxillary palpi broadest at basal third, strongly narrowed toward apex. Thorax robust, PW 0.62 x WL. In profile, posterior half of mesoscutum flat, apical margin below level of convex scutellum; scutellum and metanotum forming continuous convex surface. Petiole in profile compressed-cuneate, crest sharp; distinctly notched; from above, about three times wider than long. V estiture. — ^VQCi hairs present on all parts of face, least abundant on cheeks immediately below eyes and between eyes and ocelli; occipital hairs irregular in length, longest about as long as eye length, hairs on frons and clypeus equally variable, but a little shorter. Hairs abundant on thoracic dorsum and sides highly variable in length, some longer than eye length (especially on sides); basal third of propodeum with abundant long hair, apical two-thirds with very short erect hairs, especially toward apex and around gland opening. Crest and sides of petiolar scale with numerous long erect hairs, gastric tergites with abundant fully erect hairs on disc, separated 1971 Studies on California Ants 13 by less than their own lengths, mostly about half as long as eye length. Fore femora without conspicuous erect hair on inner face. All tibiae with abundant suberect hairs which are about as long as minimum thickness of hind tibia. Scapes with abundant suberect, short hairs on outer and lower faces. Forewing without marginal fringe, hind wing fringed on posterior margin. Pubescence long, notably dense only on first three gastric tergites, variably fully appressed to subappressed, especially on head. lntegument.~}^Qdid moderately shiny, surface micro-reticulate; frontal lobes with close, fine punctures of variable size, round to ovoid, separated by about a puncture diameter; genae more sparsely, coarsely punctate. Center of mesoscutum impunctate, median area laterally with scattered fine punctures, becoming closer and more distinct in lateral thirds and apically; anteriorly, median portion with sparse micropunctures; punctures denser and coarser laterad of parapsides. Punctures of scutellum finer than of adjacent portion of scutum, sparse in middle, denser laterad; mesopleura above minutely roughened between coarse, close punctures, below equally coarsely, more closely punctate; metapleura and propodeum similar to lower half of mesopleura. Color. — Orange ferruginous, the following brownish : rectangular median mark on anterior half of mesoscutum, broad lateral stripes on posterior four- fifths of mesoscutum, irregular blotch on mesopleura above and irregular blotch on mesepisternum. Apical gastric segments lightly infuscated. Wings whitish hyaline, radial vein and stigma brownish, remainder of veins yellowish. MALE. Measurements. -m. 0.86-0.90; HW 0.83-0.90; SL 1.03-1.10; EL 0.30-0.33; WL 2.0-2.2; PW 1.16-1.23. Margins of head distinctly convergent toward mandibular insertions; head as broad as long or slightly longer (Cl 96-100), distinctly shorter than scape; SI 119-128. OMD 0.80-0.90 x EL. Anterior ocellus % diameter of lateral ocelli; lOD 2.5-3. 0 x OD; OOD 2.5-3. 0 x OD. Mandible without basal teeth. Clypeus with short transverse depression below middle. Petiole, in profile, higher than long, narrowed above, crest convex; in frontal view, evenly convex from side to side, except for vague to prominent median notch; in dorsal view about twice as wide as long. Terminalia: Fig. 3 Cc-e. Vestiture. — ^XQoX hairs abundant on body, those of scutellum as long as eye diameter, length elsewhere generally shorter but variable. Erect hairs of hind tibiae about as long as thickness of scape. Pubescence sparse on head and thorax, abundant on propodeum and first four tergites. Forewing without marginal fringe, hind wing with fringe on posterior margin only. Integument. — ModQV3XQ\y shiny, closely coriaceous, with scattered fine piligerous punctures. Co/or. — Uniformly blackish, appendages light brownish. Type material. — TioXoiypQ worker, allotype male, 193 worker and two 14 Contributions in Science No. 214 1971 Studies on California Ants 15 female paratypes, 6 mi SE of Pearblossom, 3500 ft, Los Angeles Co., 15 August 1965 (R. R. Snelling). Holotype, allotype and most paratypes in LACM; six paratypes in each of the following: AMNH, MNHG, USNM and the collections of A. C. Cole, W. S. Creighton, and G, C. Wheeler. Etymology.— This ant is dedicated to the late William Morton Wheeler, whose career included the first comprehensive study of these remarkable ants. Habitat.— This species occurs from the central San Joaquin Valley (Merced County) southward to San Diego County, in habitats ranging from oak grasslands to juniper-pinon. The workers forage diurnally and frequent blossoms for nectar in addition to being assiduous scavenger-predators of other small insects. Repletes are unknown. Discussion.— This species was erroneously identified by Wheeler (1908, 1912) as M. melliger semirufus var. testaceus Emery. Emery’s species is un- related and belongs to the mexicanus group, where it is a senior synonym of M. mexicanus mojave Wheeler (Snelling, 1969). The concolorous orange- ferruginous body of the worker caste is usually sufficient to separate M. wheeleri from all other species. Some samples of M. kennedyi Cole (=M. melliger semirufus of authors, not of Emery; see Snelling, 1969) may have the first two gastric segments orange-ferruginous. These may be separated from M. wheeleri by the lack of dense appressed hairs on the third tergite. Minor workers of some populations of M. wheeleri from Mojave desert areas of Los Angeles County have the entire gaster, legs and much of the thorax extensively infuscated. The media and large workers are normally colored. These minors may be separated from those of M. kennedyi and M. flaviceps Wheeler by the combination of densely pubescent third tergite and long, flexuous hairs on the pronotum. The female is best recognized by the largely orange-ferruginous color and the shape of the penultimate segment of the maxillary palpi. The shape of the male aedeagus is unique and is the best recognition character for this caste. Figure 3. Myrmecocystus wheeleri, new species. A, female, lateral view; Aa, female head, frontal view; Ab, female mesoscutum. B, worker major, lateral view; Ba, worker major, head, frontal view; Bb, worker minor, head, frontal view; Be, worker petiole, profile and posterior views; Bd, worker, portion of third tergite to show distribution of erect and appressed vestiture. C, male, lateral view; Ca, male head, frontal view; Cb, male mandible; Cc, male ninth sternite; Cd, male volsella, inner view; Ce, male aedeagus, lateral view. 16 Contributions in Science No. 214 Literature Cited Cole, A. C., Jr. 1968. Pogonomyrmex harvester ants. A study of the genus in North America. Univ. Tenn. Press, x + 222 p. Creighton, W. S. 1950. The ants of North America. Mus. Comp. Zool., Bull. 104:1-585 1956. Notes on Myrmecocystus lugubris Wheeler and its synonym, Myrme- cocystus yuma Wheeler. Amer. Mus. Novitates. 1807:1-4. Snelling, R. R. 1969. Taxonomic notes on the Myrmecocystus melliger complex. Los Angeles County Museum, Contrib. Sci. 170: 1-9 Wheeler, W. M. 1908. Honey ants, with a revision of the American Myremecocysti. Amer. Mus Nat. Hist., Bull. 24:345-397. 1912. Additions to our knowledge of the ants of the genus Myrmecocystus Wes- mael. Psyche 19:172-181. Wilson, E. O. 1955. A monographic revision of the ant genus Lasius. Mus. Comp. Zool., Bull. 113:1-205 Accepted for publication November 10, 1970 J "4 4 ■ i- i t "'■1 Printed in Los Angeles, California by Continental Graphics LOS j ANGELES COUNTY MUSEUM [umber 215 CONTRIBUTIONS IN SCIENCE June 9, 1971 STATUS OF CERTAIN FROGS OF THE GENUS COLOSTETHUS, WITH DESCRIPTIONS OF NEW SPECIES By Philip A. Silverstone ■iUn* I 5 ’■9/1 Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 CONTRIBUTION S page size. Consult the 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 STATUS OF CERTAIN FROGS OF THE GENUS COLOSTETHUS, WITH DESCRIPTIONS OF NEW SPECIES By Philip A. Silverstone^ Abstract: Three new species of dendrobatid frogs are described from Antioquia, Colombia: Colostethus ramosi, C. lehmanni, and C. fraterdanieli. Lectotypes are designated for C. pratti (Boulenger), 1899 and C. kingsburyi (Boulenger), 1918. The holotype of Dendrobates ranoides Boulenger, 1918 is a Colostethus, specifically distinct from C. brunneus (Cope), 1887. The dendrobatid genus Colostethus is one of the largest and poorly known groups of frogs in the Neotropics. During the course of my work with frogs of the genus Dendrobates, I had the opportunity to study frog populations referable to Colostethus in Colombia and to examine extant types for most of the nominal species. Some of the results of these observations are presented here in an attempt to clarify the status of some populations and names. Three populations encountered in Colombia represent currently un- recognized species, and I propose they be named as follows: Colostethus ramosi, new species Figure 1 Holotype— L ACM 44147, Colombia: Antioquia: Alto de la Honda (hill near San Rafael), 1240 m; collected 8 June 1968 by P. A. Silverstone, J. E. Ramos, and C. Jimenez. Topoparatype.—L ACM 44148. Diagnosis— Colostethus ramosi differs from other species of the genus in the following combination of characters: complete light lateral diagonal stripe from groin to eye; venter lacking spotting or marbling; third finger of male not swollen; no light vertebral stripe; no toe webbing; throat of male dark; and no dark longitudinal line on anterior surface of thigh. Description of holotype —Adult female, 20 mm snout-vent length; skin smooth, except slightly granular on posterior belly; first finger equal to second; tarsal fold present; tarsal tubercle present; digits lack webbing and fringes; maxillary and premaxillary teeth present, no vomerine teeth; posterodorsal portion of tympanum concealed; diameter of eye 1.7 times diameter of tym- panum; diameter of eye greater than distance from eye to nostril; snout sub- ^Research Associate in Herpetology, Los Angeles County Museum of Natural History. 1 2 Frogs of the Genus COLOSTETHUS No. 215 truncate in dorsal aspect, rounded in lateral aspect; canthus rostralis rounded; interorbital space 1.7 times diameter of upper eyelid. Color in life: iris black with gold ring around pupil; head and back dull brown; no dorsolateral stripes; sides of head and body black, crossed by complete gold diagonal lateral stripe from groin to eye, along canthus rostralis, and around snout tip; white ventro- lateral stripe (disappears in preservative) from hind limb insertion along upper lip to snout tip; dorsal surface of hind limbs dull orange, of forelimbs gold; light area subtended by brown mottling on posterior surface of thigh; longitudinal dark brown stripe on anterior surface of upper arm, posterior Figure 1. Colostethus ramosi, holotype (LACM 44147). surface of forelimb, and dorsal surface of thigh, but not on anterior surface of thigh; ventral surface of limbs dull orange; throat, chest, and belly yellow- ish white, latter with metallic gold tinge. Paratype—AdvXt male, 18.5 mm snout-vent length; third finger not swol- len; similar to female, except (in life) forelimb brown above and gray below, throat and chest dark gray, and ventrolateral stripe broken into white spots posterior to forelimb insertion. Habits and habitat were heard calling on fallen leaves on ground in second growth forest and in tall grass on slope near forest, not near water. Call consisted of one note, repeated one minute or more, averaging 86 notes per minute, and 1.4 notes per second. Specimens were taken between 0800 and 1030 hrs. Mean annual temperature at San Rafael reported as 23° C 1971 Contributions in Science 3 (Colombian Dept, of Statistics), and thus is drier and warmer than Santa Rita (collecting site for following two species). Type locality— Sdin Rafael is 16 km east of Guatape, on Guatape-Rio Magdalena highway. Etymo/ogy.— Named after Jorge Eduardo Ramos P. of Cali, Colombia. Colostethus lehmanni, new species Figure 2 Holotype— L ACM 44156, Colombia: Antioquia: Santa Rita (near Rio Nare), 1890-1910 m; collected 9 June 1968 by P. A. Silverstone and J. E. Ramos. Topoparatypes.—h ACM 44154-55, 44157-61. Diagnosis.— Colostethus lehmanni differs from other species of the genus by the following combination of characters: complete light lateral diagonal stripe from groin to eye; venter lacking spotting or marbling; third finger of male not swollen; no light vertebral stripe; no toe webbing; throat of male often bearing a vague dull wash of gray, but not being conspicuously dark, and thus not contrasting with a light venter as in ramose, dark longitudinal line on anterior surface of thigh present. Description of holotype.— Adult male, 18 mm snout-vent length; third finger, not swollen; skin smooth except few tubercles on posterior back and dorsal surface of hind limbs; first finger equal to second; tarsal fold present; tarsal tubercle present; digits lack webbing and fringes; maxillary and pre- maxillary teeth present, no vomerine teeth; posterodorsal portion of tym- panum concealed; diameter of eye twice diameter of tympanum; diameter of eye greater than distance from eye to nostril; snout sub truncate in dorsal aspect, rounded in lateral aspect; canthus rostralis rounded; interorbital space 1.7 times diameter of upper eyelid. Color in life: iris black; head and back brown; no dorsolateral stripes; upper sides of head and body black, crossed by complete gold diagonal lateral stripe from groin to eye, along canthus rostralis, and around snout tip; lower sides of body brown with white spots; light upper lip stripe; dorsal surface of limbs brown, banded darker brown; dark brown longitudinal stripe on anterior surface of thigh and upper arm; light longitudinal stripe on posterior surface of thigh, subtended by brown mottling; ventral surface grayish, except belly yellowish; ventrodistal upper arm and ventroproximal forearm dark, former swollen. Paraty pes. -Snout-\ent length in mm (average in parentheses): five adult males: 15.5-18 (16.7); two juveniles: 9.5-14.5; LACM 44154-55, 44157-58 similar to holotype, but 44154 paler, 44155 lacks white spots on lower sides, 44158 lacks dark swollen areas on ventral surface of forelimb, and 44157 has dark markings on back; some individuals lack posterior thigh stripe; some have slightly granular belly; LACM 44159-61 have lighter ground color than holotype, shorter snout, dark back markings, second, lower light 4 Frogs of the Genus COLOSTETHUS No. 215 Figure 2. Colostethus lehmanni: left, paratype (LACM 44160); right, holotype (LACM 44156). lateral stripe, dark mottling on upper lip, and lack dark swollen areas on ventral surface of forelimb; LACM 44157 is transitional, with dark ground color as in holotype, but also dark spots on upper lip and dark median back stripe ending anteriorly in Y-mark, as in LACM 44159, which has light ground color. Habits and Specimens were taken on moist ground between 0600 and 1330 hrs in disturbed, primary forest. The habitat contained abun- dant mosses, palms, and bromeliads. Mean annual rainfall 5000 mm, mean annual temperature 20°, maximum 30°, minimum 10-12° C (Empresas Municipals, Medellin). Type locality Rita is a dam-site on Rio Nare, on road north from San Rafael-Guatape highway, near fauatape. Fry mo/ogy.— Named after F. Carlos Lehmann V. of Cali, Colombia. Colostethus fraterdanieli, new species Figure 3 Holotype— E ACM 44164, Colombia: Antioquia: Santa Rita (near Rio Nare), 1890-1910 m; collected 9 June 1968 by P. A. Silverstone and J. E. Ramos. Paratypes— AW from Colombia: LACM 44165-69, topoparatypes; AMNH 39179, 39187, 39189, 39195, 39198, and five untagged, Antioquia: 1971 Contributions in Science 5 Medellin; USNM 150715-17; "'Caldas: Montanita” (east of Rio Cauca, be- tween Salamina and San Felix, 2300-2400 m, fide Rvdo. Hno. Niceforo Maria); USNM 150718, “Caldas: 9 km east of Santa Rosa de Cabal” (Santa Rosa de Cabal is southwest of Manizales, on Manizales-Pereira road, eleva- tion at collecting site is 2000-2500 m, fide Rvdo. Hno. Niceforo Maria). Diagnosis.— Differs from C. nubicola in larger size and basal toe webbing (none in C. nubicola) ; differs from C. infraguttatus in basal toe webbing (none in C. infraguttatus) and ventral pattern dark spots on light ground (light spots on dark ground in C. infraguttatus)', differs from C. subpunctatus in having swollen third finger in males; differs from C. kingsburyi in having basal toe webbing and complete light lateral stripe, and lacking light ventrolateral stripe (C. kingsburyi lacks toe webbing and has dorsolateral and ventrolateral stripes); differs from other Colostethus in having complete light lateral stripe, basal toe webbing, and males with swollen third finger and dark ventral spotting. Description of holotype.—Adu\i male, 23 mm snout-vent length; head relatively broad; third finger swollen; skin smooth except few tubercles on Figure 3. Colostethus fraterdanieli, holotype (LACM 44164). posterior back; first finger slightly shorter than second; tarsal fold present; tarsal tubercle present; fingers lack webbing and fringes, toes basally webbed but little or no fringing, webbing formula (Savage and Heyer, 1967) I 2-3 6 Frogs of the Genus COLOSTETHUS No. 215 II 2-3.5 III 3-4 IV 4.3-3 V; maxillary and premaxillary teeth present, no vom- erine teeth; posterodorsal portion of tympanum concealed; diameter of eye twice diameter of tympanum; diameter of eye greater than distance from eye to nostril; snout rounded in dorsal and lateral aspects; canthus rostralis rounded; interorbital space twice diameter of upper eyelid. Color in life: iris black with gold ring around pupil; head and back light brown; no dorsolateral stripes; sides of head and body dark brown, olive posterolaterally; interorbital area and partial median back stripe darker brown than rest of dorsum; com- plete diagonal lateral gold stripe from groin to eye, along canthus rostralis, and around snout tip; groin, axilla, and posterior tip of lateral stripe yellow; upper lip stripe and diagonal stripe from eye to forelimb insertion gold; dorsal surface of forelimbs orange, of hind limbs yellow-orange with brownish cast; posterior surface of thigh has narrow, wavy yellow-orange stripe subtended by mottling of yellow-orange and dark brown; dark brown longitudinal stripe on anterior and posterior surfaces of forelimb and anterior and posterodorsal surfaces of thigh; dorsal surface of tibia banded dark brown; ventral surface of limbs yellow-orange with brownish cast; throat, chest, and belly yellowish white. Paraty pes. -Snout-Yent lengths in mm (average in parentheses) : seven adult males: 20.5-24 (22.3); eight adult females: 24-27 (25.8); 15 adults: 20.5-27 (24.2) ; five juveniles: 16-22; third finger of males swollen, of females not swollen; prominent tubercles on posterior back except in LACM 44166-69; female USNM specimens have no ventral spotting, males have small dark spots on throat and chest; AMNH females vary from heavy ventral spotting (39187) to few dull spots, males have dark spotting on chest and anterior belly and marbling on throat; LACM specimens have dull spotting on throat and chest only, darker in males than in females, but not as dark as in USNM and AMNH specimens; posterior thigh pattern in AMNH specimens similar to that in holotype, but light stripe sometimes broken into row of spots; in USNM specimens, posterior light thigh stripe conspicuous against dark brown back- ground; all specimens have dark brown interorbital area, contrasting with lighter brown back color; AMNH specimens have small dark spots on back. Etymology.— Nstmed after Reverendo Hermann Daniel of the Christian Brothers. The type series of the following two taxa contain more than one species. In order to facilitate future work, I propose here to make the taxa objective by designating lectotypes. Colostethus pratti (Boulenger) Phyllobates pratti Boulenger, 1899: 274-275, pi. 11, Fig. 3. Type material.— Tyfo syntypes, BM 1947.2.13.94-95, includes two species. I designate BM 1947.2.13.94 (figured by Boulenger), 20 mm female, Colom- bia: Santa Ines, north of Medellin, 3800 ft ( 1159 m), collected by A. E. Pratt, 1971 Contributions in Science 7 as lectotype of Phyllobates pratti Boulenger. It has incomplete light lateral stripe. BM 1947.2.13.95 differs from lectotype in having complete light lateral stripe (i.e., stripe reaches eye anteriorly). Colostethus kingsbunji (Boulenger) Phyllobates kingsburyi Boulenger, 1918: 427-428. Type material— Pour syntypes, BM 1947.2.14.3-6, includes two species; BM 1947.2.14.3 is not conspecific with other three and does not agree with Boulenger’s original description. It has basal toe webbing, complete light lateral diagonal stripe, and is fat-bodied; the other three specimens lack toe webbing, have ventrolateral stripes, and are relatively narrow-bodied. I desig- nate BM 1947.2.14.5, 25 mm female, Ecuador: El Topo, Rio Pastaza, 4200 ft (1281 m), collected 1912 by M. G. Palmer, as lectotype of Phyllobates kings- buryi Boulenger. After examining the type, I propose the following new combination : Colostethus ranoides (Boulenger) Dendrobates ranoides 1918: 428-429. Type material— I agree with Dunn ( 1957) that the holotype, BM 1947.2.- 15.35, Colombia: Villavicencio, Guatiquia River, 400 ft (122 m) has teeth and is not Dendrobates. It is Colostethus, but not C. brunneus (Cope), 1887, as currently considered. The type material of C. brunneus (ANSP 11241- 61) is in such poor condition that it is useless for comparison. Topotypes of C. brunneus (KU 93154-55) have immaculate venter and dorsolateral stripe. The holotype of C. ranoides differs from these topotypes in having spotted venter and complete light lateral diagonal stripe. Acknowledgements I thank J. M. Savage and I. R. Straughan, University of Southern Cali- fornia and J. W. Wright, Los Angeles County Museum of Natural History (LACM) for criticizing the manuscript. I thank the following for permission to examine specimens in their charge: A. G. C. Grandison and staff, British Museum (Natural History, BM); J. W. Wright and A. H. Brame, Jr. (LACM); J. A. Peters and staff. United States National Museum (USNM); R. G. Zweifel and C. W. Myers, American Museum of Natural History (AMNH); W. E. Duellman, University of Kansas Museum of Natural His- tory (KU) ; and J. E. Bohlke and E. V. Malnate, Academy of Natural Sciences of Philadelphia (ANSP). Rvdo. Hno. Niceforo Maria, Instituto de La Salle, Bogota provided information on localities. J. E. Ramos P., H. Trapido, R. Diaz V., Universidad del Valle, Cali, F. Carlos Lehmann V., Museo Departamental de Historia Natural, Cali, and Rvdo. Hno. Daniel and col- leagues, Colegio San Jose, Medellin facilitated my work in Colombia in 1968. Studies were partially aided by a National Defense Education Act Title IV s Frogs of the Genus COLOSTETHUS No. 215 Fellowship and by Biomedical Sciences Support Grant FR-070 12-02 from the National Institute of Health. Resumen Se describen tres nuevas especies de ranas (Dendrobatidae) procedentes de Antioquia. Colombia. Colostethus ramosi, C. lehmanni, y C. fraterdanieli. Tanto los sintipos de C. pratti (Boulenger) como los de C. kingsburyi (Bou- lenger) representan dos especies; por consiguiente se designa un lectotipo de cada serie. De acuerdo con Dunn (1957), el holotipo de Dendrobates ranoides Boulenger es un Colostethus con dientes; este nombre no es un sinonimo de C. brunneus (Cope) como ha sido considerado, sino que repre- senta una especie distinta. Literature Cited Boulenger, G. A. 1899. Descriptions of new batrachians in the collection of the British Museum Natural History). Ann. Mag. Nat. Hist., Ser. 7, 3: 273-277. 1918. Descriptions of new South- American batrachians. Ann. Mag. Nat. Hist., Ser. 9, 2: 427-433. Cope, E. D. 1887. Synopsis of the Batrachia and Reptilia obtained by H. H. Smith, in the province of Mata Grosso, Brazil. Proc. Am. Phil. Soc. 24: 44-60. Dunn, E. R. 1957. Neotropical frog genera: Prostherapis vs. Hyloxalus, with re- marks on Phyllobates. Copeia 1957: 77-78. Savage, J. M., and W. R. Heyer. 1967. Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central America. Beitr. Neotrop. Fauna 5: 111-131. Accepted for publication July 9, 1970 L I LOS !, ANGELES I COUNTY i MUSEUM ij jMBER 216 CONTRIBUTIONS IN SCIENCE July 8, 1971 PHYLOGENETIC RELATIONSHIPS OF CERTAIN NEOTROPICAL |)ADS WITH THE DESCRIPTION OF A NEW GENUS (ANURA: BUFONIDAE) I I' By Linda Trueb ! jl i I: ii i Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 PHYLOGENETIC RELATIONSHIPS OF CERTAIN NEOTROPICAL TOADS WITH THE DESCRIPTION OF A NEW GENUS (ANURA: BUFONIDAE) By Linda Trueb' Abstract: A new bufonid genus, Rhamphophryne, is de- scribed. This genus contains five species (Atelopus festae Peracca, Bufo rostratus Noble, Bufo nicefori Cochran and Goin, and two new species, R. acrolopha and R. macrorhina) , and ranges from extreme eastern Panama to southern Ecuador. The genus is characterized by a reduced number of presacral vertebrae, a pro- tuberant snout, highly modified hands and feet, and few, large, unpigmented eggs. In addition to these characters, the condition of the pectoral girdle and the cranial osteology seem to relate Rhamphophryne most closely to Oreophrynella and Atelopus in the new World. Morphological and zoogeographic evidence suggests that Rhamphophryne may have been derived from an ancestral Bufo valliceps-Yikt stock. Rhamphophryne shares many specialized characters with Old World bufonids such as Necto- phryne, Pelophryne, Mertensophryne, and Laurentophryne. It is suggested that these resemblances are the result of convergence rather than close phylogenetic relationship. Among anurans, the bufonids have undergone an almost unparalleled, nearly cosmopolitan radiation; they are represented on all continents except Australia and Antarctica. As Tihen (1962) has pointed out, the family itself is difficult to define, and parallel differentiation within the family has consistently frustrated attempts to establish meaningful taxonomic subdivisions limited to either the New World or Old World. Within the New World, Tihen (1962) has proposed a systematic arrangement of five species groups of Bufo based princi- pally on osteological criteria. This study is a valuable contribution to an understanding of the more generalized, widespread New World representatives of Bufo. There is, however, a significant number of more specialized bufonids (including Atelopodidae auctorum) of less widespread geographical distribu- tions. Of particular interest are those restricted to montane environments. Some of these taxa are presently included in the genus Bufo (rostratus, periglenes, fastidiosus, holdridgei, simus, coerulescens); their inclusion within this genus probably represents a taxonomic convenience rather than a significant phylo- genetic placement. In contrast, other montane taxa, more recently investigated or having more obvious morphological specializations (e.g. Oreophrynella, Crepidophryne) , have been relegated to separate genera. The relationships of these taxa to one another and to other bufonids of the New World and Old ^Research Associate, University of Kansas Museum of Natural History, Lawrence, Kansas 66044. 1 2 Contributions in Science No. 216 Figure 1. Dorsal view of paratype of Bufo rostratus (=Rhamphophryne macror- hina), $ (AMNH 1384). X 2. World have been largely ignored. The present paper presents the descriptions of a new genus and two new species, and offers comments on the relationships and zoogeography of some of the Neotropical, montane bufonids. Noble (1920) described a new species of small toad {Bufo rostratus) on the basis of three specimens from the southern part of the Departamento de Antioquia, Colombia. The type series consists of the holotype, an adult male, and two paratypes, an adult male and a juvenile of undetermined sex. Noble distinguished B. rostratus from other species of Bufo on the bases of the fol- lowing characters: (1) snout produced into a three-edged rostrum; (2) paro- 1971 Relationships of Neotropical Toads 3 Figure 2. Dorsal view of holotype of Bufo rostratus (=Rhamphophryne rostrata), $ (AMNH 1359). X 1.5. toid glands triangular, not longer than greatest diameter of the eye; (3) tympanum hidden; (4) fingers one-third, and toes one-fourth webbed; (5) tar- sal fold absent; (6) prominent series of large tubercles along each side of back. More recently, Cochran and Coin (1970) redescribed Bufo rostratus and named a new subspecies. Bufo rostratus nicefori, based on five specimens, also from the Departamento de Antioquia, Colombia. My re-examination of Noble’s type series revealed that the adult male paratype (Fig. 1) of Bufo rostratus is clearly a different species than the holotype (Fig. 2) and juvenile paratype. Comparison of these specimens with the redescription of Bufo rostratus rostratus provided by Cochran and Coin (1970) indicates that they based their redescription on the adult paratype, rather than on the holotype of Bufo rostratus. There is no doubt that both species represented in the type series of Bufo rostratus are closely related and that Cochran and Coin’s Bufo rostratus nicefori is closely allied to them. A series of small toads, recently acquired from eastern Panama, and A telopus festae, known from southern Ecuador, represent 4 Contributions in Science No. 216 two other specifically distinct forms which show striking similarities to the Bufo rostratus group from Colombia. Many characteristics shared by these five taxa are unique among the cur- rently recognized bufonids. Adults are all relatively small, ranging from 32 to 51 mm in snout-vent lengths. There are seven or eight presacral vertebrae included in the vertebral column, and the sacrum is either fused to the coccyx Figure 3. Lateral views of heads of Rhamphophryne: (a) R. acrolopha, $ (KU 76965); (b) R. festae, $ (CAS 11417); (c) R. macrorhina, $ (LACM 44394); (d) R. nicefori, 9 (USNM 163476); (e) R. rostrata, $ (AMNH 1359). X 1.6. 1971 Relationships of Neotropical Toads 5 or articulated with it by a very weak bicondylar or a monocondylar articulation. The adductor longus muscle is absent, as are the annulus tympanicus and columella. All species except one (for which data are not available) have large, unpigmented eggs, which are few in number. Members of this group have heavily ossified, triangular heads which are produced into distinctive, anteriorly protruding snouts. The foregoing combination of characters indicates a close Figure 4. Dorsal views of heads of Rhamphophryne: (a) R. acrolopha, $ (KU 76965); (b) R. festae, $ (CAS 11417); (c) R. macrorhina, $ (LACM 44394); (d) R. nicefori, $ (USNM 163476); (e) R. rostrata, $ (AMNH 1359. X 1.6. 6 Contributions in Science No. 216 relationship among the four taxa mentioned and clearly distinguishes them as a group from the genus Bufo, which is characterized by eight presacral verte- brae, a bicondylar sacral-coccygeal articulation and many, small, pigmented eggs. On the bases of these distinctions and with reference to the peculiar cranial condition shared by these five taxa, they are here referred to a new genus. Rhamphophryne, new genus Type species— Rhamphophryne acrolopha, new species. Diagnosis. —These small bufonids can be distinguished from other mem- bers of the family by the following combination of characters: (1) seven or eight independent presacral vertebrae; (2) sacrum fused to coccyx or sacrum having a monocondylar or very weakly bicondylar articulation with coccyx; (3) cervical and second presacral vertebrae not fused; (4) omosternum absent; (5) protruding snout produced by extended ossification of anterior part of sphenethmoid and nasal cartilages; (6) presence of partial integumentary- cranial co-ossification; (7) tips of phalanges not expanded; (8) parotoid gland present; (9) adductor longus muscle absent; ( 10) eggs large, unpigmented, few in number; (11) annulus tympanicus and columella absent; and ( 12) arciferal pectoral girdle. Content.— Five species, as follows : Rhamphophryne acrolopha, new species, R. macrorhina, new species, Bufo rostratus Noble, Bufo rostratus nice fori Cochran and Coin, and Atelopus festae Peracca. Description. — General morphology. The snout in profile (Fig. 3) pro- trudes beyond the upper jaw and in dorsal aspect is acuminate (Fig. 4). Loreal region is concave, and nostril is lateral, protuberant and located ventral to the canthal ridge. Pupil is horizontally elliptical. The tongue is longer than wide and free posteriorly. Choana is small, ovoid and lateral in position. Prevomerine teeth are absent. Vocal slits are variable in occurrence. All members of this genus have well defined canthal ridges and supraorbital crests. Postorbital, supratympanic, pretympanic and occipital crests vary in appearance and degree of development. The forelimbs and hind limbs vary from slender to robust; all lack folds. Axillary membranes are absent. Although the hands and feet have the common phalangeal formulae of 2-2-3-3 and 2-2-3-4-3, respectively, they are highly variable with regard to other characters (Figs. 5 and 6). The digits are short and robust in nice fori and elongate and slender in rostrata and acrolopha. Fingers and toes are nearly fully webbed in some species and vestigially webbed in others. Supernumerary tubercles are present in all species, but variable in their abundance. Subarticular tubercles, large and well developed in three species, are replaced by lamellar, padlike swellings in macrorhina and nice fori. Palmar and metatarsal tubercles are characteristic of all species but variable in their occurrence and appearance in nice fori. 1971 Relationships of Neotropical Toads 7 The cloacal openings are directed posteriorly or posteroventrally at the mid- or upper levels of the thighs. Only one species, nicefori, has an cloacal sheath. Parotoid glands vary in appearance from scarcely discernible in festae to large and smooth in nicefori. Tubercles are small and conical in macrorhina, acrolopha, and festae, but large and rounded in rostrata and nicefori. All species have characteristic dorsolateral rows of tubercles extending from the posterior margin of the head or parotoid gland posteriorly along the flanks, and small, round tubercles covering the eyelids. They tend to have a prolifera- tion of tubercles on the dorsal surfaces of the limbs. Osteology.— ThQ skulls (Figs. 7 and 8) of Rhamphophryne are triangular in dorsal and ventral views and broader than long. Only one specimen of each of three species, viz. Rhamphophryne acrolopha, macrorhina, and festae is available for detailed ostelogical study; other osteological data were obtained by dissection and/or examination of stereo radiographs. The cranial measure- ments and ratios (Table 1 ) show that the shapes of the skulls of these species Table 1. Skull measurements in Rhamphophryne Species HL (Head Length) mm HW (Head Width) mm H (Height) mm H/HL H/HW HL/HW acrolopha 16.8 18.6 9.0 0.536 0.484 0.903 macrorhina 13.8 14.1 8.0 0.570 0.567 0.979 festae 10.3 11.4 6.3 0.612 0.553 0.904 conform closely to one another. Both acrolopha and festae have skulls which are obviously wider than long, whereas in macrorhina, a species nearly equiva- lent in size to acrolopha, the skull length nearly equals the width. Similarly, the skulls of festae and macrorhina are somewhat narrower with respect to their heights than is that of acrolopha. These flgures tentatively suggest that acro- lopha has, proportionally, the broadest skull of the three species. Complete integumentary-cranial co-ossification does not occur in this genus. The skin is movable on those smooth areas of the skull which are not exostosed. Skin overlying rugose, exostosed areas such as the cranial crests, canthal ridges, and anterior margins of the orbits is partially adherent. It is likely that a similar, partially co-ossified situation prevails in Rhamphophryne as was described for the partly co-ossified hylid frog, Osteocephalus taurinus (Trueb, 1970). If this assumption is correct, the exostosed portions of the skulls of Rhamphophryne have probably been laid down in the lower, dense connective tissue layer of the overlying skin, thereby causing partial adherence of the overlying dermis to the skull beneath. 8 Contributions in Science No. 216 Figure 5. Palmar views of right hands of Rhamphophryne: (a) R. acrolopha, $ (KU 76965); (b) R. festae, $ (USNM 167175); (c) R. macrorhina, $ (LACM 44394); (d) R. nicefori, $ (USNM 163476); (e) R. rostrata, $ (AMNH 1359). Lines equal 2 mm. 1971 Relationships of Neotropical Toads 9 Figure 6. Plantar views of right feet of Rhamphophryne: (a) R. acrolopha, S (KU 76965); (b) R. festae, $ (USNM 167175); (c) R. macrorhina, 9 (LACM 44394); (d) R. nicefori, $ (USNM 163476); (e) R. rostrata, $ (AMNH 1359). Lines equal 2 mm. 10 Contributions in Science No. 216 The premaxillaries are separated medially by connective tissue. A con- spicuously long, pointed palatine process is present posteromedially on each premaxillary (Fig. 7b). The alary processes of the premaxillaries are widely separated medially by the anterior end of the sphenethmoid; they are inclined anteriorly (Fig. 8), and laterally deflected in at least two species (acrolopha and macrorhina). The length of the alary processes varies from two times {acrolopha and macrorhina) as long as the depth of the pars dentalis of the premaxillary to two and one-half times (festae). In ventral view (Fig. 7b), the prevomers lie well lateral to the midlines of the skulls. The anterior ends lie adjacent to the anterior ends of the partes palatinae of the maxillaries. The posterior ends of the prevomers lie ventrally adjacent to the sphenethmoid in festae and acrolopha; in macrorhina the entire prevomer lies laterally adjacent to the anterolateral margin of the sphenethmoid. The anterolateral wings of the prevomers form the anteromedial margins of the choanae; this wing is reduced in size in macrorhina. The postero- medial margins of the choanae are formed partially by the sphenethmoid; the posterolateral prevomerine wings are medially deflected in macrorhina and acrolopha and reduced to tiny spurs in festae. The prevomers of these three species are edentate; prevomerine teeth are also absent in rostrata and nice fori. Palatine bones are present in all species of Rhamphophryne; the bones are narrow and lie posterior to the internal nares. In macrorhina, acrolopha, and festae the distal ends of the palatines are only slightly expanded and articulate with the posterior processes of the partes facialae of the maxillaries and the maxillary processes of the nasals. Proximally the palatines articulate with the sphenethmoid. The palatine is smooth in macrorhina, bears a weak longitudinal ridge in festae and a slightly irregular ridge in acrolopha. The nasals (Figs. 7a and 8) are large, extending anteriorly beyond the level of the mandibular arch to terminate at about the level of the anterodorsal tips of the alary processes. In macrorhina and acrolopha, the nasals converge medially throughout most of their lengths and are entirely underlain by the ossified sphenethmoid. In festae the nasals are medially separated throughout their lengths and extend considerably beyond the anterior terminus of the bony sphenethmoid. The nasals arch posterodorsally over the external nares and terminate posteroventrally in slender maxillary processes which articulate with the palatines and posterior processes of the partes faciales of the maxillaries. The posterior nasal margins bordering the orbits are slightly exostosed in acrolopha and festae, but smooth in macrorhina. Dorsally, the posterior edges of the nasals of all three species articulate with the anterior margins of the frontoparietals. The maxillary is moderately robust in acrolopha and less well developed in macrorhina and festae. The partes faciales are moderate in size and do not articulate with the nasals, except posteriorly in macrorhina and acrolopha; in festae, the pars facialis is reduced with only the posterior process remaining. 1971 Relationships of Neotropical Toads 11 Ventromedially, the partes palatinae of the maxillaries of all three species are inconspicuous. The maxillaries articulate firmly with the short, stout quadra- tojugals posteriorly at approximately the level of the prootic foramen. The frontoperietals (Fig. 7a) converge medially throughout their lengths. Anteriorly, they articulate with the posterior margins of the nasals and join with the posterodorsal corners of the nasals to form upturned supraorbital crests. Posteriorly, the frontoparietals extend laterally over the epiotic eminence areas of the cristae paroticae to form the medial portions of the postorbital crests. The occipital ridge is scarcely discernible in festae; it is represented by a low thickening which borders the medial edge of the open carotid canal and is formed in the dorsal surface of the frontoparietal. The occipital ridge is somewhat better developed in acrolopha; irregularly exostosed spurs of bone arch over the carotid canal and partially roof it. In macrorhina the occipital crest is prominent; it forms a complete roof over the carotid canal. The fronto- parietals articulate with the squamosals dorsal to the cristae paroticae and seem to be fused with the prootics in each species, although this character is difficult to assess without examination of sectioned material. The most striking cranial feature of Rhamphophryne is the development of the sphenethmoid and its associated cartilages. Again, the nature of this unusual characteristic is difficult to describe adequately without having studied serial cross-sections. In lieu of such an opportunity, one is forced to extrapolate on the bases of previous studies. In the majority of anuran skulls, the endo- chondral sphenethmoid is in synosteotic continuity with the septum nasi anteriorly. Associated with the septum, which forms the medial wall of the nasal cavities, are various other cartilages such as the tectum nasi (roof of the nasal cavities) and solum nasi (floor of the nasal cavities). The cartilage forming these elements is continuous with that in which the sphenethmoid is laid down. Usually the anterior parts associated with the nasal cavities remain cartilaginous, whereas the posterior part ossifies to varying degrees, forming a casing for the anterior part of the brain; this is commonly known as the sphenethmoid. In Rhamphophryne acrolopha and macrorhina (and, pre- sumably, also in nice fori and rostrata) the ossification of the sphenethmoid is continuous anteriorly with completely or nearly completely ossified nasal septa and their associated cartilages. There seems to have been a proliferation of the cartilaginous anterior end of the nasal septum, producing the protuberant snout characteristic of this genus. This area is subsequently ossified in acrolopha and macrorhina to produce the bony snout of the adult. A similar situation prevails in festae, except that the anterior end of the snout remains cartilaginous in all but the largest adults. The anterior extension of the snout in this genus has resulted in some secondary modifications of cranial structure. As the snout grows forward during development, it evidently causes the anterior rotation of the alary processes of the premaxillaries so that the processes come to be anteriorly 12 Contributions in Science No. 216 Figure 7. Skull of Rhamphophryne festae, $ (USNM 167168); (a) dorsal; (b) ventral. X 7. 1971 Relationships of Neotropical Toads 13 Figure 8. Skull of Rhamphophryne festae, $ (USNM 167168) : in lateral view. X 7. inclined {macrorhina and acrolopha) or nearly horizontally oriented {festae; Fig. 8). In macrorhina and acrolopha, the alary processes are laterally de- flected, presumably as a result of the thickening of the anterior end of the septa nasi. Examination of preserved specimens of this genus shows that the nostrils are located anterior to the position one would normally anticipate with respect to the anterior limit of the upper jaw. Because the external nares usually lie posterior to the anterior levels of the nasal cavities, the position of the nostrils suggests that the nasal structures have expanded or moved forward to occupy a more anterior position in the extended snouts. Modifications of this kind are not unknown among other anurans. Similar phenomena have been described by Lynch (1971) for the fuscus group of Leptodactylus and by Trueb (1970) in the casque-headed hylids. There is, however, one striking difference which should be noted in the hylids. The addition of new bones to produce protuberant snouts in the hylids and the proliferation of existing cranial structures have been derived by modifications of the highly plastic dermal elements. The kinds of changes which have oc- curred in Rhamphophryne have been produced by proliferation and increased ossification of endochondral elements. The sphenethmoid (Fig. 7b) is not extensively ossified posteriorly, especially in macrorhina. In this species, the bony part of the sphenethmoid terminates a short distance posterior to the level of the palatines, thereby 14 Contributions in Science No. 216 creating a particularly large, unossified area around the optic foramen. The sphenethmoid is better ossified posteriorly in festae and acrolopha. The parasphenoids of all three species are very similar. The bones are edentate and truncate anteriorly. In festae and acrolopha the anterior end of the bone lies ventral to the sphenethmoid, just posterior to the level of the palatines, whereas in macrorhina the anterior terminus lies more posterior and articulates with the posterior bony margin of the sphenethmoid. The squamosals (Fig. 8) are robust. The anterior arms are short, broad spurs which project one-fourth, or less, of the distance from the dorsal union of the three squamosal arms to the maxillary. An inconspicuous pretympanic crest in some specimens of acrolopha and macrorhina forms by thickening of this squamosal element. The posterior arms lie adjacent to, and do not extend beyond, the distal edges of the cristae paroticae. In dorsal view the posterior squamosal arm broadly overlaps the crista parotica and articulates at least partially with the frontoparietal. Along the anterior edge of the crista, the overlapping portion of the squamosal is exostosed to form a postorbital ridge; similarly, along the lateral margin of the crista, the squamosal forms a supra- tympanic crest. The ventral squamosal arms are particularly robust. Each arm bears a broad articulation with the posterior ramus of the pterygoid medially and the quadratojugal laterally. There is little variation among the pterygoids. Each species has a well- developed pterygoid (Fig. 7b), in which the posterior ramus articulates with the squamosal, and the medial ramus broadly overlaps the parasphenoid posterolaterally. The anterior pterygoid rami of festae and acrolopha articulate laterally with the maxillaries and anteriorly with the distal end of the palatines. The anterior pterygoid ramus of macrorhina is shorter, articulating only with the maxillary laterally. The prootic and exoccipital bones are well ossified and, as previously mentioned, apparently fused with the frontoparietal. Bony oculomator, prootic, and jugular foramen are evident. All vestiges of the columella are lost, and the lateral portions of the otic capsule are cartilaginous. The occipital condyles are closely juxtaposed rather than widely separated. Similarly, the cervical condyles are closely juxtaposed as in all other bufonids (Lynch, 1969). There are seven or eight presacral vertebrae which bear mod- erately long transverse processes (Fig. 9). In an anterior to posterior sequence, the widths of the transverse processes from narrowest to widest are 2-7-6-S-3-4 in those species having seven presacral vertebrae. In macrorhina the widths of the transverse processes are 2-8-7-6-5-S-4. The transverse pro- cesses of the second presacral vertebrae are directed anteriorly. Those of vertebrae three through six in nicefori, festae, and acrolopha, three through seven in macrorhina, and three through five in rostrata are directed posteriorly. Transverse processes of vertebrae six and seven are directed anteriorly in rostrata. Those of the seventh vertebrae in nicefori, festae, and acrolopha and 1971 Relationships of Neotropical Toads 15 Figure 9. Ventral view of vertebral column of Rhamphophryne festae, 9 (USNM 167168). X 7. 16 Contributions in Science No. 216 the eighth in macrorhina are directed anteriorly. The sacral diapophyses of all species are expanded. In festae and rostrata the sacrum and coccyx are fused. In nicefori and acrolopha the sacrum bears a monocondylar articulation with the coccyx, whereas in macrorhina the articulation is very weakly bicondylar. The coccyx bears no noticeable lateral expansion in any of the species. The omosternum is absent in this genus, as it is in all other bufonids ex- cept N ectophrynoides and toads of the Bufo haematiticus group. The pectoral girdle is arciferal but tends to have a broad epicoracoid bridge. The sternum is cartilaginous. The ilia bear no notable prominences or crests. The limb ele- ments are relatively slender. The phalanges are markedly reduced in length in festae, but of normal extent in other species. Myology.— Th& adductor longus muscle is absent in the thighs of all species of Rhamphophryne. The length of the tensor fasciae latae muscle is somewhat variable. The muscle is short in acrolopha, macrorhina, and rostrata; it originates from a point about one-fourth the distance from the proximal end of the ilium and inserts on the surface of the cruralis-gluteus complex. In festae and nicefori the tensor is longer; it originates at approxi- mately the midlength of the ilium. Natural history. — Little is known concerning the habits and life history of this genus. All species (with the possible exception of Rhamphophryne rostrata for which there are no data available) have few, large, unpigmented eggs. No tadpoles have been associated with the genus. Although vocal slits are present in the males of three species {nicefori, macrorhina, and rostrata), none has been observed to call. The available field notes suggest that members of this genus are diurnal; all specimens have been obtained from the forest floor by day. Significantly, none (including gravid females) has been recorded as having been found near water. Etymology.— ThQ generic name is derived from the Greek words rhampho, meaning beak, and phryne, meaning toad, with reference to the protuberant snout characterizing this group of bufonids. Distribution. — Members of this genus occur at moderate elevations in the mountains of northern Colombia and extreme eastern Panama, and on the lower Andean slopes and in the upper Amazon Basin of Ecuador (Fig. 10). Remarks. — On the basis of external characters, members of the genus Rhamphophryne are easily confused with juveniles of Bufo typhonius. Toads of the latter species are widespread, occurring throughout the Amazon Basin, in northwestern South America, and eastern Panama. Bufo typhonius generally is found at moderate to low elevations (up to 800 m). All specimens of Rhamphophryne have been collected at elevations exceeding 800 m except festae, which apparently occurs sympatrically with Bufo typhonius in Amazonian Ecuador. Adult Bufo typhonius are easily distinguished from Rhamphophryne by their much greater size, less protuberant snouts, broad alate processes of the squamosal dorsally, and conspicuous tympani. How- ever, in smaller individuals of this species, the snouts are proportionally 1971 Relationships of Neotropical Toads 17 Figure 10. Map of Panama, western Colombia and Ecuador, showing locality records of Rhamphophryne. 18 Contributions in Science No. 216 more protuberant, the squamosal processes poorly developed, and the tympani much less evident. Although inconspicuous in some individuals, the presence of a tympanum consistently distinguishes Bufo typhonius from all members of the genus Rhamphophryne. Some comment should be made concerning the relationship of Rhampho- phryne with a group of small, earless, montane bufonids found at elevations above 1000 meters in Costa Rica and western Panama. Included in this group are Bufo periglenes, fastidiosus, holdridgei, and Crepidophryne epioticus. Two other species {Bufo coerulescens and simus) often have been mentioned in conjunction with these taxa. Bufo coerulescens is known only from two type specimens of doubtful maturity, and the type of simus is apparently lost. A great deal of confusion exists concerning the relationships among all of these species and of these with other bufonids. Hopefully, a thorough study will elucidate these problems; however, until such work appears, it should be pointed out that Rhamphophryne is morphologically distinct from these toads. Key to the Species of RHA MPHOPHR YNE 1. Hands with long, free fingers 2 Hands palmate 3 2. Dorsolateral tubercles small, depressed, in continuous row; snout directed anteroventrally; cranial crests moderately developed acrolopha Dorsolateral tubercles large, conical, in discontinuous row; snout directed anteriorly; cranial crests weak or absent rostrata 3. Dorsolateral tubercles small, depressed in continuous row; occipital crests well developed macrorhina Dorsolateral tubercles moderate-sized, conical, not in continuous row; occipital crests weak 4 4. Third finger much longer than others festae Third finger only slightly longer than others nice fori Accounts of Species Rhamphophryne acrolopha, new species Holotype. — Adult male. University of Kansas Museum of Natural History (KU) 76965 from Cerro Mali, Darien Province, Panama, elevation 1410 m; obtained on July 12, 1963, by William E. Duellman. Paratypes. — KU 76961-4 and UMMZ 129694-5 obtained with the type, MLS 1240 and LACM 54337-8 obtained on July 10 and 11, 1963, by William E. Duellman; USNM 151075-6 from elevations of 1265 and 1451 m on Cerro Mali, and USNM 151101, 151104-5, and 151107 from Cerro Tacarcuna, Darien Province, Panama, between elevations of 1265 and 1481 m, obtained between January 29 and March 1, 1964, by Charles O. Handley, Jr. Diagnosis.— A moderate-sized Rhamphophryne (^38 mm, $ 45 mm) which can be distinguished from other members of the genus by the following 1971 Relationships of Neotropical Toads 19 combination of characters: (1) seven presacral vertebrae; (2) sacral vertebra not fused with coccyx; (3) snout long, protuberant, and directed anteroven- trally; (4) cranial ornamentation moderately developed; (5) supraorbital, postorbital and supratympanic crests present; (6) pretympanic crest variable in occurrence; (7) occipital crest present but indistinct; (8) dorsal tubercles small and conical; (9) dorsolateral tubercle row small, depressed, extending from posterior margin of parotoid gland posteriorly to a point about two-thirds of the distance between axilla and groin; (10) hands and feet with long digits and reduced webbing; (11) subarticular tubercles present; (12) supernumer- ary tubercles present but poorly developed; (13) subdigital lamellar pads absent; (14) tensor fasciae latae not extended; (15) vocal slits absent; (16) cloacal sheath absent. Description of /lo/o/ype.— Snout-vent length (S-VL) 41.9 mm; tibia length 15.4 mm, 36.8 per cent of S-VL; foot length 23.3 mm, 55.6 per cent of S-VL; head length 14.7 mm, 35.1 per cent of S-VL; head width 16.4 mm, 39.1 per cent of S-VL; snout in dorsal aspect acuminate (Fig. 4a), in profile (Fig. 3a) long, directed anteroventrally, truncate terminally; distance from nostril to tip of snout equal to distance between nostril and eye, 3.3 mm, constituting 22.4 per cent of head length; tip of snout bulbous, connected by broad ventral keel to upper jaw; loreal region concave; nostrils pro- tuberant, lateral, at level anterior to terminus of upper jaw, slightly ventral to canthal ridge; diameter of eye 4.0 mm, 27.2 per cent of the head length; pupil horizontally elliptical; interorbital distance 7.1 mm, 43.3 per cent of head width; ear absent; tongue twice as long as wide, widest at two-thirds its length, pointed anteriorly and posteriorly; tongue at greatest width about two-thirds as wide as mouth; posterior two-thirds of tongue free; choanae minute, round, lateral in position; prevomerine teeth and vocal slits absent. Canthal ridges angular in section, converging anteriorly to form acumi- nate snout, terminating posteriorly in slight swellings at anterodorsal corner of orbit; crown of head with moderately developed, dorsally depressed cranial ridges; broad supraorbital ridge bordering medial edge of orbit, continuous with narrow postorbital crest; supratympanic crest continuous with postorbital ridge, narrow anteriorly, broad and depressed posteriorly adjacent to anterior margin of parotoid gland; occipital crest scarcely discernible spur produced at junction of supraorbital and postorbital ridges; pretympanic crest poorly- developed spur projecting ventrally from junction of postorbital and supra- tympanic ridges at posterior, dorsolateral margin of orbit. Arms moderately robust and lacking folds; axillary membrane absent; fingers (Fig. 5a) long, slender; length of fingers from shortest to longest 1-2-4-3; phalangeal formula of hand 2-2-3-3; fingers vestigially webbed; sub- articular tubercles of hand large but depressed; supernumerary tubercles of hand few and small in size; inner palmar tubercle round, flat; medial palmar tubercle large, one and one-half times as large as inner palmar tubercle, ovoid, less depressed than inner palmar tubercle; prepollical protuberance and nuptial 20 Contributions in Science No. 216 excresence absent; hind limbs relatively long, lacking folds; toes long, slender (Fig. 6a); length of toes from shortest to longest 1-2-3-5-4, third and fifth about equal in length, phalangeal formula of foot 2-2-3-4-3; webbing extend- ing to bases of penultimate phalanges of toes one and two; third toe one-half webbed; fourth toe one-fourth webbed; fifth toe one-half webbed; subarticular tubercles large, ovoid, depressed; supernumerary tubercles few in number, small, greatest proliferation between inner and out metatarsal tubercles; inner metatarsal tubercle large, flat, and irregularly shaped; out metatarsal tubercle round, slightly depressed, one-half size of inner metatarsal tubercle. Cloacal opening directed posteroventrally at mid-level of thighs; no noticeable anal sheath; parotoid glands small, rounded in dorsal and lateral views; skin of dorsum with small, conical tubercles; scattered, indistinct, larger tubercles on eyelid, at angle of jaw and along flanks; skin of throat, chest, belly and posterior surfaces of thighs granular; dorsolateral row of small, rounded, depressed tubercles extending posteriorly from posterior margin of parotoid gland to a point about two-thirds distance between axilla and groin. Dorsal ground color pale creamy tan with darker markings; reticulated dorsal dark brown marks forming interorbital bar with dark marks extending onto eyelids; V-shaped mark medial to each parotoid extending from occiput to posterior area of pectoral region; sacral area traversed by broad chevron; dorsal surface of limbs with narrow transverse bands proximally and wide transverse bands distally; hands and feet dark dorsally except for first and second fingers and first, second, and third toes which are marbled with creamy tan; flanks and postympanic areas predominately dark brown and grayish tan; belly and ventral surfaces of limbs creamy tan marbled with dark brown; palmar and plantar surface dark brown with grayish tan palmar and metatarsal tubercles; ventral ground color creamy tan with bold, dark brown marbling on limbs and belly; coalescence of marbling anteriorly to form nearly uniform dark brown throat region; scattered, small, white tubercles over entire venter. Description and variation. — Eleven adult males from Cerro Mali and Cerro Tacarcuna, Panama, have snout-vent lengths of 31.5 to 41.9 mm (mean 37.8 mm). Eight adult females from the same localities have snout-vent lengths of 40.6 to 49.1 mm (mean 44.9 mm). There is no marked intersexual or intra- sexual variation in the measurements and ratios compared with those of the holotype. In preservative, the dorsal ground coloration of Rhamphophryne acro- lopha varies from a uniform dark brown to a medium or grayish brown. The former is less common; in these specimens the row of dorsolateral tubercles is grayish tan. There is an absence of any distinctive markings on the dorsal surfaces of the limbs or the side of the head. The posterior surfaces of the thighs and all ventral surfaces are heavily marbled with dark brown. Paler individuals are marked with a variety of dorsal spots and blotches ranging from gray to black in color. A partial or complete vertebral stripe and a light colored labial stripe may be present; occasionally, a mid-ventral stripe is present. 1971 Relationships of Neotropical Toads 21 In life, dorsal colors have been observed to vary from yellowish tan to reddish and dark brown. The iris is bronze with black reticulations. There is some variation among specimens in the development of cranial crests and the snout. In general, larger individuals, such as mature females, tend to have the most pronounced development of both crests and snout. This is to be expected as a result of continued ossification. In subadult specimens of Rhamphophryne acrolopha, the snout is more nearly truncate in profile; with increased size, the snout develops its typical anteroventral orientation. Etymology.— The specific name is derived from the Greek akrolophos meaning crest of a mountain or ridge, with reference to the isolated occurrence of this species at high elevations in the Serrania del Darien. Distribution.— Rhamphophryne acrolopha is known only from the Ser- rania del Darien (Cerro Tacarcuna and Cerro Mali) in Panama and adjacent Colombia (Fig. 10). Remarks. — Externally Rhamphophryne acrolopha most closely resembles, and most easily is confused with, macrorhina from Colombia. Both are rela- tively large members of the genus, and both are characterized by extremely protuberant snouts and small, depressed series of dorsolateral tubercles. It should be noted that the cranial crests and tubercles on the eyelids are more extensively developed in macrorhina, and the parotoid glands in that species are less evident than in acrolopha. The hands and feet are less specialized in acrolopha than in macrorhina; the tendency for formation of lamellar pads is less marked, webbing is comparatively reduced, and sub articular and super- numerary tubercles are more distinct in acrolopha. The inner and outer palmar tubercles are obvious in acrolopha but barely evident in macrorhina. Despite the external similarities of these species, they are quite distinct internally. Rhamphophryne acrolopha lacks vocal slits, whereas macrorhina retains them. Furthermore, macrorhina has eight presacral vertebrae and a weakly bicon- dylar sacral-coccygeal articulation in contrast to the seven presacral vertebrae and monocondylar articulation of acrolopha. Virtually nothing is known concerning the habits and life history of this species. All specimens (adult males, females, and juveniles) were found by day amidst leaf litter on the forest floor between the months of January and July. The forest at the Cerro Mali site contains many small trees, including palms, reaching heights of about 12 m; larger trees, with heights to 30 m, are scattered throughout the forest. There is an understory of bushes and ferns, and most of the trees have a thick covering of moss on the trunks. The forest floor is characterized by a thick, mulch layer and many rotting logs. Bromeli- ads abound, but orchids and tree ferns are rare (Field notes, W. E. Duellman, July 10, 1963). Gravid females were obtained during the months of February, March, and July. One female (KU 76964) from Cerro Mali contained a total of 145 unpigmented eggs on one side of the body cavity (eggs present on both sides of the body cavity for the species) . Of this total, 50 of the eggs have an average 22 Contributions in Science No. 216 diameter of 2.5 mm, whereas the remainder have diameters of less than 1.0 mm. Tadpoles are unknown. Specimens examined.— {6 \) COLOMBIA: Choco: East slope Cerro Mali, 1470 m, KU 76984, 76986, USNM 150941. PANAMA: Darien: Cerro Mali, 1265-1451 m, KU 76941 (juvenile), 76944 (juvenile), 76945, 76946-54 (juveniles), 76955-6 (skeletons), 76957-60 (juveniles), 76961-5, 76968-83 (juveniles), 76984, 76985-6 (juveniles), LACM 54337-8, MLS 1240, UMMZ 129694-5, USNM 151075-6; Cerro Tacarcuna, 1265-1481 m, USNM 151100 (juvenile), 151101, 151102-3 (juveniles), 151104-5, 151106 (skeleton), 151107; headwaters of Rio Pucro, USNM 151071 (juvenile). Rhamphophryne festae, new combination Atelopus festae Peracca, 1904: 18 [Syntypes. — IZS 2903 from Gualaquiza and 2904 from “Rio Santiago” (= lower Rio Zamora, fide Peters, 1955), Morona-Santiago Province, Ecuador; Enrico Festa, collector]. Noble, 1920:446. Peters, 1955:347. Diagnosis. — A moderate-sized Rhamphophryne (^ 42.0 mm, $ 45.8 mm) which can be distinguished from other members of the genus by the following combination of characters: (1) seven presacral vertebrae; (2) sacral vertebra fused with coccyx; (3) snout long, protuberant, directed anteroventrally; (4) cranial ornamentation moderately well developed; (5) su- praorbital, postorbital, and supratympanic crests present; (6) pretympanic crest present; (7) occipital crest moderately developed; (8) dorsal tubercles scattered, minute, conical; (9) dorsolateral row of slightly enlarged, conical tubercles extending from posterior margin of skull to a point about three- fourths distance between axilla and groin; (10) hands and feet with some shortened digits and no marked reduction of webbing; (11) subarticular tubercles present; (12) supernumerary tubercles present; (13) subdigital la- mellar pads absent; ( 14) tensor fasciae latae extended; (15) vocal slits absent; (16) cloacal sheath absent. Description and variation . — Average measurements of three adult females and three adult male Rhamphophryne festae are, as follows (corresponding figures for males are included in parentheses after females): snout-vent length (S-VL) 37.2 mm (33.2 mm) ; tibia 12.1 mm ( 10.9 mm), 31.9 (33.4) percent of S-VL; foot and tarsus length 17.1 mm ( 15.8 mm), 46.6 (48.2) per cent of S-VL; head length 12.7 mm ( 1 1.3 mm), 34.0 (34.2) per cent of S-VL; head width 13.6 mm (12.6 mm), 38.2 (38.1) per cent of S-VL. The snout in dorsal aspect (Fig. 4b) is acuminate, and in profile (Fig. 3b) is moderately long, directed anteroventrally, and acuminate terminally. The distance from the nostril to the tip of the snout ( $ 2.3 mm, of 5 2.1 mm) is noticeably less than the distance from the nostril to the eye ( 9 2.9 mm, $ 2.6 mm). The tip of the snout is pointed, not bulbous, and a sharp, ventral keel runs from the tip of the snout to the upper jaw. The loreal region is concave. Nostrils are slightly protuberant and lateral; they are located at a level anterior 1971 Relationships of Neotropical Toads 23 to the terminus of the upper jaw and ventral to the canthal ridge. The diam- eter of the eye is 3.7 mm (3.3 mm), 29.9 (29.6) per cent of the head length. The pupil is horizontally elliptical. The interorbital distance is 6.1 mm (5.9 mm), 44.9 (46.8) per cent of the head width. The ear is absent. The tongue is small, pointed anteriorly and round posteriorly. It is three-fourths as wide as long, widest at one-half its length, and slightly more than one-third as wide as the mouth. The posterior half of the tongue is free. Choanae are minute, round, and lateral in position. Prevomerine teeth are absent. The canthal ridges are angular in section and converge anteriorly to form the acuminate snout. Cranial ridges are present and moderately well developed in larger individuals (Fig. 4b). The supraorbital, postorbital and supratym- panic crests are narrow; in smaller specimens, they bear series of small, conical tubercles throughout their lengths. The pretympanic crest does not bear tubercles. The occipital crest is absent in smaller individuals (Fig. 11b); in place of the crest, there is a row of small tubercles. Larger individuals have a moderately well-developed occipital crest which is most obvious near its junc- tion with the supraorbital and postorbital cranial crests. The arms are slender and lack folds. An axillary membrane is absent. Some of the fingers are reduced in length (Fig. 5b), their lengths from shortest to longest being 2-4- 1-3. The phalangeal formula of the hand is 2-2-3-3. Web- bing extends to the bases of the penultimate phalanges of fingers one, two, and four. The third finger is basally webbed. The inner palmar tubercle is moderate in size, elliptical and flat. Supernumerary and subarticular tuber- cles are present. The medial palmar tubercle is twice as large as the inner tubercle, ovoid, and flat. Males lack prepollical protuberances and nuptial excrescences. The hind limbs are relatively long, slender, and lack folds. Some of the toes are reduced in length, their lengths from shortest to longest being 1-2-3-5-4; the third and the fifth toes are about equal in length (Fig. 6b). The phalangeal formula of the foot is 2-2-3-4-3. Webbing extends to the bases of the penultimate phalanges of all toes except the fourth which is webbed basally. Subarticular and supernumerary tubercles are present. The inner metatarsal tubercle is moderate in size, irregularly shaped and flat, whereas the outer metatarsal tubercle is small (one-half the size of the inner metatarsal tubercle), ovoid and depressed. The cloacal opening is directed posteriorly at the upper level of the thighs. A cloacal sheath is absent. The parotoid glands are moderately well developed in larger specimens; however, in smaller specimei^ they are small, barely evident dorsally, and most obvious in lateral view 'as long, narrow swellings ventral to the dorsolateral tubercle row (Fig. 11a and b). There is a proliferation of minute, conical tubercles scattered over the dorsal surfaces of ^ body; these tubercles are especially concentrated on the dorsal surfaces of the limbs. Small tubercles cover the eyelids and line the cranial ridges. A series of very slightly enlarged tubercles runs in a posterodorsal direction from the angle of the jaw, and dorsolaterally from the posterior margin of 24 Contributions in Science No. 216 a Figure 11. Immature Rhamphophryne festae, $ , 26.5 mm snout-vent length (USNM 167175): (a) dorsal; (b) lateral. X 3. the supratympanic ridge, posteriorly, to a point about three-fourths of the distance between the axilla and the groin. The skin of the throat, chest, belly, flanks, ventral surfaces of the limbs, and the anterior and posterior surfaces of the thighs is granular. In preservative, the dorsal ground color of Rhamphophryne festae varies from a uniform dark brown to yellowish tan. Darker mottling and transverse bars are evident on paler specimens. The ventral surfaces vary from yellowish tan to dark brown in color. In some, the venters are uniform, whereas in others, they are variously mottled with dark brown. The palmar and plantar tubercles and digital pads of all specimens are creamy tan in color. Rhamphophryne festae, in life, is reddish brown above with slightly darker cross-banding, according to color notes recorded by James A. Peters (personal communication). The tubercles forming the dorsolateral tubercle 1971 RELATIONSfflPS OF NEOTROPICAL TOADS 25 row have orange-red heads. The belly is a dull, dusty red with bluish white mottling posteriorly. The venters of the digits are pale pink, and the iris is black with a narrow ring of gold surrounding the pupil. Distribution— Rhamphophryne festae is known from moderate and low elevations ( 100-700 m) on the Atlantic Andean slopes and upper Amazon Basin of Ecuador (Fig. 10). Although one of the syntypes (IZS 2903) of Rhamphophryne festae is a gravid female, it lacks certain diagnostic features evident on larger, more mature specimens (compare Figs. 3b, 4b, and 11). The cranial crests are much more evident on the larger specimens and the snout more pro- tuberant. The hands and feet, and other characteristics described by Peracca ( 1904) are virtually the same. Rhamphophryne festae is possibly the most specialized member of the genus. The highly modified feet render festae readily distinguishable from other species of Rhamphophryne. The triangular head and the physiognomy of the hands and feet are probably responsible for the original placement of this species in the genus Atelopus\ it should be noted that the cervical and first presacral vertebrae are not fused in festae, whereas these elements are char- acteristically fused in all A telopus. The specialization of Rhamphophryne festae is most evident internally. Rhamphophryne rostrata and festae are the only two members of the genus having a fused coccyx and sacrum, which presumably is an advancement over the weakly bicondylar articulation of macrorhina. Like acrolopha, and in contrast to rostrata, nice fori, and macrorhina, festae has lost the vocal slits. The second and fourth fingers and the first, second, third and fifth toes are notably short; the phalangeal formulae are normal, although the indi- vidual phalangeal elements of the digits are shortened and poorly, if at all, ossified. Very little natural history information is available. Peters observed that a subadult specimen obtained in July was found on a log in the jungle, quite high above any of the streams. A gravid female (USNM 167174) contains relatively few, large (1.8 mm), unpigmented eggs. Specimens examined.— (IS) ECUADOR: Napo: near Arajuno, 537 m, KU 124949-50, USNM 167176; Loreto, CAS 11414-7, USNM 167175; Pastaza: Alto Rio Pucayacu, USNM 167168 (skeleton), 167169; Don Tomas, 5 km S Montalvo, USNM 167172; Rio Bobonaza, 703 m, USNM 167167; Rio Solis (headwaters), USNM 167174; Rio Capahuari (headwaters), USNM 167171; Rio Rutuno (tributary of Rio Bobonaza), USNM 167170; Rio Vil- lano USNM 167173; Morona-Santiago : Gualaquiza, IZS 2903; “Rio Santiago” (= lower Rio Zamora, fide Peters, 1955), IZS 2904. Rhamphophryne macrorhina, new species Bufo rostratus (part) Noble, 1920:445. Gallardo, 1962:96. Cochran and Goin, 1970:93. 26 Contributions in Science No. 216 Holotype— Adult female, Los Angeles County Museum of Natural His- tory (LACM) 44394 from Santa Rita, Departamento de Antioquia, Colom- bia, between elevations of 1890 and 1910 m, one of a series obtained on June 9, 1968, by P. A. Silverstone and J. Ramos. Paratypes: LACM 44395 and KU 124946 obtained with the type by P. A. Silverstone and J. Ramos; KLF 124947 (skeleton) from the type locality at an elevation of 1930 m on June 8, 1968, by P. A. Silverstone and J. Ramos. Diagnosis —A large Rhamphophryne ($ 43.4 mm, $ 51.0 mm) which can be distinguished from other members of the genus by the following com- bination of characters: (1) eight presacral vertebrae; (2) sacral vertebra not fused with coccyx, with a weakly bicondylar articulation; (3) snout long, pro- tuberant, and directed anteroventrally; (4) cranial ornamentation well devel- oped; (5) supraorbital, postorbital, and supratympanic crests present; (6) pretympanic crest present; (7) occipital crest present and distinct; (8) dorsal tubercles scattered, small and rounded; (9) dorsolateral row of small, de- pressed tubercles extending from posterior margin of parotoid gland pos- teriorly to a point about one-half distance between axilla and groin; (10) hands and feet with moderate-sized digits and no marked reduction of web- bing; (11) subarticular tubercles absent; (12) supernumerary tubercles pres- ent; (13) tendency towards presence of subdigital lamellar pads; (14) tensor fasciae latae not extended; (15) vocal slits present in males; (16) cloacal sheath absent. Description of holotype.Suout-VQnt length (S-VL) 51.4 mm; tibia 16.0 mm, 31.1 per cent of S-VL; foot length 26.0 mm, 50.6 per cent of S-VL; head length 18.2 mm, 35.4 per cent of S-VL; head width 18.8 mm, 36.6 per cent of S-VL; snout in dorsal aspect (Fig. 4c) acuminate, in profile (Fig. 3c) long, directed anteroventrally, acuminate terminally; distance from nostril to tip of snout about equal to distance between nostril and eye, 4.0 mm, constituting 22 per cent of head length; tip of snout bulbous, not connected by distinct ventral keel to upper jaw; loreal region concave; nostrils slightly protuberant, lateral, at level anterior to terminus of upper jaw, ventral to canthal ridge; diameter of eye 4.8 mm, 26.4 per cent of head length; pupil horizontally elliptical; interorbital distance 8.2 mm, 43.6 per cent of head width; ear absent; tongue slightly less than twice as long as wide, widest at one-half its length, pointed anteriorly, rounded posteriorly; tongue at greatest width about one-half as wide as mouth; posterior five-eighths of tongue free; choanae minute, round, lateral in position; prevomerine teeth absent; vocal slits present in males. Canthal ridges angular in section, converging anteriorly to form acumi- nate snout (Figs. 3c and 4c), terminating posteriorly in slight swelling at anterodorsal corner of orbit; crown of head with well-developed cranial ridges; distinct supraorbital ridge bordering medial edge of orbit, continuous with well-developed postorbital crest; supratympanic crest continuous with post- orbital ridge, narrow anteriorly, expanded posteriorly and depressed postero- medially; posteromedial portion of supraorbital crest bordering anterior margin 1971 Relationships of Neotropical Toads 27 of parotoid gland; occipital crest well-developed ridge, produced at junc- tion of supraorbital and postorbital crests; pretympanic crest moderately well-developed spur projecting ventrally from union of postorbital and supra- tympanic ridges at posterior, dorsolateral margin of orbit. Arms moderately slender and lacking folds; axillary membrane absent; fingers of moderate length, not especially slender (Fig. 5c); length of fingers from shortest to longest 2- 1-4-3, first and third about equal in length; pha- langeal formula of hand 2-2-3-3; webbing extending from ultimate phalange of first finger to distal part of penultimate phalange of second finger, from base of ultimate phalange of second finger to base of antepenultimate pha- lange of third finger, from base of antepenultimate of third finger to base of penultimate of fourth finger; subarticular tubercles absent; series of sub- digital, lamellar, pad-like swellings on each finger; supernumerary tubercles scattered, few in number; inner palmar tubercle absent; medial palmar tuber- cle ovoid, depressed, indistinct; hind limbs relatively long, lacking folds; toes of moderate length, not especially slender (Fig. 6c); length of toes from shortest to longest 1-2-3-5-4, third and fifth about equal in length; phalangeal formula of foot 2-2-3-4-3; first two toes fully webbed; second and fifth toes three-fourths webbed; third toe about one-third webbed; subarticular tubercles absent; series of subdigital, lamellar, pad-like swellings on each toe; super- numerary tubercles scattered principally on central plantar surface; inner metatarsal tubercle elliptical, flattened; outer metatarsal tubercle about two- thirds size of inner metatarsal tubercle, ovoid, depressed. Cloacal opening directed posteriorly at mid-level of thighs; cloacal sheath absent; parotoid glands small, indistinct, and diffuse dorsally, scarcely evident laterally; skin of dorsum with a few scattered, depressed tubercles; round, densely distributed tubercles covering eyelid; three larger, conical tubercles arranged in posterodorsally oriented row at angle of jaw; other conical tuber- cles on limbs and around anus; skin of throat, chest, belly and posterior surfaces of thighs granular; dorsolateral row of small, rounded, depressed tubercles extending posteriorly from posterior margin of parotoid gland to about mid-point of distance between axilla and groin. Dorsal ground color medium brown; indistinct and variable creamy tan mottling on back and first two digits of hand and foot; delicate, incomplete vertebral line on posterior one-third of dorsum; dorsolateral tubercle row grayish tan; flanks and sides of head mottled brown; limbs dark brown dor- sally with light colored tubercles along lateral edges; distinct, pale, creamy tan area anterior to forelimb and at angle of jaw on upper lip; less distinct pale spot ventral to eye; venter yellowish tan delicately mottled with dark brown; concentrations of darker pigment around mandible and in gular and pectoral regions; scattered, cream colored tubercles on venter, most numer- ous at angle of jaw and in pectoral region. Description and variation.— Four adult males have snout-vent lengths of 35.6, 42.1, 43.4, and 43.7 mm (mean 41.2 mm). All measurements and ratios 28 Contributions in Science No. 216 are comparable to those of the holotype except one. The ihterorhital distance of the four males is 45.3 per cent (42.9-52.9) of the head width, whereas the interorbital distance of the female is only 43.6 per cent of the head width. Interorbital distances were measured between the medial margins of the supra- orbital crests, thus the apparent variation probably only reflects the greatly thickened condition of the crests in the large female. Similarly, the other cra- nial crests are less well developed and the snouts straighter in the smaller and less extensively ossified males, females, and the single juvenile specimen. In preservative, the only color variation noted is in the dorsal marking. The males tend to have distinct creamy tan dorsal spots which are absent on the large female. In life, a subadult female (LACM 44393) was noted to be dull, dark brown above and light brown marbled with dark brown below; there were some traces of ruddy on the venter and sides. The iris was yellow, heavily reticulated with black and with a ring of bright yellow around the pupil. A horizontal, brown line divided the iris. Limbs were dull buff ven- trally, marbled with brown. Etymology. — The specific name is derived from the Greek prefix macro, meaning large, and the Greek word rhinos, meaning nose, with reference to the inordinately long snout of this species. Distribution.— Rhamphophryne macrorhina is known only from four localities in Colombia (Fig. 10). It has been obtained at Santa Rita north of Medellin, at localities 6 km ESE of Villa Maria and 8 km S Valdivia, and from Santa Rita Creek, 14 miles N of the village of Mesopotamia (south of Medel- lin). There is some question as to the validity of the latter locality. Santa Rita lies to the east of Mesopotamia, rather than to the north, according to Richard Newcomer (Stephen R. Edwards, personal communication) . Remarks.— Rhamphophryne macrorhina is superficially most similar to R. acrolopha; for a comparison of these two species, refer to the remarks section in the account of acrolopha. Morphologically, macrorhina combines highly specialized external characters with some of the least specialized internal characters found in the genus. The tendency toward shortened digits, forma- tion of subdigital lamellar pads, reduction of palmar tuberculation and parotid glands, and proliferation of cranial crests obviously represent derived char- acter states. In contrast to the latter, macrorhina is the only member of Rhamphophryne which retains eight presacral vertebrae. The sacral-coccygeal articulation is very weakly bicondylar, that is, the separation between the sacral condyles is reduced to a shallow groove. This condition seems to be a modification of the bicondylar articulation characteristic of most other bufonids. Cochran and Goin’s ( 1970) redescription of Bufo rostratus was based on a specimen of Rhamphophryne macrorhina (AMNH 1384, a paratype of Bufo rostratus). The specimen redescribed is a male, not a female as stated. The dorsolateral tubercles are not conical, but depressed. The upper jaw does not extend beyond the lower jaw, although the anterior position of the nostrils might create this illusion. The tympanum is not hidden; it is absent. 1971 Relationships of Neotropical Toads 29 P. A. Silverstone (personal communication) collected four individuals (an adult male, subadult and gravid female, and a juvenile) of Rhampho- phryne macrorhina in June. Each was obtained by day on the ground of a forested hillside not close to any body of water. Although males were not observed to call, one individual (a young female) uttered a “sad, small squeak” when handled. Noble (1917) provided a description of the habitat in which one individual was found (refer to the remarks section in the account of Rhamphophryne rostrata) . A gravid female has relatively few, large (about 2 mm), unpigmented eggs. Males lack prepollical protuberances and nuptial excrescences. Specimens examined: (9) COLOMBIA: Antioquia: Santa Rita, 1890- 1910 m, KU 124946, 124947 (skeleton), LACM 44394-5; Santa Rita Creek, 14 miles N Mesopotamia, 1890-2424 m, AMNH 1384; 8 km S. Valdivia, 1450 m, FMNH 63838-40; Caldas: 6 km ESE Villa Maria, 2130 m, KU 129904. Rhamphophryne nicefori, new combination Bufo rostratus nicefori Cochran and Coin, 1970:95 [Holotype.— USNM 163476 from Hacienda Palmas, El Chaquiro, Departamento de Antioquia, Colombia, elevation 2670 m; obtained in December, 1966, by Hno. Niceforo Maria]. Diagnosis.— A small species of Rhamphophryne {$32 mm, $ 33 mm) which can be distinguished from other members of the genus by the following combination of characters: (1) seven presacral vertebrae; (2) sacral vertebra not fused with coccyx, with a monocondylar articulation; (3) snout moderately short, protuberant and directed slightly anteroventrally; (4) cranial ornamen- tation evident; (5) supraorbital crest present and postorbital and supratym- panic crests absent; (6) pretympanic crest barely evident; (7) occipital crest present; (8) dorsal tubercles scattered, large, depressed; (9) dorsolateral row of enlarged, depressed tubercles extending from posterior margin of parotoid gland to a point about three-fourths the distance between the axilla and groin; (10) hands and feet with relatively short, heavy digits and reduced webbing; (11) subarticular tubercles absent; (12) supernumerary tubercles present; (13) subdigital lamellar pads present; (14) tensor fasciae latae extended; (15) vocal slits present in males; (16) cloacal sheath present. Description and variation.— Ay erage measurements of two adult female and measurements of one adult male Rhamphophryne nicefori are as follows (corresponding figures for male are included in parentheses after females): snout-vent length (S-VL) 32.9 mm (31.9 mm); tibia length 9.0 mm (10.2 mm), 29.1 (32.0) per cent of S-VL; foot length 16.1 mm (15.0 mm), 48.8 (47.0) per cent of S-VL; head length 10.1 mm (10.0 mm), 30.7 (31.3) per cent of S-VL; head width 11.4 mm (11.1 mm), 34.5 (34.8) per cent of S-VL. The snout in profile (Fig. 3d) is moderately short, directed slightly anter- oventrally, and rounded terminally and in dorsal aspect (Fig. 4d) is acumi- nate. The distance from the nostril to the tip of the snout ( $ 1.7 mm, $ 1.9 mm) is equal to or less than the distance from the nostril to the eye ( $ 2.2 30 Contributions in Science No. 216 mm, $ 1.9 mm). The tip of the snout is rounded; there is no evidence of a ventral keel connecting the tip to the upper jaw. The loreal region is concave. Nostrils are slightly protuberant and lateral; they are located at a level obvi- ously posterior to the terminus of the upper jaw and ventral to the canthal ridge. The diameter of the eye is 3.2 mm (3.3 mm), 31.7 (33.0) per cent of the head length. The pupil is horizontally elliptical. The interorbital distance is 4.0 mm (4.0 mm), 34.9 (36.0) per cent of the head width. The ear is absent. The tongue is pointed anteriorly and round posteriorly. It is one-half as wide as long, widest at two-thirds its length, and at its widest point, about one-half as wide as the mouth. The posterior one-third of the tongue is free. Choanae are small, round and lateral in position. Prevomerine teeth are absent. Vocal slits are present in males. The canthal ridges are angular in section and converge anteriorly to form the acuminate snout. Some of the cranial ridges are obvious. The supratym- panic and occipital crests are very well developed. The postorbital crest is absent. The presence of a supratympanic crest is questionable; in preserved specimens this area is obscured by an anterolateral, finger-like projection of the parotoid gland. However, a skeletal preparation might reveal the presence of a small supratympanic ridge. The arms are robust and lack folds. An axillary membrane is absent. The fingers are short and thick (Fig. 5d) ; their lengths from shortest to longest being 1 -2-4-3; the phalangeal formula of the hand is 2-2-3-3. The fingers are about one-fourth, or less, webbed. Supernumerary tubercles are present. Sub- articular tubercles are absent; instead, the digits bear lamellar, pad-like swellings. Males apparently lack prepoll ical protuberances and nuptial ex- crescences. The inner palmar tubercle is indistinct or absent in females; in males it is distinct and ovoid. The medial palmar tubercle is round and in the male, about the same size as the inner tubercle. The hind limbs are short, robust and lack folds. The toes are short and thick (Fig. 6d), their lengths from shortest to longest being 1-2-3-5-4; the phalangeal formula of the foot is 2-2-3-4-3. The first three toes are about three-fourths webbed. The fourth and fifth toes are one-third webbed. A few supernumerary tubercles are present. Subarticular tubercles are absent; the toes bear subdigital, lamellar, pad-like swellings. The inner metatarsal tubercle is ovoid, depressed, and variously indistinct or distinct depending on the specimen. The outer metatarsal tubercle, if present, is round and indistinct. It often is absent. The cloacal opening is directed posteroventrally at the mid-level of the thighs. A relatively long cloacal sheath is present. The parotoid glands are large, rounded and obvious both dorsally and laterally. An anterior, dorso- lateral extension of the parotoid gland extends from the main body of the gland to the corner of the eye. Scattered, depressed tubercles occur over the dorsum; these are densely distributed on the limbs and the flanks. Small, round tubercles cover the eyelids. A dorsolateral series of larger, depressed tubercles extends posteriorly from the parotoid gland to a point about three-fourths the 1971 Relationships of Neotropical Toads 31 distance between the axilla and the groin; occasionally, adjacent tubercles coalesce to form a long, irregularly shaped swelling. The entire skin is rather coarsely granular. In preservative, both females are uniform dark brown above and slightly lighter brown below with no visible markings. The male has a dark brown dorsal ground color, and a tan vertebral stripe; all dorsal surfaces are marked with a fine reticulate network of light tan. The ventral surfaces are light brown in color, marbled with a slighly darker brown. There is an indication of an incomplete mid-ventral stripe. The subadult female and the juvenile specimens are marked similarly to the adult male, except that the dorsal light markings are more extensive. There is no information available on the colors in life. Distribution. —Rhamphophryne nice fori is known only from the type locality at Hacienda Palmas, El Chaquiro, Departamento de Antioquia, Colombia (Fig. 10). Remarks.— Rhamphophryne nice fori is intermediate in terms of special- ization and characters shared with other members of the genus. Development of cranial crests is less obvious than in acrolopha and macrorhina, and more obvious than in rostrata and festae. The presence of relatively large, widely spaced conical tubercles and a less protuberant snout relates nice fori with rostrata. Like acrolopha, nicefori has a monocondylar sacral-coccygeal artic- ulation, but the hands and feet are most similar to macrorhina. The digits are relatively short and there is a tendency toward formation of subdigital, lamellar pads; palmar and metatarsal tubercles are much less evident in nicefori than in macrorhina. During the month of December, three females (two adults and one young), an adult male, and a juvenile were collected by day. The gravid female contains relatively few, large (1.8 mm), unpigmented eggs. Specimens examined. — {5) COLOMBIA: Antioquia: El chaquiro. Ha- cienda Palmas, 2760 m, KU 124948, MLS 449, 451-2, USNM 163476. Rhamphophryne rostrata, new combination Bufo rostratus (part) Noble, 1920:445 [Holotype. — American Museum of Natural History (AMNH) 1359 from Santa Rita Creek, 14 miles north of the village of Mesopotamia in the Department of Antioquia, Colombia; R. D. O. Johnson, collector]. Gallardo, 1962:96. Cochran and Coin, 1970:93. Diagnosis. — A large Rhamphophryne { $ 42 mm) which can be distin- guished from other members of the genus by the following combination of characters : ( 1) seven presacral vertebrae; (2) sacral vertebra fused with coccyx; (3) snout moderately long, protuberant, not directed posteroventrally; (4) cranial ornamentation scarcely evident; (5) supraorbital and postorbital crests absent, and supratympanic crest present; (6) pretympanic crest present; (7) occipital crest absent; (8) dorsal tubercles scattered, large, rounded; (9) dorsolateral row of seven large tubercles extending between posterior margin of parotoid gland and groin; ( 10) hands and feet with long, slender digits, 32 Contributions in Science No. 216 and no marked reduction of webbing; (11) subarticular tubercles present; (12) supernumerary tubercles present; (13) subdigital lamellar pads absent; ( 14) tensor fasciae latae not extended; (15) vocal slits present in males; ( 16) cloacal sheath absent. Description.— Rhamphophryne ro strata is known only from two speci- mens, the holotype, an adult male (AMNH 1359; Fig. 2), and a juvenile (AMNH 1362). Measurements of the holotype are as follows : snout-vent length (S-VL) 42.0 mm; tibia 15.3 mm, 36.4 per cent of S-VL; foot length 25.1 mm, 59.8 per cent of S-VL; head length 13.6 mm, 32.4 per cent of S-VL; head width 14.3 mm, 39.1 per cent of S-VL. The snout in dorsal aspect (Fig. 4e) is acuminate, and in profile (Fig. 3e) is moderately long, straight, and slightly rounded terminally. The distance from the nostril to the tip of the snout (2.4 mm) is noticeably less than the distance from the nostril to the eye (3.3 mm). The tip of the snout is not bulbous; a sharp ventral keel runs from the tip of the snout to the upper jaw. The loreal region is concave. Nostrils are slightly protuberant and lateral; they are located at a level anterior to the terminus of the upper jaw and ventral to the canthal ridge. The diameter of the eye is 4.6 mm, 33.8 per cent of the head length. The pupil is horizontally elliptical. The interorbital distance is 6.5 mm, 45.5 per cent of the head width. The ear is absent. The tongue is slightly less than twice as long as wide; it is widest at one-half its length, pointed anteriorly, and rounded posteriorly. The tongue is one-half as wide as the mouth and free posteriorly for one-fourth of its length. Choanae are minute, round and lateral in position. Prevomerine teeth are absent. Vocal slits are present in males. The canthal ridges are angular in section and converge anteriorly to form the acuminate snout. Cranial ridges are scarcely evident on the crown of the head. Supraorbital, occipital, and postorbital crests are absent. Supratympanic and pretympanic crests are obvious in lateral view. The arms are moderately slender and lack folds. An axillary membrane is absent. The fingers are long and slender (Fig. 5e), their lengths from shortest to longest being 1 -2-4-3; the phalangeal formula of the hand is 2-2-3-3. The first and second fingers are one-half webbed, whereas the third and fourth are only about one-fourth webbed. Supernumerary and subarticular tubercles are present. The inner palmar tubercle is elliptical, flat and indistinct, whereas the medial palmar tubercle is larger, round and distinct. The hind limbs are relatively long and lack folds. The toes are long and slender (Fig. 6e), their lengths from shortest to longest being 1-2-3-5-4; the phalangeal formula of the foot is 2-2-3-4-3. The first toe is fully webbed. Webbing extends from the ultimate phalange of the second toe to the base of the penultimate phalange of the third, from the base of the ultimate phalange of the third toe to the base of the antepenultimate phalange of the fourth, and from the base of the antepenultimate phalange of the fourth toe to the base of the penultimate phalange of the fifth toe. Supernumerary and subarticular tubercles are present. The inner metatarsal tubercle is moderate-sized, elliptical and flat, whereas the 1971 Relationships of Neotropical Toads 33 outer metatarsal tubercle is round, flat, and only about one-half the size of the inner metatarsal tubercle. The cloacal opening is directed posteriorly at the mid-level of the thighs. An cloacal sheath is absent. The parotoid glands are relatively large and distinct, especially in lateral view. The dorsal surfaces of Rhamphophryne rostrata have scattered, large tubercles which are rather densely distributed on the limbs. The eyelids are covered with many smaller, round tubercles. Dorsolaterally there is a series of seven, enlarged tubercles between the posterior margin of the parotoid gland and the groin. Groups of other large tubercles occur at the angle of the jaw laterally and ventrally. The skin of the throat, chest, belly and posterior surfaces of the thighs is slightly granular. The two specimens of Rhamphophryne rostrata are badly faded. In preservative, both are an almost uniform light brown color. The holotype appears to have had some darker brown mottling dorsally on the anterior and dorsal surfaces of the thighs, the flanks, and laterally, anterior to the forearm. There is a faint indication of darker transverse bars on the distal parts of the limbs. Ventrally, the belly, chest and throat are light brown with indistinct traces of delicate, darker brown mottling. Distribution. — Rhamphophryne rostrata is known only from the type locality 14 miles N of Mesopotamia, Departamento de Antioquia, Colombia (Fig. 10). For remarks about this locality, refer to the section on distribution in the R. macrorhina account. Remarks.— Tht relationships of Rhamphophryne rostrata are especially confusing because this species is represented by only one adult specimen, a male in a relatively poor state of preservation. Until a gravid female is available, it is impossible to ascertain the nature of the eggs. However, on the bases of the other morphological evidence, it seems reasonable to assume, at least for the present, that rostrata probably has large, unpigmented eggs. Externally, rostrata is the least specialized member of the genus. Cranial crests are absent, dorsal tubercles are scattered, hands and feet are generalized and the snout protrudes anteriorly only slightly. In contrast, internally, there are only seven presacral vertebrae and the sacrum is fused to the coccyx. Contrary to Noble’s ( 1920) description, the tympanum is absent, not hidden. Some information on the habitat of Rhamphophryne rostrata and R. macrorhina is available from a letter of Mr. R. D. O. Johnson quoted in part by Noble (1917). According to Johnson, the area of the type locality is cool (daytime temperature 58-68°F.), shaded, and moist. Ferns and mosses are common, and the forest floor is covered with a mass of decaying leaves, mosses and ferns, all held together by fine roots. This ground cover is locally termed “capote.” Specimens examined. — (2) COLOMBIA: Antioquia: Santa Rita Creek, 14 miles N Mesopotamia, 1890-2424 m, AMNH 1359, 1362 (juvenile). 34 Contributions in Science No. 216 Intrageneric Relationships Unfortunately the morphological features characterizing the species of Rhamphophryne are not especially suggestive of evolutionary trends and phylogenetic relationships within the genus. If one can assume that the genus has been derived from a more generalized bufonid stock such as the Bufo V allice ps line (Tihen, 1962), then the following evolutionary trends can be postulated. There is a tendency towards reduced ossification and reduction in the number of osteological elements. Some of these alterations might be associated with the relatively small sizes of the species and their inclination towards a walking rather than a hopping gait. The length of the limb bones are obviously shortened in comparison with those of a typical Bufo. The pectoral girdle, although arciferal, is modified slightly in the direction of firmisterny. The vertebral column tends to be shortened by a reduction in the number of presacral vertebrae. It would seem reasonable to assume that the replacement of a bicondylar sacral-coccygeal articulation with a monocondylar articulation would result in some loss of axial stability. Fusion of the sacrum and coccyx overcomes inherent stability problems at the expense of mobility and axial flexibility posteriorly. Although the three structural conditions, i.e. weak bicondyly, monocondyly and fusion seem to constitute a natural evo- lutionary sequence, monocondyly and fusion may actually represent two divergent structural adaptations. Cranial structure cannot be unequivocally categorized as being either one of reduced ossification (and therefore, by implication, “advanced” or “speci- alized”) or not markedly reduced ossification (“generalized”). It has become increasingly evident from recent studies on modern anurans (Trueb, 1970) that trends towards loss of elements and reduced ossification in some parts of a frog skull may be countered by increased ossification and/or the development of additional structural elements elsewhere in the skull. Present evidence suggests that cranial restructuring by increased ossification reflects structural adaptations imposed on a more general evolutionary trend towards reduced ossification. The cranial structure of species of Rhamphophryne has obvious marks of reduction in the loss of complete integumentary-cranial co-ossifica- tion, loss of the ear apparatus, and reduced size of the prevomers. In contrast, there have been marked alterations in the anterior end of the skull involving proliferation of the sphenethmoid bone to produce the protuberant snout characteristic of this genus. The characteristic is least developed in rostrata and nice fori, and maximally developed in festae and macrorhina. A structural alteration of this kind, involving modification of an endochondral structure, would seem to represent a single evolutionary trend. Characteristics of the soft anatomy are often more plastic and therefore more difficult to interpret than osteological features. In Rhamphophryne there is a tendency towards loss of vocal slits (the slits are absent in festae and acrolopha and present in the other species) which can be logically correlated with the absence of ears. Several trends seem to be evident in the structure 1971 Relationships of Neotropical Toads 35 of the hands and feet. There is a tendency towards marked shortening of the digits and reduction of webbing in festae, macrorhina, and especially nicefori. All of the digits are shortened in macrorhina and nicefori, whereas in festae the third finger and fourth toe remain long in comparison with the other digits. The digits are longer and more slender in rostrata and acrolopha; webbing is somewhat reduced in acrolopha, but not notably so in rostrata. Patterns of palmar and plantar tuberculation seem to be expressed in two general ways. Rhamphophryne rostrata, festae, and acrolopha have well-developed palmar, metatarsal, and subarticular and supernumerary tubercles typical of the more generalized bufonids. Rhamphophryne macrorhina and nicefori have less distinct palmar and supernumerary tubercles, and the subarticular tubercles have been replaced by subdigital, lamellarlike pads. Consideration of these trends suggests that there have been at least two evolutionary lines within the genus with respect to hand and foot morphology. That of rostrata and acrolopha seems to be more generalized and presumably representative of an ancestral type from which the other’s could have arisen. The hands and feet of festae are so obviously different from those of macrorhina and nicefori that it would seem that two separate structural adaptations have appeared among these three species. A summary of the individual characters possessed by each species would show that many of the proposed evolutionary trends overlap and contradict one another. As an example, on the bases of cranial and external morphology, rostrata seems to be the most generalized member of the genus. However, the presence of only seven presacral vertebrae and a fused coccyx and sacrum confuse the picture. Rhamphophryne macrorhina, on the other hand, appears to be quite specialized on the basis of external features; it is the only member of the genus having eight presacral vertebrae and a weak, but nonetheless bicondylar sacral-coccygeal articulation. It is obvious that if any intrageneric relationships are to be postulated on the bases of the present, limited amounts of evidence, equivalent consideration cannot be given to all characters. The most conservative characters available for interpretation are those involving the osteology. On this basis I would propose that rostrata and macrorhina most closely resemble the ancestral stock of the genus, as evidenced by the condition of the sphenethmoid in the former and the vertebral column in the latter. This hypothesis is corroborated by the generalized morphology of the hands and feet and the retention of vocal slits in these species. The generalized condition (i.e. with respect to other bufonids) of the vertebral column of macrorhina seems to suggest that it may be an early derivative of an evolutionary line which gave rise to the three remaining species by a reduction in the number of presacral vertebrae from eight to seven. Of these three species, nicefori and acrolopha seem to constitute one evolutionary line relatively closely related to macrorhina; festae represents a second, more divergent line. Rhamphophryne acrolopha and nicefori are alike in having a monocondylar sacral-coccygeal articulation and similarly modified hands and 36 Contributions in Science No. 216 feet. Of the two species, nicefori is somewhat less specialized in retaining vocal slits and lacking the protuberant snout characteristic of acrolopha. Rhampho- phryne festae is probably the most specialized and divergent member of the genus. It lacks vocal slits, has a highly modified cranial structure, seven presacral vertebrae and a fused sacrum and coccyx. The hands and feet are specialized and dissimilar from any other member of the genus. The nature of the diagnostic characters distinguishing species of Rham- phophryne from other bufonids and the restricted distribution of the genus seem indicative of its specialized position and probable recent origin within the family. The morphological similarities among the species and confusing combinations of characters suggest either that many of the characters are non-adaptive or that the animals have not had a long and uniform evolutionary history during which selective pressures could act. The small populations and isolated, principally montane, distributions of the species, as presently known, suggest that the animals have not had a long evolutionary history. It seems most reasonable to assume that they represent relict populations derived from an already specialized ancestral stock which was, at one time, more widely distributed on the Andean slopes. Discussion Species of Rhamphophryne possess a highly distinctive suite of characters making it especially difficult to assess the relationships of this genus with respect to other bufonids. Nearly all the characters are specialized and/or advanced, with the possible exception of the retention of certain cranial elements such as the palatine and quadratojugal. The proliferation of endo- chrondral elements to form the protuberant snout characteristic of Rham- phophryne is probably a secondary modification brought about through an increase in cranial ossification. If this is true, then the retention of cranial supportive elements may be a logical corollary to the need for more support and thus an advanced trait rather than an unspecialized character. Currently there are some 19 genera loosely referred to the family Bufonidae (including Atelopodidae, auctorum). These include five arciferal- firmisternal genera as follows: Atelopus, Dendrophryniscus, and Melanophry- niscus of the New World, and Cacophryne and Didynamipus of the Old World. The following fully arciferal genera are referred to the Bufonidae: Oreophry- nella, Crepidophryne, and Rhamphophryne of the Neotropics, and Ansonia, Laurentophryne, Mertensophryne, Nectophrynoides, Nectophryne, Pedostibes, Pelophryne, Pseudobufo, Werneria, and Wolterstorffina of the Old World. Rhamphophryne shares some striking features with several Old World bufonid genera having arciferal pectoral girdles. Rhamphophryne, Laurento- phryne, Mertensophryne, Nectophryne, and Pelophryne are alike in having a reduced number of presacral vertebrae. In addition to Rhamphophryne, a second New World genus, Oreophrynella, has a reduced number (five) of presacral vertebrae. This type of reduction has come about from a forward 1971 Relationships of Neotropical Toads 37 shift of the sacral articulation, according to Tihen (1960) which is corrobo- rated by the fact that in all of the genera where there is a reduced number of presacral vertebrae, the coccyx is either fused with the sacrum or has a monocondylar or weakly bicondylar articulation with it. There are two notable exceptions to the occurrence of this character sequence. One African genus, W olterstorffina, has a fused coccyx and sacrum but shows no reduction in vertebral number. Similarly, a single species of Rhamphophryne, R. macrorhina, has eight presacral vertebrae but a very weak bicondylar articu- lation between the coccyx and sacrum. In addition to these osteological characters, the occurrence of subdigital lamellar pads is known only in Rhamphophryne and the highly specialized African genus Nectophryne. Several characteristics of Rhamphophryne are common among arciferal- firmisternal and arciferal genera of both the old and new worlds. The pro- duction of relatively few, large, unpigmented eggs occurs in Rhamphophryne, Atelopus, Laurentophryne, Nectophryne, and Pelophryne. The Neotropical genus Crepidophryne has an intermediate condition; the eggs are unpigmented, but small and numerous. The tensor fasciae latae muscles are extended in Rhamphophryne, Atelopus, Cacophryne, Ansonia, Laurentophryne, Merten- sophryne, Nectophryne, Pelophryne, Pseudobufo, and Wolterstorffina. The loss of the adductor longus muscle of the thigh is widespread; this muscle is known to occur only in Melanophryniscus, Bufo, and Cacophryne. The loss of the columella is common among the bufonids of the New World; this element is retained in most species of Bufo. In contrast, many Old World groups have retained the columella, the notable exceptions being Laurento- phryne, Mertensophryne , Nectophryne, Werneria, and Wolterstorffina which have lost this element. To summarize the information presented above, Rhamphophryne seems to be most similar to Atelopus and Oreophrynella in the New World and the small group of highly specialized African genera consisting of Laurentophryne, Mertensophryne , and Nectophryne. Rhamphophryne is distinguished from both Atelopus and Oreophrynella by the absence of fusion of the cervical and second presacral vertebrae. Furthermore, the structure of the pectoral girdle varies among these genera. The girdle is arciferal in Rhamphophryne; however, the structure differs from that typical of Bufo in that the epicoracoids barely overlap. Oreophrynella is further removed from classical arcifery by the broad precoracoid bridge and shallow overlap of the epicoracoids. In Atelopus, broad fusion of the epicoracoids results in a condition of functional firmistery. Super- ficially, Rhamphophryne seems to be more similar to the African genera; however, there are differences which should be noted. Of the four arciferal African genera mentioned, Nectophryne, and Laurentophryne are character- ized by having the tips of the phalanges expanded. The latter character is unknown among New World bufonids, but quite common among the Old World genera. Rhamphophryne is most similar to Mertensophryne of the Old World. In view of these data, at least two alternative hypotheses concerning 38 Contributions in Science No. 216 the relationships of Rhamphophryne are available, as follows: (1) Rhampho- phryne is geographically isolated, but phylogenetically close to Mertensophryne and its African allies; or (2) Rhamphophryne, Oreophrynella, and Atelopus constitute a phylogenetic New World group which has evolved in a fashion closely paralleling that of an Old World line composed of Laurentophryne, Nectophryne, Pelophryne, and Mertensophryne. Both of the above hypotheses are amenable to Tihen’s (1960) postulate that two principal lines were established very early in the history of the genus Bufo. The two lines are designated as the “regularis” and “angusticeps” complexes; the former is cosmopolitan, whereas the latter occurs in the Old World. The distinctions between these two lines rests on osteological criteria. Fusion of the frontoparietal and prootic is universal in the “angusticeps” complex and occurs in only one or two groups in the “regularis” complex. Dermal ornamentation and the presence of an enclosed canal for the occipital artery are characteristic of the latter, whereas the former lacks ornamentation and has an open canal through which the occipital artery runs. Unfortunately, Rhamphophryne is not clearly referable to either one line or the other. Dermal ornamentation is present in some species and absent in others; the occipital canal varies from open to completely enclosed. The frontoparietal and prootic are fused, but this is a character common to either complex. There are at least three available hypotheses to explain the present distri- bution of the bufonids on all continents except Australia and Antarctica. Classically and according to Matthewsian concepts a bufonid prototype originated in northern parts of the Old World. Subsequent differentiation and dispersal resulted in the distribution of the family in southern Asia, Africa, and via the Bering Strait land bridge into North America and thence into South America. The absence of early fossil remains in the Northern Hemi- sphere and the presence of the greatest numbers and diversity, as well as the most primitive forms in the Southern Hemisphere, throw some doubt on the credibility of this hypothesis. More recently, it has been suggested (Blair, 1969) that South America may have been the site of bufonid origins. He indicated that bufonids may have dispersed northward from South America and reached the Old World by a northern land route. This hypothesis does not satisfactorily account for the presence of a primitive bufonid genus in Africa (Tihen, 1960). Lynch (1971) hypothesized that the Bufonidae origi- nated in Africa and South America at a time when the two continents were connected. Evidence is insufficient to warrant relying on any one of these theories without reservation. However, in favor of the last of these hypotheses, there is an increasing amount of evidence, not the least of which is the accumu- lation of data confirming the occurrence of continental drift. Moreover, Lynch ( 1971) provided a convincing discussion in which he postulated a southern origin, with subsequent equability-controlled dispersal patterns for the frog family Leptodactylidae, from which the bufonids may have arisen. Within the New World “regularis” complex there are only two groups 1971 Relationships of Neotropical Toads 39 (as described by Tihen, 1962) which are characterized by fusion of the prootic and frontoparietal. These are the valliceps and spinulosus groups; Tihen has suggested that the latter may have been derived from the former. There is a tendency for reduction in the size of the frontoparietals, loss of cranial crests, deletion of the enclosed canal for the occipital artery, and depression of the skull in the spinulosus group in contrast to the valliceps group. Rhampophryne has the same tendency, as Bufo spinulosus in the reduction of the frontoparietal, but Rhamphophryne is variable with respect to both groups in the conditions of the cranial crests and the occipital canal. Furthermore, the skull of Rhamphophryne is elevated (Table 1) and characterized by the modifications of the sphenethmoid and the anterior cartilages. Comparison of these few characters indicates that Rhamphophryne cannot be directly related to the spinulosus group. Although they share some generalized evolutionary trends, they are clearly divergent with respect to a host of other characters. The evidence is especially tenuous, but it is possible that both the spinulosus group and Rhamphophryne may have been derived from an ancestral valliceps stock. In this event, Rhamphophryne must be viewed as an extremely divergent, specialized, and probably much earlier derivative of the ancestral stock than the spinulosus group. Acknowledgements Dr. Robert F. Inger (Field Museum of Natural History, FMNH), Dr. Alan E. Leviton (California Academy of Sciences, CAS), Dr. James A. Peters (United States National Museum, USNM), Dr. Charles W. Myers (American Museum of Natural History, AMNH), Prof. U. Parenti (Museo ed Instituto di Zoologia Sistematica della Universita di Torino, IZS), Dr. John W. Wright and Mr. Arden Brame (Los Angeles County Museum of Natural History, LACM), Hno. Niceforo Maria (Museo de la Salle, MLS), and Dr. William E. Duellman (University of Kansas Museum of Natural History, KU) provided the specimens and data on which this study is based. I am grateful to each of the above for his co-operation and also to Dr. Coleman J. Coin, Mr. P. A. Silverstone, and Mr. Stephen R. Edwards who furnished essential information. Particular thanks are extended to Dr. Joseph Tihen who critically reviewed the manuscript and to Dr. John D. Lynch for his technological aid and provocative observations, both of which figured significantly in this work. This study was supported by a grant (GB-8139) from the National Science Foundation, William E. Duellman, principal investigator. Resumen Un nuevo genero de bufonidos, Rhamphophryne, se describe. Este genero contiene cinco especies (Atelopus festate Peracca, Bufo rostratus Noble, Bufo nicefori Cochran and Coin, y dos especies nuevas— R. acrolopha y R. macro- rhina), y se extiende desde el extreme este de Panama hasta el sur de Ecuador. El genero es caracterizado por un numero reducido de vertebras presacrales. 40 Contributions in Science No. 216 un hocico protuberante, manos y patas altamente modificadas, y algunos hue- vos grandes sin pigmentacion. Ademas de estos caracteres, la condicion de la cintura pectoral y la osteologia del craneo, parecen relacionar a Rhampho- phryne mas cercanamente a Oreophrynella y AteJopus en el Nuevo Mundo. La evidencia morfologica y zoogeografica sugiere que Rhamphophryne puede haberse derivado de una linea ancestral similar a Bufo valliceps. Rhampho- phryne comparte muchos caracteres especializados, con bufonidos africanos, como por ejemplo Nectophryne, Mertensophryne y Laurentophryne. Se sugiere que estas semejanzas son el resultado de convergencia mas que de una relacion filogenetica estrecha. Literature Cited Blair, W. F. 1969. Influencia de la cuencia [sic] del Rio Amazonas sobre la evolu- cion de los sapos del genero Bufo. [abstract] II Simposio Biol. Trop. Ama- zonica, 2 p. Cochran, D. L. and C. J. Coin. 1970. Frogs of Colombia. Bull. LF.S. Nat. Mus. 288, l-xii+655 p. Gallardo, Jose M. 1962. A proposito de Bufo variegatus (Gunther), sapo del bosque humedo antartantico, y otras especies de Bufo Neotropicales. Physis 23(64) :93-102. Lynch, J. D. 1971. Evolutionary relationships, osteology and zoogeography of leptodactyloid frogs. Univ. Kans. Mus. Nat. Hist., Misc. Publ. 53:1-238. Noble, G. K. 1917. The systematic status of some batrachians from South America. Bull. Amer. Mus. Nat. Hist. 37(30) :793-814. 1920. Two new batrachians from Colombia. Bull. Amer. Mus. Nat. Hist. 42(9):441-446. Peracca, M. G. 1904. Viaggio del Dr. Enrico Festa nell’Ecuador e regioni vicine. Reptile ed Amfibii. Bolletino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino 19(465) : 1-41. Peters, J. A. 1955. Herpetological type localities in Ecuador. Rev. Ecuatoriana Ent. Par. 2(3-4): 335-352. Tihen, j. a. 1960. Two new genera of African bufonids, with remarks on the phyto- geny of related genera. Copeia (3) :225-233. 1962. Osteological observations on New World Bufo. American Midi. Nat. 67(1): 157-183. Truer, L. 1970. Evolutionary relationships of casque-headed tree frogs with co- ossified skulls (Family Hylidae). Univ. Kans. Publ. Mus. Nat. Hist. 18(7): 547-716. Accepted for publication November 23, 1970 1 LOS ANGELES CONTRIBUTIONS COUNTY 1 MUSEUM IN SCIENCE JMBER 217 November 12, 1971 ; son. 13 1 c a PLIOCENE AVIAN REMAINS FROM BAJA CALIFORNIA By Hildegarde Howard Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. Issues are numbered sep- arately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. 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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 PLIOCENE AVIAN REMAINS FROM BAJA CALIFORNIA By Hildegarde Howard^ Abstract: Seven species are represented in an Early Plio- cene avifauna from marine deposits in three canyons in the southeastern part of Cedros Island. Predominant is a new species of flightless auk, Mancalla cedrosensis, represented by four in- complete skeletons as well as disassociated elements. Other new species described are Puffinus tedfordi and Cerorhinca minor. The primitive flamingo, Megapaloelodus opsigonus Brodkorb, is tentatively identified. A shearwater, gannet, and murrelet (listed by genus only) complete the avifauna. Introduction In 1964 and 1965 the University of California, Riverside, in cooperation with the University of Baja California, Ensenada, collected vertebrate fossils on Cedros Island. Through the kindness of Dr. Richard Tedford, who headed the field work, the bird remains were presented to the Los Angeles County Museum of Natural History. Ledford’s notes accompanying the collection indicate that the fossil- bearing marine deposits involved are “in the basal part of the northwesterly dipping Pliocene section as it crops out in deep canyons and ridges near the base of the escarpment which forms the high northern wall of Valle Blanca.” This prominent valley, he states, runs east and west, draining much of the southeast corner of Cedros Island, with its mouth opening at the sea about two miles south of the “cannery village.” The fossils were collected from marine sandstones that overlie unconformably the Tortuga Formation of Late Miocene (Temblor) age, and because of this relationship, they have been referred to the Almejas Formation of Mina (1957), whose type section occurs at Tortuga Bay on the nearby Viscaino Penninsula. Dr. Frank Kilmer, of the Geology Department of Humboldt State Col- lege, California, who studied the invertebrate beds on Cedros Island, believes (personal communication) that the vertebrate-bearing sands may be Early to Middle Pliocene in age. Although these beds are so far not known to contain invertebrates, he notes that they rest on diatomaceous strata of probable Upper Middle Miocene (Luisian) age, and underlie coarsely clastic invertebrate- bearing beds of probable mid-Pliocene age. With respect to the Miocene-Pliocene contact, most of the bird remains occurred in the lower 10-70 feet of the Pliocene deposits. They were taken from approximately 22 field collecting sites over a half-mile of strike in three canyons. From west to east the canyons were named by the field party: ^Research Associate in Vertebrate Paleontology, Los Angeles County Museum of Natural History. 1 2 Contributions in Science No. 217 Arroyo Esqueletos, Arroyo Tiberon, and Arroyo Delphin. Los Angeles County Museum of Natural History (LACM) locality numbers have been assigned to the collecting sites within these canyons based, in general, on their depth within the section. These are as follows (from lowest to highest within each canyon or ridge, and including from one to five field numbers each) : Arroyo Esqueletos, loc. nos. 65144, 65145, 65146, and 65147; Arroyo Tiberon, loc. nos. 65152, 65151, 65150; Arroyo Delphin, loc. nos. 65148, 65149; ridge between Arroyo Tiberon and Arroyo Delphin, loc. nos. 65154 and 65153. Avifauna Predominant among the approximately 100 avian bones found are those of a heretofore undescribed species of the flightless auk, Mancalla. Other spe- cies (all extinct) represented in the collection by fragmentary elements are two shearwaters, a gannet, a flamingoid species of the extinct family Palaelodi- dae, an auklet, and a murrelet. The taxa represented are as follows : Puffinus tedfordi, n. sp. Puffinus sp. Moms sp. ?Megapaloelodus opsigonus Brodkorb Cerorhinca minor, n. sp. Endomychura (?) sp. Mancalla cedrosensis, n, sp. Analysis of Species Order Procellariiformes Family Procellariidae-Shearwaters Puffinus tedfordi, new species Figure 1, A, B, E, F Holotype: Proximal end of left tarsometatarsus, LACM 15386. Collected by Whistler and Tedford, August, 1965. Locality: LACM locality 65151, Arroyo Tiberon, approximately 70 feet above Miocene-Pliocene contact; north side of Valle Blanca, IV2 miles south of cannery village, southeast corner of Cedros Island, Baja California, Mexico. Formation and Age: Almejas Formation; Early Pliocene. Diagnosis: Resembling Puffinus opisthomelas in gradual merging of inter- nal calcaneal ridge with internal edge of shaft, but shaft broader, and proxi- mal end relatively narrower and more evenly flared; anteriorly both edges of Figure 1. A, B, E, F, Puffinus tedfordi, n. sp. tarsometatarsi; A, F, paratype; B, E, type. C, Puffinus sp. D, G, H, J, Cerorhinca minor, n. sp.; D, tarsometatarsus 15407; G, ulna 15406; H, coracoid 15421; J, type humerus. I, ^Endomychura sp., cranium 15426. K, M, N, IMegapaloelodus opsigonus tarsometatarsus 15423, distal, internal and anterior views. L, Mancalla cedrosensis. n. sp., tarsometatarsus 15425. Figs. A, B, D, E, F, G, H, I, J, approx. X 1% ; figs. C, K, L, M, N, approx. X 1. 1971 Pliocene Avian Remains 3 i •i 4 Contributions in Science No. 217 shaft forming well-defined ridges, with external edge more extended anteriorly; internal face of shaft sloping laterally so as to be plainly visible in anterior view in proximal half of bone. Paratype: Incomplete tarsometatarsus LACM 15387; same data as for holotype. Although both proximal and distal ends are incomplete in this specimen, the contact of the internal calcaneal ridge with the shaft and the anterior shaft ridges are preserved and conform in character with the holo- type. The paratype is important in providing a measurement of length between the distal and proximal foramina, showing the specimen to be within the range of 13 tarsometatarsi of P. opisthomelas in this dimension, but 28 per cent broader of shaft than the maximum for that species. Without this dimension of length, the stoutness of the shaft would suggest considerably greater length of the element.. Measurements: Holotype, breadth of proximal end 6.8 mm; least breadth of shaft 3.3 mm; greatest depth of external side of shaft 4.0 mm; depth of internal side of shaft below internal calcaneal ridge 2.7 mm. Paratype, length between proximal and distal foramina 32.2 mm; breadth of shaft 3.0 mm; depth of external side of shaft 3.9 mm; depth of internal side of shaft 2.7 mm. Comparison with Other Fossil Shearwaters: Only two other Pliocene shearwaters are known, both from California, Puffinus kanakoffi from San Diego and P. felthami from Corona del Mar. Tarsometatarsi of both are at hand. P. tedfordi resembles P. kanakoffi in the merging of the internal cal- caneal ridge with the postero-internal edge of the shaft and approximates this species in length of the tarsometatarsus. It differs, however, in 20 per cent broader shaft, more evenly developed proximal end, and visibility of the inter- nal side of the shaft in anterior view. P. felthami differs from P. tedfordi in having the internal calcaneal ridge slightly inset mediad from the internal edge of the shaft and the posterior surface of the shaft adjacent to the ridge swollen, not flat as in P. tedfordi. P. felthami is broader proximally than P. tedfordi, the shaft is deeper but narrower; no measurement of length is possible. Three Miocene west-coast shearwaters in which the tarsometatarsus is identified are at hand {P. diatomicus, cast; and P. calhouni and P. priscus, original material). All have the calcaneal ridge inset and the swollen shaft adjacent as in P. felthami. To judge from the figured type tarsometatarsus of P. arvernensis from the Miocene of France (Shufeldt, 1896, pi. 24, figs. 1, 2) this specimen is shorter and stockier even than in P. tedfordi, but the external side of the shaft is not as deep and the proximal end is more flared. Other Miocene shearwaters are known only from the humerus. Using P. opisthomelas and P. griseus as points of reference for size, P. micraulax from Florida (as recorded by Brodkorb, 1966:161) would have been markedly smaller than P. tedfordi, whereas P. mitchelli and P. inceptor (both studied from casts) from California, and P. conradi (cast) from Maryland would have been larger. Lambrecht’s ( 1933:273) reference to P. aquitanicus and P. antiquus from the Miocene of France suggests that these species also were larger than P. tedfordi. 1971 Pliocene Avian Remains 5 Puffinus sp. Figure 1, C A distal fragment of a humerus (LACM 15404) and a nearly complete ulna (LACM 15403) from the ridge between Arroyo Tiberon and Arroyo Delphin (LACM locality 65153) represent a larger species of shearwater than P. tedfordi. Except for the fact that the humerus shows the marked lateral compression of the shaft characteristic of the subgenus Puffinus, no reliable diagnostic characters remain on this fragment. The few measurements that can be made in comparison with the type humerus of the California Pliocene P. felthami indicate close similarity in size. The fragment is too poor, how- ever, to assign or describe. The ulna is well preserved, with only a small section of the shaft incom- plete. The proximal end is narrow relative to its depth through the olecranon as in members of the subgenus Puffinus. The brachial impression is long (13.5 mm) and deeply depressed adjacent to the prominence for the ante- rior articular ligament. The bone is slightly more slender than the ulna of P. griseus and markedly shorter, being about midway between this species and P. kanakoff. As the ulna is not known in most of the previously described fossil shearwaters, it would be unwise to attempt assignment of the Cedros Island bone. Measurements of ulna LACM 15403: Greatest length (with possibly a section of shaft missing) 86.7 mm; breadth across proximal cotylae 7.8 mm; depth from cotylae through olecranon 9.2 mm; breadth of distal trochlea 5.1 mm; depth of external crest of trochlea 6.9 mm; dimensions of middle of shaft 4.1 X 4.8 mm. A very broad distal half of a tibiotarsus (LACM 15378) from Arroyo Tiberon (LACM locality 65151) exceeds even P. griseus in shaft breadth. It is doubtful that it is allied with the wing bones here discussed. Order Pelecaniformes Family Sulidae-Boobies and Gannets Morus sp. A distal end of a left humerus (LACM 15413) from Arroyo Delphin (LACM locality 65148) is crushed and broken in several important diag- nostic areas. However, the long attachment of the anterior articular ligament and the positions of the attachments on the entepicondyle and ectepicondyle suggest alignment with the gannets (genus Morus) rather than with the boobies (genus Sula). The olecranal fossa, also, is seemingly less depressed than in Sula although the broken borders of the fossa may create a false visual impression of the area. One portion of the specimen, fortunately preserved, is the well-developed entepicondylar area, which serves immediately to distinguish it from Sula humeralis from the San Diego Pliocene. S. humeralis is similar in size to the 6 Contributions in Science No. 217 Cedros Island bone, but is described (Miller and Bowman, 1958:10) as being underdeveloped in the entepicondylar area. Size alone separates the Cedros bone from west-coast Tertiary members of the genus Morns (larger than Morns vagabnndns and smaller than Morns lompocanns) . However, it closely approximates in breadth a humerus from the east coast referred to Morns loxostyla (Wetmore, 1926:467), a species known only from the Miocene. Assignment of the imperfect Cedros bone is inadvisable. Measnrements: Breadth of distal end 20.5 mm; length of attachment of anterior articular ligament 9.8 mm. Order Ciconiiformes: Suborder Phoenicopteri-Flamingos Family Palaelodidae-Primitive Flamingos IMegapaloelodns opsigonns Brodkorb Figure 1, K, M, N A distal end of a left tarsometatarsus (LACM 15423) and a fragment of the distal end of a tibiotarsus (LACM 15422), both from Arroyo Delphin locality 65148, 10 feet above the Miocene/ Pliocene contact, are assigned to the family Palaelodidae. In the tarsometatarsus the palaelodid characters are seen in the shape and position of the internal and external trochleae as com- pared with the tarsometatarsus of Phoenicopterns, as follows: (1) internal trochlea arising from the shaft well in front of the posterior margin of the middle trochlea and rounding upward at the tip; (2) external trochlea also rounding upward posteriorly and its external lip more elevated than its inter- nal lip. The contours of the tibiotarsus are badly worn, but the specimen is referred to the Palaelodidae on the basis of the following characters : ( 1 ) distal articular surface with low median ridge, and deep, pit-like depression on each side posterior to the condyles; (2) small, lateral papilla present above the external condyle anteriorly, but no large anterior intercondylar tubercle as in Phoenicopterns. The family Palaelodidae is best known from the several species of the genus Palaelodns recorded from the Miocene of France and Germany. Previ- ous records in North America are assigned to the genus Megapaloelodns, with two species: M. connectens Miller (genotype) and M. opsigonns Brodkorb. The former was described (A. Miller, 1944:86) on the basis of a large distal end of tarsometatarsus from the Miocene of South Dakota. Later, L. Miller (1950:70) referred a distal end of a tibiotarsus from the Miocene of Cali- fornia to the same species. M. opsigonns was described (Brodkorb, 1961:173) from a proximal end of a tarsometatarsus found in lower Pliocene deposits in Oregon. Characters of the Cedros Island tarsometatarsus are as follows: internal trochlea well elevated above middle trochlea and posteriorly rotated toward the center of the shaft; proximo-distal dimension (height) 90 per cent of its antero-posterior depth; internal surface flat and bordered posteriorly by a 1971 Pliocene Avian Remains 7 long, heavy ridge, deflected away from the shaft. External trochlea laterally compressed; proximo-distal dimension 89 per cent of its antero-posterior depth; external surface with deep depression near posterior tip. The specimen was compared with casts of the comparable portion of the tarsometatarsus of Palaelodus ambiguus and Megapaloelodus connectens, the types of their respective genera. Resemblance is closer to P. ambiguus in the elevation of the internal trochlea and the lateral compression of the external trochlea, but closer to Megapaloelodus in the relative height to depth of the internal and external condyles. The internal surface of the internal trochlea is flatter than in either of the other forms, but is furthest removed from P. ambiguus, in which this surface is deeply depressed. The degree of deflection of the pos- terior tip of the internal trochlea away from the shaft is, perhaps, closer to the condition in Palaelodus than in Megapaloelodus although this area in M. connectens is incomplete. Miller, however (1944:86), notes less abrupt deflection of this area as one of the characters distinguishing M. connectens from Palaelodus. The other character that he emphasizes concerns more pro- nounced development of the outer rim of the external trochlea. The angular measurement that he gives (formed with the axis of the shaft by a line from the inner to the outer rim of this trochlea) I have been unable to duplicate. Possibly, however, the greater inflation of this trochlea as compared with that of the Cedros specimen is a reflection of this character. In breadth of the distal end, taken at right angles to the axis of the shaft, the Cedros tarsometatarsus is 10 per cent narrower than that of M. connec- tens, and 6 per cent broader than that of the largest species of Palaelodus {P. goliath). Compared with the proximal end of the tarsometatarsus that forms the type of Megapaloelodus opsigonus, the Cedros bone is approxi- mately the same size. Both specimens measure 19.6 mm in breadth of their respective articular ends. There is some variation in relative breadth of articu- lar ends of this element as noted in Palaelodus and Phoenicopterus. Milne- Edwards (1867-1871, 2:78) records the proximal ends of P. gracilipes and P. ambiguus as 2 mm broader than the distal, but shows the proximal end of P. crassipes to be 0.8 mm narrower than its distal end. In two modern tarsometatarsi of Phoenicopterus ruber the proximal ends are narrower than the distal; in another P. ruber and in a specimen of P. antiquus both ends are the same breadth. The agreement in size between the Cedros tarsometatarsus and that of M. opsigonus, coupled with the fact that both come from deposits of early Pliocene age, strongly suggests that they are representative of the same species. The Cedros tarsometatarsus, therefore, is tentatively referred to M. opsigonus. The fact that the Cedros bone does not appear to agree with Megapaloelodus in the two characters particularly emphasized in Miller’s (1944:86) description of M. connectens is not considered a deterrent in view of the overall comparison with both P. ambiguus and M. connectens. Milne- Edwards notes considerable variability among the several species of Palaelo- 8 Contributions in Science No. 217 dus, and similar variability may have existed in Megapaloelodus. Furthermore, there is also a possibility that Megapaloelodus itself is but another specific variant of Palaelodus. The fragmentary distal portion of tibiotarsus from Cedros Island is com- mensurate in size with the tarsometatarsus, and is, therefore, also tentatively assigned to P. opsigonus. The only character of any possible importance below the family designation in this poorly preserved specimen is the large oval opening at the distal margin of the supratendinal bridge. The tibiotarsus re- ferred to M. connectens is said to have a round, rather than an oval opening (L. Miller, 1950:70). Measurements of tarsometatarsi of Megapaloelodus and Palaelodus Cedros Id. LACM 15423 M. connectens P. ambiguus P. goliath Breadth of distal end at right angles to axis of shaft 19.6 mm 21.7 mm 12.0 mm 18.5 mm Height of internal trochlea 12.8 12.2 6.8 _ Depth of internal trochlea 14.2 14.2 8.9 _ Height of external trochlea 12.0 12.1 6.0 _ Depth of external trochlea 13.5 14.0 8.7 Order Charadriiformes : Suborder Alcae Family Alcidae, Subfamily Alcinae-Auks, etc. Of eight small alcid bones, six show relationship to the group which includes the Rhinoceros Auklet and the Puffins. The three genera involved (Cerorhinca, Lunda, and Fratercula) so closely resemble each other in skeletal characters that the term “puffin” might properly be applied to all three. Similar characters are: humerus with long, hooked tip of internal tuberosity, deeply undercut head, and medially expanded pectoral attachment proximally; ulna with abrupt ffare from shaft to internal cotyla, narrow notch between olecranon and tricipital attachment (viewed proximally), area anconally bordering attachment of anterior articular ligament uneven in profile as viewed from anconal side (more expanded proximally than distally), distally, carpal tuberosity long and prominently projected laterally; coracoid with long procoracoid and foramen very low; tarsometatarsus with broad, flat shaft. It is only by weighing a number of small details that the fossils are assigned to Cerorhinca. All elements are 25-30 per cent smaller than those of Cero- rhinca monocerata, and the fossil species is described as new. 1971 Pliocene Avian Remains 9 Cerorhinca minor, new species Figure 1, D, G, H, J Holotype: Proximal end of right humerus, LACM 15408, collected by Whistler and Jefferson, July, 1964. Locality: LACM locality 65153, ridge between Arroyo Tiberon and Arroyo Delphin; 45-60 feet above Miocene-Pliocene contact; north side of Valle Blanca, Wi miles south of cannery village; southeast corner of Cedros Island, Baja California, Mexico. Formation and Age: Almejas Formation; Early Pliocene. Diagnosis: Head smoothly rounded on lower margin; deep channel undercutting head and proximo-medial edge of pectoral attachment; internal edge of channel limiting medial extent of capital groove and aligned with median crest; pneumatic fossa more round than oval; approximately three- fourths the size of C. monocerata humerus. Referred Specimens: Proximal half of ulna, LACM 15406 and incom- plete tarsometatarsus, LACM 15407 from the type locality; incomplete right proximal end of humerus, LACM 15420 and upper end of coracoid, LACM 15421, from Arroyo Delphin (LACM locality 65148). Tentatively referred distal half of ulna, LACM 26572 from Arroyo Tiberon (LACM locality 65151). Description of Referred Material: Humerus LACM 15420 is broken but resembles the type in all areas preserved. Proximal end of ulna with shorter, straighter, more blunt olecranon than in C. monocerata, attachment of anterior articular ligament shorter and broader, and internal side of shaft more rounded, resulting in less bladelike shaft; 72 per cent of size of C. monocerata in breadth of proximal end. Tentatively assigned distal half of ulna (LACM 26572) larger than proximal half in shaft breadth and approxi- mately 82 per cent of the size of C. monocerata in depth through the external crest of the trochlea. Coracoid with straight upper border of long procoracoid as in Cerorhinca {Lunda and Fratercula with upcurved tip); triosseal canal broadly rounded (lacking keellike raised area slightly anterior of center found in Lunda and Fratercula) ; three-fourths the size of C. monocerata in breadth and further distinguished by less expanded head, less undercut by triosseal canal; the foramen was evidently low on the procoracoid as at least half the procoracoid is preserved and the foramen is not in evidence. Tarsometatarsus broken above the tibialis anticus tubercles and through the upper portion of the distal trochleae; shaft broad and flat and well depressed on anterior surface; resembling Cerorhinca in distinction to Lunda and Fratercula in less internal flare of trochlear region and presence of small, slitlike depression separating base of internal trochlea from middle trochlea; approximately three- fourths the size of C. monocerata in breadth of shaft at upper level of distal foramen. Measurements: Type humerus, breadth proximal end 10.5 mm; dimen- sions of shaft 3.0 X 4.2 mm. Ulna LACM 15406, breadth across proximal 10 Contributions in Science No. 217 cotylae 5.0 mm; depth from internal cotyla to olecranon 5.5 mm; dimensions of shaft 2.3 X 3.3 mm. Ulna LACM 26572, depth external crest of distal trochlea 5.4 mm; dimensions of shaft 2.8 x 3.8 mm. Coracoid LACM 15421, length from below scapular facet to head 9.6 mm; breadth through triosseal canal 2.9 mm; breadth of head 3.9 mm. Tarsometatarsus LACM 15407, least breadth of shaft 2.7 mm; depth of shaft 1.9 mm; breadth of shaft at upper level of distal foramen 4.2 mm; length from distal foramen to top of tibialis anticus tubercles 12.8 mm. Comparison with previously recorded fossil Cerorhinca: Cerorhinca dubia from the Miocene of Lompoc, California is known only from the leg bones. The tarsometatarsus (as recorded by Miller, 1925:116, and as examined on a cast of the type) is only 2 per cent shorter than that of C. monocerata, hence notably larger than that of C. minor. The only other Tertiary record of Cerorhinca is from the late Miocene of Laguna Hills, Cali- fornia. A proximal end of an ulna and a fragment of a humeral shaft were assigned only as Cerorhinca sp. (Howard, 1968:16). The ulna (LACM 18274) is 30 per cent larger than in C. minor. lEndomychura sp.-Murrelet Figure 1, I The two remaining small alcid bones consist of a dorsal cap of a cranium (LACM 15426) from Arroyo Esqueletos (LACM locality 65144) and a proximal fragment of a humerus (LACM 26571) from Arroyo Tiberon (LACM locality 65151). Both are commensurate in size with the elements assigned to C. minor. However, they show closer relationship to the mur- relets. On the cranium, the depressions above the orbits form evenly rounded, short arches, and the frontal area is well vaulted adjacent to them. This con- dition appears in both Endomychura and Synthliboramphus. The humerus is closer to that of Endomychura in the very shallow depression below the head (even shallower than in either E. craverii or E. hypoleuca) . Neither specimen is sufficiently complete to permit of definite assignment. Subfamily Mancallinae-Flightless “Auks” Bones of the flightless auk, Mancalla, occurred at all LACM localities, and from approximately 5 feet (one bone) to approximately 120 feet (one bone) above the Miocene-Pliocene contact, with greatest concentration between 10-70 feet. Four incomplete skeletons and disassociated elements are represented, making a total of more than 80 bones. Previously described species of this genus (see Howard, 1970, for review) have come from the Pliocene of California: Mancalla californiensis, M. diegense, and M. milleri. A possible ancestral form, Praemancalla lagunensis, is recorded from the late Miocene of California. The bones from Cedros Island are close in size to those of M. diegense, but structural peculiarities 1971 Pliocene Avian Remains 11 serve to distinguish most of the elements. The Mexican form is, therefore, described as a distinct species. Mancalla cedrosensis, new species Figure 2, A-K and Figure 1, L Holotype: Associated skeletal elements of one individual, LACM 15373, including essentially complete right scapula, coracoid, humerus, ulna, femur, tibiotarsus, left ulna, radius, and carpometacarpus; also including distal ends of right radius, right carpometacarpus, and left tibiotarsus, extreme tip of proximal end of right tarsometatarsus, proximal three-fourths of left femur, and several fragmentary thoracic and cervical vertebrae. Collected by R. Tedford, 1965. Locality: LACM locality 65147 (Tedford field no. 1296), east branch of Arroyo Esqueletos, approximately 70 feet above Miocene/ Pliocene con- tact; 2 miles south of cannery village, southeast corner of Cedros Island, Baja California, Mexico. Formation and Age: Almejas Formation, Early Pliocene. Referred Material: Three associated skeletons: LACM 15425 from LACM locality 65144 (Arroyo Esqueletos, approximately 25 feet above Miocene/ Pliocene contact) including proximal ends of left femur and tibio- tarsus, distal end of right tibiotarsus, complete right tarsometatarsus, fragment of pelvis and vertebrae; LACM 23739, also from LACM locality 65144, including right and left femora, left tibiotarsus (distal end), and complete left tarsometatarsus; and LACM 15410 from LACM locality 65150 (Arroyo Tiberon, approximately 70 feet above Miocene/ Pliocene contact) including right and left scapulae and coracoids, furcula, crushed sternum, and a car- pometacarpus. Also unassociated elements from all localities, consisting of a rostrum, 2 scapulae, 1 coracoid, 10 humeri, 14 ulnae, 3 radii, 5 carpometa- carpi, 1 femur, 4 tibiotarsi, 6 tarsometatarsi, and 3 vertebrae. Diagnosis: Coracoid relatively short and heavy (ratio of breadth below furcular facet to length more than 1 2 per cent, ratio of length of coracoid to length of humerus 59 per cent) ; tip of furcular facet blunt but upturned. Humerus with internal contour from shaft through bicipital crest broadly and gradually curved; area below head, between pectoral attachment and pneu- matic fossa oval and deeply depressed. Olecranon of ulna extending beyond shaft in ulnar contour. Length of process of metacarpal 1 of carpometacarpus more than 44 per cent of total length of bone. Femur with obturator ridge relatively short, straight in internal contour, and with well-marked depression internal to ridge. Tibiotarsus with distal internal ligamental prominence a distinct, rounded papilla below the level of the proximal edge of internal condyle; external ligamental prominence negligible. Tarsometatarsus (based on referred material) flared proximally and distally, shaft depressed anteriorly with bordering ridge sharply defined. 12 Contributions in Science No. 217 Comparisons: In all qualitative characters separating M. diegense and M. milleri from M. calif orniensis (Howard, 1970, Table I), M. cedrosensis is closer to M. diegense and M. milleri. In the characters that distinguish M. diegense from M. milleri (Howard, op. cit.. Table IV), M. cedrosensis resembles M. diegense in the projecting anterior edge of the furcular facet of the coracoid and in the proximal flare of the tarsometatarsus, but the borders of the metatarsal shaft are more sharply defined. In the femur, the depth of the depression internal to the obturator ridge resembles the condi- tion in M. milleri, but the ridge is shorter. In the tibiotarsus, the distal internal ligamental prominence is more prominent than in either M. diegense or M. milleri, but seemingly is closer to the latter in placement below the level of the proximal border of the condyle; the poor preservation of the San Diego specimens somewhat obscures the exact position of the prominence; the external ligamental prominence is negligible as in M. milleri. M. cedrosensis is distinct from both M. diegense and M. milleri in the contour of the humerus from shaft to bicipital crest, being more broadly and gradually rounded even than in M. milleri', also distinct is the oval shape of the area of origin of the external head of the triceps below the humeral head; in both California species this area is broader and of generally angular contour at the junction of the head and pectoral attachment. Distinction is noted, also, in the ulna in which the olecranon is more prominently developed and extends slightly beyond the shaft in ulnar contour. The type of M. cedrosensis provides, for the first time, an accurate guide to the proportionate dimensions of the skeleton in a species of Mancalla. It is 12-16 per cent longer than the average for M. milleri in all represented elements except the coracoid, and is 8 per cent longer in this element. It is essentially equal to the average of M. diegense in length of ulna, carpo- metacarpus, and femur, 5-7 per cent shorter in humerus, tibiotarsus and, probably, tarsometatarsus (this element is incomplete in the type), and 12 per cent shorter in the coracoid. Compared with the 59 per cent ratio of length of coracoid to length of humerus in the type of M. cedrosensis, the ratio in both M. diegense and M. milleri is 64 per cent, based on averages. Compared with a specimen of present-day Uria aalge (LACM Bi480, which is of minimum size for the species in most elements), the type of M. cedrosensis shows greatest difference in 16 per cent longer coracoid and femur, but 13 per cent shorter humerus, 18 per cent shorter carpometacarpus and 53 per cent shorter ulna. The tibiotarsus is 2 per cent longer. The tar- sometatarsus in the type is incomplete but estimating its length from the breadth of the proximal end, it was probably very nearly the same length as in U. aalge. Figure 2. Skeletal elements from type specimen of Mancalla cedrosensis LACM 15373: A, D, tibiotarsus internal and anterior views; B, humerus; C, G, J, coracoid posterior, internal and anterior views; E, ulna; F, radius; H, scapula; I, carpometa- carpus; K, femur. All figures slightly more than natural size. 1971 Pliocene Avian Remains 13 14 Contributions in Science No. 217 These proportions differ somewhat from those computed for the San Diego Mancalla material (Miller and Howard, 1949:207) which, we now know, were based on two species (Howard, 1970). With unequal repre- sentation of the large and small species, these prior figures overemphasized the large size of the coracoid on one hand, and the small size of the humerus, carpometacarpus, and ulna on the other; the length of the femur was also underestimated. Measurements of Type, LACM 15373: Coracoid, length (from head to sternal facet) 43.2 mm, breadth across furcular facet 7.3 mm, distance from below scapular facet to head 15.6 mm, least breadth of shaft below procora- coid 5.3 mm, breadth immediately below furcular facet 5.3 mm. Scapula, breadth across articular end 12.6 mm. Humerus, length from head to internal condyle 72.4 mm, breadth of proximal end 17.9 mm, breadth of distal end 7.2 mm, length from head to distal edge of deltoid crest 48.8 mm. Ulna, length from intercotylar ridge to internal condyle 29.5 mm, depth of proximal end through olecranon 9.5 mm, breadth across proximal cotylae 6.5 mm, breadth of shaft at middle 4.2 mm, depth of shaft 6.2 mm. Radius, greatest length 29.0 mm. Carpometacarpus, length to end of metacarpal II 35.8 mm, length of process of metacarpal I 16.0 mm (est.). Femur, greatest length 54.7 mm, depth through trochanter and obturator ridge 9.2 mm, greatest breadth of distal end 10.6 mm. Tibiotarsus, length from proximal articular surface to distal condyles 83.9 mm, breadth distal condyles 9.4 mm, depth distal condyles 8.8 mm. Tarsometatarsus, breadth of proximal end 10.2 mm. Summary and Conclusions The seven species represented in the Early Pliocene avifauna of Cedros Island are undoubtedly all extinct. Three, however, are represented by material that is unsatisfactory for assignment beyond the generic level. One is tenta- tively allocated to Megapaloelodus opsigonus, a primitive flamingo known only from the Early Pliocene of Oregon. Three are described as new: Puffinus tedfordif Cerorhinca minor, and Mancalla cedrosensis. The genus Puffinus is well represented in the Tertiary of the west coast, with five Miocene and two Pliocene species previously recorded from California. Cerorhinca has only two previous fossil records, both from the Miocene of California. C. minor is the smallest known species of the genus. Mancalla cedrosensis adds a fourth species to those recorded for this genus of flightless auks; previous records are all from California, and presumably all Pliocene. The avifauna of Cedros Island, although not nearly as abundantly repre- sented as that of the somewhat later Pliocene of San Diego, California, is similar in the predominance of Mancalla. The absence of immature bones indicates that the area may have been a congregating site but not a nesting site for these flightless birds. This is thought to be true, also, of the San Diego site. The Cedros bones, however, are much better preserved than most of those from San Diego, suggesting that they were less subject to weathering 1971 Pliocene Avian Remains 15 Size Range (in mm) of Mancalla cedrosensis, Compared with M. diegense and M. milleri Coracoid Length Breadth below furcular facet Scapula Breadth proxi- mal end Humerus Length Breadth proxi- mal end Ulna Length Breadth proxi- mal end Radius Length Greatest depth shaft Carpometacarpus Length Length process metacarpal I Depth proximal end through metacarpal I Femur Length Breadth of distal end Tebiotarsus Length Breadth of distal end Tarsometatarsus Length Breadth proxi- mal end M. cedrosensis Max. Mean Min. 47.5 45.3 43.2 6.0 5.5 5.2 14.2 13.7 12.6 80.0 73.5 69.5 20.1 17.9 17.0 31.7 30.0 28.3 7.2 6.6 6.2 31.1 29.3 27.3 6.1 5.6 5.4 36.0 35.4 34.9 16.0 15.6 15.4 11.1 10.5 10.2 56.5 55.4 54.7 10.8 10.6 10.4 83.9 9.5 9.2 9.0 42.0 39.2 37.3 10.9 10.3 9.9 M. diegense Max. Mean Min. 54.0 49.1 47.5 6.3 5.8 5.4 14.4 13.5 12.5 85.2 76.5 71.0 20.3 18.7 17.3 32.1 29.4 28.0 6.6 6.4 5.9 31.8 30.9 29.6 6.4 6.35 6.3 37.2 35.6 35.0 15.5 15.3 15.2 11.0 10.3 9.7 57.0 55.2 53.8 11.1 10.2 9.7 98.5 90.2 81.0 10.7 9.0 8.2 43.6 39.9 38.0 10.7 9.6 8.7 M. milleri Max. Mean Min. 43.5 40.0 38.5 5.1 4.9 4.0 11.8 10.9 10.3 66.6 62.7 56.4 16.4 15.3 14.2 27.0 25.3 23.6 5.7 5.2 4.5 26.9 25.1 24.2 5.6 4.9 4.6 33.2 31.6 29.6 14.5 12.6 9.5 9.1 8.8 51.1 47.9 45.0 9.3 9.0 8.4 77.5 74.9 72.0 7.9 7.2 6.5 36.0 34.0 31.1 8.4 8.0 7.2 and may have been deposited further offshore. In addition to Mancalla, other similarities of occurrence in the two avifaunas are shearwaters, sulids, and alcids although in no instance are the species the same. Grebes and loons, present at San Diego are lacking at Cedros. On the other hand, the Cedros primitive flamingo, Megapaloelodus, represents a group that has not been noted at San Diego. 16 Contributions in Science No. 217 The Early Pliocene fauna of Juntura, Oregon (including fish, reptile, bird, and mammal) indicates a pond, lake or river community, not marine (Shotwell, 1963 : 15) . The Megapaloelodus of this fauna is the only avian form similar to the avifauna of Cedros Island. Other Juntura birds include cormo- rant, goose, ducks, and coot. The two bones from Cedros Island assigned to Puffinus sp. suggest a shearwater the size of P. felthami described from the Early Pliocene of Corona del Mar, California, but positive identification is impossible. The scant remains from Corona del Mar include but one other species, Mancalla californiensis. In three of the other Pliocene marine localities in California, Mancalla is the only bird recorded: M. californiensis from the Third Street tunnel, Los Angeles; M. diegense from Humboldt County (age not confirmed), and Laguna Hills, Orange County. A fourth locality, in Monterey County, has a single record of a loon, Gavia concinna. The addition of the Cedros avifauna to the fossil record brings to 19 the number of specifically assigned avian forms known from the marine Pliocene of the west coast. Acknowledgments The generosity of Dr. Richard Tedford, now of the Department of Vertebrate Paleontology, American Museum of Natural History, in pre- senting the Cedros Island avian material to the Los Angeles County Museum of Natural History and in providing critical data pertaining thereto for this study, is gratefully acknowledged. The species Puffinus tedfordi is named in his honor. My thanks are extended to Dr. Joel Cracraft for providing casts of Palaelodus and Mega paloelod us, and to him and to Drs. Frank Kilmer and Pierce Brodkorb for discussion and information provided. Posthumous recog- nition and appreciation go to Professor Loye Miller for reading and discussing the section on Mancalla a few weeks prior to his death. Except for the Palaelodid specimens, comparative material examined, both Recent and fossil, was available at the Los Angeles County Museum of Natural History as original material or casts. I am continually grateful to the staff of that Museum for many courtesies and kindnesses. Photographs are by Lawrence Reynolds. Resumen Siete especies se representan en una avifauna del Plioceno Temprano de depositos marinos sobre Isla Cedros. Mayor numero des huesos fosiles representan la especie nueva, Mancalla cedrosensis. Dos otras especies nuevas se describan, y el flamenco primitivo, Megapaloelodus opsigonus Brodkorb se identifica tentativamente. Tres especies se registran solamente por el genero. 1971 Pliocene Avian Remains 17 Literature Cited Brodkorb, Pierce. 1961. Birds from the Pliocene of Juntura, Oregon. Jour. Florida Acad. Sci. 24:170-184. Howard, Hildegarde. 1949. New avian records for the Pliocene of California. Carnegie Inst. Washington Publ. 584:177-199. 1968. Tertiary birds from Laguna Hills, Orange County, California. Los Angeles Co. Mus., Contrib. Sci. 142:1-21. 1970. A review of the extinct avian genus, Mancalla. Los Angeles Co. Mus., Contrib. Sci. 203:1-12. Miller, Alden H. 1944. An avifauna from the Lower Miocene of South Dakota. Univ. Calif. Publ., Bull. Dept. Geol. Sci. 27:85-100. Miller, Loye. 1925. Avian remains from the Miocene of Lompoc, California. Carnegie Inst. Washington, Publ. 349:107-117. 1950. A Miocene flamingo from California. Condor 52:69-73. Miller, Loye, and Robt. I. Bowman. 1958. Further bird remains from the San Diego Pliocene. Los Angeles Co. Mus., Contrib. Sci. 20:1-15. Miller, Loye, and Hildegarde Howard. 1949. The flightless Pliocene bird Man- calla. Carnegie Inst. Washington, Publ. 584:201-228. Milne-Edwards, Alphonse. 1867-1871. Recherches anatomiques et paleontologi- ques pour servir a I’histoire des oiseaux fossiles de la France (Paris, Victor Masson et Fils). Text, vol. 1, 474 p., Vol. 2, 632 p.; Atlas, Vol. 1, 96 pis., Vol. 2, 104 pis. Mina, Frederico. 1957. Bosquejo geologico del territoria sur de la Baja California. Assoc. Mexicana Geol. Petroleros 9: 141-269. Shufeldt, Robert. 1896. Fossil bones of birds and mammals from Grotto Pietro Tamponi and Grive-St. Alban. Proc. Acad. Nat. Sci. Phila. 1896:507-516, and pi. xxiv. Shotwell, Arnold. 1963. Pliocene mammalian communities of the Juntura Basin. The Juntura Basin: Studies in Earth History and Paleoecology. Trans. Amer. Phil. Soc. n.s. 53:7-21. Wetmore, Alexander. 1926. Observations on fossil birds described from the Miocene of Maryland. Auk 43:462-468. Accepted for publication August 30, 1971 LOS ’ ANGELES I COUNTY MUSEUM CONTRIBUTIONS Cxjuci IN SCIENCE 'umber 218 December 29, 1971 A DISTRIBUTIONAL CHECKLIST OF THE AMPHIBIANS OF HONDURAS By John R. Meyer and Larry David Wilson I II Los Angeles County Museum of Natural History • Exposition Park Los Angeles, California 90007 CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers in the fields of Biology, Geology and Anthropology, published at irregular intervals by the Los Angeles County Museum of Natural History. 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, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007. INSTRUCTIONS FOR AUTHORS Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU- TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of papers will be determined by the amount and character of new information. Al- though priority will be given to manuscripts by staff members, or to papers dealing largely with specimens in the Museum’s collections, other technical papers will be considered. All manuscripts must be recommended 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 examined for suitability by an Edi- torial Committee including review by competent specialists outside the Museum. Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must indicate that the primary type has become the property of a scientific institution of their choice and cited by name. MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological Journals is to be followed in preparation of copy. (2) Double space entire manu- script. (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 unavoidable 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 1964 AIBS Style Manual for Biological Journals in preparing illustration and legend copy for style. Submit only illustrations made with permanent ink and glossy photo- graphic 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. Changes in the manuscript after galley proof 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 DISTRIBUTIONAL CHECKLIST OF THE AMPHIBIANS OF HONDURAS By John R. Meyer^ and Larry David Wilson^ Abstract: The amphibian fauna of the Republic of Hon- duras is listed. For each of the 52 species present, brief synon- ymies, a statement of the entire range, a list of Honduran locality records, and a statement of the ecological distribution in Honduras are given. Keys for identification are provided. A list of species of questionable occurrence and a gazeteer of herpeto- logical localities in Honduras are included. Situated at the southeastern end of the Nuclear Core region (Schu- chert, 1935) of Middle America, the Republic of Honduras has remained virtually unexplored herpetologically until recent years. Middle America was recently characterized as a distinct herpetofaunal region (Savage, 1966), owing in part to the separation of the Nuclear Core region from South America during much of the Tertiary. Duellman (1966) discussed the composition, ecology, and distribution of the recent herpetofauna of Middle America and characterized the area with regard to assemblages of amphibians and reptiles. These authors’ hypotheses regarding the characteristics of the recent herpeto- fauna of Middle America, the role of the Nuclear Core region in the develop- ment of the Mesoamerican herpetofauna, and subsequent movements of extralimital elements into the area were of necessity made with only minimal distributional data for the amphibians and reptiles of Honduras. Previous treatments of the herpetofauna of the country have been limited to reports of collections by Werner (1896), Dunn and Emlen (1932), Schmidt (1933, 1936), Lynch and Fugler (1965), Meyer (1966), and Echternacht (1968) and a scattering of notes by a few other authors. Since Honduras comprises nearly half the landmass involved in the Nuclear Core region, data on the composition and distribution of the herpeto- fauna of the country are desirable to evaluate present concepts and hypotheses regarding the Middle American herpetofauna. The present work is the first in a series of papers planned to elucidate the composition and relationships of the Honduran herpetofauna. Methods and Materials Meyer first became involved with the Honduran herpetofauna during a trip to that country in 1963. Field studies in Honduras were undertaken by both of us in 1967, 1968, and 1971 ; additional trips were made by Wilson in 1969 and 1970. ^The Research Ranch, Elgin, Arizona 85611; and Research Associate, Los Angeles County Museum of Natural History, Los Angeles, California 90007. ^Horizon School, 9025 Sunset Drive, South Miami, Florida 33143. 1 2 Contributions in Science No. 218 In addition to field studies, herpetological collections from Honduras in major museums have been examined. The total number of specimens examined (amphibians and reptiles) was approximately 7000. Of this total, about 3000 specimens representing 130 of the 196 species known from the country were collected by us. 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. The complete range for each species is given, utilizing the elevational designations of Stuart (1963). These categories are: Low— 0 to 600 meters; Moderate— 600 to 1500 meters; Intermediate— 1500 to 2700 meters; High- above 2700 meters. Locality records and museum numbers given are for specimens examined by us, unless otherwise noted. The distribution of each species in Honduras is given in terms of the ecologic formations (bioclimates) of Holdridge (1964). The extent of the various formations in Honduras is indicated in Figure 1 (pages 24 and 25). 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 mem- bers, Basil Nafpaktitis, Findlay Russell, John Wright, and Russell Zimmer. Curators of collections from which material was examined are Walter Auffenberg, Florida State Museum (UF) ; James E. Bohlke, Academy of Natural Sciences of 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, Carnegie Museum (CM) ; the late Michael Ovchynnyk, Michi- gan State University (MSU) ; James A. Peters, United States National Museum (USNM) ; Douglas A. Rossman, Museum of Zoology, Louisiana State Uni- versity (LSUMZ) ; Dorothy M. Smith, Museum of Natural History, Univer- sity of Illinois (UIMNH) ; Charles F. Walker, Museum of Zoology, Univer- sity of Michigan (UMMZ) ; Ernest E. Williams, Museum of Comparative Zoology, Harvard University (MCZ) ; John W. Wright, Los Angeles County Museum of Natural History (LACM) ; Richard G. Zweifel, American Mu- seum of Natural History (AMNH). In addition, we would like to thank John Dickson, La Lima, Honduras (JD) for the use of his private collection and Ernest A. Liner, Houma, Louisiana for the donation of a portion of his private collection to a permanent institution at our request. 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; 1971 The Amphibians of Honduras 3 faculty and students of the Escuela Nacional de Agricultura, Catacamas; personnel 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 (Wil- son), and in 1968 by a Graduate School Travel Fellowship from the Uni- versity 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, Meyer’s wife, Terry, in 1967, 1968, and 1971, and Wilson’s wife, Betty, in 1969 and 1970. Order GYMNOPHIONA Family Caeciliidae KEY TO THE GENERA OF CAECILIANS IN HONDURAS Primary annuli less than 115; secondary annuli less than 75 Dermophis Primary annuli more than 120; secondary annuli more than 95 Gymnopis Genus Dermophis Peters Dermophis mexicanus (Dumeril and Bibron) Siphonops mexicanus Dumeril and Bibron, 1841 : 284. Dermophis mexicanus: Peters, 1879: 937; Werner, 1896: 351; Dunn, 1928: 74. Gymnopis clarkii Barbour, 1926: 191. Gymnopis mexicana clarki: Dunn, 1942: 477. Dermophis mexicana clarkii: Meyer, 1966: 172. Dermophis mexicanus mexicanus: Taylor, 1968: 486. Dermophis mexicanus clarkii: Taylor, 1968: 483. Range.— Los/^ and moderate elevations from Veracruz to Tabasco, Mexico on the Atlantic versant and from Oaxaca, Mexico to western Panama on the Pacific; also on the Atlantic versant in northwestern Honduras and in Costa Rica. Locality Records.-ATLA^TIDA : Tela, MCZ 11047, 11779. CORTfiS: San Pedro Sula, AMNH 33386; 2 mi W San Pedro Sula, TCWC 19159-60; La Lima, JD (4); 4.5 mi ENE Villanueva, LACM 47201. VALLE: Amapala, USNM 51380. Ecologic Distribution.— From sea level to 100 meters in the Tropical Moist Forest and Tropical Dry Forest formations. Individuals were found in and under rotting logs in gallery forest along streams. 4 Contributions in Science No. 218 Genus Gymnopis Peters Gymnopis multiplicata Peters Gymnopis multiplicata Peters, 1874: 616. Gymnopis multiplicata multiplicata: Dunn, 1942: 464. Range.— and moderate elevations from Guerrero, Mexico to Guatemala on the Pacific versant and from Guatemala to western Panama on the Atlantic; also on the Pacific versant in Costa Rica and western Panama. Locality Records.-EL PARAISO: Arenal, LACM 10979. OLANCHO: 4.5 km SE Catacamas, LSUMZ 21324. YORO: Progreso District, MCZ 1 1048. Ecologic Distribution.— From near sea level to 400 meters in the Tropical Moist Forest and Tropical Dry Forest formations. One individual was found beneath a rotting log in gallery forest along a stream. Order CAUDATA Family Plethodontidae KEY TO THE GENERA OF SALAMANDERS IN HONDURAS 1 . Costal grooves more than 16 Oedipina 1. Costal grooves not more than 14 2 2. A well developed sublingual fold; hands and feet narrow, width less than length Chiropterotriton 2. No sublingual fold; hands and feet broad, width greater than length Bolitoglossa Genus Bolitoglossa Dumeril, Bibron, and Dumeril KEY TO THE SPECIES OF BOLITOGLOSSA IN HONDURAS 1. Toes fully webbed (webbing reaching di^M end of terminal phalanx 3 1. Toes not fully webbed 2 2. Large species, reaching 105 mm snout-vent length; vomerine teeth 18-20 in series schmidti 2. Medium sized species, not over 60 mm snout-vent length; no more than 1 5 vomerine teeth in series dunni 3. Belly striped, yellow and blackish brown striatula 3. Belly not striped 4 4. Tail noticeably longer than snout-vent length mexicana 4. Tail noticeably shorter than snout-vent length 5 5. Maxillary teeth absent rufescens 5. Maxillary teeth present 6 6. Small species, not exceeding 40 mm snout-vent length; dorsal pattern of dark streaks and pepperings on light background occidentalis 6. Large species, exceeding 60 mm snout-vent length; dorsal pattern of dark mottling on lighter background dofleini 1971 The Amphibians of Honduras 5 Bolitoglossa dofleini (Werner) Spelerpes dofleini Werner, 1903: 352. Bolitoglossa doffleini: (sic) Taylor, 1944: 219. Range.— Low and moderate elevations of the Atlantic versant from Alta Verapaz, Guatemala to north central Honduras. Locality Records.— YORO: Portillo Grande, FMNH 34681. Ecologic Distribution.— Known only from about 1300 meters in the Sub- tropical Wet Forest Formation. Bolitoglossa dunni (Schmidt) Oedipus dunni Schmidt, 1933 : 16. Bolitoglossa dunni: Wake and Brame, 1963: 386; Brame, 1967: 18. Oedipus morio: Dunn, 1926: 387 (in part). Magnadigita dunni: T&yXor^ 1944: 218. Range.— ModQXditQ to high elevations of western Honduras. Locality Records.— COKTLS: mountains W of San Pedro Sula, FMNH 4544-45, 4547, 4550-51, 4553-54, 4559-60, 4562, UMMZ 80933. OCO- TEPEQUE: 12.5 mi E Nueva Ocotepeque, LACM 46940. YORO: Portillo Grande, MCZ 21245-46. Ecologic Distribution.— From 1500 to 1900 meters in the Lower Montane Wet Forest and Lower Montane Moist Forest formations. A single individual was found beneath a log in a clearing in the forest, and Schmidt ( 1942) reported finding this species in epiphytic bromeliads in the forest. Bolitoglossa mexicana Dumeril, Bibron, and Dumeril Bolitoglossa mexicana Dumeril, Bibron, and Dumeril, 1854; 93; Taylor, 1944: 224; Wake and Brame, 1963: 384. Spelerpes variegatus var. C: Werner, 1896: 351. Oedipus salvinii: Dunn, 1926: 408 (in part). Oedipus platydactylus: Dunn, 1926: 400 (in part). Bolitoglossa moreleti: Stuart, 1963 : 17. Range.— Low and moderate elevations of the Atlantic versant from Veracruz, Mexico to Honduras; also possibly occurs on the Pacific versant in Honduras. Locality R^cor^/5.— ATLANTIDA: Tela District, MCZ 10214. COMAYA- GUA: 8 km S La Mision, AMNH 54920, La Mision, MCZ 26422-23. CORTES: El Jaral, FMNH 4539-42; 1 mi W El Jaral, LACM 45300-04, 47620; La Lima, LACM 47621; Agua Azul, AMNH 54952; Cofradia, AMNH 45337-43. OLANCHO: 26 km NE Catacamas, LACM 45168, 45180. YORO: Subirana Valley, FMNH 21825 (3), MCZ 21235-41; Portillo Grande, MCZ 21242; Coyoles, LSUMZ 21325. Ecologic Distribution.— Soa. level to about 1400 meters in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, and Subtropical Moist Forest formations. This species was found in epiphytic 6 Contributions in Science No. 218 bromeliads, in bunches of bananas, and under rotting logs on the ground. In the drier formations it occurs along watercourses. Bolitoglossa occidentalis Taylor Bolitoglossa Occident alis TdiyXov, 1941: 145. Low and moderate elevations of the Pacific versant from Chiapas, Mexico to Honduras; also on the Atlantic versant in Honduras. Locality Records —COMAY AG\J A : Barranca de Trincheras, FMNH 64647. Ecologic Distribution— only from about 1300 meters in the Sub- tropical Moist Forest Formation. According to data in the museum catalog, the single specimen from the country was found in an epiphytic bromeliad in pine-oak forest. Bolitoglossa rufescens (Cope) Oedipus rufescens Cope, 1869b: 104; Dunn and Emlen, 1932: 22. Bolitoglossa rufescens: Taylor, 1941 : 147. Range.— Low and moderate elevations of the Atlantic versant from San Luis Potosi, Mexico to north central Honduras. Locality ATLANTIDA: mountains above Corozal, LACM 46941; Lancetilla (Dunn and Emlen, 1932). YORO: Subirana Valley, MCZ 21244; Portillo Grande, MCZ 21243, FMNH 34682. Ecologic Distribution.-Erom near sea level to about 1400 meters in the Tropical Moist Forest and Subtropical Wet Forest formations. One individual was found in undisturbed forest at night, on the vegetation about two meters above ground. Bolitoglossa schmidti (Dunn) Oedipus schmidti Dunn, 1924: 96; Dunn, 1926: 361 ; Dunn and Emlen, 1932: 22. Bolitoglossa schmidti: Taylor, 1 944 : 219; Wake and Brame, 1963 : 386; Brame, 1967: 19. Range.— Moderate elevations of northwestern Honduras. Locality Records.— COKTLS : mountains W of San Pedro Sula, MCZ 16287, FMNH 4538. Ecologic D is t rib u tion . —Known only from about 650 meters in the Subtropical Wet Forest Formation. Bolitoglossa striatula (Noble) Oedipus striatulus Noble, 1918: 344. Bolitoglossa striatula: Taylor, 1944: 219; Wake and Brame, 1963 : 386; Brame, 1967: 19. Range.— Low and moderate elevations of the Atlantic versant from north- eastern Honduras to Costa Rica. 1971 The Amphibians of Honduras 7 Locality Records —GKACl AS A DIOS: Cropunta, MCZ 21251. Ecologic Distribution. —Y^no^Nn only from about 50 meters in the Tropical Moist Forest Formation. Genus Chiropterotriton Taylor KEY TO THE SPECIES OF CHIROPTEROTRITON IN HONDURAS Nostrils greatly enlarged; 2-3 costal folds between adpressed limbs nasalis Nostrils not noticeably enlarged; 4 costal folds between adpressed limbs barbouri Chiropterotriton harhouri (Schmidt) Oedipus barbouri Schmidt, 1936: 43. Chiropterotriton barbouri: BvdimQ, 1967: 20. ?Pseudoeurycea barbouri: Tsiylov, 1944: 212. Range.— Moderate and intermediate elevations of north central Honduras. Locality Records.— CORTES \ Agua Azul, AMNH 54949. YORO: Portillo Grande, MCZ 21247-50, FMNH 21866-67. Ecologic Distribution.— Tvom 750 to 1700 meters in the Subtropical Wet For- est and Lower Montane Wet Forest formations. According to Schmidt ( 1936), individuals were found in epiphytic bromeliads. Chiropterotriton nasalis (Dunn) Oedipus nasalis Dunn, 1924: 97; Dunn, 1926: 377. Chiropterotriton nasalis: T?iy\or, 1944: 216; Rabb, 1960: 304; Brame, 1967: 20. Range.— Intermediate and high elevations of northwestern Honduras. Locality Records.— CORTES: mountains W of San Pedro Sula, FMNH 4564, 4568, 4571, 4578, 4581-84, 4586, 4589-90, 91462-63, UMMZ 80926 (2), USNM 103336-37. SANTA BARBARA: Cerro Santa Barbara, E slope, 1900- 2850 m, AMNH 55296-308, LACM 47195-200. Ecologic Distribution.— From 1500 to about 2900 meters in the Lower Mon- tane Wet Forest and Montane Rainforest formations. All individuals of this species have apparently been found in epiphytic bromeliads in the forest. Genus Oedipina Keferstein KEY TO THE SPECIES OF OEDIPINA IN HONDURAS Dermal glands on head and dorsum profuse and conspicuous; snout-vent length greater than 50 mm stuarti Dermal glands absent or inconspicuous; snout-vent length less than 45 mm cyclocauda 8 Contributions in Science No. 218 Oedipina cyclocauda Taylor Oedipina cyclocauda Taylor, 1952a: 764; Brame, 1967: 20; Brame, 1968: 30. Range— Low and moderate elevations of the Atlantic versant from northern Honduras to western Panama. Locality i?ecorum6n\, 1853: 169. Agalychnis moreletii: 1860: 110; Werner, 1896: 351. Agalychnis callidryas taylori: Lynch and Fugler, 1965: 10 (in part). Phyllomedusa moreletii: Savage and Heyer, 1967: 112. Moderate elevations of the Atlantic versant from Veracruz, Mexico to Honduras. Locality Records —COKYES: Agua Azul, AMNH 54914. Ecologic Distribution —YnoyNW only from about 800 meters in the Subtropical Wet Forest Formation. 1971 The Amphibians of Honduras 15 4. 5. 8. KEY TO THE SPECIES OF HYLA, PTYCHOHYLA, AND SMILISCA IN HONDURAS A broad axillary web attaching to upper arm almost at the elbow Hyla loquax Axillary web absent or inconspicuous 2 Upper jaw projecting strongly shelflike over lower jaw anteriorly Hyla staufferi Upper jaw not projecting shelflike over lower jaw 3 Greatest diameter of tympanum at least two-thirds the length of eye; thighs generally mottled, spotted, or banded ................. 4 Tympanum less than two-thirds (generally less than one-half) the length of eye; thighs immaculate, peppered with dark, or striped 5 Lips barred; flanks cream colored with bold brown or black mottling in groin; posterior surfaces of thighs brown with cream colored flecks Smilisca baudinii Lips not barred; a narrow white labial stripe present; flanks cream colored with fine black venation; posterior surfaces of thighs pale brown, with or without darker flecks or small cream colored spots Smilisca phaeota A dark stripe, generally with a narrow light border above, extending from eye above or through tympanum to varying lengths posteriorly 6 No dark stripe extending posteriorly from eye 7 Upper surface of thighs peppered with dark; dorsum yellow to reddish with scattered darker spots Hyla picta Upper surface of thighs immaculate or with dark peppering restricted to knee region Hyla microcephala Sides never mottled, but frequently with light brown pepperings Hyla bromeliacea Sides with brown mottling on a lighter background . 8 Males with lateral glands; nuptial tuberosities on thumbs composed of large discrete horny spines; rostral keel present . Ptychohyla spinipollex Males without lateral glands; nuptial tuberosities on thumbs composed of horny pads; no rostral keel Hyla Salvador ensis Genus Hyla Laurenti Hyla bromeliacea Schmidt Hyla bromeliacea Schmidt, 1933: 19; Stuart, 1948: 29; Lynch and Fugler, 1965: 11. Range.— Moderate and intermediate elevations of the Atlantic versant from northeastern Guatemala to northern Honduras. Locality ATLANTIDA: mountains above La Ceiba, MCZ 21301. 16 Contributions in Science No. 218 CORTES: mountains W of San Pedro Sula, FMNH 4718, 4720-22, MCZ 17431,21302-03. Ecologic D is t rib ution. —Known only from about 1500 meters in the Lower Montane Wet Forest Formation. Schmidt (1942) reported finding this species in epiphytic bromeliads, where eggs also had been laid. Hyla loquax Gaige and Stuart Hyla loquax Gaige and Stuart, 1934: 1; Taylor, 1952b: 868; Lynch and Fug- ler, 1965: 1 1 ; Duellman, 1966: 271. Hyla stadelmani Schmidt, 1936: 45; Lynch and Fugler, 1965: 11. Range —Eow and moderate elevations of the Atlantic versant from Veracruz, Mexico to Costa Rica; also on the Pacific versant in Honduras. Locality Records.-ATLAmiDA: Corozal, LSUMZ 21316. COPAN: 8 mi ENE Copan, LACM 47360-61; 12 mi ENE Copan, LACM 47362-64; 4-5 mi SE Santa Rosa de Copan, LACM 47365-69. CORTES : Lago de Yojoa, AMNH 54943-46, 54948, 54956; 1 mi NW El Jaral, LACM 47370-82, 47388-411; 2.5 mi E La Lima, LACM 47383-87. FRANCISCO MORAZAN: Cerro Uyuca, AMNH 54847-52; Montana de Guaimaca, AMNH 54905-08, UMMZ 123104. OLANCHO: 4.5 km SE Catacamas, LACM 45218. YORO: Subirana Valley, MCZ 21310-11, FMNH 21842. Ecologic D istrib u tion .—Sea level to 1000 meters in the Tropical Moist Forest, Tropical Dry Forest, Subtropical Wet Forest, and Subtropical Moist Forest formations. This species was found breeding in temporary ponds in cleared and brushy areas in June, July, and August. Hyla microcephala Cope Hyla microcephala Cope, 1886: 281. Hyla underwoodi: Dunn and Emlen, 1932: 25; Lynn, 1944: 189. Hyla microcephala martini: Lynch and Fugler, 1965: 10; Meyer, 1966: 173. Hyla microcephala underwoodi: Duellman and Fouquette, 1968: 529. Range.— Low and moderate elevations from Veracruz, Mexico to Honduras and from Panama to Venezuela on the Atlantic versant; occurs from Honduras to Panama on the Pacific versant. Locality Records. -ATLAmiD A: 2.5 mi S La Ceiba, LACM 47477-78. COLON: Trujillo, LACM 47424. COMAYAGUA: 2 km E Siguatepeque, LSUMZ 22602. COPAN: 8 mi ENE Copan, LACM 47425-33; 12 mi ENE Copan, LACM 47434-44; 5 mi SE Santa Rosa de Copan, LACM 47445-48. CORTfiS: Agua Azul, AMNH 54917-19; Lago de Yojoa, AMNH 54957, 55134, MSU 4593-607, KU 64565-77; 1 mi NW El Jaral, LACM 47459-99, 47479-85; 2.5 mi E La Lima, LACM 47467-76. EL PARAISO: Valle de Jamastran, AMNH 54807-12. FRANCISCO MORAZAN: Zamorano, AMNH 54879-81, KU 103223, TCWC 19173-77; Santa Clara, AMNH 54873-78; Rancho San Diego, AMNH 54939; Montana de Guaimaca, AMNH 54900-04. INTIBUCA: Viejo Intibuca, AMNH 54912-13. ISLAS DE LA 1971 The Amphibians of Honduras 17 BAHIA: Isla de Roatan, 0.5 km N Roatan, LSUMZ 21315; Isla de Guanaja, La Playa Hotel, LACM 47412-19. OLANCHO: 26 km NE Catacamas, LACM 45170; 1 km NW Catacamas, LSUMZ 21300-09; 4.5 km SE Cataca- mas, LSUMZ 21310-14, LACM 45217. SANTA BARBARA: 5 mi S Santa Barbara, LACM 45311-13; 14 mi SW Sula, LACM 47449-58; 8 km E Qui- mistan, LSUMZ 22628-29; 28 km SW Chamelecon, LSUMZ 22624-27. YORO: Coyoles, LACM 47420; 1.5 km W Olanchito, LACM 47421-23. Ecologic Distribution-Sea. level to 1000 meters in the Tropical Moist Forest, Tropical Dry Forest, Tropical Arid Forest, Subtropical Wet Forest, and Subtropical Moist Forest formations. This species was found breeding in temporary ponds in cleared and brushy areas in June, July, and August. Hyla picta (Gunther) Hylella picta Gunther, 1901: 286; Lynn, 1944: 190. Hyla picta: Smith and Taylor, 1948: 85. Range— Low elevations of the Atlantic versant from San Luis Potosi, Mexico to northern Honduras. Locality RpcorJ^.-ATLANTIDA: La Ceiba, USNM 117603; Lancetilla, MCZ 17982-87; 20 mi E Tela, LACM 47244. COPAN: 12 mi ENE Copan, LACM 47231-43. Ecologic Distribution.— Sea level to about 600 meters in the Tropical Moist Forest and Subtropical Wet Forest formations. This species was found calling from brush around a pond in a cleared area in June, and one individual was taken on the forest floor in late afternoon in July. Hyla salvadorensis Mertens Hyla salvadorensis Mertens, 1952: 169. Ptychohyla spinipollex: Lynch and Fugler, 1965: 11 (in part). Range.— Intermediate elevations of the Pacific versant from northwestern El Salvador to southeastern Honduras. Locality Recor^/5. -FRANCISCO MORAZAN: Cerro Uyuca, AMNH 54823, 54827-39, KU 103033, 103255-60. INTIBUCa: La Esperanza, LACM 47230. OCOTEPEQUE: 9 mi E Nueva Ocotepeque, LACM 47226-29, 47253; 12.5 mi E Nueva Ocotepeque, LACM 47224. Ecologic Distribution.— From 1600 to 1900 meters in the Lower Montane Moist Forest Formation. This species was found calling in June along a small stream in a clearing and in a roadside seep. One individual was found calling from a wet rock along a stream in August, and larvae were found in a pool nearby. Hyla staufferi Cope Hyla staufferi Cope, 1865a: 195; Lynn, 1944: 189; Taylor, 1952b: 865; Lynch and Fugler, 1965: 11; Meyer, 1966: 173; Duellman, 1966: 274. Hyla culex Dunn and Emlen, 1932: 24; Dunn, 1933: 61. Range.— Low and moderate elevations from Tamaulipas, Mexico to Panama 18 Contributions in Science No. 218 on the Atlantic versant and from Guerrero, Mexico to Panama on the Pacific. 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