JSiu A.
December 14, 1962
TABLE OF CONTENTS
AND
AUTHOR INDEX Nos. 35-63 1960-1962
Los Angeles County Museum
Exposition Park
Los Angeles 7, Calif.
?
Contributions in Science
TABLE OF CONTENTS
No. 35. The Machris Brazilian Expedition. Botany: Pteridophyta, by C. V. Morton. 7 pp. Jan. 20, 1960.
No. 36. The Machris Brazilian Expedition. Entomology: Two new species of Anacroneuria (Plecoptera) from Goias, Brazil, by Stanley G. Jewett, Jr. 4 pp., 2 figs. May 19, 1960.
No. 37. A new genus and species of glossophagine bat from Colima, Mexico, by W. J. Schaldach and Charles A. McLaughlin. 8 pp., 3 figs. May 19,
1960.
No. 38. A census of the abundant large Pleistocene mammals from Rancho La Brea, by Leslie F. Marcus. 1 1 pp., 2 figs. May 19, 1960.
No. 39. A skull of the grizzly bear ( Ursus arctos L.) from pit 10, Rancho La Brea, by Bjorn Kurten. 7 pp., 2 figs. Dec. 15, 1960.
No. 40. The Machris Expedition to Tchad, Africa. Amphibians and reptiles, by David B. Wake and Arnold G. Kluge. 12 pp., 1 fig. May 1 1, 1961.
No. 41. The Machris Brazilian Expedition. Ornithology: Non-passerines, by Kenneth E. Stager. 27 pp., 2 figs. May 11, 1961.
No. 42. A new Geomys from the Vallecito Creek Pleistocene of California, with notes on variation in recent and fossil species, by John A. White and Theodore Downs. 34 pp., 17 figs. June 30, 1961.
No. 43. New light on the flightless goose, Chendytes lawi, by Loye Miller, Edw. D. Mitchell and Jere H. Lipps. 11 pp., 2 pis. June 30, 1961.
No. 44. A new walrus from the Imperial Pliocene of southern California: with notes on odobenid and otariid humeri, by Edw. D. Mitchell, Jr. 28 pp., 13 figs. Oct. 26, 1961.
No. 45. A study of variation and evolution in Miocene Merychippus, by Theo- dore Downs. 75 pp., 22 figs. Dec. 21, 1961.
No. 46. A new bird of the genus Picumnus from eastern Brazil, by Kenneth E. Stager. 4 pp., 1 fig. Dec. 21, 1961.
No. 47. The Machris Brazilian Expedition. Botany: A new Brazilian Begonia, by Lyman B. Smith and Bernice G. Schubert. 3 pp., 1 fig. Dec. 21,
1961.
No. 48. The echinoid Mellita in the Pacific coast Cenozoic, by J. Wyatt Dur- ham. 12 pp., 2 pis. Dec. 21, 1961.
No. 49. A new species of salamander from Colombia and the status of Geo- triton andicola Posada Arango, by David B. Wake and Arden H. Brame, Jr. 8 pp., 1 fig. Feb. 26, 1962.
1962
Table of Contents
3
No. 50. Growth measurements of young captive Atlantic sea turtles in tem- perate waters, by David K. Caldwell. 8 pp. Feb. 26, 1962.
No. 51. A new fish of the genus Coleotropis, family Atherinidae, from Carib- bean Costa Rica, by David K. Caldwell. 8 pp., 2 figs. Feb. 26, 1962.
No. 52. A second record of Osteodontornis, Miocene “toothed” bird, by Hilde- garde Howard and John A. White. 12 pp., 5 figs. Feb. 26, 1962.
No. 53. Postlarvae of the blue marlin, Makaira nigricans, from off Jamaica, by David K. Caldwell. 1 1 pp., 2 figs. May 1 1 , 1962.
No. 54. A new bat of the genus Glossophaga from Mexico, by Alfred L. Gard- ner. 7 pp., 4 figs. May 11, 1962.
No. 55. The Machris Brazilian Expedition. Entomology: Belostomatidae (Hemiptera), by Arnold S. Menke and David R. Lauck. 8 pp., 5 figs. Nov. 16, 1962.
No. 56. A walrus and a sea lion from the Pliocene Purisima formation at San- ta Cruz, California: with remarks on the type locality and geologic age of the sea lion Dusignathus santacruzensis Kellogg, by Edw. D. Mitch- ell, Jr. 24 pp., 12 figs. Dec. 14, 1962.
No. 57. A morphological comparison of Pholisma arenarium Nuttall and Pholisma paniculatum Templeton (Lennoaceae) , by Bonnie C. Tem- pleton. 29 pp., 25 figs. Nov. 16, 1962.
No. 58. A comparison of avian assemblages from individual pits at Rancho La Brea, California, by Hildegarde Howard. 24 pp., 5 figs. Dec. 14, 1962.
No. 59. The Machris Expedition to Tchad, Africa. Birds, by Herbert Fried- mann. 27 pp., 1 fig. Dec. 7, 1962.
No. 60. Factors in the ability of the northeastern Pacific green turtle to orient toward the sea from the land, a possible coordinate in long-range navi- gation, by Melba C. Caldwell and David K. Caldwell. 27 pp., 8 figs. Dec. 7, 1962.
No. 61. Sea turtles in Baja Californian waters (with special reference to those of the Gulf of California), and the description of a new subspecies of northeastern Pacific green turtle, by David K. Caldwell. 31 pp., 5 figs. Dec. 7, 1962.
No. 62. Carapace length— body weight relationship and size and sex ratio of the northeastern Pacific green sea turtle, Chelonia mydas carrinegra, by David K. Caldwell. 10 pp. Dec. 7, 1962.
No. 63. The Machris Brazilian Expedition. Botany: Various families, coordi- nated by E. Yale Dawson. 9 pp. Dec. 14, 1962.
4
Contributions in Science
AUTHOR INDEX
Brame, Arden H., Jr. Caldwell, David K. Caldwell, Melba C. Dawson, E. Yale Downs, Theodore Durham, J. Wyatt Friedmann, Herbert Gardner, Alfred L. Howard, Hildegarde Jewett, Stanley G. Kluge, Arnold G. Kurten, Bjorn Lauck, David R.
Lipps, Jere H. McLaughlin, Charles A. Marcus, Leslie F. Menke, Arnold S. Miller, Loye Mitchell, Edw. D., Jr. Morton, C. V. Schaldach, W. J. Schubert, Bernice G. Smith, Lyman B.
Stager, Kenneth E. Templeton, Bonnie C. Wake, David B.
White, John A.
No. 49
Nos. 50, 51, 53, 60, 61, 62
No. 60
No. 63
Nos. 42, 45
No. 48
No. 59
No. 54
Nos. 52, 58
No. 36
No. 40
No. 39
No. 55
No. 43
No. 37
No. 38
No. 55
No. 43
Nos. 43, 44, 56 No. 35 No. 37 No. 47 No. 47 Nos. 41 , 46 No. 57 Nos. 40, 49 Nos. 42, 52
BER 35
January 20, 1960
rov, 73
THE MAGHRIS BRAZILIAN EXPEDITION
BOTANY: Pteridophyta
By C. V. Morton
\ngeles County Museum • Exposition Park • Los Angeles 7, Calif.
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. Issues are numbered separately and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientists and scientific institutions on an exchange basis. Copies may also be purchased at a nominal price.
The MACHRIS BRAZILIAN EXPEDITION from the Los Angeles County Museum was sponsored by Mr. and Mrs. Maurice A. Machris and Mrs. Maybell Machris Low. It was conducted under the auspices of the Museu Nacional do Brasil. Botanical and zoological collections were made from April through June, 1956, in the region of the headwaters of the Rio Tocantins in the state of Goias. General accounts and intineraries are given in papers 1 and 2 of this series. Technical type specimens of new entities are deposited in the Museu Nacional in Rio de Janeiro.
Hildegarde Howard Editor
E. Yale Dawson Associate Editor
THE MACHRIS BRAZILIAN EXPEDITION
BOTANY : Pteridophyta By C. V. Morton1
The Pteridophyta collected by Expedition Botanist E. Yale Dawson were sent to the United States National Museum for determination. This collection, listed below, proved to contain several new records for the state of Goias, so far as published accounts go. The region is not rich in Pteridophyta, and relatively little has been published on its species.
The collections are cited by Dawson’s field numbers. The correspond- ing localities can be found in the general account of the botany of the Expedition.2 All of the materials, however, came from the region of the Chapada dos Veadeiros (nos. 14133-14815) or the region of the Serra Dourada and immediately north (nos. 14816-15236). The first set of specimens is in the herbarium of the Los Angeles County Museum. A partial second set is in the U. S. National Herbarium.
LYCOPODIACEAE
Lycopodium alopecuroides L. 14652
Lycopodium carolinianum var. meridionale (Underw. & Lloyd) Nessel & Hoehne 14746
Lycopodium cernuum L. 14837
SELAGINELLACEAE
Selaginella ery thro pus (Mart.) Spring. 14849; 14850; 14927
Easily distinguished from other Brazilian species by the erect, red lower axes.
Selaginella marginata (Humb. & Bonpl.) Spring. 15066
Selaginella simplex Baker, ex. char. 14482 Apparently
the only Brazilian species with dimorphic sporophylls, according to the treatment of Brazilian selaginellas by Alston (Fedde, Repert. Sp. Nov. 40: 303-319, 1936). Baker’s original plants were much smaller, and perhaps depauperate.
Selaginella sp. cf. S. tenuissima Fee 14767 Identified
from description.
SCHIZAEACEAE
Lygodium venustum Swartz 14984; 15181
Anemia Swartz3
The highlands of Brazil are extremely rich in species of Anemia , and any expedition into the interior of this region is sure to be highly
^-Curator, Division of Ferns, U. S. National Museum, Smithsonian Institution, Wash- ington, D. C.
2Dawson, E. Yale. 1957. The Machris Brazilian Expedition. Botany: General. Los An- geles Co. Mus. Contr. Sci. (2) : 1-20.
3The determinations and notes on this genus were provided by John T. Mickel, Depart- ment of Botany, University of Michigan, Ann Arbor, Michigan.
«*
4
Contributions in Science
No. 35
rewarding. Probably the most extensive collection in southeastern Brazil was made by Ynez Mexia in 1931 in the neighboring state of Minas Gerais, but few collections have been made in Goias itself. Nearly two-thirds of the species of Anemia have been found in Brazil, and the great diversity of form within the genus is shown in Dr. Dawson’s collections, which include nine or, perhaps, ten species, and represent six different sections.
Several sections of the genus, such as the Tomentosae, Oblongifoliae, Hirsutae, and Phyllitides, are very problematic due to the great confusion in nomenclature and in the interpretation of subtle morphological differ- ences. For this reason, many identifications must be tentative until more detailed studies on these groups can be completed. Such work is currently in progress.
Anemia buniifolia (Gardner) Moore 14588 Although most specimens of this species are from Venezuela, Colombia, and Matto Grosso, Brazil, there is no doubt of the identity of this one from Goias. The architecture of the specimen is more regularly pinnate than usual, and the fertile fronds are more strict. It is interesting from a morpho- logical standpoint in its intemediate manner of displaying its fertile and sterile fronds. The fronds of the species are typically entirely dimorphic, but this specimen is unusual in exhibiting several intermediate stages, from completely sterile, to fronds with fertile basal pinnae, to half or three-quarters fertile, to completely fertile.
Anemia mille folia Gardner 14712 An average specimen
of this distinctive species. It has been collected before in Goias (Ule 360 (UC) ). As in the preceding species, collections have been more frequent in Matto Grosso and farther north in Venezuela, Colombia, and Panama.
Anemia anthrisci folia Schrad. 15111 This species is
extremely variable and widespread, ranging from northern Mexico to Argentina. The specimen is a perfect match of Rosenstock 18344 (US) from Goias, and closely resembles other specimens from southeastern Brazil and Uruguay.
Anemia fulva (Cav.) Swartz 14475; 15115 In cutting
and texture these two specimens closely resemble several others from this region of Brazil: Chase 11433 (US) from Goias, and Chase 9254.5 (US), Chase 10629 (US), Macedo 2334 (US), and Regnell 1480 (US) from Minas Gerais.
Anemia tenella (Cav.) Swartz 14250; 14427 The ex-
tremely fine dissection of the pinnae and the apparently limited range (southeastern Brazil) suggest that this taxon be upheld as a distinct species. However, with further study it may prove to be a variety of A. hirsuta, as was suggested by Baker (Hooker and Baker, Synopsis Filicum p. 433, 1868).
Anemia oblongifolia (Cav.) Swartz 14268; 14572; 14713;
14714; 15114 Collections 14714 and 15114 are much smaller than
1960
Morton: Brazil, Botany
5
the rest, both having fronds one to four centimeters long. From a comparison with juvenile and dwarfed forms of A. oblongifolia, these appear merely to be depauperate.
Anemia pastinacaria Moritz 14737 Goias is near the
southern limit of the range of this species; the majority of collections come from Central America and the West Indies, and a few from Colombia, Venezuela, Peru, Bolivia, and Brazil. Although one of the fronds somewhat resembles A. oblongifolia, characters of the pinnae and spores distinguish it from that species.
Anemia ouropretana Christ 14332 This specimen is a
good example of the species. It is apparently endemic to southeastern Brazil, for all previous specimens have been collected in Minas Gerais by Damazio and Mexia.
Anemia phyllitidis (L.) Swartz 14413; 14889; 14928 It
is not surprising to find this species among the collections since it is probably the most widespread species of Anemia in America. The pinnae are rather narrow in these specimens, but the species as a whole is extremely variable.
Anemia sp. nov. ? 15180 This specimen is definitely
related to the A. phyllitidis group in having reticulate venation, but its extremely narrow pinnae (0.5 by 3.5 cm.) are distinctive. The somewhat rounded apices of the pinnae suggest A. tweediana of Uruguay and northern Argentina, although some specimens of the West Indian A. underwoodiana and juvenile forms of A. phyllitidis show the same condi- tion. The pinnae are much narrower than any found in A. tweediana, and even taking into consideration the extreme variation of A. phyllitidis, there are no specimens that I have seen with such slender pinnae. A detailed study of the entire complex must be undertaken before the identity of this specimen can be determined.
GLEICHENIACEAE
Dicranopteris flexuosa (Schrader) Underw. 14206
Gleichenia pennigera (Mart.) Moore 14656 I am tenta-
tively following Holttum (Reinwardtia 4: 257-280. 1957) in regarding the genus Sticherus as not generally separable from Gleichenia.
HYMENOPHYLLACEAE
Trichomanes pellucens Kunze 14738
Trichomanes pinnatum Hedwig 14269; 14869a
CYATHEACEAE
Alsophila paleolata Mart. 14653 A collective species as
currently recognized.
Alsophila villosa (Humb. & Bonpl.) Desv. 14790
Cyathea sternbergii Pohl 14968
6
Contributions in Science
No. 35
POLYPODIACEAE
Adiantopsis radiata (L.) Fee 14501; 14506
Adiantum delicatulum Mart. 14925
Adiantum intermedium Swartz 14335; 14530; 14929 This species has been, and still is, somewhat dubious. The specimens cited agree with a photograph of the type in the herbarium at Stockholm.
Adiantum poiretii Wikstr. 15113 Probably the first
record from Goias of this widely distributed species.
Adiantum serrato-dentatum Willd. 14868; 14869
Adiantum sinuosum Gardner 15112; 14873 (?)
Asplenium formosum Willd. 14503
Bakeropteris pinnata (Kaulf.) Kuntze ( Cassebeera pinnata Kaulf. ; Pellaea pinnata Prantl) 14711 Probably known previously
only from Minas Gerais. The ultimate generic disposition of this species remains to be determined. It has been considered allied to Pellaea. Cf. Tryon, Contr. Gray Herb. 143:67. 1942.
Blechnum asplenioides Swartz 14492; 14573; 15067
Blechnum brasiliense Desv. 15105
Blechnum fraxineum Willd. 14573a Theoretically differs
from B. occidentale and its allies in having a conform terminal pinna, but there are intermediate conditions in which it is difficult to decide if the apex is conform or pinnatifid. The present collection is one of these uncertain intermediates.
Blechnum imperiale (Fee & Glaziou) Christ ? 14665
This collection is slightly different from collections from Rio de Janeiro and Minas Gerais. Further study might show it to be separable. It is a Brazilian endemic.
Blechnum lanceola Swartz 14739
Blechnum occidentale L. 14251; 14334; 14861; 14891; 14924
Blechnum regnellianum (Kunze) C. Chr. 14744 Perhaps
new to Goias. A Brazilian endemic.
Ctenitis deflexa (Kaulf.) Copel. 14932
Doryopteris ornithopus (Mett.) J. Smith 14589 The only
collection previously known from Goias is Ule 798 (p.p.) from Serra Dourada, the type of D. ornithopus var. pygmaea Brade, which is not considered by Tryon (Contr. Gray Herb. 143: 29. 1942) to be separable from the typical variety.
Dryopteris meniscioides var. conjerta (Kaulf.) Morton 14893
Elaphoglossum burchellii (Baker) C. Chr. 14507 The
determinations of elaphoglossums must be considered tentative, pending a revision of the genus.
Elaphoglossum dusenii Christ 14574
Elaphoglossum glabellum J. Smith 14741 Elaphoglossum macahense (Fee) Rosenst. ?
14508
1960
Morton: Brazil, Botany
7
Elaphoglossum scalpellum (Mart.) Moore 14359; 14477
Lindsaea guianensis (Aubl.) Dryand. subsp. lanceastrum Kramer 14870; 14866 This is the common subspecies in Brazil, the subsp.
guianensis being restricted to Amazonas. It has been collected once previously in Goias, at Sucuri, Rio das Femeas, Luetzelburg 13749a. Pityrogramma calomelanos (L.) Link 14651; 14987
Polypodium aureum L. 14333; 14766
Polypodium latipes Langsd. & Fisch. 14683
Pteridium aquilinum var. arachnoideum (Kaulf.) Herter 14529
Pteris quadriaurita Retz. 14985; 14986
Thelypteris angustifolia (Willd.) Proctor 14930 This
specimen from deep forest along the Ribeirao Cristalino, 25 km. east of Formoso, Serra Dourada, Goias, is a new record for Brazil. For distribution see Maxon & Morton, Bull. Torrey Bot. Club 65: 361. 1938.
Thelypteris opposita (Vahl) Ching var. rivulorum (Raddi) Morton, comb. nov.
Polypodium rivulorum Raddi, Plant. Bras. 1 : 23, pi. 35. 1825.
Dry o pteris opposita var. rivulorum C. Chr. ex Rosenst., Hedwigia
46: 120. 1906.
14497 ; 14939 The variety is characterized by Christensen
(Dansk. Vid. Selsk. Skrift. vii., Naturvid. Afd. iv, 4: 289. 1907).
Thelypteris salzmannii (Fee) Morton, comb. nov.
Meniscium salzmannii Fee, Gen. Fil. 223. 1853.
Dryopteris salzmannii Maxon & Morton. Bull. Torrey Bot. Club
65: 357. 1938.
14493 Another collection of this alliance, perhaps different,
is 14931.
LOS ANGELES COUNTY MUSEUM CONTRIBUTIONS IN SCIENCE
The Machris Brazilian Expedition
No. 1. General Account, by Jean Delacour.
No. 2. Botany: General, by E. Yale Dawson.
No. 3- Botany: A New Dodder from Goias, by T. G. Yuncker.
No. 4. Botany: The Lichens, by Carroll W. Dodge.
No. 5. Botany: Cyanophyta, by Francis Drouet.
No. 6. Botany: A New Mint from Goias, by Carl Epling.
No. 7. Botany: Phanerogamae, various smaller families, edited by E. Yale Dawson. No. 10. Botany: A New Columnar Cactus from Goias, by E. Yale Dawson No. 11. Botany: Chlorophyta; Euglenophyta, by G. W. Prescott
No. 12. Entomology: General; Systematics of the Notonectidae (Hemiptera), by Fred S. Truxal.
No. 13. Botany: Phanerogamae, Leguminosae, by Richard S. Cowan.
No. 14. Entomology: Gelastocoridae (Hemiptera), by E. L. Todd.
No. 17. Botany: Phanerogamae, Bromeliaceae and other smaller families, by Lyman B. Smith.
No. 18. Botany: Musci, by Howard Crum.
No. 21. Botany: Phanerogamae, Euphorbiaceae, Lentibulariaceae, Rubiaceae, by Julian A. Steyermark.
No. 22. Botany: Gramineae, by Jason R. Swallen.
No. 23. Botany: Phanerogamae, Alstroemeriaceae and other families, by Lyman B. Smith and collaborators.
No. 24. Botany: Fungi, by G. W. Martin and collaborators.
No. 26. Botany: Hepaticae, by Margaret Fulford.
No. 28. Botany: Phanerogamae, Melastomataceae and Polygalaceae, by J. J. Wurdack. No. 30. Botany: Phanerogamae, Amaranthaceae and other families, by Lyman B. Smith and collaborators.
No. 32. Botany: Phanerogamae, Acanthaceae, by Emery C. Leonard.
No. 33. Ornithology: Two new birds from Central Goias, Brazil, by Kenneth E. Stager. No. 35. Botany: Pteridophyta, by C. V. Morton.
Other Subjects
No. 8. Notes on Eastern Pacific Insular Marine Algae, by E. Yale Dawson.
No. 9. A New Species of Passerine Bird from the Miocene of California, by Hildegarde Howard.
No. 15. Marine Algae of the Pacific Costa Rican Gulfs, by E. Yale Dawson.
No. 16. A Classification of the Oscines (Aves), by Jean Delacour and Charles Vaurie.
No. 19. A New Race of the Pocket Gopher Geomys bursarius from Missouri, by Charles A. McLaughlin.
No. 20. Further Bird Remains from the San Diego Pliocene, by Loye Miller and Robert 1. Bowman.
No. 25. Miocene Sulids of Southern California, by Hildegarde Howard No. 27. Marine Algae from the 1958 Cruise of the Stella Polaris in the Gulf of Cali- fornia, by E. Yale Dawson.
No. 29. Quaternary Animals from Schuiling Cave in the Mojave Desert, California, by Theodore Downs, Hildegarde Howard, Thomas Clements and Gerald A. Smith. No. 31. Late Pleistocene Invertebrates of the Newport Bay area, California, by George P. Kanakoff and William K. Emerson.
No. 34. A new Giant Water Bug from Mexico, by Arnold S. Menke.
[ber 36 May 19, 1960
7ojJ 7 3
\£L%i>X THE MACHRIS BRAZILIAN EXPEDITION
ENTOMOLOGY: Two New Species of Anacroneuria (Plecoptera) from Goias, Brazil
By Stanley G. Jewett, Jr.1
Stoneflies of the genus Anacroneuria occcur commonly throughout most of Central and South America. There are numerous species, and many of them are difficult to identify. In two earlier papers (Jewett, 1958 and 1959) comments are offered on some of the systematic problems involved in classifying members of this genus. It is sufficient to state here that the color pattern of head and pronotum and the shape of the distal border of the female subgenital plate appear to be the most reliable morphological characters to distinguish the species.
Stonefly material taken by the Machris Expedition2 contains two species of Anacroneuria which do not fit existing descriptions. These are described below.
Unless otherwise noted all specimens have been softened after having been pinned and are now preserved in alcohol. Specimens are deposited in the Museu Nacional do Brasil (MN), the Los Angeles County Museum (LACM), and the writer’s collection (SGJ).
Anacroneuria dourada n. sp.
(Figs. 1, 1A)
Length of forewing, 15.5 to 16 mm. in female, 11 mm. in male.
Head mostly brown, females showing pattern as follows in specimens relaxed and placed in alcohol after originally being pinned: entire frons brown; M-line faintly discernible as lighter shade; area lateral to and behind ocelli light brown, almost yellow; lightest area adjacent to and of
*7742 S.E. 27th Avenue, Portland 2, Oregon.
2See L. A. Co. Mus. Contrib. Sci. no. 12 for general account of the entomology of the expedition.
SMITHSONIAN ^
INSTITUTION JOfi I IP
2
Contributions in Science
No. 36
2
Figs. 1-1 A. Anacroneuria dourada n. sp. 1, Head and Pronotum. 1A, Eighth and ninth sternites. Fig. 2. Anacroneuria galba n. sp., eighth and ninth sternites.
1960
Jewett: Brazil Plecoptera
3
about same size as ocelli; lappets very dark distally; head of males almost uniformly brown. Pronotum brown with narrow, median, light-brown or yellow stripe occupying possibly an eighth of pronotal width. Legs with outer faces light to dark brown, the tibiae darker than femora. Tarsi brown. Tails distinctly bicolored except first two or three segments; wings tinged lightly brown; veins brownish, the subcostal vein darkest.
Female: — In specimens cleared in KOH and somewhat flattened, the subgenital plate four-lobed, the median notch V-shaped and deep compared to shallow lateral notches. Sclerotized area of ninth sternite in typical T-shape, the stem long and narrow, the entire area covered with fine short hairs.
Male: — Smaller and darker than female and with small conical nail on ninth sternite.
The head and pronotal color pattern, and the shape of the female subgenital plate form a combination of characters that separate this species from described species of Anacroneuria.
Collection Data: — Holotype female, 24 kilometers east of Formoso, Goias, Brazil, 9-VI-56, F. S. Truxal (MN). Allotype male, same data except 23-V-56 (MN). Paratypes as follows: Same data as allotype, female (pinned, LACM) ; same data except 26-V-56, male, 2 females (SGJ) ; same data except 29-V-56, 2 females (female, SGJ ; pinned female, LACM); same data except 9-VI-56, 2 females (female, SGJ; pinned female, LACM) ; 20 kilometers north of Sao Joao da Alianga, Goias, Brazil, 28-IV-56, F. S. Truxal, female (pinned, LACM).
Anacroneuria galba n. sp.
Fig. 2
Length of forewing, 14 to 17.5 mm. in female, 10 to 11 mm. in male.
Head yellow, darkest on lappets, without pattern, the ocelli ringed in black. Pronotum yellow with irregular, broad, brown stripe on either side occupying about half total width of each disc; lateral borders of pronotum yellow; central yellow stripe occupying about one third of pronotal width. Legs and tails yellow. Antennal segments bicolored, but dark areas not sharply delineated. Wings tinged lightly with yellow, the costal vein not darker than others.
Female: — In specimens cleared in KOH and somewhat flattened, the subgenital plate four-lobed, the notches of about equal depth. Sclerotized area of ninth sternite in typical T-shape, the arms thick and covered with long hairs.
Male :— Smaller than female and with conical nail or ninth sternite.
This species resembles several others with similarly marked heads and pronota described from Mexico, Central America, and northern South America. In both color pattern and shape of the female subgenital plate it is, for example, similar to A. crenulata Jewett described from Mexico
4
Contributions in Science
No. 36
and Central America. It differs from this and other species, however, in details of the shape of the subgenital plate and apparently in being yellower in color.
Collection Data: — Holotype female, Veadeiros, Goias, Brazil, 30-IV-56, F. S. Truxal (MN). Allotype male, 24 kilometers east of Formoso, Goias, Brazil, 26-V-56, F. S. Truxal (MN). Paratypes as follows: Same data as allotype except 19-V-56, female (SGJ) ; same data except 26-V-56, 2 males, female (male, female, SGJ; pinned male, LACM) ; same data except 29-V-56, female (LACM) ; same data except 6- VI-56, male (SGJ); same data except 9- VI-56, male (pinned, LACM).
Provisionally identified with this species are the following specimens which were kindly made available for study by the Museu Nacional do Brasil: Serra da Bocaina, 1300 meters, parquede criagao de trutas, Sao Paulo, Brazil, III-54, D. Albuquerque e Rego Barros, male (MN) ; Reserva do Museu, Santa Tereza, Espirito Santo, Brazil, 13-XI-55, N. Santos, J. Machado, A. Barros, 3 males, 4 females (male, 2 females, SGJ; 2 males, 2 females, MN) ; Alem Paraiba, Minas Gerais, Brazil, J. Morjen, female (MN). These differ in being darker in color with brownish wing veins, the costal veins darkest, and in having the median notch in the subgenital plate of the female somewhat greater in depth than the lateral notches.
Literature Cited
Jewett, S. G., Jr.
1958. Stoneflies of the genus Anacroneuria from Mexico and Central America (Plecoptera). Amer. Midi. Nat. 60(1): 159-175.
1959. Some stoneflies from Santa Catarina, Brazil (Plecoptera). Ibid. 61(1): 148-161.
Los Angeles County Museum
Exposition Park
Los Angeles 7,
MBER 37
0 7. 73 7-L^k
May 19, 1960
A NEW GENUS AND SPECIES OF GLOSSOPHAGINE BAT FROM COLIMA, MEXICO
By
W. J. SCHALDACH1 AND CHARLES A. McLAUGHLIN2
During the course of field work in the State of Colima, western Mexico, in August, 1958, three adult male long-nosed bats resembling Choeronycteris mexicana Tschudi were taken by Schaldach near Pueblo Juarez. Subsequent collections by A. L. Gardner in August and September, 1959, yielded ten more specimens of this bat. Close examination has shown these specimens to belong to a heretofore unknown genus and species, which may be described as follows:
SUBFAMILY GLOSSOPHAGINAE Musonycteris gen. nov.3
Genotype: Musonycteris harrisoni sp. nov.
General characters: Allied with Choeronycteris , which it resembles superficially, but with distinct cranial characters.
Diagnosis: Skull among the longest for the subfamily due to great elongation of the rostrum; cheek teeth narrow and delicate in structure, with reduction of lingual elements; nasals elevated dorsally; hamular processes of pterygoids enlarged, extending onto ventral surface of auditory bullae to level of anterior margin of meatus; basioccipital and basisphenoid produced into high, thin median ventral crest; teeth widely separated, the space between p2 and p3 approximately length of p2.
Dental formula: 2123
— - — - — - — - X 2 = 30 0 13 3
Comparisons: The rostrum of Musonycteris represents more than one-half the total length of the skull, and is longer than that of any other
Research Investigator, Los Angeles County Museum.
2Associate Curator, Ornithology and Mammalogy, Los Angeles County Museum. 3Derived from the generic names of the banana {Musa) and a bat (Nycteris) .
INSTITUTION JON I
2
Contributions in Science
No. 37
genus of glossophagine bat save the South American genus Platalina. Platalina , however, is a larger bat (Sanborn, Zool. Ser. Field MNH, vol. 24, no. 25, Jan. 6, 1943) and is further distinguished from Musonycteris by having two lower incisor teeth. Musonycteris is apparently most closely related to Choeronycteris as indicated by size, general appearance, general skull structure, and structure of the wing skeletal elements. It differs from that genus, however, as follows (see figs. 1-3 and table 1) : longer rostrum, representing more than 50% of the total length of the skull as contrasted with approximately 40% in Choeronycteris ; greater elevation of the dorsal surface of the nasals, which are almost flat in Choeronycteris; more delicate structure of the teeth; wider diastemae between the teeth; a high, thin crest on the mid-ventral line of the basioccipial and basisphenoid rather than a low rounded crest; more greatly elongate and inflated hamular processes which extend posteriorly to a line connecting the
Fig. 1. Lateral view of the skulls, X 3: A. Musonycteris harrisoni , adult male, holotype, LACM No. 11480. B. Choeronycteris mexicana , adult male, LACM No. 9971 from Mitla, Oaxaca, Mexico.
1960
SCHALDACH & McLaUGHLIN: MEXICAN BAT
3
anterior margins of the meati, rather than merely touching the bullae; lack of a mid-ventral, posterior continuation of the internasal septum as a low ridge on the roof of the interpterygoid fossa as found in Choeronyc- teris; anteriorly “V”-shaped rather than “U”-shaped interpterygoid fossa.
Musonycteris harrisoni sp. nov.4
Type: Los Angeles County Museum No. 11480, adult male, skin with skull, collected by A. L. Gardner (orig. no. 245) from 2 km. southeast of Pueblo Juarez (formerly Hacienda La Magdalena), Colima, Mexico, Sept. 5, 1959. Measurements of type: total length, 85 mm.; tail length, 9 mm.; hind foot length, 10 mm.; ear from notch, 18 mm.; tragus, 9 mm.; forearm length, 42.9 mm.; greatest length of skull, 35.2 mm.; length of maxillary tooth row, 14.1 mm.; dorsal length of rostrum (anterior midline of nasals to line connecting supraorbital foramina), 18.3 mm.
Range: So far as known, only from the type locality and one adjacent area (see table 1).
Diagnosis: Same as for genus.
Description: Color: Posterior dorsal region between Mummy Brown5 and Clove Brown, lightening on middle back and shoulder region toward brownish light drab; bases of hairs between Avellaneous and white; underparts like shoulder region. Size (average of 11 adult males from the type locality): total length (10 individuals), 85.2 mm.; tail length, 9.7 mm.; hind foot length, 10.5 mm.; ear length, 16.6 mm.; tragus length 5.6 mm.; forearm length, 42.29 mm.; greatest length of skull, 34.46 mm.; maxillary tooth row, 13.36 mm.; interorbital breadth, 3.89 mm.; mastoidal breadth, 9.71 mm.; palatal length, 22.71 mm.; dorsal length of rostrum, 17.52 mm.; percentage dorsal length of rostrum to greatest length of skull, 50.96 per cent; mandibular length, 25.80 mm. Skull: rostrum extremely long, averaging over one-half of the greatest length of the skull; nasals long and nearly parallel-sided, arched above the molars to form a prominent hump; cheek teeth delicately formed with much reduction of lingual elements; teeth well separated; interpterygoid space “V”-shaped anteriorly; internasal septum completely internal, without a ridge continuing posteriorly onto the roof of the interpterygoid fossa; hamuli enlarged, in close contact with the auditory bullae, with tips extending noticeably posterior to a line connecting the anterior margins of the auditory meati; basioccipital and basisphenoid produced into a high, thin median ventral crest; lower jaw elongate with teeth well separated; lower incisors absent.
Comparisons: Same as for genus.
Remarks: All the specimens of Musonycteris were caught in nylon “mist” nets set across a small irrigation ditch that runs through a banana
4Named in honor of Ed N. Harrison, in recognition of his interest in, and generous support of Schaldach’s field work.
“Names of colors are capitalized when direct comparison has been made with Ridgway’s “Color Standards and Color Nomenclature,” 1912.
TABLE 1
q}Su8T; uijb9.io,£
OS o csi csi
CO o csi csi d '3-
Iinqs jo qjSuoT; }S9jB9Jf)/q}Su9q[ jBJjsoy; 9Sb}U99J9<i
t>- CO
o d
vo vn
|
cq |
CO |
vq |
CSI |
|
|
aiqtpuBK |
cd |
cd |
vd |
cd |
|
cq |
CSI |
csi |
CSI |
■*-*
o
s
o
k.
©
"Si
O
03
S
cd
qjSuoT; IBaisaa; iBsaoa
qjSuaT IBJBIBd
qjp"B9Ja lopiojSBH
qjpBaag; jBjiqjoaajui
CO os cd oi csi csi
CO
d d
N. O
CO
CSI CO
00 t>-
csi oi csi csi
o o d d
qjSu9T; avoj qjooj .£jBnixBj\[
d P
rj- CO
cq vq co* co
53
-si
IinqS
jo qjSuoT; }S9JB9J£)
qjSu9i; ijbj,
O 00
d P
CO CO
qjSu9T IBjox
X9S
VO o 00 00
CSI
00 00
2 s s s
c d
13
V. r.
« N CD
O
a
jo
o
o
ffi
d pH
£2 Cj O
8 S
a P
U U < <1
u u <! <
AJIIBOOT PUB UOXBJ,
LACM 11481 (Paratype) M 88 10 34.5 13.2 4.1 9.6 22.7 17.6 26.0 51.1 42.6
LACM 11482 (Paratype) M 87 10 34.4 13.7 3.8 9.7 22.6 17.7 25.7 51.5 41.8
LACM 11483 (Paratype) M 89 10 34.2 13.3 3.8 9.6 22.3 17.6 25.0 51.5 42.8
LACM 11484 (Paratype) M 89 10 34.0 13.1 3.9 9.4 22.5 17.5 25.6 51.5 42.2
LACM 11485 (Paratype) M 84 9 34.4 13.3 4.1 9.9 22.6 17.2 25.5 50.0 41.8
|
in |
o |
cq |
co |
CO |
—1 |
oo |
p |
p |
CO |
p |
tN |
P |
p |
P |
O |
||
|
ci |
CO |
P |
CM |
P |
cd |
id |
id |
P |
cd |
id |
cd |
id |
id |
P |
cd |
P |
ci |
|
Tj- |
-'f- |
P |
Th |
Xh |
-'3- |
Tj- |
■'}■ |
-'f- |
^3- |
P |
"j- |
||||||
|
tN |
o |
o |
00 |
p |
co |
CO |
p |
io |
co |
o |
CO |
P |
00 |
tN |
tN |
||
|
o' |
d |
d |
d |
d |
P |
CO |
CO |
i |
00 |
d |
P |
p |
d |
00 |
d |
00 |
d |
|
in |
"3- |
^3- |
m |
X |
CO |
^3- |
tF- |
CO |
CO |
co |
co |
co |
|||||
|
cq |
in |
O) |
p |
CO |
o |
05 |
tN |
o |
p |
p |
05 |
||||||
|
in |
CO |
CO |
cd |
p |
P |
tH |
P |
p |
CO |
d |
P |
p |
cd |
P |
P |
ci |
d |
|
CM |
CM |
co |
co |
co |
CM |
CO |
CO |
CO |
co |
co |
co |
CO |
co |
CO |
CO |
co |
co |
|
cq |
o |
00 |
cq |
co |
CO |
tN |
CO |
tN |
CO |
co |
CO |
In |
tN |
p |
|||
|
In |
CO |
in |
tN- |
oo |
ci |
CO |
co |
co |
p |
CO |
ci |
CO |
ci |
P |
ci |
p |
P |
|
CO |
05 |
00 |
05 |
CO |
p |
p |
tN |
p |
p |
cq |
oq |
tN |
■'J; |
p |
|||
|
csi |
d |
o |
co |
00 |
oo |
00 |
00 |
00 |
00 |
oo" |
00 |
00 |
d |
In |
tN |
00 |
In |
|
CO |
CO |
co |
co |
||||||||||||||
|
in |
CO |
CO |
o |
00 |
p |
"3- |
in |
00 |
|||||||||
|
d |
d |
d |
d |
d |
d |
d |
d |
1 |
o |
d |
d |
d |
P |
d |
d |
d |
d |
|
'r_l |
T"H |
rH |
'r— 1 |
||||||||||||||
|
tN |
cq |
cq |
in |
p |
oo |
p |
cq |
cq |
, |
co |
. |
p |
p |
p |
|||
|
CO |
CO |
P |
cd |
cd |
cd |
cd |
cd |
cd |
cd |
P |
P |
P |
cd |
P |
P |
P |
|
|
l |
m |
o |
CO |
CO |
cq |
co |
cq |
o |
in |
00 |
P |
05 |
p |
05 |
|||
|
c6 |
CM |
CO |
cd |
P |
TH |
P |
P |
P |
P |
P |
d |
P |
P |
d |
|||
|
T— 1 |
T-1 |
||||||||||||||||
|
CO |
o |
CO |
o> |
o |
o |
cq |
-'f- |
00 |
00 |
CO |
co |
co |
o |
||||
|
p |
co |
CM |
P |
d |
d |
d |
d |
i |
d |
d |
d |
d |
P |
d |
d |
d |
d |
|
CO |
co |
CO |
co |
co |
co |
co |
co |
co |
co |
co |
co |
co |
CO |
co |
CO |
co |
|
|
CO |
CO |
co |
co |
co |
In. |
00 |
05 |
00 |
co |
||||||||
|
in |
In. |
CO |
o |
o |
CO |
00 |
o |
00 |
CO |
in |
co |
tN |
05 |
-'f- |
|||
|
00 |
00 |
00 |
CO |
CO |
05 |
00 |
tN |
00 |
tN |
tN |
0-5 |
00 |
00 |
tN |
In |
00 |
|
|
§ |
Ph |
Ph |
s |
p |
Ph |
Ph |
Ph |
pH |
Ph |
s |
Ph |
P P P
S3 8
05
a
p
CO
f-H
co
Oh
|
N |
N |
N |
£> |
|
"k |
* Jh |
’k |
|
|
<! |
<1 |
< |
|
|
to |
<j |
|
u |
°s “5 «. . CO |
S u |
k •N O X |
o U <0 V5 |
o U Q) C/5 |
o u <0 C/5 |
o U 05 C/5 |
fl ci o CJ |
d o u |
d d o u |
SI d k U |
QJ d |
|
<1 |
g.S |
< |
R O |
P |
P |
P |
P |
co |
co r-< |
CO |
CO |
k O |
|
p |
1-^ |
p |
k |
CJ |
CJ |
CJ |
CJ |
s |
S |
s |
d |
d |
|
tJ-U |
o |
u |
u |
u |
CJ |
id |
p |
p |
cO CO |
o CO |
S S S S £
o o
x x
CO CO
O O
6
Contributions in Science
No. 37
grove. This grove lies about 2 kilometers to the southeast of Pueblo Juarez, Colima, Mexico. The grove is situated in Arid Thorn Forest, but is adjoined closely by an extensive stand of climax Tropical Deciduous Forest. A large, open corn field, or milpa, forms one edge of the grove. The banana trees in the grove were in flower at the time that the first three specimens were collected by Schaldach (August, 1958), and it is assumed that these bats were feeding on the nectar of these flowers.
The net wherein the first three specimens were caught was set at right angles to the stream flow through the thickest patch of banana trees. Other nets (5 in all) were set the same and subsequent nights, both upstream and downstream from this site, but they took only Glossophaga soricina leachii and Artibeus j. jamaicensis. Five Glossophaga and seven Artibeus were also taken in the same net with the new bat. The Musonyc-
Fig. 2. Dorsal view of the skulls, X 3: A. Musonycteris harrisoni, adult male, holotype, LACM No. 11480. B. Choeronycteris mexicana , adult male, LACM No. 9971, from Mitla, Oaxaca, Mexico.
1960
ScHALDACH & McLAUGHLIN: MEXICAN Bat
7
teris were entangled quite high in the net, about five feet above the ground. All were taken about 10 p.m.
During the process of preparation of these specimens, the extremely long tongue of these animals was noted. The tongue of one of the animals measured 76 mm. from the gape of the jaws to the outstretched tip (held by a pair of forceps). The total length of the bat itself measured only 80 mm. One of Schaldach’s Mexican assistants immediately applied the name “ murcielago trompudo ” to these bats. The authors think, however, that this was a newly coined name to describe the trumpet-shaped snout of the animal, rather than a previously accepted vernacular name.
For the loan of comparative material of Choeronycteris we are indebted to Dr. E. Lendell Cockrum of the University of Arizona (U.A.). Grateful acknowledgment is also made to Ing. Juan Lozano Franco, of the Direccion General Forestal y de Caza, Mexico, D.F., under whose kind
Fig. 3. Ventral view of the skulls, X 3: A. Musonycteris harrisoni , adult male, holotype, LACM No. 11480. B. Choeronycteris mexicana, adult male, LACM No. 9971, from Mitla, Oaxaca, Mexico.
8
Contributions in Science
No. 37
auspices our federal collecting permit was obtained. All specimens without designation are in the collection of the Los Angeles County Museum.
Specimens examined: Thirteen Musonycteris harrisoni from Colima: type locality, 12; 4 kms. so. of Cerro de Ortega, 1. Sixteen Choeronycteris mexicana from the following localities: Arizona, Cochise County, 1 mi. north of Paradise, 3 (U.A.) ; South Fork Cave Creek, 1 (U.A.) ; Pima County, vicinity of Tucson, 4 (U.A.) ; Santa Cruz County, 5 mi. north, 2 mi. west of Patagonia, 1 (U.A.) ; Mexico, Sinaloa, 6 mi. east of Santa Lucia, 1; Sonora, 11 mi. by road northeast of Imuris, 1 (U.A.) ; Oaxaca, 6 mi. west of Mitla, 5.
Los Angeles County Museum
Exposition Park
Los Angeles 7, G
tfBER 38
May 19, 1960
>7- 73
A CENSUS OF THE ABUNDANT LARGE PLEISTOCENE MAMMALS FROM RANCHO LA BREA1
By Leslie F. Marcus2
Introduction
The Pleistocene fossil deposits at Rancho La Brea, now famous throughout the world, are chiefly represented by specimens in the Los Angeles County Museum. This large assemblage of materials makes possible the taking of a census of the representations from the several different pits.
Early excavators observed apparent differences in the faunas and characteristics of the separate pits of Rancho La Brea, and critical study has subsequently underscored several of these features. For example, the abundance of Proboscidea in Pit 9 (Fig. 1) compared to their near exclusion in other pits has been pointed out (Stock, 1956) as well as the presence of human remains in Pit 10. Such observations have indicated the desirability of thoroughgoing pit censuses, since small differences between the several individual pits may be detected by comparing the numbers of individuals of various species that are preserved. Thus far, only two reports have appeared in which comparisons of vertebrate remains from separate pits have been made: Howard & Miller’s (1939) bird census of the Los Angeles County Museum pits 3, 4 and 10 ; Bratt- strom’s (1953) comparison of the lower vertebrate faunas.
The present census was undertaken to discover whether there are detectable differences, among the major pits, in the faunas of the most
^his paper was prepared with the partial support of the Office of Naval Research (Nonr -222-43). This paper in whole or in part may be reproduced for any purpose of the United States Government.
2University of California, Berkeley.
SMftTHSMNiftN
imwmtmm
JUK I
2
Contributions in Science
No. 38
abundant large mammals. The results have indicated that faunal differ- ences from pit to pit do exist. Furthermore, the differences appear too large to be accounted for by chance, and may more reasonably be explained as the result of differences between pit activities, such as temporal differences or pit preferences by certain species. More complete censuses of the separate pits for all elements of the faunas, as well as detailed studies of the faunas and their component species, will be necessary before a realistic mathematical model can be proposed to explain the pit differences.
Dr. Theodore Downs and Dr. Hildegarde Howard kindly made available to me for study the Rancho La Brea collection and records of the Los Angeles County Museum. I am indebted to them for many helpful suggestions.
Available Data
The present census is based entirely on the collection of the Los Angeles County Museum.3 Only the more abundant species, represented by 18 or more individuals, were usable in this comparative analysis of the separate pits. These species, all of them extinct, are Paramylodon harlani, N othr other ium shastense, Cards dims, Canis orcutti, Panthera atrox, Smilodon calif ornicus, Camelops hesternus, Breameryx minor. Bison antiquus, and Equus occidentalis. Canis furlongi, the extinct timber wolf, is distinguishable with difficulty on other than cranial characters, and only eight skull specimens are known (Stock, 1956 and Nigra and Lance, 1947). The numbers of individuals of the rarer large species, not included here, are given by Stock (1929). The numbers of individuals of T remar ctotherium simum and Mammut americanus may exceed 18 individuals, but they are most abundant in pits other than those from which the more abundant materials were collected (see below). About half of the species of large Rancho La Brea mammals are catalogued in a series of permanent ledgers, which indicate catalogue number, species identification, skeletal element, pit number and coordinates within pit for each specimen. Some of the rarer large mammals are catalogued in an older, card-catalogue. Both catalogues were used to make the pit censuses. A direct count of bones was made only for Bison, which is incompletely catalogued, and for Camelops as a verification of the catalogue. Paramy- lodon, N othrotherium, and Breameryx are listed only in the older cata- logue. It would be a prodigious task to count the numbers of individuals for every species using the collection itself. Stock (1929) did this for his total census, except that the numbers of the individuals for the two largest groups, the dire wolves and the sabre tooth cats, were only approximated. In Stock’s census only adult animals, in general, were considered; in the present census, juveniles may have been included, since the catalogue
3The University of California collection is incompletely catalogued, and a portion of it has been sent to other institutions.
1960
Marcus: Pleistocene Mammal Census
3
does not distinguish them from adults. The census figures were also compared to the field notes where, in most cases, the number of skulls discovered in each separate pit was recorded. In only a few cases is there a discrepancy towards a higher number in the field notes. In some cases, specimens were recorded in the notes as being discovered, but were not collected because of poor preservation.
The term pit, as used in this report, refers to the 96 numbered excavations made by the Los Angeles County Museum from July, 1913, to September, 1915. Over fifty of these excavations were mere test holes, completely unproductive; others, such as Pits 1, 2, and 90 were pockets in the walls of the earlier excavations made by the University of Cali- fornia; Pits 61 and 67 were found to unite into one large pit, and several numbered areas represented test digs at different spots in previously tested outcrops. Only fourteen to sixteen of the ninety-six excavations, according to the field notes, appear to have been actual self-contained deposits which probably represented tar traps for the larger mammals
Fig. 1. Topographic map of Rancho La Brea showing location of principal excavations. Diagram modified after Stock.
4
Contributions in Science
No. 38
sometime in the past (see Fig. 1). In seven of these, Pits 9, 17, 43, 44, 57, 65 and 72, the bone was of such poor preservation, much of it water soaked and rotten, that collecting was unsatisfactory and only a few specimens were taken. A complete record of the species from these pits is not indicated in the catalogue. Pits 81 and 91 were kept intact for display purposes and no complete count was made of the contained animals. Pit 36 was a rather special, shallow deposit measuring only 2 ft. X 4 ft. at a depth of 6 feet. Its fauna is so small that its inclusion in the discussion was considered impractical. Pit 10 is also excluded from the discussion because, for the most part, it contained small forms suggesting Recent age. Only the birds of Pit 10 have been studied in detail (Howard and Miller, 1939) ; the few isolated bones of large mammals (bear, wolf, horse and deer) need further study for accurate identification (see Merriam, 1914, and Stock, 1956).
It may now be recognized that only Pits 3, 4, 13, 16, 60, 61-67, and 77 are useful for comparison of census figures. These seven pits, here to be analyzed, will hereafter be referred to as the “major pits” by reason of the abundance of material contained. In some of these pits, the upper few feet of material was discarded before complete collection was attempted. During excavation the bones were segregated as to depth and position below a three-foot grid at the surface. Accordingly, the deposit may be reconstructed in detail from the catalogue or the museum collection, and more detailed census studies of the fauna may be made utilizing these data. Depths in these major pits ranged from 17 feet in Pit 13, to 27 feet in Pit 3;in Pits 16, 60, and 61-67 the greatest depth was 20 feet, in Pit 77, 21 feet, and in Pit 4, 25 feet.
Procedure
The present census was made in the following way. For each species in each pit, a tally was made of one skeletal element, for example, the right calcaneum. This count was repeated for all other elements for which there seemed to have been a good chance of preservation and collection and a small chance of loss or breakage in accumulation. The tallies for the separate elements, or bones, were reviewed and the total number of individuals in a pit was estimated from the number of specimens of the most frequently preserved element in that pit. For example, for Smilodon in Pit 3, there were 339 atlases and 254 right calcanea, whereas in Pit 4 there were 94 atlases and 105 right calcanea. Thus, the estimates for the total number of individuals collected from Pits 3 and 4 were taken
aTotal census, using most common element per pit as explained in text. bTotal number of most common element over all excavations.
() Numbers in field notes.
*01d catalogue.
** Actual counts.
£
o
^ C/5
CO
3
co
vn
oo
vn
s
3
05
cC
o
"cC
U
CO 05
O VO
CM
05 <0
oo
CM
o
X
00
c n
ca
O) tJ- -3- 05
oo oo o
w
w
H
s
%
I>
Tf
VO
#
CO
O OO
CM oo
1-0
O 05 t^.
CM <o OO
CM 05
Ct 05 xo
00
CO 00
CO ^3-
cm
o
CO VO
oo
CM
OO
CM co oo
Tj- tJ- ^ CO
oo oo et ^ CM CM
oo CM OO CO vn
CO
s
U
W
Ph
CO
V-5
Co
Total individuals 1040 798 495 422 313 102 86 3256 3447 3052
6
Contributions in Science
No. 38
as 339 and 105, respectively. Such inconsistencies between different elements of a species within a pit were noted in several cases and may, perhaps, be accounted for by incomplete preservation of individuals or by breakage or loss in collection. The method used, however, should provide a dependable count of the number of individuals actually preserved in a pit, provided that one individual did not have parts of its skeleton preserved in more than one pit. The census for each species in each major pit, using this method, is given in Table 1. These are the estimates used in the discussion to follow. As a matter of interest, the total census of each species for all the seven selected pits (using the same method) is given in the column “All Excavations, Total”, and may be compared (in the column “Most Common Element, Total”) with the census of each species for all pits based on the most frequent element throughout the pits. These latter estimates are not affected by the preservation of individuals in more than one pit, and, in most instances, give a lower total census figure. Stock’s (1929) total estimates were made in this way.
Results
For a meaningful comparison of the numbers of individuals from different pits, it must be assumed either that estimated numbers of individuals collected, housed, and catalogued in the Museum are very near the actual numbers of individuals trapped and preserved in the pits, or that they faithfully represent the relative abundance of species. This assumption implies that after entrapment there was no selective loss of individuals, and that in collection and curation there was no selection of specimens of one species over those of another in different pits. Counts based on less obvious elements such as carpals, tarsals, metapodials, and vertebrae should be less subject to such selection if it existed.
In Table 1, the estimated total number of individuals of the abundant species of large mammals is entered for the seven major pits. Counts based on field notes are entered in parentheses.
The predominance of Canis dims over Smilodon calif ornicus is evident in each of the major pits, though the relative proportions of these two to each other and to the rest of the fauna are quite variable. These two species, plus Panthera atrox , outnumber the herbivores from 5.5 to 1 (in Pit 4) to 7.8 to 1 (in Pit 61-67). Canis dims outnumbers Smilodon californicus by ratios from 1.1 to 1 (in Pit 77) to 3.0 to 1 (in Pit 16). Canis orcutti is especially abundant in Pit 16, and N othrotherium is relatively abundant in Pit 61-67. Other comparisons can be made by examining Table 1 and figure 2.
A map of the area is given in figure 1. No consistent pattern of relationship between pit faunas has been discovered that can be correlated with pit location. The seven separate major pits are considered as individual locations of accumulation in the discussion which follows.
1960
Marcus: Pleistocene Mammal Census
7
Discussion
Merriam, Stock, and others viewed the tar traps as places where unwary herbivores, especially the very young and the very old, were trapped while in search of water in the open pits, or while crossing innocuous looking, dust-covered pits. Once an animal was trapped, its cries attracted carnivores. These in turn were trapped in greater numbers than their prey. Later, carrion feeders came on the scene, and they, too, fell victim to the deceptive pits.
The above is the only reasonable explanation that has been offered for the preponderance of carnivorous animals in the pits. Before the present census counts were made, this explanation was elaborated into a mathematical model, which, it was hoped, could be used to explain the results of the census and permit estimates of the relationships between predators and prey. The testing of the model had to be abandoned when the present study revealed that of the 96 excavations, only five, or at most seven, were useful for the purpose. Instead, the hypothesis of similar proportional faunal composition among the several pits was tested using the X2 test for deviation, corresponding to the test of association in Simpson and Roe (1939, p. 290). If all pits attracted all species equally, we would expect to find approximately the same propor- tional relationship of the separate species in each pit, and a consequent low factor of deviation (X2). For example, Canis dirus should be in nearly the same dominant proportion in each pit. The pits with the greater numbers of individuals would be expected to be more similar in proportions. The large observed deviations from equal proportionality, however, lead one to favor alternative hypotheses, such as preferential attraction of some species to certain pits, or differences in times of activity for the different pits. A large number of one species in a pit might be correlated with its abundance in the area during the time of major activity of the pit.
The observed value of X2 for the ten most abundant large species in the seven major pits is 349.2. A deviation value as large as this, under the hypothesis of similar proportionality, would be expected rarely (P < .00001, Hartley and E. S. Pearson, 19504), and reflects considerable dissimilarity in accumulation of the faunas of the separate pits. Similarities of proportions for the herbivores and carnivores were tested separately. The six herbivores gave a X2 equal to 55.7 for the seven major pits, again a large deviation under the hypothesis of similarity (P = .003). When N othr other ium is removed from the analysis, however, X2 is equal to 27.4 which is not significant (P = .29). Therefore, the proportions of large herbivores other than Nothrotherium are not significantly different, which may mean that abundance and entrapment of these species were not different during the times of activity of the several pits. The presence or absence of Nothrotherium is probably significant in itself, considering
4The lowest probability figure used statistically.
1040
86
paramylodon
NOTHROTHERIUM
CAMELOPS
BREAMERYX
BISON
EQUUS
PANTHERA ATROX SMILODON CANIS ORCUTTI CANIS DIRUS
Fig. 2. Pie diagrams giving the proportions of the ten most abundant species for the seven major pits. The number below each pie indicates the total number
313
of individuals of these species from that pit. The areas of the pies are proportional to these total pit numbers, (see Table 1).
10
Contributions in Science
No. 38
the size of the faunas of the large pits. The four carnivores give a X2 equal to 281.3 for the seven major pits, and this is highly significant (P < .00001). Even when Canis orcutti is removed from the analysis a highly significant X2 value of 65.3 is obtained (still P < .00001). The carnivores contribute a major part to the deviation from the expected numbers under the hypothesis of similarity. They do not seem to follow the variation in the herbivores in any systematic manner, except for a possible association of dire wolves and bison.
Summary and Conclusions
The present census does not alter the general picture of accumulation at Rancho La Brea. It does indicate that the separate pits had nearly the same fauna, but differed significantly in numbers of individuals of some of the species. The proportion of carnivores to herbivores is consistently greater, but varies from pit to pit. In no instance, however, do the present figures indicate as great a predominance of carnivores over herbivores as noted by Stock (1929), even allowing for the fact that several of the small carnivores used in Stock’s totals, are not included here.
Differences in physical characteristics of the pits are probably insufficient to explain the census differences pointed out here, although the associated plant life might have influenced the variable abundance of N othr other ium. Rather, I favor the hypothesis that the pits were not all continuously active for the same period of time. Either each pit was continuously active for a different interval of time, or the separate pits’ periods of quiescence and entrapment were not coincident even though their total times of activity may have been roughly equivalent. Complete censuses of all faunal and floral elements for all pits will cast more light on this subject. The smaller mammals, the birds and the reptiles should be particularly useful. The majority of them have not become extinct as have the larger mammals. The value of a bird census has already been pointed out by Howard and Miller (1939) in supporting a younger age for Pit 10, which contains human remains. At the same time, these authors presented avian census differences between Pits 3 and 4 (op. cit., figs. 1 and 2) that indicate probably significant differences between these pits. Herpeto- logical material has not been studied in the major pits other than Pit 3 (Brattstrom, 1953). Brattstrom did, however, suggest an age difference in the pits he studied, placing Pit 3 and two other lesser Los Angeles Museum excavations together with the University of California excavation no. 2051 as probably older than L. A. Museum Pits A and B and Univ. of Calif, excavation no. 2052.
The work of Nigra and Lance (1947) and Menard (1947) on measurements of length of the metapodials of the dire wolf and sabre tooth cat, respectively, contributes another interesting facet to the evidence for inter-pit differences. These authors show that in average length of
1960
Marcus: Pleistocene Mammal Census
11
metapodials, the animals from Pits 4 and 13 show consistent separation, with noticeable, though less consistent, differences between the individuals of Pits 77, 61-67 and 3. If the time differences between the pits can be determined, the collection from Rancho La Brea will provide a unique opportunity for the study of evolution in several species over a a relatively short time.
LITERATURE CITED
Brattstrom, Bayard H.
1953. The amphibians and reptiles from Rancho La Brea. Trans, of San Diego Soc. Nat. Hist. 11 (4) : 365-392, Figs. 1-4.
Hartley, H. O., and E. S. Pearson
1950. New tables of statistical functions. Biometrika 37: 168-225.
Howard, Hildegarde, and A. H. Miller
1939. The avifauna associated with human remains at Rancho La Brea, California. Carnegie Inst, of Wash., Pub. No. 514: 39-48. Cont. to Paleo. III.
Menard, Henry W., Jr.
1947. Analysis of measurements in length of the metapodials of Smilodon. So. Calif. Acad. Sci., Bull. 46: 127-134.
Merriam, John C.
1914. Preliminary report on the discovery of human remains in an asphalt deposit at Rancho La Brea. Science, N.S. 40 (1023): 198-203.
Nigra, John O., and John F. Lance
1947. Analysis of measurements in length of the metapodials of Canis dirus. So. Calif. Acad. Sci., Bull. 46: 26-34.
Simpson, G. G., and Anne Roe
1939. Quantitative zoology. McGraw Hill, New York and London. XYII -j- 414 pp.
Stock, Chester
1929. A census of the Pleistocene mammals of Rancho La Brea, based on the collections of the Los Angeles Museum. Jour, of Mammalogy, 10 (4): 281-289. 3 Figs.
1956. Rancho La Brea — A record of Pleistocene life in California. Los Angeles County Museum, Science Ser. No. 4, Paleo. No. 4.
Wyman, L. E., et al
1915. A record of excavations from July, 1913 to September, 1915. Field Notes, Rancho La Brea. (Manuscript on deposit in Los Angeles County Museum.)
LOS ANGELES COUNTY MUSEUM CONTRIBUTIONS IN SCIENCE
The Machris Brazilian Expedition
No. 1. General Account, by Jean Delacour.
No. 2. Botany: General, by E. Yale Dawson.
No. 3. Botany: A New Dodder from Goias, by T. G. Yuncker.
No. 4. Botany: The Lichens, by Carroll W. Dodge.
No. 5. Botany: Cyanophyta, by Francis Drouet.
No. 6. Botany: A New Mint from Goias, by Carl Epling.
No. 7. Botany: Phanerogamae, various smaller families, edited by E. Yale Dawson.
No. 10. Botany: A New Columnar Cactus from Goias, by E. Yale Dawson.
No. 11. Botany: Chlorophyta; Euglenophyta, by G. W. Prescott.
No. 12. Entomology: General; Systematics of the Notonectidae (Hemiptera), by Fred S. Truxal.
No. 13. Botany: Phanerogamae, Leguminosae, by Richard S. Cowan.
No. 14. Entomology: Gelastocoridae (Hemiptera), by E. L. Todd.
No. 17. Botany: Phanerogamae, Bromeliaceae and other smaller families, by Lyman B. Smith.
No. 18. Botany: Musci, by Howard Crum.
No. 21. Botany: Phanerogamae, Euphorbiaceae, Lentibulariaceae, Rubiaceae, by Julian A. Steyermark.
No. 22. Botany: Gramineae, by Jason R. Swallen.
No. 23. Botany: Phanerogamae, Alstroemeriaceae and other families, by Lyman B. Smith and collaborators.
No. 24. Botany: Fungi, by G. W. Martin and collaborators.
No. 26. Botany: Hepaticae, by Margaret Fulford.
No. 28. Botany: Phanerogamae, Melastomataceae and Polygalaceae, by J. J. Wurdack.
No. 30. Botany: Phanerogamae, Amaranthaceae and other families, by Lyman B. Smith and collaborators.
No. 32. Botany: Phanerogamae, Acanthaceae, by Emery C. Leonard.
No. 33. Ornithology: Two new birds from Central Goias, Brazil, by Kenneth E. Stager.
No. 35. Botany: Pteridophyta, by C. V. Morton.
No. 36. Entomology: Two New Species of Anacroneuria (Plecoptera) from Goias, Brazil, by Stanley G. Jewett.
Other Subjects
No. 8. Notes on Eastern Pacific Insular Marine Algae, by E. Yale Dawson.
No. 9. A New Series of Passerine Bird from the Miocene of California, by Hildegarde Howard.
No. 15. Marine Algae of the Pacific Costa Rican Gulfs, by E. Yale Dawson.
No. 16. A Classification of the Oscines (Aves), by Jean Delacour and Charles Vaurie.
No. 19. A New Race of the Pocket Gopher Geomys bursarius from Missouri, by Charles A. McLaughlin.
No. 20. Further Bird Remains from the San Diego Pliocene, by Loye Miller and Robert I. Bowman.
No. 25. Miocene Sulids of Southern California, by Hildegarde Howard.
No. 27. Marine Algae from the 1958 Cruise of the Stella Polaris in the Gulf of California, by E. Yale Dawson.
No. 29. Quaternary Animals from Schuiling Cave in the Mojave Desert, Cali- fornia, by Theodore Downs, Hildegarde Howard, Thomas Clements and Gerald A. Smith.
No. 31. Late Pleistocene Invertebrates of the Newport Bay area, California, by George P. Kanakoff and William K. Emerson.
No. 34. A new Giant Water Bug from Mexico, by Arnold S. Menke.
No. 37. A New Genus and Species of Glossophagine Bat from Colima, Mexico, by W. V. Schaldach and Charles A. McLaughlin.
No. 38. A Census of the Abundant Large Pleistocene Mammals from Rancho La Brea, by Leslie F. Marcus.
Vmber 39
December 15, 1960
A SKULL OF THE GRIZZLY BEAR (URSUS ARCTOS L.)
FROM PIT 10, RANCHO LA BREA
By Bjorn Kurten
os Angeles County Museum
Exposition Park
Los Angeles 7, Calif.
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. Issues are numbered separately and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientists and scientific institutions on an exchange basis. Copies may also be purchased at a nominal price.
Hildegarde Howard Editor
E, Yale Dawson Associate Editor
A SKULL OF THE GRIZZLY BEAR ( URSUS ARCTOS L.) FROM PIT 10, RANCHO LA BREA
By Bjorn Kurten1
The grizzly bear, Ursus arctos L. (“ Ursus horribilis ” Ord2), has been regarded as a member of the Rancho La Brea late Pleistocene fauna since the report by Merriam (1912) of a large ursine atlas vertebra from the tar pits. The description of this specimen (op. cit ., p. 41) reads: “In form, size and position of the opening of the vertebrarterial canal the atlas specimen from Rancho La Brea is nearer to the black bear than it is to the grizzly . . . The animal represented by the ursine atlas from Rancho La Brea was about as large as a grizzly of average size . . Thus, it will be seen that the determination as a
grizzly bear is based on the size of the specimen, rather than on its morphological characters, which suggest Ursus americanus Pallas. The greatest transverse diameter across the anterior articular faces is given as 65.5 mm., which is in excess of the size of the living black bears, and much the same as in the grizzlies. The writer has observed, however, that the fossil forms of Ursus americanus reached much greater dimen- sions than the living ones. In three fossil skulls (from Papago Springs Cave, Arizona; Friesenhahn Cave, Texas; and Samwel Cave, California) the width across the occipital condyles is 60, 64, and 66 mm., respectively. The size of the Rancho La Brea atlas is, thus, within the range found in fossil Ursus americanus as well as that of the grizzlies. The black bear is also represented by other specimens from the tar pit fauna. There is, for instance, an excellent skull and jaw of a juvenile individual from Rancho La Brea. Though this skull, being immature, is of small dimen- sions, the large size of the teeth shows that it belongs to the typical Pleistocene form. The Rancho La Brea black bear was identified by Schultz (1938) with his Ursus optimus from McKittrick. This form can only be regarded as a subspecies of the black bear, and at present the best course is probably to unite all the large Ursus americanus of Wisconsin age under the oldest name, or Ursus americanus amplidens Leidy, which may thus be placed on record for Rancho La Brea.
There is now, however, an indubitable specimen of Ursus arctos from the Brea pits. This is a skull. No. 133, in the Los Angeles County Museum. Because of its relatively small size, the skull might be passed off as Ursus americanus , but a closer study leaves no doubt that it is
Zoological Institute, Helsingfors, Finland.
2On the specific identity of the grizzly and brown bears see, e.g. Erdbrink, 1953, and Rausch, 1953.
4
Contributions in Science
No. 39
Ursus arctos. The specimen, however, does not belong to the typical Rancho La Brea fauna; it comes from Pit 10, the locality at which human remains were discovered (Merriam, 1914). Since this appears to be the only certain record of the grizzly from the tar pits, and it may have a bearing on the age of the Pit 10 assemblage, it seems appropriate to offer some observations on the specimen.
The skull represents a relatively small individual of Ursus arctos , and the size of the canine teeth indicates that it is a female. The specimen is of a young adult with somewhat worn cheek teeth. The carnassials have been lost on both sides. The three anterior premolars are totally absent, and there is no trace of an alveolus. The complete absence of all three is rare in the grizzly, but it does occur in some specimens. According to a compilation in Erdbrink (1953, p. 372; original data from Hall, von Middendorff, and Degerb0l), this condition was found in one out of 81 skulls of Alaskan Ursus arctos , and in two out of 66 Eurasian Ursus arctos. The modal condition is the presence of P1, P3, and P4. In Ursus americanus , not a single specimen (out of a total of 100) was found to lack all the anterior premolars (Erdbring, op. cit ., p. 305, on data from Hall), and modally, all four premolars are present.
The skull from Pit 10 exceeds in size the living Ursus americanus , but a number of fossil black bear skulls attain similar or greater dimensions. The relative proportions of the skull and the teeth are, however, quite different from those found in the fossil black bears. This is perhaps most clearly demonstrated in a ratio diagram (fig. 1). The measurements are recorded in Table 1. The Recent grizzly is represented by five female specimens, of which four are from Canada and one from Wyoming; the means and observed ranges are given. The table also records the mean values for a small sample of fossil Ursus americanus amplidens, which is being discussed in more detail elsewhere (Kurten, MS), and the means for four Recent black bear skulls from the southern part of the species range (2 from Florida and 2 from Coahuila). The values for the Rancho La Brea grizzly have been used as a standard in the diagram.
It is evident that the Recent grizzly sample agrees very closely in dimensions with No. 133, and all the measurements of the latter fall within the observed range of variation in the Recent female grizzlies, except the rostral breadth; and this is but slightly greater in the fossil skull.
In the fossil Ursus americanus amplidens the length of the skull is about the same, but the cheek teeth are smaller, especially M2, and the width measurements between the orbits and over the postorbital processes are much greater. It is interesting to note that the relative proportions in the small. Recent Ursus americanus are somewhat more “arctos-like”
1960
Kurten: Fossil Bear
5
LOG DIFFERENCE SCALE
Fig. 1. Ratio diagram of dimensions of bear skulls and teeth. Standard (straight vertical), no. 133, Rancho La Brea Pit 10, Ursus arctos. Solid circles and horizontal bars show means and observed ranges of variation in 5 Recent female grizzly bears. Open circles represent Ursus americanus means; dashed lines connect those for the fossil sample; continuous lines, those for the four Recent samples.
Table 1
|
Dimensions of bear skulls and upper dentitions (in millimeters). GRIZZLY BEARS BLACK BEARS Fossil 5 Recent females Sample means Observed Measurement No. 133 Mean Range Fossil Recent |
||||||
|
Cs, width |
13.2 |
13.54 |
12.8 |
- 14.1 |
13.50 |
10.50 |
|
M1, length |
23.0 |
22.78 |
21.9 |
- 23.4 |
21.19 |
17.73 |
|
M1, width |
17.0 |
17.30 |
16.8 |
- 17.7 |
_ |
_ |
|
M2, length |
34.7 |
34.64 |
33.9 |
- 36.0 |
30.56 |
26.30 |
|
M2, width |
18.8 |
18.00 |
16.8 |
- 19.2 |
_ |
_ |
|
Skull, basal length |
290 |
284.8 |
272.0 |
- 293.0 |
301.0 |
244.0 |
|
Rostral width (at Cs) |
75 |
70.0 |
68.0 |
- 73.0 |
78.5 |
57.6 |
|
Muzzle width (at M2) |
83 |
82.4 |
78.0 |
- 86.0 |
81.4 |
67.8 |
|
Interorbital width |
67 |
66.4 |
64.0 |
- 70.0 |
82.0 |
58.8 |
|
Postorbital processes |
95 |
98.3 |
94.0 |
- 107.0 |
116.0 |
83.0 |
|
Postorbital constriction |
75 |
70.4 |
67.0 |
- 75.0 |
75.0 |
64.8 |
6
Contributions in Science
No. 39
than in the fossil (as shown by the closer approximation to a straight vertical in the diagram). The differences between the fossil and Recent black bears may partly be due to allometric growth.
The comparison between the avifaunas of Pit 10 and the other Rancho La Brea localities suggests that the assemblage of this pit is of somewhat later date, perhaps post-Pleistocene (Howard and Miller, 1939). The presence of an Ursus arctos indistinguishable from the living grizzlies may well be of importance in this context. There appears to be no convincing evidence that the grizzly existed here during the Rancholabrean sensu stricto, now known to extend to the late Wisconsin. It is indeed questionable whether the species entered temperate North America during the Wisconsin. The two grizzly skulls from Oklahoma described by Stovall and Johnston (1935) are stated to come from river gravels of Pleistocene age, but no more precise date was established. Ursus procerus Hay, considered by Stovall and Johnston to be a grizzly, is also an isolated find without any certain clue as regards the age. On the other hand, there are some finds of Ursus arctos which are definitely postglacial. It may also be noted that the Californian cave and tar pit faunas of late Pleistocene date, though rich in bears of the species Ursus americanus and Arctodus pristinus (“ Arctotherium simum \ etc.), do not yield any grizzlies. It seems, therefore, as if the grizzly may have entered this area only at the end of the Pleistocene or in postglacial time.
Literature Cited
Erdbrink, D.P.
1953. A review of fossil and recent bears of the old world. Deventer, 2 parts.
Howard, Hildegarde, and Alden H. Miller
1939. The avifauna associated with human remains at Rancho La Brea, California. Carnegie Inst. Publ. no. 514: 39-48.
Kurten, Bjorn
MS Fossil bears from Texas.
Merriam, John C.
1912. Recent discoveries of Carnivora in the Pleistocene of Rancho La Brea. Univ. California Publ., Bull. Dept. Geol., 7 (3): 30-46.
1914. Preliminary report on the discovery of human remains in an asphalt deposit at Rancho La Brea. Science, n.s. 40 (1023): 198-203.
Rausch, Robert
1953. On the status of some Arctic mammals. Arctic 6 (2):91-148.
Stovall, J. Willis, and C. Stuart Johnston
1935. Two fossil grizzly bears from the Pleistocene of Oklahoma. Jour. Geol. 43 (2): 208-21 4.
Fig. 2. Ursus arctos skull, L.A.C.M. no. 133 from Pit 10, Rancho La Brea; lateral, ventral and dorsal views, approximately 0.40 times natural size. Photo by Armando A. Solis.
LOS ANGELES COUNTY MUSEUM CONTRIBUTIONS IN SCIENCE
Nos. 1-34 Table of Contents available.
No. 35. The Machris Brazilian Expedition. Botany: Pteridophyta, by C. V. Morton.
No. 36. The Machris Brazilian Expedition. Entomology: Two new species of Anacroneuria (Plecoptera) from Goias, Brazil, by Stanley G. Jewett, Jr.
No. 37. A new genus and species of glossophagine bat from Colima, Mexico, by W. J. Schaldach and Charles A. McLaughlin.
No. 38. A census of the abundant large Pleistocene mammals from Rancho La Brea, by Leslie F. Marcus.
No. 39. A skull of the Grizzly Bear ( Ursus arctos L.) from Pit 10, Rancho La Brea, by Bjorn Kurten.
BER 40
THE MACHRIS EXPEDITION TO TCHAD, AFRICA
Amphibians and Reptiles
i
May 11, 1961
By David B. Wake and Arnold G. Kluge
Angeles County Museum
Exposition Park
Los Angeles 7, Calif.
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. Issues are numbered separately and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientists and scientific institutions on an exchange basis. Copies may also be purchased at a nominal price.
Hildegarde Howard Editor
E. Yale Dawson Associate Editor
THE MACHRIS EXPEDITION TO TCHAD, AFRICA
Amphibians and Reptiles By David B. Wake1 and Arnold G. Kluge1
During the dry season in the winter of 1960, a Los Angeles County Museum expedition under the sponsorship of Mr. and Mrs. Maurice A. Machris of Los Angeles visited the Republic of Tchad, Africa. The expedi- tion was headed by Mr. Machris, and Dr. Charles A. McLauglin, Associate Curator of Ornithology-Mammalogy at the Museum, and had as its prime purpose the collecting of birds and mammals. Incidental to his other work, Dr. McLaughlin obtained a small but important series of amphibians and reptiles. Virtually all collecting centered in the semi- desert and savanna country of east-central and southern Tchad. The localities where specimens were collected are indicated on the accompanying map (Fig. 1). Herpetological records for Tchad are few and the specimens collected by Dr. McLaughlin contribute to our knowledge of the distribution of various species, while several constitute major range extensions. The material from the expedition is deposited in the collections of the Los Angeles County Museum.
Acknowledgments
We wish to extend our appreciation to Mr. and Mrs. Maurice A. Machris who sponsored the field work and to Dr. Charles A. McLaughlin for the privilege of reporting on the collection and for providing valuable information concerning the specimens. We thank Dr. Georges Pasteur for information on the species of Tarentola and Mr. David DuVal for information on Geochelone. We especially extend our appreciation to Dr. Jay M. Savage for his aid in identification of the specimens and for the use of his personal library. Dr. Savage also read the manuscript and made many valuable criticisms.
Collecting Localities
Koro Toro — a desert outpost located in a sand dune area about 350 miles northeast of Fort Lamy. The dunes are covered and held by fine bunch grass and melon vines. With the exception of the last listed locality (Gongo), all are in the Sudanese Arid Zone. The elevation is about 700 feet (225 m.).
Fad A — a town located in the Ennedi Mountains. The surrounding area is pure sand, and the vegetation consists of bunch grass and scattered low acacias. Limestone and volcanic outcroppings are common. Water is found in small stream beds and in artificial ponds. The elevation of Fada is about 1800 feet (550 m.).
Oued Archei — -a wadi in the Ennedi Mountains near Fada. Vegeta- tion is scarce, but permanent ponds of water do exist.
department of Biology, University of Southern California.
oiyi Its loUIW-,IM
INSTITUTION
MAY
2
% 196
4
Contributions in Science
No. 40
Fig. 1. Map of the Republic of Tchad showing the localities at which amphibians and reptiles were collected.
1961
Wake & Kluge: Amphibians and Reptiles
5
Oum el Adam — a village located at an elevation of about 1300 feet (385 m.) in the desert in an area of drifting dunes with some rocky outcrops. The vegetation consists of scattered bunch grass and acacias.
Oum Chalouba — a town located in a semi-desert situation with overgrazed bunch grass as the dominant vegetation. Rocky outcrops are common in the area. The elevation is about 1400 feet (435 m.).
Arada- — a town located in an area with vegetation similar to that of the Oum Chalouba region. The elevation is about 1624 feet (497 m.).
Abeche — a town located on a rolling plain at about 2000 feet eleva- tion (600 m.). The semi-desert region is dominated by scattered patches of short grass and thorny acacias.
Gongo — a small village located on the north edge of the Aouk River (Chari River drainage) at a point 45 miles east-southeast of Fort Archam- bault. Collections were made in the surrounding Guinean Savanna, a tall grass prairie interspersed with fingers of gallery forest and scrub thicket. The very tall, dense grass is constantly burned by the natives to permit agriculture and grazing. The area is in an ecotone between forest and savanna. The elevation of Gongo is about 1300 feet (385 m.).
Species Account
AMPHIBIA
Bujo regularis Reuss Four immature specimens of this wide-
spread African toad were collected at Fada between January 26 and February 1, 1960. The toads range from 42 to 57 mm. in snout- vent length.
Seventeen ranid frogs representing two genera (Hemisus and Ptychadena) and the following five species were found in swampy vegetation surrounding a pond located at the transition from gallery forest to tall grass savanna at Gongo.
Hemisus marmoratus (Peters) A single small (25.0 mm. snout-
vent length) specimen was collected on February 23, 1960 at the Gongo pond. This is a wide-spread savanna form known from Ethiopia and Gambia in the north to Angola and Bechuanaland in the south.
Ptychadena oxyrhyncha (A. Smith) One adult female of P.
oxyrhyncha (51.4 mm. snout-vent length) was taken in the Gongo pond mentioned above. The species is widespread in forested and semi-forested regions of Africa from Portuguese Guinea and Ethiopia in the north to South Africa. It has been recorded from Fort Crampel, Central African Republic, approximately 150 miles to the south of Gongo.
The single specimen is in a poor state of preservation and characters associated with the web and foot of the hindlimb are not easily discerned. External metatarsal tubercles and tarsal tubercles appear to be absent. No supernumerary metatarsal or tarsal tubercles are present. The web is almost complete. The terminal phalanx of toe 4 is free from the webs on both sides, slightly more than one phalanx is free from the inner web of
6
Contributions in Science
No. 40
toe 3, and slightly more than one phalanx is free from the inner web of toe 2. The hind limb is long (slightly more than twice the body length), and the tibio-tarsal joint of the appressed limb extends far beyond the end of the snout. Three pairs of regular glandular ridges extend almost the full length of the body posterior to the eye. The color pattern consists of dark round spots arranged in longitudinal rows on a lighter background. The femur and tibia bear dark bars dorsally. The posterior border of the femur is reticulated with dark pigment. An interorbital dark bar is present but there is no vertebral stripe. The venter is immaculate, with the exception of some dark pigment on the mandibular border.
Ptychadena trinodis (Boettger) Two female frogs taken in the
Gongo pond are assigned to Ptychadena trinodis , although this locality is far beyond the known range of the species. The larger specimen is 51.0 mm. in snout-vent length and the smaller is 50.4 mm. The internal metarsal tubercle is large and very well-developed. A large and well-developed external metatarsal tubercle is present in both specimens. A small but well-developed tarsal tubercule is present near the proximal end of the tarsus. No supernumerary metatarsal tubercles are present. The web is only slightly reduced and extends virtually to the tip of toes 5 and 1, and to the tip on the external margins of toes 2 and 3. Two phalanges are free of the webs of toe 4, two phalanges are free from the inner web of toe 3, and slightly over one and one-half phalanges are free from the inner web of toe 2. The web exists as a very narrow fringe almost to the tip of toe 4. The robust hind limb is of moderate length (1.4 - 1.6 times body length). The tibio-tarsal joint of the appressed limb reaches the snout in the larger specimen and the nostril in the other. A number of irregular glandular ridges are present on the back. The most lateral are the longest, and are whitish in color. The color pattern consists of longitudinal series of dark spots arranged on the lighter ground color of the back to form transverse series approximating bars. A dark spot is located on each eyelid, but no interorbital bar is present. A well defined narrow, light vertebral stripe extending from the tip of the snout to the anus is present in the smaller individual, but is lacking in the other. Three to four dark bars are present on the femur and tibia. The posterior surface of the tibia is marked with alternating irregular bands of light and dark. The ventral surface is immaculate with the exception of some small dark spots in the lateral gular region and irregular spotting on the border of the mandible.
The presence of external and internal metatarsal tubercles and of a well developed tarsal tubercle is diagnostic of P . trinodis. These conditions are found among Ptychadena only in P. trinodis , P. pumilio , and P. boettgeri , with rudimentary tarsal tubercles reported in some P. uzung- wensis. Our specimens agree with P. trinodis and are distinguished from the other species by the characters mentioned above. Guibe and Lamotte (1957) state that P. trinodis is found in the savanna zones of Senegambia,
1961
Wake & Kluge: Amphibians and Reptiles
7
French Sudan, and French Guinea. Lamotte, Dzieduszycka, and Lauwarier (1958) record it from the region of Diafarabe in French Sudan. Diafarabe is about 1600 miles west and a little north of Gongo, and Gongo is 1100 miles from the nearest part of French Sudan. Guibe and Lamotte (1958) predicted that P. trinodis would be found in neighboring Cameroons. Early records of its occurrence in East Africa are evidently in error (Loveridge, 1957).
Ptychadena maccarthyensis (Anderson) A series of 12 small
frogs from the Gongo pond is tentatively assigned to P. maccarthyensis. Although some adult frogs are represented in the series, the general size is small (40.7-27.0 mm. snout-vent length). A moderately large internal metatarsal tubercle and a smaller, round external metatarsal tubercle are present. These specimens are noteworthy for the large number of super- numerary metatarsal and tarsal tubercles. The distribution, range, and means of the supernumerary tubercles are as follows:
metatarsal 2 0-4 (2.1) metatarsal 5 0-6 (2.4)
metatarsal 3 2-8 (5.1) tarsal 3-8 (6.1)
metatarsal 4 5-10 (7.1)
No supernumerary tubercles are present on metatarsal 1. Super- numeraries are present on one or more of metatarsals 2, 3, and 4 in all 12 frogs, and on metatarsal 5 in 11 frogs. Tarsal supernumeraries are present on the surface of the tarsus, opposite the well defined tarsal fold, in all 12 frogs. In ten frogs the tarsal supernumerary nearest the heel is the largest, and in these specimens the tubercle is somewhat reminiscent of the tarsal tubercle of P. trinodis , although not nearly as large or well developed. Guibe and Lamotte (1957) report a more complete web in P. maccarthyensis than in our specimens, but webbing appears to be slightly variable intraspecifically in Ptychadena. None of the toes are fully webbed. About one-half of a phalanx is free from the web of toe 5, two phalanges are free of the outer web and two and one-half of the inner web of toe 4, one from the outer web and two and one-half from the inner web of toe 3, one from the outer and two from the inner web of toe 2, and one and one-half phalanges are free of the web of toe 1. The fairly robust limb is of moderate, but somewhat variable, length (1.5 - 1.8 times the body length). The tibio-tarsal articulation of the appressed limb generally reaches the nostril, but in some indivduals it reaches the snout and in others falls slightly short of the nostril. Eight pairs of regular glandular ridges are present dorsally. The median pair is very short and falls far short of the eye. The ridging pattern is more similar to that illustrated by Schmidt and Inger (1959, p. 71) for P. maccarthyensis than the illustration of Guibe and Lamotte (1957, p. 956) for the same species. The arrangement of the pustules at the corner of the mouth and in the tympanum region is similar to that illustrated by Guibe and Lamotte (1957, p. 956) although the pustules are somewhat smaller. One male has two uniform black vocal sacs located below and extending toward the
8
Contributions in Science
No. 40
lower insertion of the forelimb postero-obliquely from the mandibles. The distance between the vocal sacs is slightly less than that from the anterior edge of the sacs to the tip of the snout. The ground color of the dorsum is light and is marked with a variable number of rather irregularly placed dark spots that are of the same size or smaller than the tympanum. The spots are arranged in transverse rows in some individuals, but in others in longitudinal rows. The dorsal surface of the head is immaculate, but some small spots may be present on either eyelid. In three of the specimens a well defined, light line is present mid-dorsally. The line is broad anteriorly and becomes narrowed posteriorly. Alternating longitudinal irregular bands of light and dark are present on the posterior border of the femur. The dorsal surfaces of the femur and tibia are marked by three dark bars. The ventral surface is immaculate with the exception of the border of the mandibles, the throat, and the pectoral regions which are marked with black punctations.
Our specimens differ from the P. maccarthyensis of Guibe and Lamotte (1957) in the following characters: less extensive webbing of toes 1 and 4 ; presence of supernumerary metatarsal tubercles on metatarsals 2 and 5 in addition to metatarsals 3 and 4; presence of tarsal super- numerary tubercles; and smaller size. It should be mentioned that the illustration of the foot of P. maccarthyensis in Guibe and Lamotte (1957, p. 955) shows supernumerary tubercles on metatarsal 5, but these are not mentioned in the text. Ptychadena maccarthyensis is known from forest, gallery forest and savanna regions of Africa from Gambia to the Ivory Coast, according to Guibe and Lamotte (1957). Schmidt and Inger (1959), however, state that P. maccarthyensis is represented in the specimens from Belgian Congo identified by Noble (1924) as Ptychadena mascareniensis , and it appears that P. maccarthyensis is more widely dis- tributed than was once thought. Significantly, P. maccarthyensis and P. trinodis have been taken together at Diafarabe, French Sudan (Lamotte, Dzieduszycka, and Lauwarier, 1958).
Ptychadena sp. One small (31.6 mm. snout-vent length) frog
is unassignable to any described species as nearly as can be ascertained from the literature. This immature female, taken in the Gongo pond, lacks tarsal and supernumerary tarsal and metatarsal tubercles. The external metatarsal tubercle is very small and poorly distinguished. The webbing of the hind foot is greatly reduced. Slightly more than one phalanx is free from the web of toe 5, three phalanges are free from the web of toe 4, one and one-half from the outer and two and one-half from the inner web of tone 3, one from the outer and two from the inner web of toe 2, and one and one-half phalanges are free from the web of toe 1. The robust hind limb is short (1.4 times body length), and the tibio- tarsal articulation of the appressed hind limb reaches to the center of the eye. The first finger of the fore limb is slightly longer than the second. The subarticular tubercles of the fore limb are very large. The tympanum
1961
Wake & Kluge: Amphibians and Reptiles
9
is smaller than the eye. The nostril is located nearer the snout than the eye. Dorsal glandular ridges are present, but they are very irregular. The most regular are the lateral whitish ridges. The dorsal ground color is rather dark, with a number of small darker spots. The spots are mostly lateral in position and are generally longer than broad. Several small spots are located on either eyelid. The dorsal surfaces of the femur and tibia are inconspicuously barred with dark spots, and the posterior surface of the femur is marked with irregular dark vermiculations. The ventral surface is immacualte and white, with pigment along the borders of the mandible. One conspicuous dark bar is located on either mandibular ramus directly under the eye.
The single specimen has a shorter, broader, and less pointed head than any of the species described above, and its body form is more robust than any of the sympatric species. Similarities between this individual and certain described species have been ascertained from the literature. According to Laurent (1954), P. frontalis is a short-limbed frog in which the tibio-tarsal articulation of the appressed limb reaches to a point between the tympanum and the nostril. Laurent’s description is partially corroborated by Schmidt and Inger (1959) who commented on the short legs of P. frontalis. However, the latter authors also noted the presence of supernumerary tubercles on the metatarsals of P. frontalis , in contrast to the situation in our specimen. The Gongo example has a fuller web than P. frontalis, and the color pattern of the posterior portion of the femur and the back, and the pattern of the dorsal glandular ridges also differ from the situation described for P. frontalis. Laurent (1954) examined the types of P. abyssinica and compared this species with P. frontalis. He noted the short limbs and the similarity of the webbing in the two species. The external metatarsal tubercle of P. abyssinica is poorly developed in contrast to P. frontalis. The degree of webbing in P. frontalis, and pre- sumably in P. abyssinica, differs from that in our specimen. Parker (1930) assigned P. abyssinica to the synonymy of the long-legged P. oxyrhyncha, but as pointed out by Laurent (1954) the former is a short-legged species. Our specimen is certainly not referable to P. oxyrhyncha. Only sketchy descriptions of P. abyssinica are available, and the limited information makes it impossible for us to determine whether or not the specimen under consideration should be assigned to that species.
Rana occipitalis Gunther Seven examples of this widely dis-
tributed river frog were taken in stream beds and artificial ponds at Fada between January 26 and February 1, 1960. The specimens range from 49.6 to 70.6 mm. in snout-vent length.
REPTILIA
Ptyodactylus hasselquisti hasselquisti (Donndorf) Four geckos
were taken at Fada on February 1, 1960. Pertinent anatomical features of the two males and two females are as follows: snout-vent length 72.0 mm.,
10
Contributions in Science
No. 40
65.0 mm., 59.0 mm., and 53.5 mm. respectively; internasals 1; supralabials 9-11, mean 10; infralabials 11; tubercle rows on dorsum (rows somewhat irregular) 13-15, mean 13.7; scansors under fourth toe 10-13, mean 11.5; uniform fawn in color.
Ptyodactylus h, hasselquisti ranges from Asia to Niger and Algeria across northern Africa. The nearest known record to Fada is from the Tibesti Mountains of northern Tchad.
Tarentola neglecta Strauch A female of this very distinctive
gecko was taken at Fada on February 1, 1960, and one female was taken at Gongo on February 23, 1960. The salient characters of the Gongo and Fada specimens respectively are as follows: snout-vent length 69.6 mm. and 73.2 mm.; internasals 0 and 1; interorbitals 13 and 12; supralabials 8 and 8; infralabials 8 and 8; breadth of mental divided into its length
I. 3 and 1.4; dorsal tubercle rows 13 and 14; scansors under fourth toe 14 and 17. The dark brown eye bar is very obvious and continues posteriorly to form the dorsal body reticulation.
Loveridge (1947) states that T . neglecta is known from the Algerian Sahara east to Libya. The Gongo locality is approximately 1200 miles southeast of the nearest locality recorded by Loveridge (Serdeles, Libya) .
Tarentola annularis (Geoffroy) One specimen was collected at
Oum Chalouba, one at Arada, and one at Gongo on February 8, February
II, and February 23, 1960, respectively. The salient characters of the Oum Chalouba (male), Arada (male), and Gongo (female) specimens respectively are as follows: snout-vent length 89.1 mm., 66.6 mm., and 53.2 mm.; internasals 2, 1, 1; interorbitals 18, 18, 16; supralabials 10, 11, 10; infralabials 9, 9 ,10; breadth of mental divided into its length 1.5, 1.6, 1.8; dorsal tubercle rows 12, 12, 12; scansors under fourth toe 19, 21, 20. The male specimen from Arada has a dark brown ground color. The female is very light.
Loveridge (1947) considered Tarentola annularis and T. ephippiata to be closely related, and included T . ephippiata as a subspecies of T. annularis. His decision has led to some confusion, and, on the basis of our material, we find it impossible to consider these forms conspecific. The most obvious character differentiating the two species is the shape of the tail. In T. annularis the tail is greatly depressed, with a lateral serration formed by enlarged, laterally projected scales. In T . ephippiata the tail is rounded and without an obvious lateral serration. The distinctness of these two species has been reiterated by Dr. Georges Pasteur (in litt.) .
Tarentola annularis is known from Arabia west to Libya and south to Sudan, Ethiopia, and British Somaliland, according to Loveridge (1947). Our records appear to be the first for Tchad, and the Gongo locality is apparently at the southwest border of the known range.
Chamaeleo gracilis Hallowell One chamaeleon was taken in the
gallery forest near Gongo on February 22, 1960. The tarsal spur is particu- larly evident in this individual. The species is a savanna form found
1961
Wake & Kluge: Amphibians and Reptiles
11
along the edges of the rain forest zone in Africa, from Senegal and French Somaliland probably to Angola.
Uromastix acanthinurus Gray Two specimens were taken at
Fada on January 29, 1960. A male of 175 mm. (snout-vent length) is dark in coloration with black head, shoulders, and throat, and black blotches between the whorls on the dorsal surface of the tail. The ventral surfaces, other than the throat, are white with scattered black bars and blotches. The 187 mm. (snout-vent length) female is lighter than the male with a light head covered with brown reticulations. The ventral surfaces are an immaculate and uniform whitish color. Black blotches are present only between the whorls on the dorsal surface of the tail. Uromastix is a north African element and has been found in several saharan mountain ranges, but has not been recorded previously from the Ennedi region.
Agama agama Linne Two specimens of this wide ranging
African species were taken at Abeche on February 10 and 12, 1960, and 12 specimens were taken at Gongo on February 20, 1960. Geographic variation in this species is not well understood, and subspecific determina- tion of these specimens has not been attempted.
Mabuya quinquetaeniata (Lichtenstein) Two male skinks were
taken at Fada on February 1, 1960. The characteristics of the specimens are as follows: snout-vent length 83.5 mm. and 78.0 mm.; supraciliaries 5 and 6; scales around middle of body 36 and 36; dorsal and lateral scales tricarinate; anterior nuchals mostly tricarinate, a few quinque- carinate. Three indistinct light bluish lines are located on the dorsum of both specimens, with a very distinct whitish lateral line extending from the tip of the snout to the groin on either side. This widespread savanna species reaches the Mediterranean Sea along the Nile River drainage, but apparently has not been recorded before from the semi-desert mountain areas of central Africa.
Tarbophis obtusus (Reuss) A single snake was taken at Oum el
Adam on January 31, 1960. The characters of this juvenile male are as follows: supralabials 10 (4, 5, and 6 enter eye) ; infralabials 11 (first four in contact with anterior chinshield) ; temporals 24-3 + 4(5) ; scale rows 20-20-15; ventrals 265; subcaudals 65 + 1; snout- vent length 255 mm.; tail length 39 mm. The dorsal body coloration consists of 79 rather well defined quadrangular light brown bars; the venter is immaculate. The distinctness of the dorsal body blotches may be due to the age of the specimen.
Tarbophis obtusus is a wide ranging colubrid snake of the arid semi- desert regions from Mauritania to Somalia, but has not been recorded previously from Tchad.
Geochelone sulcata (Miller) One tortoise was taken at Koro
Toro; another was purchased from a convict at Fada and almost certainly was taken at that place. Carapace length of these two specimens is 400.2 mm. and 201.8 mm. respectively. According to Loveridge and Williams
12
Contributions in Science
No. 40
(1957), G. sulcata is known from desert regions across Africa at about 15° N Latitude, but the species evidently has not been recorded previously from Tchad.
Crocodilus niloticus Laurenti The common African crocodile
was sighted, but not collected, in Oued Archei near Fada, in the Ennedi Mountains. This is the sole species of Amphibia or Reptilia recorded from the Ennedi region by Angel and Lhote (1938). It apparently survives in permanent ponds in the wadi.
LITERATURE CITED
Angel, F. and H. Lhote.
1938. Reptiles et Amphibiens du Sahara central et du Soudan. Bull. Com. Et. histor. et scient. de l’Afrique Occidentale Frangaise, 21: 345-384. Guibe, Jean and Maxime Lamotte.
1957. Revision systematique des Ptychadena (Batraciens Anoures Ranides) d’Afrique occidentale. Bull. Inst. Frang. Afrique Noire, 19(3): 937-l°03.
1958. Les Ptychadena (Batraciens Ranides) du Cameroon. Bull. Inst. Frang. Afrique Noire, 20(4): 1448-1461.
Lamotte, Maxime, Sophie Dzieduszycka, and Guy Lauwarier.
1958. Contribution a F etude des Batraciens de l’Ouest africain VIII. Les formes larvaires de Ptychadena submascareniensis, Pt. tournieri et Pt. trinodis. Bull. Inst. Frang. Afrique Noire, 20(4): 1464-1482.
Laurent, R.
1954. Etude de quelques especes meconnes de genre Ptychadena. Ann. Mus. Roy. Congo Beige, Ser. 8, 34: 1-34.
Loveridge, Arthur.
1947. Revision of the African lizards of the family Gekkonidae. Bull. Mus. Comp. Zool., 98(1): 1-469.
1957. Checklist of the Reptiles and Amphibians of East Africa (Uganda; Kenya; Tanganyika; Zanzibar). Bull. Mus. Comp. Zool., 117(2): 153-362.
Loveridge, Arthur and Ernest E. Williams.
1957. Revision of the African tortoises and turtles of the Suborder Crypto- dira. Bull. Mus. Comp. Zool., 115(6): 163-557.
Noble, G. K.
1924. Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909-1915. Part III. Amphibia. Bull. Amer. Mus. Nat. Hist., 49(2): 147-347. Parker, H. W.
1930. Report on the Amphibia collected by Mr. J. Omer-Cooper in Ethiopia. Proc. Zool. Soc. London, 1930: 1-6.
Schmidt, Karl P. and Robert F. Inger.
1959. Exploration du Parc National de l’Upemba. I. Mission G. F. de Witte. Amphibiens. Inst. Parcs Nat. du Congo Beige, 56: 1-264.
MBER 41
May 11, 1961
THE MACHRIS BRAZILIAN EXPEDITION
ORNITHOLOGY : Non-Passerines
By Kenneth E. Stager
os Angeles County Museum
Exposition Park
Los Angeles 7, Calif.
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. Issues are numbered separately and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientists and scientific institutions on an exchange basis. Copies may also be purchased at a nominal price.
The MACHRIS BRAZILIAN EXPEDITION from the Los Angeles County Museum was sponsored by Mr. and Mrs. Maurice A. Machris and Mrs. Maybell Machris Low. It was conducted under the auspices of the Museu Nacional do Brasil. Botanical and zoological collections were made from April through June, 1956, in the region of the headwaters of the Rio Tocantins in the state of Goias. General accounts and intineraries are given in papers 1 and 2 of this series. Technical type specimens of new entities are deposited in the Museu Nacional in Rio de Janeiro.
Hildegarde Howard Editor
E. Yale Dawson Associate Editor
THE MACHRIS BRAZILIAN EXPEDITION
ORNITHOLOGY: Non-passerines By Kenneth E. Stager1
The Machris Brazilian Expedition of 1956 to Central Goias made possible the ornithological investigation of a portion of the Planalto Central of the Brazilian interior that has heretofore received relatively little attention from biologists (Fig. 1). This lack of attention has been due largely to the remoteness of the area and its difficulty of access. With the gradual extension of a road net from the sea coast northwestward toward the site of the new federal capital of Brasilia, accessibility to the vast regions of Central Goias has been improved.
For a general account of the expedition’s itinerary and accomplish- ments see Delacour (1957). The vegetative aspects of the survey area are well covered by Dawson (1957). Information presented in the two above-mentioned papers will not be repeated here except when it bears specifically on avifaunal considerations.
Ornithological exploration of southern Goias appears to have begun with the visit of Auguste de Saint Hilaire in 1816 (Pinto, 1936), followed by the work of Count Castlenau and Monsieur Deville in 1844 and 1845. In 1893, Natterer collected at Jaragua, Boa Vista, and Inhumas (Pelzeln, 1868). The collections made in 1906 by Monsieur G. A. Baer have been reported on in detail by Hellmayr (1908). Extensive field work was conducted by Pinto and Garbe in the Rio das Almas and Inhumas area in 1934 and fully reported on by Pinto (1936). The activities of the above-mentioned persons were concentrated in the southern portion of the state and included very little of the more remote areas to the north that were investigated by our field party. Castlenau and Deville ventured as far north as Amaro Leite (see Fig. 2). J. Blaser apparently spent considerable time on the Rio Sao Domingo and on the Canna Brava of central Goias, as shown by numerous specimens collected between December 1931 and June 1933 and listed by Pinto (1938) in his “Catalogo das Aves do Brasil.” Dr. Rudolf Pfrimer is known to have collected birds at Santa Maria de Taguatinga on the east central border of Goias sometime prior to 1920, but I am not aware of the exact dates or his collecting itinerary.
On April 12, 1956, the Machris Brazilian Expedition crossed the Parana-Amazonian divide and established its first base camp 20 kms. north of Sao Joao da Alianga at the headwaters of the Rio Tocantins (Fig. 2). This portion of the Planalto Central is known as the Chapada dos Veadeiros and consists of gently rolling hills covered with “cerrado” forest or open grassland, intersected by numerous small streams bordered with dense stands of gallery forest (Dawson, 1957). Extensive field work
Turator of Ornithology and Mammalogy, Los Angeles County Museum.
ii i
INSTITUTION 2 4
CD
4
Contributions in Science
No. 41
iflha da Banatyal }$ S&U7A SSA8gt.fi pkm
•^CAVAkCAHTi f, 4 mo J0AQ 6A AUAN^Aj . J ♦jPORMOSA I
y%ANAWt«A J
!«©&A$tA /
fUBiRiANPfA I %RS8gs8A0 pmw/ %mumuAs/
S*0pAUW%-^flOOg JANEIRO
SA.NTOS
Fig. 1. Map of South America showing study area of Machris Brazilian Expedition.
1961
Stager: Brazil, Ornithology
5
Fig. 2. Detail map showing collecting localities of Machris Brazilian Expedition.
6
Contributions in Science
No. 41
was conducted in this area until May 7, 1960, when operations were shifted to the second base camp located in the Serra Dourada area (Fig. 2), 20 kins, east-southeast of Formoso. In this latter area the terrain was found to be more hilly with some “cerrado” forest. Gallery forest occurred along many of the stream courses, but to the east of the base camp area the dense forest growth proved to be continuous between stream courses and of such magnitude and composition as to be designated as primary forest (Dawson, 1957). Field work was continued in the Serra Dourada area from May 12 to June 15, when activity ceased and preparations were made for the return to Sao Paulo and Los Angeles.
At both study areas on the Planalto Central the vegetative associations were well defined and each was characterized by an equally well differen- tiated avifauna. The following avian species have been selected as representative of the various plant associations investigated during our stay in central Goias.
Open
Rhea americana Rhynchotus rufescens Nothura maculosa Theristicus caudatus Buteo magnirostris Milvago chimachima Falco sparverius Cariama cristata Belanopterus chilensis Aratinga aurea Guira guira
Cerrado-Savannah
Eupetomena macroura Colibri serrirostris Ny status chacuru Colaptes campestris Lepidocolaptes angustirostris Geobates poecilopterus Furnarius rufus Xolmis velata Mimus saturnius N eothraupis fasciata Saltator atricollis
Dense
Crypturellus parvirostris H eterospizias meridionalis Gampsonyx swainsonii Penelope superciliaris Ara nobilis Amazona xanthops Piaya cayana
Cerrado
Glaucidium brasilianum Nyctidromus albicollis Baryphthengus rujicapillus Ramphastos toco Leuconerpes candidus Icterus cayanensis Cypsnagra hirundinacea
Gallery Forest
Crypturellus undulatus Mesembrinibis cayennensis Ar amides cajanea Trogon surrucura Baryphthengus rujicapillus Columba plumbea Ara ararauna Veniliornis passerinus
Dryocopus lineatus Dendro colaptes platyrostris Sclerurus scansor Thamnophilus caerulescens Antilophia galeata Cyanocorax cyanopogon Tangara cayana Arremon flavirostris
1961
Stager: Brazil, Ornithology
7
Primary Forest
Crypturellus soui Sarcoramphus papa Chondrohierax uncinatus Spizaetus ornatus Crax fasciolata Aratinga jandaya Trogon curucui Trogon strigulatus Momotus momota Monasa nigrijrons
Ramphastos vitellinus Phloeoceastes rubricollis Phloeoceastes melanoleucos Xiphorhynchus guttatus Automolus leucopthalamus Thamnophilus punctatus Oxyruncus cristatus Pipra jasciicauda Thryothorus genibarbis Xanthornus decumanus
A total of 40 collecting days in central Goias resulted in a collection of 859 specimens of birds. A detailed study of this material, aided by the vast reference collections at the American Museum of Natural History in New York, has resulted in many new avian records for the state of Goias, numerous range extensions, and the description of two new subspecies (Stager, 1960). The study area proved to be a zone of intergradation for a large number of polytypic species, and many confusing taxonomic problems have been clarified as a result of this field work. During the course of this cooperative project between the Los Angeles County Museum and the Museu Nacional do Brasil. I was aided throughout by the able assistance of Herbert F. Berla, ornithologist on the staff of this latter institution in Rio de Janeiro. I cannot speak too highly of the help I received from Mr. Berla, for it was his excellent knowledge of the avifauna of Brazil that made our ornithological efforts in Goias a real success. Dr. Jean Delacour participated actively in the field work in the Chapada dos Veadeiros area and not only aided in the task of specimen preparation, but was also responsible for securing many of the more desirable species. Throughout the course of the work in Goias, Mr. and Mrs. Maurice A. Machris served as untiring collectors, and a large portion of the collection is the result of their enthusiastic work.
All specimens mentioned are in the collection of the Los Angeles County Museum except for types, which are deposited in the Museu Nacional do Brasil.
The non-passerine birds of the Goias collection are treated in this paper. An account of the passerines will appear in a second paper to follow shortly.
SPECIES ACCOUNT
Rhea americana americana (Linnaeus) American Rhea
Rheas were frequently encountered in small bands of two to four birds throughout the Chapada dos Veadeiros. In this region they were found in fairly open cerrado forest and grassy savannah areas. Although generally quite wary, it was often possible to approach within close range of them.
8
Contributions in Science
No. 41
An adult $ (LACM 32373) collected in open grassland, 20 kms. north of Sao Joao da Alianga, April 18, 1956, weighed 35 lbs. The tarsus measures 345 mm., and the culmen 76.5 mm.
Crypturellus soui albigularis (Brabourne and Chubb) Pileated TlNAMOU Although an abundant species in the forest area of the Serra Dourada, this tinamou was not encountered in the gallery forest of the Chapada dos Veadeiros. Comparison of Serra Dourada specimens with a large series in the American Museum of Natural History from various localities south of the Amazon discloses no appreciable variation. Measurements: 2 cf cf, wing 126, 127 mm., culmen 20, 16.4 mm.
Crypturellus undulatus vermiculatus (Temminck) Banded Tinamou
Found abundantly in both study areas. Although noted most frequently in primary forest, it was also found to frequent the narrow gallery forest of small stream courses. Collections: 3 cf cf, 20 kms. N. of Sao Joao da Alianga; 1 cf, 48 kms. S. of Peixe; 1 $,24 kms. S.E. of Formoso.
Crypturellus parvirostris (Wagler) Small-billed Tinamou
These small tinamou were collected in the vicinity of base camp #1 where they were encountered in gallery forest along the small stream courses. The species was not met with in the Serra Dourada area, but may have been overlooked. Examination of a large series of skins of this species in the collections of AMNH indicates that the species parvirostris is badly in need of revision. The specimens taken by us tend towards the dark coloration of birds from the south in contrast to lighter colored birds from northeastern Brazil. On the other hand, they tend to be larger than specimens from the Rio Madeira area to the west.
1 cf and 1 $ , 20 kms. N. of Sao Joao da Alianga.
Rhynchotus rufescens rufescens (Temminck) Rufous Tinamou
A common species in the cerrado and open grasslands of the Chapada dos Veadeiros. Less abundant in the cerrado areas of the Serra Dourada.
Specimens from the Chapada dos Veadeiros are large and light colored and resemble typical skins of R. r. rufescens from the Sao Paulo area, while the specimen from the Serra Dourada is smaller and darker and approximates R. r. catingae of the Rio Madeira and Bahia areas. It would seem that a critical study of this species is badly needed.
1 $,20 kms. N. of Sao Joao da Alianga; 1 cf, 24 kms. S.E. of Formoso.
Nothura maculosa major (Spix) Spotted Tinamou
Spotted Tinamou occur abundantly in the Chapada dos Veadeiros where they frequent open grassy savannahs and scattered cerrado. The species was not met with in the Serra Dourada area. All specimens taken agree with Conover’s (1942) differentiation of the races maculosa and major. All specimens are paler than maculosa and the neck markings are streaks rather than spots.
1961
Stager: Brazil, Ornithology
9
1 cf and 1 $,20 kms. N. of Sao Joao da Alianga; 1 cf, near Braslandia (west of Planaltina).
Podiceps dominicus speciosus (Arribalzaga) Least Grebe
The least grebe was the only species of grebe encountered during the course of field work in Goias and it was found on only one occasion. The scarcity of quiet ponds and lagoons in the study area readily explains the absence of birds of this type. However, a small pond situated among several acres of cleared land, approximately 28 kms. southeast of Formoso, supported a small number of grebes. On June 10, 1956, an adult cf (LACM 32289) was collected from this pond.
Butorides striatus striatus (Linne) Striated Heron
Striated herons were generally to be found in and around any sizeable pond or marsh area. An adult $ was collected May 24, 1956 from the margin of the pond mentioned above.
Theristicus caudatus caudatus (Boddaert) White-throated Ibis
A common species on open grassy savannahs and about clearings on the Chapada dos Veadeiros. In this area small flocks of ten to twenty birds were frequently found foraging for insects.
Mesembrinibis cayennensis (Gmelin) Cayenne Ibis
Solitary individuals and single pairs of this large, dark colored ibis were often flushed from the deep shaded banks of stream courses in the gallery forests of the Chapada dos Veadeiros. The species was never noted as common in the above area. An adult cf and $ were collected April 21, 1956, 20 kms. N. of Sao Joao da Alianga.
Neochen jubata (Spix) Orinoco Goose
The only Anserine collected during the course of our field work in Goias was an adult cf of this species, taken by Maurice A. Machris, June 1, 1956 on the Rio Araguaia, near Ilha do Bananal. In this area Machris noted the species as common. Ducks and geese were scarce on the Chapada dos Veadeiros, and the only species observed was Cairina moschata, which was frequently encountered singly and in pairs along the forested stream courses but was not collected.
Sarcoramphus papa (Linne) King Vulture
Although noted frequently in the gallery forests of the Chapada dos Veadeiros and the primary forest of the Serra Dourada, this large species was never found in large numbers. An adult cf was collected on June 20, 1956, 20 kms. N. of Sao Joao da Alianga. Two other species of vultures, Coragyps atratus and Cathartes aura were present in both areas in considerable numbers, but no specimens were collected of either species.
Chondrohierax uncinatus uncinatus (Temminck) Hooked-billed Kite On May 25, 1956, while collecting in heavy primary forest of the Serra Dourada, 28 kms. S.E. of Formoso, I attracted an adult cf of this
10
Contributions in Science
No. 41
species while “squeaking” for birds. Five days later on May 30, 1956 at the same spot in the forest I lured a second bird into range with the same technique. The second specimen was an adult 9 and possibly the mate of the former bird. In plumage, both specimens are quite uniform for the species, which is one that frequently shows considerable individual variation.
Harpagus diodon (Temminck) Rufous-thighed Hawk
An adult cf (LACM 32320) of this species was collected on April 18, 1956, in heavy gallery forest, 20 kms. north of Sao Joao da Alianga. According to Delacour, the bird responded to “squeaking” at the edge of a road clearing. This specimen evidently constitutes the first record of the species for Goias, for Pinto (1938) does not record it from Goias, and Hellmayr (1908) does not list it among the species taken by Baer. Measurements are: wing 201, tail 143, culmen (from cere) 16.4 mm.
Accipiter bicolor pileatus (Temminck) Pileated Accipiter
A single specimen (LACM 32311) was collected on April 16, 1956, in heavy gallery forest, 20 kms. north of Sao Joao da Alianga. The bird is a juvenile cf with a few dorsal feathers of the back showing adult coloration.
Accipiter erythronemius erythronemius Kaup Red-thighed Accipiter A fairly common species in both study areas. An adult cf was taken in gallery forest adjacent to our base camp north of Sao Joao da Alianga and two additional specimens (an adult 9 and a juvenile c? ) were secured in the Serra Dourada, 24 kms. S.E. of Formoso. These specimens are apparently the first records for Goias, although the species is to be expected there within its wide range from Central America to the Argen- tine. The above specimens compare favorably in size and coloration with a series of the nominate race in the American Museum of Natural History.
Heterospizias meridionalis (Latham) 1 Savannah Hawk
A single specimen of this wide-spread species was collected at the margin of a large marsh in heavy cerrado, 48 kms. north of Porangatu, between the Serra Dourada and Peixe, on June 7, 1956. The specimen, an adult 9 (LACM 32299), has a wing of 405 and tail of 200 mm.
Buteo magnirostris natter eri (Sclater and Salvin) Large-billed Hawk Small buteos of this species were found to be exceedingly common in the cerrado of both study areas. They were frequently encountered on the margin of gallery forest and primary type forest, but never appeared to penetrate either of these latter forest types. The individuals encountered were not the least bit shy and could be approached quite closely. A total of six specimens was secured, and examination of this series shows them to be intermediate between subsp. nattereri and magniplumis of the south. All specimens have the short wing of nattereri as well as the variegated whitish throat of this race. The breasts of the Goias series
1961
Stager: Brazil, Ornithology
11
are not as uniformly rufous as in nattereri, but are more striated as in magniplumis. In dorsal aspect all birds are intermediate between the two forms. In evaluating all differences, the series of specimens tends to resemble nattereri more closely than magniplumis.
Buteo nitidus pallidus (Todd) Gray Hawk
The gray hawk may be more abundant in Goias than our observations indicate, for the species was encountered but once during the course of our field work. On June 5, 1956, a juvenile cf (LACM 32301) was collected at the edge of a fazenda clearing, in heavy cerrado 40 kms. south of Peixe. In ventral aspect the specimen is very light with white rather than buff markings. There is some buff on the sides of the head as well as on the primary wing coverts. The wing measures 250, and the tail 168 mm.
Busarellus nigricollis nigricollis (Latham) Black-collared Hawk
A common species in the cerrado, especially around the margins of swampy areas. A single adult cf (LACM 32300) was secured on June 7, 1956, 48 kms. north of Porangatu: wing 398, tail 187 mm.
Spizaetus ornatus ornatus (Daudin) Hawk Eagle
Hawk eagles were encountered only in the Serra Dourada area. The species was not common, but individuals would occasionally be seen soaring over clearings in the heavy primary forest. The natives of the area were very familiar with the species, and on May 19, 1956 an adult cf was brought to us alive by a caboclo who stated that he had trapped it near its nesting site. A second specimen, also an adult cf , was secured on May 31, 1956, in heavy primary forest. Both specimens are in fresh plumage and are typical of this wide ranging species. Measurements: wing 350, 325, tail 243, 232, culmen (from cere) 28, 26.3 mm.
Herpetotheres cachinnans cachinnans (Linne) Laughing Hawk
Laughing hawks were frequently heard calling in the Serra Dourada area, but seldom seen. An adult cf was taken June 7, 1956, in heavy gallery forest, 18 kms. east of Formoso: wing 267, tail 202, culmen (from cere) 22.2 mm.
Micrastur ruficollis ruficollis (Vieillot) Red-necked Harrier-Hawk
Although there appear to be very few previous records for the species’ occurrence in Goias, it has been recorded from Matto Grosso to the southwest as well as to the east. A single adult cf was taken on April 21, 1956, in heavy gallery forest 20 kms. north of Sao Joao da Alianga. The specimen is in the “plumbeous phase” with but a faint wash of rufous on the throat and wings. The head is gray and the black striations of the ventral area are very narrow. Measurements: wing 168, tail 166, culmen (from cere) 14.2 mm.
Daptrius americanus americanus (Boddaert) Red-throated Caracara Noted as common in the Serra Dourada area. These strikingly- colored caracaras most generally were observed travelling in pairs and,
12
Contributions in Science
No. 41
in contrast to the common Caracara plancus, were quite wary and difficult to approach. An adnlt $ in molt was taken on June 7, 1956, in the Serra Dourada: wing 365, tail 250, culmen (from cere) 25.5 mm.
Milvago chimachima chimachima (Vieillot) Chimachima Caracara
An exceedingly common species in both study areas, but most frequently noted in cerrado type associations.
Caracara plancus plancus (Miller) Common Caracara
Abundant on the Chapada dos Veadeiros and in the Serra Dourada. Caracaras were encountered most frequently in the cerrado and in the vicinity of cleared forest areas of the Serra Dourada.
Gampsonyx swainsonii swainsonii Vigors Pearl Kite
The small pearl kite was not encountered in the vicinity of the base camp on the Chapada dos Veadeiros, but was fairly common in the Serra Dourada area and the cerrado region south of Peixe. Individuals were found to frequent the margins of forest clearings where they foraged from vantage spots atop isolated trees left standing in the fazenda clearings. Their behavior and flight is much like that of the white-tailed kite (Elanus leucurus) . Measurements of specimens collected are as follows: cf, Serra Dourada, wing 155, tail 95, culmen (from cere) 13.2 mm.;
$, Serra Dourada, wing 157, tail 94, culmen (from cere) 13.1 mm.;
cf , 24 kms. south of Peixe, wing 149, tail 88, culmen (from cere) 13.2 mm.
Falco sparverius cearae (Cory) Sparrow Hawk
Exceedingly common on the Chapada dos Veadeiros in cerrado
forest areas. Two cf cf taken 20 kms. north of Sao Joao da Alianga
show five black bars on the inner web of the outermost rectrix. Both specimens also possess the grayish tail-tips of race cearae.
Crax fasciolata jasciolata Spix Curassow
Although seldom seen, this large curassow is apparently fairly common in the heavy primary forest of the Serra Dourada and is well known to the local caboclos who hunt them for food. This is accomplished by the hunter locating a fruiting tree where the curassows come to feed, secreting himself below and shooting the birds as they quietly fly into the tree for fruit. Two specimens, an adult cf and $ , were secured in this manner by a local hunter and brought to us. The specimens compare favorably with a large series of the nominate race fasciolata from the Rio Araguaia in the American Museum collection. Measurements: cf wing 393, tail 367 ; $ wing 350, tail 321 mm.
Penelope superciliaris jacupemba Spix Spix’s Guan
A common and wide-spread species in central Goias. Guans were noted in gallery forest and dense adjacent cerrado on the Chapada dos Veadeiros.
In the Serra Dourada area the species occurs commonly in heavy
1961
Stager: Brazil, Ornithology
13
primary forest, where it feeds in the tops of tall fruiting trees.
The four specimens secured by us in central Goias agree with a large series of jacupemba in the American Museum. The superciliary stripe in all four specimens is grayish rather than ochraceous, and the frontal grayish band joining the superciliaries, which Neumann (1932) gives as the principal character for his race argyromitra, is present in three (Serra Dourada) of the four LACM specimens. As pointed out by Hellmayr (1942), however, this character is highly variable, and he synonymized argyromitra with jacupemba. The single LACM specimen from the Chapada dos Veadeiros area has the ochraceous margins of the wing feathers much narrower than as shown by the three specimens from the Serra Dourada. Measurements of the four specimens are as follows: cf, 20 kms. N. of Sao Joao da Alianga, wing 251, tail 285 mm.; cf, Serra Dourada, wing 253, tail 299 mm.; cf, Serra Dourada, wing 241, tail 283 mm. ; 9 , Serra Dourada, wing 235, tail 290 mm.
Aramides cajanea cajanea (P. L. S. Muller) Wood Rail
Wood rails occur commonly in the gallery forests of the Chapada dos Veadeiros and in the primary forests of the Serra Dourada. A total of seven specimens were obtained (4 Chapada dos Veadeiros and 3 Serra Dourada). All specimens were purchased alive from local caboclos who trap them with great ease.
The LACM series from both areas in Goias are remarkably uniform, indicating that a careful study of large series of South American specimens might enable someone to bring order out of the confused subspecies problem which surrounds this species in the South American portion of its range. Hellmayr and Conover (1942), after examining a series of 80 specimens from all parts of its range, concluded that there was too much individual variation among the South American specimens to warrant any decision other than lumping all of them under the nominate subspecies cajanea. This conclusion may be correct, but the problem may deserve re-examination. Wing measurements of the LACM Goias series are 4 cf cf 195, 3 9 $ 185, 189, 190 mm.
Cariama cristata (Linne) Seriema
Seriemas or cariamas are very abundant in the cerrado forest of the Chapada dos Veadeiros and in cerrado areas of the Serra Dourada. The species was exceedingly abundant in the vicinity of our base camp, 20 kms. north of Sao Joao da Alianga, and the morning and evening chorus was of spectacular volume. Individuals continuously circled our camp areas and exhibited considerable curiosity, much as did the Rheas. An adult 9 (LACM 32335) taken at the above locality on April 28, 1956, measures: wing 367, tail 317 mm.
Belanopterus chilensis lampronotus (Wagler) Brazilian Lapwing
A very common species in suitable open areas on both the Chapada dos Veadeiros and in the Serra Dourada. An adult cf and 9 were
14
Contributions in Science
No. 41
collected on April 13, 1956, 20 kms. north of Sao Joao da Alianga.
Hoploxypterus cay anus (Latham) Cayenne Plover
A single specimen was taken on June 7, 1956, 99 kms. north of Porangatu, on the road to Peixe. At this locality the species was noted as common about the margins of a large lagoon.
Columba speciosa Gmelin Scaled Pigeon
Scaled pigeons were not recorded from the Chapada dos Veadeiros, but the species was found to be common in the heavy primary forest areas of the Serra Dourada. Two specimens were collected at this latter locality.
Columba cayennesis sylvestris Vieillot Rufous Pigeon
A common species about fazenda clearings in close proximity to gallery forest areas on the Chapada dos Veadeiros and in primary forest in the Serra Dourada. Two specimens taken at the above localities have the bicolor rectrices characteristic of the subspecies sylvestris.
Columba plumbea baeri Hellmayr Plumbeous Pigeon
Although present in the Serra Dourada area, the species was encountered much more frequently in the Chapada dos Veadeiros. In this latter locality plumbeous pigeons were found abundantly about fazenda clearings in gallery forest. A taken on May 1, 1956, 20 kms. north of Sao Joao da Alianga, was compared with the type of subsp. baeri and found to agree very well in size and color. The pale gray coloration of baeri immediately separates central Goias specimens from the much darker nominate race plumbea.
Scardafella squamata squamata (Lesson) Scaled Dove
An exceedingly common species in central Goias, occurring in many types of habitat with the exception of heavy primary forest. Even within such forest, the species is abundant in fazenda clearings and along road margins which are used as avenues of penetration.
Columbigallina talpacoti talpacoti (Temminck) Talpacoti Dove
An abundant species in the Serra Dourada area, but relatively scarce on the Chapada dos Veadeiros, at least in the area of our greatest activity, 20 kms. north of Sao Joao da Alianga. An adult c? and $ were collected May 17, 1956, 24 kms. S.E. of Formoso (Serra Dourada).
Claravis pretiosa (Ferrari-Perez) Cinereous Dove
Although it ranges widely over South America, this small gray dove was encountered only in the Serra Dourada area where it is fairly abundant in suitable habitat. In the Serra Dourada it was noted most frequently in areas of intergradation between cerrado and primary forest. In such situations this dove commonly feeds in clearings, on oxcart roads and foot trails. Two c? cf and one $ were secured 24 kms. S.E. of Formoso.
1961
Stager: Brazil, Ornithology
15
Leptoptila verreauxi decipiens (Salvadori) White-fronted Dove
A common and wide-spread species in central Goias. Specimens were taken in both study areas. Central Goias birds appear to agree more closely with subsp. decipiens in coloration, but approach subsp. approxi- mans in size. Measurements of three cf cf are as follows: wing 138, 140, 142, tail 107, 110, 110, culmen 14.5, 15.0, 15.5, mm.
Leptoptila ruf axilla reichenbachii (Pelzeln) Gray-fronted Dove
Gray-fronted doves are common on the Chapada dos Veadeiros, but were not noted in the Serra Dourada area. Two specimens, taken on April 23, 1956, 20 kms. north of Sao Joao da Alianga, agree with a large series of subsp. reichenbachii in the American Museum. One specimen is slightly lighter ventrally, tending towards subsp. bahiae, but the second bird, collected at the same locality on the same date, is darker as in subsp. reichenbachii. Dorsally, both specimens are identical.
Anodorhynchus hyacinthinus (Latham) Hyacinthine Macaw
The large hyacinthine macaw was not met with on the Chapada dos Veadeiros nor in the Serra Dourada area. It was observed in small flocks of three to six individuals in the cerrado country lying between the Serra Dourada and Peixe. In this region there are stands of heavy cerrado intersected by stream courses bordered with gallery forest or stands of the large fan-leaved Burity Palm, and it was in the close proximity of these stream courses that all small flocks of this macaw were observed. On June 6, 1956, one cf and two $ $ were collected from a flock
of four birds, 50 kms. south of Peixe. All specimens agree in color with a series of skins in the American Museum. The measurements are as follows: wing cf 434, $ $ 416, 395; tail cf 555, $ $ 522, 490; culmen cf 90.5, $ $ 89.9, 83.5 mm.
A careful watch was kept for individuals of the rare Anodorhynchus leari , whose habitat is yet to be discovered, but no trace of the species was found in central Goias.
Ara ararauna (Linne) Blue and Yellow Macaw
Blue and yellow macaws are very abundant in the cerrado of the Chapada dos Veadeiros and the Serra Dourada.
These noisy macaws usually traveled in pairs and could generally be seen overhead at any time of day. They frequent stream courses lined with the tall Burity Palm, feeding upon the palm nuts and using the palm trunks for excavation of nest sites. Three specimens, a cf and two $ $ , were collected on the Chapada dos Veadeiros.
Ara sever a castaneifrons Lafresnaye Brazilian Macaw
A single cf specimen was collected by Maurice A. Machris, June 5, 1956, on the Ilha do Bananal, Rio Araguaia. Machris reported the species as being common in that area, but it was not met with in the Serra Dourada, nor on the Chapada dos Veadeiros.
16
Contributions in Science
No. 41
Ara nobilis longipennis (Neumann) Noble Macaw
These small macaws were not recorded from the area of our first base camp on the Chapada dos Veadeiros, but were found to be very common in the Serra Dourada area where they showed a decided prefer- ence for the cerrado areas and fazenda clearings at the edge of primary forest. These gregarious birds travel about in flocks of 15 to 25.
Neumann’s race longipennis is based on size of wing and tail, with no apparent color difference. Examination of a large series of Ara nobilis (AMNH) from Brazil shows the presence of a good size-cline from south to north as is to be expected, with size decreasing from south to north. Birds of the nominate race nobilis north of the Amazon are much smaller than the birds of Para and Maranhao which are assignable to cumanensis. The difference in size between southern birds from Matto Grosso ( longipennis ) and typical cumanensis to the north is not as great. LACM birds from the Serra Dourada are intermediate in size between longipennis and cumanensis and could be assigned to either of these two races. Due to the close proximity of Neumann’s type locality on the Rio Sao Miguel to the Serra Dourada, assignment is made to longipennis. Measurements are as follows: cf cf, wing 182-196, tail 165-183 mm.; 9 9 , wing 175-186, tail 160-162 mm.
Aratinga jandaya (Gmelin) Janday Paroquet
The distribution of this brilliantly colored paroquet in Goias is evidently rather spotty, as it seems to have been missed by most workers in that area. The only records other than those of the LACM are three specimens listed by Pinto (1937) as taken by Blaser on the Rio Sao Domingo and the Canna Brava in 1932. During the course of our field work in Goias, the species was encountered only in the Serra Dourada area where two cf cf and two 9 9 were taken. The species was not found to be common, but would occasionally be met with in small flocks of ten to fifteen birds, or as solitary pairs and individuals. In all instances, the species was found in primary forest areas at the margin of clearings.
Aratinga aurea aurea (Gmelin) Golden-crowned Paroquet
A common cerrado species of both the Chapada dos Veadeiros and the Serra Dourada. Flocks of 30 to 50 birds were encountered daily throughout the area. LACM Goias birds are slightly smaller and paler than specimens from Sao Paulo and Matto Grosso.
Specimens collected: 3, 20 kms. north of Sao Joao da Alianga; 1, 7 kms. south of Veadeiros; 1, Serra Dourada.
Brotogeris versicolorus chiriri (Vieillot) Yellow-winged Paroquet
An abundant species on the Chapada dos Veadeiros in gallery forest association and in primary forest areas of the Serra Dourada. Generally noted in very large flocks, sometimes numbering hundreds of individuals. LACM Goias birds and those from Piauhy and Maranhao are slightly
1961 Stager: Brazil, Ornithology 17
smaller and paler than specimens from farther south, as is to be expected. Specimens collected: 3, 20 kms. north of Sao Joao da Alianga; 1, Serra Dourada.
Pionus maximiliani maximiliani (Kuhl) Maximilian’s Parrot
Pinto (1936) assigned birds from the Rio das Almas to the southern race siy on the basis of size. A study of all LACM Goias birds, however, shows them to be intermediate in size between birds from Bolivia (LACM) and Matto Grosso (AMNH) in the south and Piauhy and Bahia to the north and east. On the basis of smaller wing length and overall lighter coloring, I feel that all LACM Goias birds should be assigned to the nominate race maximiliani.
Parrots of this species occur commonly in the Serra Dourada and were also found to be fairly plentiful in gallery forest areas of the Chapada dos Veadeiros. They are relatively quiet in flight, traveling in small groups of two or four. Specimens collected: 4, Serra Dourada; 2, 20 kms. north of Sao Joao da Alianga.
Amazona aestiva aestiva (Linne) Blue-fronted Parrot
Common throughout the Serra Dourada and Chapada dos Veadeiros areas. Small flocks were present at all fazenda clearings. This species is the common cage bird of the caboclos, and every family seems to have at least one in captivity.
Amazona xanthops (Spix) Orange-faced Parrot
A species of rather local distribution in central Goias, the orange- faced parrot was noted only on the Chapada dos Veadeiros. A solitary cf was collected on April 16, 1956, 20 kms. north of Sao Joao da Alianga, atop a dead tree in a fazenda clearing in gallery forest. On April 30, 1956, four additional specimens were secured by Mr. and Mrs. Machris from a flock of approximately 35 birds found feeding in the fruit trees of an abandoned fazenda 7 kms. S.W. of Veadeiros. Two cf cf of this latter group are obviously old birds, showing considerable yellow and orange on the breast and abdomen as well as much more extensive yellow on the head.
Piaya cay ana cabanisi J. A. Allen Squirrel Cuckoo
A common species of the gallery forests of the Chapada dos Veadeiros and the primary forest areas of the Serra Dourada. A series of 8 specimens (six from the Chapada dos Veadeiros and two from the Serra Dourada) resemble most closely the race cabanisi of central Matto Grosso, but show a tendency for intergradation with the race hellmayri to the north. Dorsally, the Goias birds are closest to hellmayri but the throats are paler as in cabanisi.
Crotophaga ani Linne Ani
A common species in suitable habitat of both study areas, often in close association with the preceeding species. Specimens collected: 1 cf,
18
Contributions in Science
No. 41
2 $ 9 , 24 kms. S.E. of Formoso, Serra Dourada; 1 cT, 20 kms. N. of
Sao Joao da Alianga.
Guira guira (Gmelin) Guira Cuckoo
An abundant and gregarious species throughout the cerrado areas of both the Chapada dos Veadeiros and the Serra Dourada. Although small flocks of from 5 to 10 birds were to be noted at almost any time of day, they nevertheless were always quite shy and would maintain considerable distance between themselves and the observer. An adult 9 was collected on May 4, 1956, 20 kms. north of Sao Joao da Alianga.
Glaucidium minutissimum minutissimum (Wied) Minute Pygmy Owl A search of the literature indicates that there are very few records of this small owl from Southern America. Whether this situation indicates that the species is rare in South America or has just been overlooked by ornithologists, is not known.
An adult c? in fresh plumage was secured in the Serra Dourada on June 1, 1956, 24 kms. southeast of Formoso. During mid-morning the bird was heard calling from the crown of first class forest, but Herbert Berla and I spent a full two hours in our efforts to collect the specimen. The owl responded readily to our decoying call and was repeatedly seen flying back and forth from the crown of one heavily leafed tree to another. It was only when it made the mistake of alighting in an area of bare branches that we were able to shoot successfully. The species was not seen nor heard by us on any other occasion.
This specimen apparently represents the first record of the species for Goias, as both Baer (Hellmayr, 1908) and Pinto (1937) fail to record it for this state. Measurements: wing 85.0 mm.; tail 52.0 mm.
Glaucidium brasilianum brasilianum (Gmelin) Brazilian Pygmy Owl An abundant and widespread species in both cerrado and second class forest of the Chapada dos Veadeiros and the Serra Dourada. Four specimens secured are as follows: 1 9 , 18 kms. S.E. of Formoso, Serra Dourada — Brown phase; 1 cf, 99 kms. N. of Porangatu, on road to Peixe — Brown phase; 1 9 , 24 kms. S.E. of Formoso, Serra Dourada — Red phase; 1 9 , 20 kms. N. of Sao Joao da Alianga — Red phase.
Speotyto cunicularia gr allaria (Temminck) Burrowing Owl
A common species of the open grassland (campo limpo) areas of the Chapada dos Veadeiros. Burrowing owls were not encountered in the Serra Dourada region. One c? and two 9 9 were secured 20 kms. N.
of Sao Joao da Alianga, April 15, 1956.
Chordeiles pusillus pusillus Gould Least Nighthawic
Noted only on the Chapada dos Veadeiros where it was a common species in the air at dusk in the vicinity of base camp no. 1. Two male specimens were secured 20 kms. N. of Sao Joao da Alianga, April 29, 1956.
1961
Stager: Brazil, Ornithology
19
Nyctidromus albicollis derby anus Gould White-throated Paraque
A common species in central Goias. All individuals secured by our field party were eye-shined at night on the roads and trails: 2 cf cf , 1 $ , 20 kms. N. of Sao Joao Alianga; 4 cf cf, 24 kms. S.E. of Formoso, Serra Dourada, May 15, 1956. Six of the seven specimens collected are in the gray phase, and one is decidedly red. The series agrees very well with a large series of the race derby anus in the collections of the AMNH.
Hydropsalis brasiliana brasiliana (Gmelin) Forked-tailed Nighthawk A single individual of this species was eye-shined and collected in the Serra Dourada on May 23, 1956. The specimen, an adult cf (LACM 32420), agrees in color and size with a series of the nominate race from the Matto Grosso: wing, 161 mm., tail 330 mm.
Glaucis hirsuta hirsuta (Gmelin) Hairy Hermit
Although a widespread species in Brazil, it was apparently missed in Goias by Baer (Hellmayr, 1908) and by Pinto (1936). A single specimen taken by W. Garbe on the Rio das Almas in October of 1934, is listed for Goias by Pinto (1937). An adult cf (LACM 32433) was taken at our study area in the Serra Dourada on May 23, 1956: wing 61.5 mm., tail 42.0 mm.
Phaethornis pretrei (Lesson and DeLattre) Pretre’s Hermit
Two male specimens of this large, solitary species were secured on the Chapada dos Veadeiros, April 26 and 27, 20 kms. N. of Sao Joao da Alianga, where they were found in the shady margins of gallery forest.
Phaethornis ruber ruber (Linne) Red-vented Hermit
A common species in the Serra Dourada, but not met with on the Ghapada dos Veadeiros. A male taken on May 26, 1956, 24 kms. S.E. of Formoso, measures as follows: wing, 33.4 mm., tail 29.8 mm.
Eupetomena macroura macroura (Gmelin) Brazilian Swallow-tail
An exceedingly abundant and obvious species of the cerrado tracts of the Chapada dos Veadeiros. A series of five males collected 20 kms. N. of Sao Joao da Alianga between April 15, and Ma y4, 1956, have the following measurements: wing 68.6-75.8 mm. (71.9), tail 75.1-93.7 mm. (88.0).
Colibri serrirostris (Vieillot) Brazilian Violet-ear
Abundant throughout the cerrado areas of the Chapada dos Veadeiros. A series of five males and one female collected between April 14 and May 1, 1956, have the following measurements: 5 cf cf, wing 67.3-72.5 mm. (70.9), tail 44.1-49.0 mm. (45.2) ; 1 9 , wing 61.5 mm., tail 40.0 mm.
Anthracothorax nigricollis nigricollis (Vieillot) Black-throated Mango A common species in the Serra Dourada area but less so on the Chapada dos Veadeiros. A series of 3 cf cf and 3 $ 9 were secured in the former area, where the species was encountered in the vicinity of
20
Contributions in Science
No. 41
clearings in primary forest. On the Chapada dos Veadeiros 2 cf cf were collected along the periphery of gallery forests.
Lophornis magnified (Vieillot) Frilled Coquette
Individuals of this minute and beautiful species were noted on several occasions in clearings of primary forest of the Serra Dourada area. On June 11, 1956, an adult cf was collected 24 kms. S.E. of Formoso.
Thalurania furcata baeri Hellmayr Baer’s Wood Nymph
An exceedingly abundant species in both study areas. A series of 3 cf cf and 1 9 from the Chapada dos Veadeiros and 3 cf cf from the Serra Dourada area agree in size and coloration with the description of the race baeri as given by Hellmayr.
Heliactin cornuta (Wied) Sun Gem
A single adult cf was taken 20 kms. north of Sao Joao da Alianga, on April 30, 1956. Baer (Hellmayr, 1908) took several specimens of this species at various localities in Goias.
Calliphlox amethystina (Boddaert) Amethyst Hummingbird
A common species in the Serra Dourada and on the Chapada dos Veadeiros. A series of 2 cf cf and 1 9 from 20 kms. north of Sao Joao da Alianga, and 1 $ from 24 kms. S.E. of Formoso, agree in size and coloration with the large series in the AMNH from numerous and wide- spread localities in South America.
Trogon strigilatus strigilatus Linne Yellow-breasted Trogon
Apparently not previously recorded from Goias, as Pinto (1936) did not meet with the species in southern Goias, and Baer (Hellmayr, 1908) did not encounter it during his travels in this state. The species was found to be common in the Serra Dourada area, but was not met with in the gallery forests of the Chapada dos Veadeiros. In the Serra Dourada area this trogon was frequently encountered in heavy primary forest and a series of 2 cf cf and 3 9 $ was secured.
The above mentioned series agrees with specimens of the nominate race rather than with melanopterus of Bahia, as all three females show greater amounts of white tipping in the rectrices.
Trogon surrucura surrucura (Vieillot) SuRRUCURA Trogon
A single specimen of this species was collected on May 3, 1956, 20 kms. north of Sao Joao da Alianga. This adult cf was the only trogon secured by us on the Chapada dos Veadeiros, although continuous search was made for trogons in the gallery forests of the area. The species was not met with in the Serra Dourada area. The above specimen is in fresh plumage and possesses the bright red breast and abdomen of the nominate race. Measurements for this specimen (LACM 32470) are wing 132 mm., tail 138 mm.
Trogon curucui curucui Linne Curucui Trogon
A number of individuals of this species of small trogon were observed
1961
Stager: Brazil, Ornithology
21
in primary forest areas of the Serra Dourada, but only one specimen was secured. On May 19, 1956, an adult 9 in fresh plumage (LACM 32469) was secured 24 kms. S.E. of Formoso.
Chloroceryle americana americana Little Green Kingfisher
The zone of intergradation of the race americana with that of mathewsii coincides roughly with the Serra Dourada and Chapada dos Veadeiros regions in Central Goias. Consequently, the single specimen secured by us from 20 kms. north of Sao Joao da Alianga is difficult to assign as to race. Comparison with material of both subspecies in question indicates that this particular specimen is most closely allied to the northern form.
Chloroceryle amazona amazona (Latham) Amazon Kingfisher
A single specimen was secured on May 24, 1956, 24 kms. S.E. of Formoso, in the Serra Dourada. Other individuals of the species were observed on numerous occasions in this area.
Baryphthengus rujicapillus berlai Stager Rufous-headed Motmot
As this species had not been previously recorded from Goias, it proved of considerable interest to find it present in sizeable numbers in both collecting areas. On the Chapada dos Veadeiros it occurs abundantly in the heavy gallery forest bordering the stream courses. In the Serra Dourada area the species was noted as common throughout the tracts of primary type forest. It responded readily to “squeaking” and to an imitation of its call note.
The race berlai shows strong affinities with the nominate form of the coast rather than with B. r. martii to the northwest (Stager, 1959). Goias birds are distinguished from the nominate race by the overall more pallid color-tone of the crown, nuchal area, throat, breast and abdomen. The rufous band of the lower breast is wider and paler. Specimens secured: 3 cf cf, 2 9 9 , 20 kms. N. of Sao Joao da Alianga; 4 $ 9 ,
24 kms. S.E. of Formoso, Serra Dourada.
Momotus momota simplex (Chapman) Blue-crowned Motmot
Motmots of this species were not observed or collected on the Chapada dos Veadeiros. In the Serra Dourada area the species was relatively abundant in the primary type forest at lower elevations at the west base of the hill range. On May 27, 1956, I succeeded in decoying three specimens into collecting range by imitating their tremulous call note. These three specimens (2 c? c?, 1 9 , 18 kms. S.E. of Formoso, Serra Dourada) agree most closely with examples of the race simplex. The chestnut nape patch is only slightly evident rather than pronounced as in parensis. The back is grass green. The ventral surface is more rufescent as in simplex rather than green as in the race nattereri.
Brachygalba lugubris melanosterna Sclater Black-breasted Jacamar Two specimens of this small jacamar were collected on June 6, 1956,
22
Contributions in Science
No. 41
in gallery forest bordering a stream course in heavy cerrado, 67 kms. north of Porangatu, on the road to Peixe. The species was not encountered on the Chapada dos Veadeiros, nor in the Serra Dourada region.
Galbula rujicauda rujoviridis Cabanis Rufous-tailed Jacamar
An abundant species on both the Chapada dos Veadeiros and in the Serra Dourada area where it occurs in gallery and primary type forest. The series of specimens collected in both of the above areas agree with examples of the wide-spread race rujoviridis.
Ny status chacuru chacuru (Vieillot) White-eared Puff-bird
A characteristic and abundant species throughout the cerrado areas
of both the Chapada dos Veadeiros and the Serra Dourada area. A series
of six specimens from the two collecting areas agree with a large
comparative series in the AMNH selected from widespread localities south of the Amazon in Brazil.
Nonnula rubecula rubecula (Spix) Reddish-breasted Nonnula
A single specimen of this species of small puff-bird was taken
(LACM 32501) on April 13, 1956, in marginal area of cerrado and gallery forest area, 20 kms. N. of Sao Joao da Alianga. Pinto (1936), records several specimens from near Jaragua and Baer (Hellmayr, 1908), collected the species in the vicinity of the old city of Goyaz.
Monasa nigrijrons nigrijrons (Spix) Black-fronted Nun Bird
A dominant and widespread species in the primary type forest areas of the Serra Dourada. The speies was not, however, encountered in the gallery forests of the Chapada dos Veadeiros. The species responds readily to “squeaking” and demonstrates considerable curiosity to unusual activity in its immediate area.
A large series of specimens collected in the Serra Dourada area shows no appreciable difference from specimens from other parts of the species’ range.
Chelidoptera tenebrosa tenebrosa (Pallas) Swallow- wing
These small bucconids were fairly abundant in the Serra Dourada area, but were not noted or collected on the Chapada dos Veadeiros. In the former region the species was most frequently observed at the margins of fazenda clearings in primary type forest. They were gregarious and often seen in flocks of ten to thirty individuals perched in the tops of dead forest trees from which they would make soaring flights out over the clearings. On the wing they looked like tiny miniatures of the black vulture (Coragyps) , which undoubtedly accounts for their Brazilian name of Urubuzinho or “little vulture.”
Three specimens from the Serra Dourada, agree in color and size to the nominate race rather than with the coastal form brasiliensis. Com- pared with a series of tenebrosa from Surinam in the Los Angeles County
1961
Stager: Brazil, Ornithology
23
Museum, the Goias specimens appear to have a smaller and more delicately formed bill, but otherwise agree with the nominate race.
Pteroglossus castanotis australis Cassin Chestnut-eared Aracari
On the Chapada dos Veadeiros these small toucans were observed sparingly in heavy gallery forest, but none were collected. In the primary type forest of the Serra Dourada, however, the species was relatively common and several specimens were secured.
Examination of a large series (57 specimens) of skins of this species in the collections of the American Museum of Natural History from localities throughout its range, show the races of castanotis to be poorly differentiated. Cassin (1867) based his description of australis on the total length of dried study skins, plus the lighter coloring of the head. Measurements of the study series of 57 specimens from throughout the range of the species show no appreciable difference in size between the nominate race and individuals within the range of the supposedly smaller race australis. With regard to color, the series shows that the distribution of the chestnut markings of the head region is very unstable. There is great variation with regard to the width of the chestnut collar and the presence or absence of the chestnut crown patch. The crown patch is more noticeable in southern birds due to the lighter tone of the chestnut marks. The lighter tone of the chestnut markings in birds to the south and east of the range of the nominate form appears to be the only valid difference between these two races. Geographical series appear to show a break in color continuity at about the Rio Madeira rather than at the Rio Negro as stated by Peters (1948).
Ramphastos vitellinus pintoi Peters Sulphur and White-breasted
Toucan
Although apparently absent in the gallery forests of the Chapada dos Veadeiros, this species of toucan is abundant in the heavy primary type forests of the Serra Dourada area. In this latter region the species was frequently observed feeding and traveling through the forest crown in flocks of from four to twenty individuals. Pinto (1936) reported the species as common in the Rio das Almas area to the south of the Serra Dourada region, but Hellmayr (1908) indicates that Baer did not encounter the species during his travels through Goias.
The presence of the white-throated race pintoi in central Goias and northern Sao Paulo is one of the puzzling facets of the entire vitellinus complex. The Goias population is separated from the other two white- throated races, culminatus and the nominate form vitellinus , by the very contrasting orange-throated race ariel. Examination of the extensive series of ariel in the American Museum of Natural History from the lower Amazonian region of Para, fails to show any intergrades tending toward the white-throat of pintoi. In the series of six skins of pintoi from the Serra Dourada area, there is a slight tinge of lemon yellow on the white throat
24
Contributions in Science
No. 41
patch of two of the specimens, while a third (LACM 32529), shows a considerable wash of yellow. There is evidence, however, among other species of toucans to indicate that the yellow wash on the white-throated birds is a juvenile characteristic.
Ramphastos toco P. L. S. Muller Toco Toucan
In central Goias this large toucan appears to be more a species of the heavy cerrado rather than of the more humid primary type forest. It was exceedingly common, both on the Chapada dos Veadeiros and the Serra Dourada. It was also noted as common in the vast tracts of cerrado forest lying between the Serra Dourada region and Peixe to the north. The species did not appear to be as gregarious as the preceding toucan and was usually observed as single indivduals or small groups of two to four birds.
Cabanis (1862:344) described the race albogularis as being, “some- what smaller, with a shorter bill and with a clear white throat, without the yellow tinge and without the fine reddish spots on the border of the throat.” Examination of a very large series of specimens of this species in the collections of the American Museum and the Los Angeles Museum, however, fails to support the validity of this southern race. Three specimens from Metan, Province of Salta, Argentina, in the collection of the American Museum of Natural History (486737-39), have a heavy wash of yellow on the throat as does a specimen from western Bahia in the Los Angeles Museum collection (31532). Each of the above mentioned four specimens is a young individual, as shown by the development of the bill. The large lateral black spot at the distal end of the bill appears to be a useful character in determining adult from juvenile birds. In the adult the spot is in sharp contrast to the yellow of the bill, whereas in juvenile birds, the black gradually shades into the yellow. Numerous southern specimens also show considerable red at the lower border of the white throat, being especially pronounced in juvenile specimens with yellow-tinged throats. With regard to size, the above mentioned large series of birds studied in the two collections shows that specimens from the southern portion of the range are equally as large as northern specimens of the nominate form. Peters (1948) questions the identity of the popula- tions of this species inhabiting Bahia and Piauy, but specimens in the collections of the American Museum from these two areas appear to be quite similar to those from adjacent parts of the range. From this study it can only be concluded that the species is monotypic and that the race albogularis should be synonymized with the nominate form to the north.
Picumnus guttifer Sundevall Spotted Piculet
Although noted as fairly common on the Chapada dos Veadeiros, the collection contains but one specimen of this small woodpecker. On April 21, 1956, an adult $ (LACM (32530) was secured in gallery forest, 20 kms. north of Sao Joao da Alianga. The specimen agrees with a large
1961
Stager: Brazil, Ornithology
25
series of guttifer in the collection of the American Museum of Natural History.
Colaptes campestris campestris (Vieillot) Pampas Flicker
A common species of the open cerrado areas of the Chapada dos Veadeiros. Three specimens, taken 20 kms. north of Sao Joao da Alianga agree with specimens of typical campestris and are larger and darker than the race chrysosternus which occurs to the northeast of this area.
Chrysoptilus melanochloros natter eri Malherbe Black and Yellow
Woodpecker
Three specimens of this woodpecker were secured in secondary forest growth in the Serra Dourada area between May 19 and 27, 1956. All three specimens are males and agree with a large series of typical nattereri in the American Museum collection. Traylor (1951), in his excellent revision of the melanochloros-melanolaimus group of woodpeckers, has at last brought understanding to a heretofore perplexing problem.
Celeus flavescens inter cedens Hellmayr Yellow Woodpecker
A common species in gallery forest of the Chapada dos Veadeiros and primary type forest of the Serra Dourada. A series of five specimens from the above two localities agree with the type of race intercedens taken by Baer at Fazenda Esperanca, some 100 kms. due south of the Serra Dourada area.
Dryocopus lineatus lineatus (Linne) Lineated Woodpecker
An abundant and widespread forest species in central Goias. The series of four specimens (1, Chapada dos Veadeiros; 3, Serra Dourada) agree well with a large series of the nominate race lineatus in the American Museum of Natural History.
Melanerpes flavifrons (Vieillot) Yellow-fronted Woodpecker
Woodpeckers of this species were met with only in the Serra Dourada area, where they were relatively common. Peters (1948) recognizes two races of the yellow-fronted woodpecker based on the descriptions of Vieillot (1818:75 and 103). According to him the nominate race flavifrons is confined to the states of Goias, Minas Gerais, Bahia, Espirito Santo and Rio de Janeiro, while rubriventris includes birds from southeastern Brazil, northeastern Argentina and Paraguay. An examination of a large series of specimens in the collections of the Los Angeles County Museum, however, indicates that the species is badly in need of critical examination before it can safely be divided into two valid races. Specimens from east central Minas Gerais are heavily suffused with red on the breast and abdomen while specimens supposedly assignable to rubriventris from Sao Paulo have the amount of red greatly restricted. Our birds from Goias which, according to Peters, should agree with the specimens from Minas Gerais, lack the suffusion of red on the breast and agree with specimens from the south. It may well be that flavifrons is divisible into two
26
Contributions in Science
No. 41
recognizable races, but it will require examination of large series of specimens before a proper delineation of racial boundaries can be established. Until such a study can be made it appears wise to recognize Melanerpes flavijrons as a monotypic species.
Leuconerpes candidus (Otto) White-bellied Woodpecker
In the Serra Dourada area, woodpeckers of this species were noted fairly frequently in stands of heavy cerrado forest. Although not observed on the Chapada dos Veadeiros, Leuconerpes probably occurs there, for the species is a widespread campo bird of central Brazil. The three specimens collected in the Serra Dourada agree with the large series of skins in the American Museum of Natural History from widespread areas of Brazil. V eniliornis passerinus trails fluvialis Hellmayr Olive Woodpecker
An extremely common species in Central Goias, occurring abundantly on the Chapada dos Veadeiros and in the Serra Dourada area.
Central Goias birds appear to be intermediate between races trans- jluvialis and olivinus of the south. They are closest to trans fluvialis, however, in that they show the back and rump very narrowly barred and spotted with pale yellow. This character is not very stable, however, as Matto Grosso (Chapada) birds show this degree of barring when examined in large series. The throats of Goias birds show the light coloring of trans fluvialis and agree in size and with this northern race. Measure- ments of five specimens in mm.:
2 cf cf
3 $ 9
Wing Tail
89, 90 51
89, 90, 93 48, 51, 52
CULMEN
18.2, 19.6 18.6, 18.9, 19
Phloeoceastes melanoleucos melanoleucos (Gmelin) Black and White
Woodpecker
Large woodpeckers of this species were frequently observed in the dense gallery forests of the Chapada dos Veadeiros where two adult specimens were secured. Individuals from central Goias are larger than specimens of race cearae to the east, but agree in size with a series from farther north in Para. The two specimens secured are uniformly darker on the lower breast and abdomen than the series examined from Para. It will be necessary to assemble a large series of specimens before a proper study of the subspecific relationships of this species can be worked out adequately. Measurements: 2 cf cf, wing 82, 85, tail 97, 103, culmen 46, 48 mm.
Phloeoceastes rubricollis olallae (Gyldenstolpe) Red-necked Woodpecker
The occurrence of this large Amazonian woodpecker has not been previously recorded for the state of Goias. The species was noted on several occasions in heavy prmary forest of the Serra Dourada region and four specimens were secured.
Gyldenstolpe (1945) separated the population of race rubricollis
1961
Stager: Brazil, Ornithology
27
south of the Amazon (Amazonas, Para and Maranhao) into the race olallae. Gyldenstolpe states in his description that specimens from Para have the outermost primary entirely black on both webs and a small rufous basal marking on the penultimate primary. This character does not hold for the series of Goias specimens, nor for a series examined from Para. In both series the rufous markings are well defined on all primaries. With regard to size, the Goias birds agree with measurements given by Gyldenstolpe for olallae, which is smaller than the nominate race rubricollis and the western form trachelopyrus. Measurements for four specimens from central Goias (Serra Dourada) : 1 cf, wing 182, tail 110, culmen 47 ; 3 9 9 , wing 178-181, tail 105-107, culmen 44.5-46 mm.
Literature Cited
Cabanis, J.
1862. Vogel von Cota Rica. Jour, fur Ornith. 10:334.
Cassin, J.
1867. A study of the Ramphastidae. Proc. Acad. Nat. Sci. Phila. 1867, pp. 100-124.
Conover, B.
1950. A study of the spotted tinamous. Fieldiana, Zool. 31(37): 339-362. Dawson, E. Yale
1957. The Machris Brazilian Expedition, Botany: General. Los Angeles Co. Mus. Contr. Sci. (2): 1-20.
Delacour, Jean
1957. The Machris Brazilian Expedition, General Account. Los Angeles Co. Mus. Contr. Sci. (1): 1-11.
Gyldenstolpe, Nils
1945. The bird fauna of Rio Jurua in West Brazil. Kungl. Svenska Veten- skapsakademiens Handlingar. Tredje Serien. 22(3): 1-338, 1 map. Hellmayr, C.E.
1908. An account of the birds collected by Mon. G. A. Baer in the state of Goyaz, Brazil. Novit. Zool. 15: 13-102.
1929. A contribution to the ornithology of northeastern Brazil. Field Mus. Nat. Hist., Zool. Ser., 12(18): 235-501.
Hellmayr, C. E. and Boardman Conover
1942. Catalogue of birds of the Americas and the adjacent islands. Field Mus. Nat. Hist., 13 pt. 1(1): 1-636.
Neumann, O.
1932. Descriptions of three new geographical races of the white-browed guan. Bull. Brit. Ornith. Club, 53: 93.
Pelzeln, A.
1868. Zur ornithologie Brasiliens. Abt. I-IV: 1-462.
Peters, J. L.
1948. Check-list of birds of the world, 6: 1-259.
Pinto, O. M. O.
1936. Contribugao a ornitholgia de Goyaz. Rev. Mus. Paulista, 20: 1-172. 1938. Catalogo das aves do Brasil (Pt. 1) Rev. Mus. Paulista, 22: 1-566. 1944. Catalogo das aves do Brasil (Pt. 2) Dept. Zool., Secret. Agric. Sao Paulo, pp. i-x, 1-700.
Stager, K. E.
1960. The Machris Brazilian Expedition, ornithology: Two new birds from central Goias, Brazil. Los Angeles Co. Mus. Contr. Sci. (33): 1-7. Traylor, M. A.
1951. A review of the woodpeckers Chrysoptilus melanochloros and Chrysoptilus melanolaimus. Fieldiana, Zool. 31(41): 412-437.
Vieillot, L. J. P.
1818. Nouv. Diet. Hist. Nat., 26, pp. 75 and 103.
LOS ANGELES COUNTY MUSEUM CONTRIBUTIONS IN SCIENCE
Nos. 1-34. Table of Contents available (1957-1959).
No. 35. The Machris Brazilian Expedition, Botany: Pteridophyta, by C. Y. Morton.
No. 36. The Machris Brazilian Expedition, Entomology: Two new species of Anacroneuria (Plecoptera) from Goias, Brazil, by Stanley G. Jewett, Jr.
No. 37. A new genus and species of glossophagine bat from Colima, Mexico, by W. J. Schaldach and Charles A. McLaughlin.
No. 38. A census of the abundant large Pleistocene mammals from Rancho La Brea, by Leslie F. Marcus.
No. 39. A skull of the Grizzly Bear ( Ursus arctos L.) from Pit 10, Rancho La Brea, by Bjorn Kurten.
No. 40. The Machris Expedition to Tchad, Africa. Amphibians and Reptiles, by David B. Wake and Arnold G. Kluge.
No. 41. The Machris Brazilian Expedition, Ornithology: Non-passerines, by
Kenneth E. Stager.
MBER 42
June 30, 1961
-
\S01. T3
p.kW
A NEW GEOMYS
FROM THE YALLECITO CREEK PLEISTOCENE OF CALIFORNIA
With Notes on Variation in Recent and Fossil Species
By John A. White and Theodore Downs
Angeles County Museum
Exposition Park
Los Angeles 7, Calif.
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. Issues are numbered separately, and numbers run consecutively regardless of subject matter. Number 1 was issued January 23, 1957. The series is available to scientists and scientific institutions on an exchange basis. Copies may also be purchased at a nominal price.
Hildegarde Howard Editor
E. Yale Dawson Associate Editor
A NEW GEOMYS FROM THE VALLECITO CREEK PLEISTOCENE OF CALIFORNIA, with notes on variation in recent and fossil species
By John A. White1 and Theodore Downs2 Introduction
The Vertebrate Paleontology Section of the Los Angeles County Museum has carried on extensive field operations during the last three years in the highly fossiliferous badlands of the Vallecito Creek valley of the Anza-Borrego desert in the western Imperial Valley of southern California. A preliminary report on the varied Middle Pleistocene verte- brate fauna collected from this area has been presented (Downs, 1957), and a complete study of this fauna, and related stratigraphy, geology, and paleoecology, is in the process of preparation.
One of the species best represented in the fauna is a pocket gopher of the genus Geomys. Approximately 75 well-preserved portions of skulls and mandibles of this gopher are present in the collection. So adequate a sample as this provides an unusual opportunity for the paleontologist to derive more biologically realistic conclusions than are usually possible in his studies. This paper, therefore, is not limited to a description of the species represented by this fossil material, hut includes: (1) analyses of the relationships of the fossil species and other known forms of Geomys , both living and extinct; (2) comments on possible paleoecological infer- ences, particularly with regard to the importance of intraspecific variation in paleozoological as well as neozoological studies.
Acknowledgments: To Harley J. Garbani we are grateful for the discovery of the fossil-bearing strata in Vallecito Creek and for his enthu- siastic aid in the field.
Although Downs initiated the study of the Vallecito Creek fauna in 1954, grants from the National Science Foundation (G-5035) to the Los Angeles County Museum Associates, in 1958 and 1960, provided him the means for more accelerated furtherance of the project.
The field work was made possible by the support and cooperation of the Department of Beaches and Parks, State of California, especially the management and personnel of the Anza-Borrego Desert State Park.
We are grateful to Charles A. McLaughlin for making the drawings and reading the manuscript critically; to Alice White and Edward D. Mitchell Jr. for preparation of the map and diagrams; and to G. Davidson Woodard for aid in stratigraphic interpretations.
Research Associate, Los Angeles County Museum, Associate Professor, Long Beach State College.
2Curator of Vertebrate Paleontology, Los Angeles County Museum.
4
Contributions in Science
No. 42
We wish to thank the following individuals and institutions for the loan of comparative material: E. Raymond Hall and Robert W. Wilson, University of Kansas Museum of Natural History (U.K.M.) ; William B. Davis, Department of Wildlife Management, Agricultural and Mechanical College of Texas (T.A.M.) ; C. Lewis Gazin and Charles Handley, United States National Museum (U.S.M.) ; Seth B. Benson, Museum of Vertebrate Zoology, University of California, Berkeley (M.V.Z.) ; W. Frank Blair, Department of Zoology, University of Texas (Z.U.T.) ; Bryan P. Glass, Department of Zoology, Oklahoma State University (O.S.U.) ; Donald F. Hoffmeister, Museum of Natural History, University of Illinois (U.I.M.) ; William D. Turnbull, Chicago Natural History Museum (C.N.M.) ; Rachel H. Nichols, American Museum of Natural History (A.M.N.), and Kenneth E. Stager and Charles A. McLaughlin, Los Angeles County Museum (L.A.M.).
The Genus Geomys
Pocket gophers are fossorial rodents that spend most of their time underground. They are, therefore, somewhat sedentary, and populations tend to be confined in a relatively small area. According to Wright (1940, “Sewell Wright Effect”), the inbreeding of populations in a small area results in a random fixation of characters that usually does not occur in populations of larger areas. This may account for the well-known intraspecific variability of pocket gophers and may explain why the various species are not easily distinguishable morphologically (see Durrant, 1953). Statistical techniques are necessary in the study of the taxonomy of these forms.
The genus Geomys is defined by the following characteristics of the skull (Merriam, 1895) : the upper incisors are doubly grooved on the anterior faces, the smaller, shallower groove being situated mediad to the larger, deeper one. P4 in adult individuals lacks enamel on the posterior border of the biting surface. Mx and M2 lack enamel on the anterior borders of the biting surfaces.
All the teeth in Geomys are rootless and ever-growing. The occlusal surface of the grinding teeth is characterized by the presence of alternating bands of enamel and dentine. These “bands” are formed by the loss of enamel from the buccal, lingual, and, less often, from the anterior or posterior surfaces of the teeth.
The occurrence of Geomys in the Vallecito Creek Pleistocene is of interest, for the genus has never before been recorded from the fossil or Recent record in California. Extant and extinct species are known in eastern, central, and southwestern United States as far west as Lordsburg, New Mexico, and in northeastern Mexico (Fig. 1).
There are seven described, living species of Geomys (Fig. 1). Three of these {G. cumberlandius, G. fontanelus , and G. colonus ) are represented by small, distinct populations found along the coast of the state of Georgia. The other four species (G. bursarius, G. personatus, G. arenarius, and
1961
White & Downs: Fossil Geomys
5
G. pinetis) are polytypic, containing two or more subspecies or geographic races.
Four extinct species have been described: G. parvidens (= G. bursarius parvidens ) from the Conard Fissure Late Pleistocene of Arkansas; G. tobinensis from the Tobin Ranch local fauna Mid Pleistocene of Kansas; G. quinni from the Sand Draw local fauna Early Pleistocene of Nebraska; and G. bisulcatus from the ?Pliocene of Nebraska (Table 8).
Two extinct genera, closely related to Geomys , have been described: Nerterogeomys ( N . minor and N. persimilis) from the Early to ?Mid Pleistocene of the San Pedro Valley of Arizona (Table 8) ; and Pliogeomys ( P . buisi) from the Late Pliocene Buis Ranch local fauna of Oklahoma.
Fig. 1. Map showing the geographic distribution of the species of Geomys and of the genera, Nerterogeomys and Zygogeomys. The encircled numbers refer to living species having a wide geographic distribution while the plain numbers refer to both living and extinct species having a limited geographic distribution.
1. Geomys bursarius 9. G. quinni and Nerterogeomys
2. G. personatus 10. G. tobinensis
3. G. arenarius 11. G. quinni
4. G. pinetis 12. N erterogeomys
5. G. fontanelus 13. Nerterogeomys
6. G. colonus 14. G. garbanii
7. G. cumberlandius 15. Zygogeomys
8. G. bursarius parvidens
6
Contributions in Science
No. 42
Materials
All comparative materials used in this study are herein listed by species and subspecies name, together with region of collection, source institution (abbreviated as indicated under acknowledgments), and number of specimens.
Geomys bursarius (Shaw)
G. b. attwateri : Texas, U.S.M. 22, L.A.M. 9. G. b. brazensis : Texas,
T. A.M. 10. G. b. breviceps : Louisiana, U.S.M. 25. G. b. dutcheri : Okla- homa, O.S.U. 28. G. b. illinoensis: Illinois, U.I.M. 25, L.A.M. 3. G. b. industrius: Kansas, U.K.M. 15. G. b. jugossicularis : Kansas, U.K.M. 11. G. b. lutescens : Kansas, U.K.M. 9. G. b. major : Kansas, U.K.M. 18; New Mexico, L.A.M. 5. G. b. majusculus : Kansas, U.K.M. 20. G. b. pratincolus : Texas, T.A.M. 10. G. b. sagittalis: Texas, T.A.M. 6.
G. personatus True
G. p. fallax : Texas, Z.U.T. 25, U.S.M. 1. G. p. megapotamus : Texas, Z.U.T. 25. G. p. personatus : Texas, L.A.M. 5.
G. arenarius Merriam
G. a. arenarius : Texas, M.V.Z. 15; New Mexico, L.A.M. 6. G. a. brevirostris: New Mexico, M.V.Z. 19, L.A.M. 5.
G. pinetis Rafinesque
G. p. austrinus : Florida, L.A.M. 1. G. p. jloridanus : Florida, U.S.M. 14. G. p. mobilensis: Florida, L.A.M. 1.
Geomys bursarius (Shaw)
G. bursarius : Kansas, Cragin Quarry local fauna, Late Pleistocene (Hibbard 1960), U.K.M. 1 cranium. G. b. parvidens : Arkansas, Conard Fissure, Late Pleistocene (Brown 1908), A.M.N. 1 cranium, 18 mandibles, 4 palates.
Geomys quinni McGrew
G. quinni : Nebraska, Sand Draw local fauna, Early Pleistocene (McGrew 1944), C. N. M. 12 mandibles and other miscellaneous elements and fragments; Kansas, Rexroad Fauna, Early Pleistocene (Franzen 1947),
U. K.M. 3 mandibles.
Geomys tobinensis Hibbard
G. tobinensis: Kansas, Tobin Fauna, Mid Pleistocene (Hibbard 1944), U.K.M. 1 mandible with P4 — M2; 1 P4.
Methods used in the study of variation
Two types of variants were studied. One type involves analysis of measurements of continuous variation, the other analysis of arbitrary categories of discontinuous variants, such as categories of shapes or degrees of expression of unmeasurable characters.
Measurements were made with dial calipers calibrated to 0.1 mm. All measurements were taken in exactly the same manner to reduce sampling error. In selecting measurements it was felt that the usual measurements of the neo-mammalogist as well as those of the students of fossil Geomys should be used in order to provide comparisons with living
1961
White & Downs: Fossil Geomys
members of the genus. Charles A. McLaughlin, Associate Curator of Mammals at the Los Angeles County Museum, who recently completed a manuscript on the taxonomy of certain groups of living Geomys , was consulted in the selection of these measurements.
The measurements of the cranium were taken as follows:
Basilar length: from the posterior surface of the upper incisor at the alveolus to the ventral, medial notch of the foramen magnum.
Length of the alveolus of the upper cheek-tooth row: from the anterior edge of the alveolus of P4 to the posterior edge of the alveolus of M3.
Length of upper diastema: from the posterior surface of the upper incisor at its alveolus to the anterior edge of the alveolus of P4.
Least interorbital constriction: self explanatory.
Breadth of rostrum at the ventral edge of the infraorbital CANALS: taken by placing the points of the calipers in each opening of the infraorbital canals and holding calipers perpendicular to the antero- posterior axis of the skull.
Breadth of rostrum at the maxillary-premaxillary suture on side of ROSTRUM: self explanatory.
Least depth of rostrum: taken at the narrowest part of the rostrum by placing one point of the calipers on the arched premaxillary (ventrally) and the other point on the dorsal surface of the nasals.
Palatofrontal depth of skull: from the ventral surface of the bony palate to the dorsal surface of the frontals.
The measurements of the mandible were taken as follows:
Length of mandible: from the posterior surface of a lower incisor at the alveolus to the posterior-most surface of the condyloid process.
Length of the alveolus of the lower cheek-tooth row: as in the upper cheek-tooth row.
Length of lower diastema: as in the upper diastema.
Distance from condyloid to angular process of mandible: from the dorso-medial surface of the condyloid process to the ventro-lateral surface of the angular process.
Tooth-row projection to mental foramen: from the “ventral” surface of a straight edge layed on the biting surface of P4 to the mental foramen. In each case the shortest distance between the mental foramen and the straight edge was estimated.
Mental foramen to posterior surface of M2: from the mental foramen to the mid-point on the posterior edge of M2.
The establishment of categories for “unmeasurable” characters is best done in the discussion of such categories.
Statistical analysis was made for each of the samples when such samples were sufficiently large to permit analysis. For each sample, the mean, standard deviation, standard error of the mean, coefficient of
8
Contributions in Science
No. 42
variability, and Student’s t-value (Simpson et al , 1960) of the 95 per cent confidence level, were obtained.
To test the significance of differences between the various samples, a modification of the Dice-Leraas (1936) method was chosen. In cases where it was deemed desirable to check further the degree of difference between two samples, a t-value was calculated and the limits indicated by it listed along with the mention of such comparisons in the text. The limits of the t-values generally accepted as indicating significant differences between two samples range between .05 and .001, which means that only 5.0 to 0.1 per cent of the time would a value fall outside of these limits (Simpson et al, 1960). For example, when the length of the lower cheek-tooth row in G. garbanii is compared with that in G. p. megapotamus , the calculated t-value is 2.570. Using the appropriate table, the t-value falls between 2.473 and 2.771 and would be expected to fall outside of these limits only 1 to 2 per cent of the time. The two samples, therefore, are probably significantly different to the .02 level of significance.
The specimens used in analyses were adults. The segregation of specimens into juveniles and adults was done somewhat subjectively and based on: (1) the degree of sutural fusion, especially between the
basisphenoids and basioccipitals, and between the supraoccipitals and exoccipitals ; (2) the degree of development of the sagittal crest or the extent to which the ridges which limit the dorsal extension of the temporal muscles approximate one another on the dorsal surface of the skull; (3) the presence or absence of enamel plates or the interruption of the enamel band around P4 and P4; (4) the relative size of the specimens within a given sample. This latter criterion is the least reliable, but nevertheless seems to serve well, and was necessarily used in dealing with fragmentary material.
Geomys garbanii3 new species
The Geomys from the Vallecito Creek fauna is distinct from all other species of the genus and, thus, is accorded a new name and described and discussed below.
Holotype: L.A.M. No. 3483; nearly complete cranium and asso- ciated mandibles.
Type locality: L.A.M. Vert. Paleo. loc. no. 1380. Vallecito Creek, Carrizo Mountain quadrangle, 1953, AMS sheet 2749 1, series V795, San Diego Co., California; grid coordinates 11,575,500 — 3,641,000.
Fauna: Vallecito Creek.
Age: Middle Pleistocene, Irvingtonian provincial age; also see Downs (1957) ; complete faunal report to be published.
Formation: Palm Springs (see Downs, in report of Continental Cenozoic Committee of the Society of Vertebrate Paleontology, in press) .
Referred material: (listed according to element, L.A.M. locality
3Named for Harley J. Garbani, the discoverer of the Vallecito Creek vertebrate fauna.
1961
White & Downs: Fossil Geomys
9
no., and catalog no.). Cranium and associated mandibles: loc. 1192, no. 3231; loc. 1314, no. 3480; cranium: loc. 1296, no. 3481; loc. 1314, no. 3525; posterior part of cranium with bullae and M2 M3: loc. 1197, no. 3229; partial cranium and associated mandibles: loc. 1192, no. 3230; loc. 1316, no. 3511; loc. 1508 or 1509, no. 4185; loc. 1548, no. 4244; partial rostra: loc. 1114, no. 1553; loc. 1191, no. 3252; loc. 1192, nos. 3352, 3491; loc. 1249, no. 3354; loc. 1296, no. 3500; loc. 1314, no. 3526; loc. 1317, no. 4186; loc. 1318, no. 3518; loc. 1386, no. 4187; loc. 1518, no. 4199; palate with both rows of cheek-teeth: loc. 1246-3, no. 3384; loc. 1514, no. 4189; undersurface of rostrum: loc. 1325, no. 4190; interorbital area: loc. 1297, no. 3524; P4: loc. 1114, nos. 3404.1, 3404.2, 3404.3; M3: loc. 1297, no. 4191; mandibles: loc. 1113, nos. 1583, 3403; loc. 1114, nos. 1551, 1552, 1554; loc. 1191, nos. 3244, 3250; loc. 1192, nos. 3352, 3492; loc. 1197, no. 3228; loc. 1248, no. 3684; loc. 1250, no. 3347; loc. 1296, no. 3499; loc. 1297, nos. 3482, 3485, 3486, 3488, 3494, 3495, 3501, 3502, 3503, 3504, 3524; loc. 1299, no. 3487; loc. 1300, no. 3496; loc. 1308, no. 3484; loc. 1313, no. 3497; loc. 1314, no. 4192; loc. 1316, nos. 3508, 3509, 3510; loc. 1317, nos. 3514, 3515; loc. 1318, nos. 4193, 3516, 3517;
loc. 1324, nos. 3519, 3520; loc. 1333, no. 3521; loc. 1336, no. 4194; loc.
1337, no. 3522; loc. 1448, no. 4198; loc. 1474, no. 4195; loc. 1500, no.
4196; loc. 1511, no. 4197; loc. 1518, no. 4188; loc. Vallecito Creek, no.
3523; partial pelvis and other doubtfully associated bones: loc. 1192, unnumbered.
Diagnosis: Large, about the size of a large Geomys bursarius or G. personatus ; rostrum deeper; tympanic bullae and opening of the external auditory meatus larger than in any other species.
Description of the holotype: A skull (Fig. 16) and associated mandibles of a fully adult animal. Left zygomatic arch present and parallel to longitudinal axis; supraorbital ridges high, delimiting a trough-like depression between them; sagittal crest not quite formed, but dorsally placed ridges bounding the temporal muscle scar about 3 mm. apart; skull table flat and rostrum sloping ventro-anteriorly ; nasals slightly flared anteriorly, nearly parallel-sided in the posterior one-third ; external auditory meatus opening dorsally, posterior to zygomatic arch and measuring 2.6 mm. in inside diameter and 4.6 mm. in outside diameter; lambdoidal crest moderately developed; anterior surface of upper incisors posterior to anterior extension of nasals; rostrum, dorsally, approximately as broad as deep; cheek-tooth rows diverging posteriorly, and posterior extent of palate distinctly posterior to right and left M3; tympanic bullae large, with inner margins diverging posteriorly to level of ventral notch of foramen magnum; tube-like structure enclosing auditory meatus extending dorso- anteriorly, terminating in a large external opening; coronoid, condyloid, and angular processes of the left mandible broken at their tips; a shallow pit present between coronoid process and capsular process of lower incisor root; valley absent between condyloid and capsular processes; mental
10
Contributions in Science
No. 42
foramen anterior to P4 and anterior to foremost extension of masseteric scar.
Detailed Description
Clearly a member of the genus Geomys: bisulcate upper incisors; enamel plate usually completely absent from posterior surface of P4 and always absent from anterior surfaces of M4 and M2 in adults.
The skull: Supraorbital ridges high and more developed than in other species, a depression in the frontal bone between them; tympanic bullae markedly inflated and larger than in any other species; external auditory meatus markedly larger than in any other species (see Fig. 13) ; length of basioccipital (although difficult to measure in adult specimens) seeming to exceed, in some specimens, the breadth of the rostrum at the maxillary-premaxillary junction, and to be less than this breadth in other specimens; zygomatic arches parallel to longitudinal axis of cranium; rostrum noticeably deeper, dorso-ventrally ; upper incisors more recurved than in other species, and their anterior surfaces posterior to anterior extension of nasals; skull table flattened and rostrum sloping slightly ventrally and anteriorly; sides of nasal bones more or less straight, conver- gent and truncated posteriorly.
The relationship of the length of the basioccipital to the breadth of rostrum at the maxillary-premaxillary junctions is difficult to determine because the suture between the basioccipital and basisphenoid is frequently absent, but the basioccipital seems longer than the rostrum is wide in the holotype and the reverse in specimen no. 3231. This character has been used to distinguish G. personatus and G. arenarius on the one hand, from G. bursarius and G. pinetis on the other hand, without any “overlap” (Davis, 1940).
Table 1
Frequency distribution of the measurements of specimens of G. garbanii n. sp. All measurable specimens are included (in millimeters). The mean and confidence limits are plotted over each frequency distribution. No confidence limits were calculated for basilar length of skull, depth of rostrum, and the distance from condyloid to angular process. The encircled capital letters correspond to skull measurements as follows: A. basilar length of skull; B. length of upper diastema; C. palatofrontal depth of skull; D. length of mandible; E. length of upper cheek-tooth row; F. breadth of rostrum at the maxillary-premaxillary junction; G. depth of rostrum; H. length of lower cheek-tooth row; I. distance from condyloid to angular process; J. distance from mental foramen to posterior surface of M2; K. least interorbital constriction; L. breadth of rostrum ventral to infra- orbital canals; M. length of lower diastema; N. lower tooth-row projection to mental foramen.
1 1
1
1
©
©
©
42.0 43.0 44.0 45.0 46.0 47.0 48.0 49.0 50.0 5L0
13.0
1
14.0
i 1 11 1 1
15.0 16.0 17.0
I | i
1111 11 1
15.0 16.0 170 18.0 19.0 20.0 21.0 22.0
18.0
1 1 1 11
24.0 25.0 26.0 27.0 28.0 29.0 30.0 31.0 32.0 33.0 34.0 35.0
©
©
11 111
1 1 1
1 1
21
1 1
©
©
©
©
©
1
1 1
8.0
1
I
1 1 11
1
211 1
1
2 11 1 1233121111 1
9.0 10.0 II. O
1 1 12.0
13.0
1 111111
r 1 1
112 11 1
I 1 1
1 1 11 11 11 121311 1 1 12 1
I 1 1
1 11 1 1 41113 12 26 1 1 112 1 2 1
5.0 6.0 7.0 8.0 9.0 10.0
12
Contributions in Science
No. 42
The mandible: The position of the mental foramen varies con- siderably from a position ventral to P4, and the anterior extension of the masseteric scar to a position anterior to these structures. This variability does not seem to be correlated with stratigraphic position in the section, but is present when a series of specimens is collected at a given locality. The position of the mental foramen is, therefore, variable and of little or no taxonomic significance. The dentary foramen is variably placed with respect to the posterior end of the pit between M3 and the ascending ramus, and may be oriented nearly at right angles with the medial border of the pit for insertion of the temporal muscle, or the angle may be as much as an estimated 135 degrees.
Table 2
Statistical data relating to fourteen measurements taken on specimens of Geomys garbanii n. sp. The capital letters refer to measurements as indicated in Fig. 2. The abbreviations are as follows: ar. m. = arithmetic mean; N = number of items in sample; st. e. m. = standard error of the mean; t = student’s t-value of confidence limits; s standard deviation; V — coefficient of variability.
|
N |
ar. m. |
Min. |
Max. |
st. e. m. |
t |
|
|
A |
4 |
45.0 |
42.5 |
50.5 |
2.37 |
6.561 |
|
B |
7 |
11.0 |
9.9 |
12.3 |
.393 |
.888 |
|
C |
8 |
16.2 |
14.0 |
18.4 |
.472 |
1.08 |
|
D |
6 |
7.4 |
6.2 |
8.1 |
.320 |
.753 |
|
E |
6 |
5.6 |
5.0 |
6.2 |
.228 |
.512 |
|
F |
5 |
11.3 |
9.8 |
12.8 |
.534 |
1.33 |
|
G |
3 |
10.9 |
10.1 |
11.5 |
— |
|
|
H |
6 |
18.3 |
14.9 |
22.0 |
1.16 |
2.72 |
|
I |
5 |
29.7 |
24.1 |
33.2 |
2.02 |
4.99 |
|
J |
10 |
10.4 |
9.0 |
12.3 |
.336 |
.724 |
|
K |
21 |
8.3 |
7.1 |
9.9 |
.164 |
.334 |
|
L |
1 |
11.3 |
— |
— |
— |
— |
|
M |
34 |
7.5 |
5.9 |
9.5 |
.159 |
.319 |
|
N |
25 |
10.2 |
8.5 |
12.0 |
.165 |
.334 |
9.1
8.7
7.7
4.4
8.8
9.4
14.1 13.6
9.7
8.8
12.2 8.0
.11
.962
1.29
.717
.487
1.07
2.59
4.03
1.01
.74
.916
.808
1961
White & Downs: Fossil Geomys
13
The diastema tends to be shorter in G. garbanii (see figs. 2-5 and table 2), when specimens of G. garbanii are compared with specimens of comparable size in any other species. No valley exists between the capsular process of the root of the lower incisor and the condyloid process. This character is as in the other species of Geomys and unlike Pliogeomys as shown by Hibbard (1954). Between M3 and the ascending ramus there is a pit for insertion of the temporal muscle; the pit seems shallower than in most species of the genus. (continue on P. 18)
Table 3
Frequency in number and per cent of variations in expression of the postero-lateral fold of M3.
|
Species |
Absent |
Slight |
Marked |
|||
|
N |
% |
N |
% |
N |
% |
|
|
Geomys garbanii |
3 |
23.2 |
5 |
38.4 |
5 |
38.4 |
|
G. bursarius |
114 |
75.6 |
34 |
22.5 |
3 |
1.9 |
|
G. personatus |
34 |
62.0 |
18 |
32.5 |
3 |
5.5 |
|
G. arenarius |
20 |
58.9 |
13 |
38.2 |
1 |
2.9 |
|
G. pinetis |
6 |
42.8 |
7 |
50.0 |
1 |
7.2 |
|
All Living Species Combined |
174 |
59.8 |
72 |
35.8 |
8 |
4.4 |
|
Table 4 |
Antero-posterior, outside diameter of the external opening of the auditory meatus in G. garbanii compared with two species that seem to have the largest such structures. For abbreviations see Table 2.
|
Name |
N |
ar. m. |
Min. Max. |
st. e.m. |
t |
V |
s |
|
|
Geomys arenarius brevirostris |
14 |
2.6 |
2.1 |
3.0 |
.083 |
.179 |
12.1 |
.312 |
|
G. personatus megapotamus |
18 |
2.9 |
2.4 |
3.3 |
.058 |
.122 |
8.6 |
.246 |
G. garbanii
3
4.6
3.9
5.3
14
Contributions in Science
No. 42
Table 5
Measurements of Geomys bisulcatus Marsh (1871) and G. garbanii (six numbered L.A. County Museum specimens). All measurements in millimeters.
G. bisulcatus Geomys garbanii
|
3483 |
3231 |
3481 3230 |
3525 |
3482 |
||
|
Transverse diameter of upper incisor |
3.4 |
2.8 |
3.2 |
2.7 |
||
|
Antero-posterior extent of upper incisor |
4.2 |
3.5 |
3.3 |
3.3 |
||
|
Depth of skull at the premaxillary suture |
12.7 |
13.3 |
14.5 |
12.7 |
||
|
Length of lower incisor on arc of curve |
31.8 |
35.6 |
33.1 |
32.9 |
||
|
Transverse diameter of apex of lower incisor |
3.2 |
2.8 |
2.9 |
2.7 |
||
|
Depth of lower jaw below first molar |
11.9 |
10.4 |
11.3 |
10.9 |
11.9 |
8.6 |
|
Antero-posterior extent of P4-M2 |
7.4 |
6.7 |
7.1 |
7.3 |
7.5 |
7.3 |
|
Table 6 |
The mean and other statistical data pertaining to the calculated incisor length in several kinds of pocket gophers. Abbreviations as in
|
Table 2. Name |
N |
ar. m. |
Min. |
Max. |
st. e. m. |
t |
V |
s |
|
Geomys bursarius |
||||||||
|
majusculus |
17 |
35.6 |
32.7 |
39.0 |
.487 |
1.003 |
5.5 |
1.95 |
|
G. personatus |
||||||||
|
megapotamus |
24 |
34.2 |
30.2 |
36.4 |
.426 |
.861 |
5.9 |
2.04 |
|
G arenarius |
||||||||
|
brevirostris |
19 |
35.4 |
31.8 |
37.8 |
.538 |
1.101 |
6.4 |
2.28 |
|
G. garbanii * |
3 |
38.5 |
32.0 |
42.2 |
— |
— |
— |
— |
|
G. quinni |
1 |
44.2 |
— |
— |
— |
. — |
— _ |
— |
*One immature specimen (measuring 32.2) has the same relative incisor length as in specimens of living species. The other two specimens are adults and have markedly longer incisors.
1961
White & Downs: Fossil Geomys
15
.18 .16 .14 .12 .10 .08 .06 .04 .02 -0
.18 .16 .14 .12 .10 .08 .06 .04 .02 -0+ .02 .04 .06 .08
Figs. 2-3. Ratio diagrams modified from Simpson et al (1960), comparing dimensions of the skull in Geomys arenarius (Fig. 2) and Geomys personatus (Fig. 3) with those in G. garbanii. The logs of the means of the dimensions in the latter species are assumed to be zero. The differences between the log of the mean in G. garbanii (standard) and species being compared are plotted to the positive (-(-) or negative ( — ) sides of the zero line. For each measurement the largest and the smallest means of the species being compared are used. The capital letters to the left of the diagrams in figures 2-5 refer to skull dimensions as follows: A. basilar length of skull; B. length of upper cheek-tooth row; C. length of upper diastema; D. least interorbital constriction; E. breadth of rostrum ventral to the infraorbital canals; F. breadth of rostrum at the maxillary-premaxillary junction; G. depth of rostrum; H. palato-frontal depth of skull; I. length of mandible; J. length of lower cheek-tooth row; K. length of lower diastema; L. distance from condyloid to angular process; M. lower tooth-row projection to mental foramen; N. mental foramen to posterior surface of M2.
16
Contributions in Science
No. 42
.24 .22 .20 .18 .16 .14 .12 .10 .08 .06 .04 .02 -0+.02.04 .06
Fig. 4. Ratio diagrams modified from Simpson et al (1960), comparing dimen- sions of the skull in Geomys bursarius and G. b. parvidens (broken line) with those in G. garbanii. For additional details see Fig. 2.
1961
White & Downs: Fossil Geomys
17
.18 .16 .14 .12 .10 .08 .06 .04 .02 -0+ .02 .04 .06 .08 .10
Fig. 5. Ratio diagram modified from Simpson et al (1960), comparing dimensions of the skull in Geomys pinetis (dots) and G. quinni (triangles) with those in G. garbanii. For additional details see Figs. 2 and 3.
18
Contributions in Science
No. 42
Dentition : The upper incisors in G. garbanii are bisulcate as in all Geomys. As in G. quinni , the upper incisors are more procumbent (re- curved) than in any other species (W. D. Turnbull examined this character for us on the type specimen of G. quinni ) . The anterior surfaces of these incisors are posterior to the anterior extension of the nasals in both latter species ( Figs. 16 and 17). No other species of geomyid has this character (Wilson 1949). The upper and lower series of grinding teeth are rootless and evergrowing. Reentrants on P4 and P4 are nearly parallel-sided, forming right angles with their innermost boundaries. In this regard G. garbanii is like all other species except G. quinni wherein sides of reentrants diverge (Franzen 1947). This latter condition is found in P4 and P4 of immature specimens of all species and may represent an embryonic char- acter which persists in adults only of G. quinni. Enamel patterns of P4 and P4 are interrupted by bands of dentine as in living species. The enamel forms a continuous layer completely surrounding the dentine of the occlusal surface in immature specimens of all species. On the sides of such teeth are inverted “V’s” (see Fig. 15) of dentine (Hibbard, 1954) which, with additional wear of the occlusal surface would cause the apex of the “V’s” to reach the occlusal surface and thus interrupt the ringing bands of enamel. W