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UTHERN CALIFORNIA ACADEMY OF SCIENCES
_ BCAS-A88(3) 93-136 (1989) DrerMprn 1989. \) ae
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| Southern California Academy of Sciences
‘ Founded 6 November 1891, incorporated 17 May 1907
© Southern California Academy of Sciences, 1989
OFFICERS.
Camm C. Swift, President
June Lindstedt Siva, Vice-President
Hans M. Bozler, Secretary
Takashi Hoshizaki, T: reasurer
Jon E. Keeley, Technical Editor
Gretchen Sibley, Managing Editor
BOARD OF DIRECTORS
1987-1989 1988-1990 1989-1991
Larry G. Allen Sarah B. George Takashi Hos!
Hans M. Bozler Margaret C. Jefferson George T. J
Allan D. Griesemer Susanne Lawrenz-Miller David L. Soltz
Peter L. Haaker John D. Soule Camm C. ‘Swi te
June Lindstedt Siva. Gloria J. Takahashi —Robert G. Zaha ary
Fellows: Elected by the Board of Directors for meritorious services.
The Bulletin is published three times each year OY the Academy. Manuscripts for publica
be sent to the appropriate editor as explained in “Instructions for Authors” on the inst de one
of each number. All other communications should be addressed to the Southern Ca nia A
of Sciences in care of the Natural History Museum of Los Angeles County, Exposit ion Par K,
Angeles. California 90007. Bis
Date of this issue 19 December 1989
THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. |
Bull. Southern California Acad. Sci.
88(3), 1989, pp. 93-102
© Southern California Academy of Sciences, 1989
Late Pleistocene Freshwater Fishes from the Rancho La Brea
Deposit, Southern California
Camm C. Swift
Section of Fishes, Natural History Museum of Los Angeles County,
900 Exposition Boulevard, Los Angeles, California 90007
Abstract. — Three species of Pleistocene freshwater fishes occur at Rancho La Brea,
the type deposit of the Rancho La Brea Age Mammalian faunas. Remains of
rainbow trout, Salmo gairdneri, (mostly isolated vertebrae) and threespine stick-
leback, Gasterosteus aculeatus (primarily pelvic spines), are both common. One
dentary and two vertebrae represent one individual of arroyo chub, Gila orcutti.
Based on pelvic and dorsal spine morphology, a relatively unarmored form of
stickleback (possibly representing the recent Gasterosteus aculeatus williamsoni)
existed in the Los Angeles Basin up to 30,000 years ago. Collectively the fish
fossils indicate local, permanent stream conditions, and not stream transport from
distant mountainous areas 10-30 km away. Absence of the more montane Ca-
tostomus santaanae and Rhinichthys osculus also argues against long-distance
transport. As with other ectothermic organisms (and possibly small endotherms),
no extinct freshwater fishes are known from the Rancho La Brea deposits.
Rancho La Brea is the largest and best known late Pleistocene terrestrial deposit
in North America. It is the type locality of Rancholabrean Age mammalian faunas
(Savage 1951), and also includes a large fauna of birds, amphibians, and reptiles
(Gehlbach 1965; Stock 1956; Harris and Jefferson 1985; Shaw and Quinn 1986).
Fine sorting for microvertebrates, invertebrates and plants has yielded three taxa
of freshwater fishes. These fishes, given preliminary notice by Akersten (1980),
Akersten et al. (1983), Marcus and Berger (1984), and Harris and Jefferson (1985),
represent living species as have most North American Quaternary freshwater fishes
thus far (Miller 1965; M. L. Smith 1981; G. R. Smith 1981).
The fish remains came from fluvial channel deposits up to 5 m deep in Pit 91
at Rancho La Brea, a rectangular pit in the northwestern area of the deposit
(Marcus and Berger 1984, Fig. 8.2). The pit was opened in 1915, but remained
unworked until 1969, when detailed excavation began, and continues today (Shaw
1982; Shaw and Quinn 1986). About 5% of the excavated sediment from Pit 91
has been sorted. A detailed stratigraphic interpretation of the Rancho La Brea
area was given by Woodard and Marcus (1973). Marcus and Berger (1984) and
Shaw and Quinn (1986) summarized much of the research to date.
Methods and Materials
The collecting methods at Rancho La Brea are published (Miller 1972; Shaw
1982). The fish fossils were compared with skeletal material of Recent taxa of the
same sizes.
Seventy fish bones (Fig. 1) have been recovered and are catalogued in the George
C. Page Museum, Natural History Museum of Los Angeles County, as follows:
93
94 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Fossil Material
Family Salmonidae
Salmo gairdneri Richardson, rainbow trout (15 bones); one ceratohyal 4.2 mm
long, R51296 (Fig. 1A), one atlas vertebra, 1.8 mm long, R45562; five abdominal
vertebrae, 1.3-1.4 mm long, R39859, R45558(2) (Fig. 1B), R45559-60; nine
caudal vertebrae, 0.9-2.0 mm long, R15824, R45551 (Fig. 1C), R45557, R45559,
R45561, R45563-64, R49533, R51295.
Family Cyprinidae
Gila orcutti (Eigenmann and Eigenmann), arroyo chub (four bones); one frag-
mentary left dentary, 3.6 mm long, R45570 (Fig. 1D, E); one anterior (first)
vertebra, 0.6 mm long, R45572 (Fig. 1F); two fifth or sixth abdominal vertebrae,
1.3 mm long, R45571 (Fig. 1G, H, I), R51293.
Family Gasterosteidae
Gasterosteus aculeatus Linnaeus, threespine stickleback (51 bones): 14 right
pelvic spines, some fragmentary, up to 4.1 mm long, R34195, R34817-19, R34822-
23, R45553-54 (Fig. 1M), R45566-67, R47902, R47904, R51291-92; 25 left
pelvic spines, some fragmentary, up to 4.8 mm long, R20861 (Fig. 10), R34807—
12, R34814-16, R34820, R36592-94, R39860-61, R46213, R46606, R47039,
R47903, R47901, R47903, R47907, R50325, R51290; six first or second dorsal
spines, 2.2—-3.6 mm high, R45552, R34813 (Fig. 1N), R21192, R45568, R45130,
R45157; four abdominal vertebrae, 0.9—1.4 mm long, R18732 (Fig. 1L), R45555,
R45565, R45569: one left pelvic girdle, R51294 (Fig. 1J); one left sphenotic,
R51294 (Fig. 1K).
Recent Comparative Material
Collections of Recent taxa are from the Natural History Museum of Los Angeles
County (Section of Fishes) (LACM) and the University of Michigan Museum of
Zoology (UMMZ); as dry skeletons (D), cleared and stained specimens (CS), bony
parts (W) dissected from, or radiographs (R) of whole preserved specimens. Recent
species were chosen from native southwestern U.S. forms (Culver and Hubbs
1917; Follett 1961; Moyle 1976), and since the fossils were indistinguishable from
some of these, a wider range of extralimital forms was not examined. Number of
specimens and size range in standard length (SL) in parentheses are for specimens
actually examined. Fish specimens were measured with dial calipers and bone
measurements were from an ocular micrometer (read to the nearest 0.1—0.04 mm,
respectively). Pelvic spine length is the maximum length, and height was the
greatest vertical distance between the proximal ends of the dorsal and ventral
flanges and perpendicular to the long axis. All measurements and serration counts
were made on pelvic spines dissected from the fish and with the skin and mus-
culature removed. Dentary length was taken from the symphysis to the anterior
origin of the ascending process. Unless otherwise noted collections are from Cal-
ifornia, Los Angeles County, and catalog numbers are LACM.
Family Salmonidae
Salmo gairdneri. 35861-1 to 4(101-155), E Fk San Gabriel R, 16 March 1973
(D); 42371-1 to 11(52—121), trib N Fk San Gabriel R, 13 October 1972 (R, CS);
LA BREA FOSSIL FISH 95
Fig. 1. Bones of: Salmo gairdneri. A. Left ceratohyal. B. Abdominal vertebra. C. Caudal vertebrae.
Gila orcutti. D. Lateral view of left dentary, anterior to left. E. Dorsal view of left dentary, anterior
to right. F. Cervical vertebrae, lateral view (left), anterior to left, dorsal view (right) anterior is up. G.
Abdominal vertebra, lateral view (left), anterior to left. H. Abdominal vertebra, dorsal view, anterior
to left. I. Abdominal vertebra, ventral view, anterior to left. Gasterosteus aculeatus. J. Lateral view
of left pectoral girdle, anterior to left. K. Dorsal view of left sphenotic, anterior to left. L. Abdominal
vertebrae. M. Right pectoral spine, anterior view. N. Dorsal spine, anterior (left) and posterior (right)
views. O. Left pectoral spine, anterior view. Scale equals 1 mm for all except L, for which it equals
0.5 mm.
35409-1, 2(123-126), trib W Fk San Gabriel R, 26 June 1975 (R); 31858-5, 4(1 19—
133), Malibu Cr, 7 March 1971 (R); 38578-1, 2(100-125), Malibu Cr, 18 February
1973 (R).
Salmo clarki Richardson, cutthroat trout. 35803-1, 2(57-64), CA, Mono Co,
96 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
N Fk Cottonwood Dr, 24 July 1974 (CS); 44404-1 to 7(90-148), OR, Linn Co,
Minto Cr, 23 August 1976 (W); UMMZ 179560-S, 1, 4, 5 (of 8) (133-175), NM,
Taos Co, La Junta Cr, 11 July 1961 (D).
Salmo aguabonita Jordan, golden trout. 35854-1(132), CA, Tulare Co, E Fk
Kaweah R, July 1973 (D); 35857-2, 3, 5(146-172), CA, Inyo Co, Hidden L, 2
September 1973 (D).
Family Cyprinidae
Gila orcutti. 35856-1 to 14(55-114), Malibu Cr, 18 November 1973 (D); 42357-
3, 25(36-89), upper Santa Clara R, 6 April 1979 (W).
Gila bicolor (Girard), tui chub. UMMZ 188955-S, 6(73-121), NV, Churchill
Co, Little Soda L, 9 mi NW Fallon, 14 August 1979 (D); 35859-1(190), 42376-
2(122), CA, Mono Co, Owens R, summer, 1970 (D); 33829-3, 4, 11 to 17(182-
235), CA, Mono Co, Crowley L, 17 May 1973 (D); UMMZ 177085-S, 8(98-200),
CA, San Bernardino Co, Soda Lake Spring at Zzyzx Ranch, 1 July 1959 (D).
Hesperoleucas symmetricus (Baird and Girard), California roach. 35429-1, 5(35-
50), CA, Santa Barbara Co, trib Sisquoc R, 16 July 1975 (W).
Rhinichthys osculus (Girard), speckled dace. 35853-1 to 6(56—-63), W Fk San
Gabriel R, 15 April 1973 (D).
Family Gasterosteidae
Gasterosteus aculeatus williamsoni (Girard). UMMZ 134677, 10(35—48), San
Gabriel R, 27 August 1941 (W); 32591-1, 8(34-39), San Gabriel R, Whittier
Narrows, 9 May 1944 (W); 43742-1, 7(43-49), and LACM JNB 10(W)74, 10(47-
56), Santa Clara R, upper Soledad Canyon, 13 April 1974 (W).
Gasterosteus aculeatus microcephalus Girard. 34070-2, 10(43—50), CA, Ventura
Co, mouth Santa Clara R, 27 July 1974 (W); 43743-1, 10(39-42), CA, Ventura
Co, Sespe Cr, 1.0 mi W Fillmore, 23 April 1981 (W).
Gasterosteus aculeatus aculeatus. 42657-2, 9(41-63), CA, Santa Cruz Co, Struve
Slough, | km N mouth Pajaro R, 9 June 1981 (W),.
Results
The fossil bones are indistinguishable from the bones of three species of fresh-
water fishes extant in Los Angeles Basin streams into historical time, namely
rainbow trout, arroyo chub, and threespined stickleback.
The rainbow trout vertebrae are very similar in both the migratory (steelhead)
and resident fish from the area today. Vertebrae of golden and cutthroat trouts
are typically longer in relation to their diameter than those of rainbow, but this
is difficult to interpret without knowing where in the vertebral column the vertebra
comes from. More diagnostic for the golden and cutthroat trouts are the ridges
interspersed between the numerous excavations on the bottom, top, and sides of
the vertebrae. In addition, these ridges are slightly to greatly longitudinal in ori-
entation. The surfaces of rainbow trout vertebrae are flat, or nearly so; the ex-
cavations appearing as holes punched in a flat surface (Fig. 1B, C). The fossil
trout ceratohyal represents a fish 61 mm SL (ceratohyal length = .057 SL + .708,
N = 12(48-155 mm SL), r = .990). Since the vertebrae are disarticulated, the
number of trout represented is not known. The largest vertebrae represent fish
about 150 mm SL.
LA BREA FOSSIL FISH 97
The three bones of arroyo chub (excluding R51293, which is fragmentary) could
also be from one fish 62 mm SL (dentary length = —1.672 + .0639 SL, N =
25(35-89 mm SL), r = .943). The anteriormost vertebra (Fig. 1 F) is not as long as
in G. bicolor, Rhinichthys osculus, or Hesperoleucas symmetricus. The prominent
circular pit on the ventral edge of this vertebra in Gila orcutti is absent or only
incipiently developed in the other three species. Also in Rhinichthys the anterior
vertebrae are wider transversely than deep, rather than circular as in the remaining
species. The anterior ramus of the dentary is moderately broader and flatter
laterally in G. orcutti than in G. bicolor and extremely more so than in Hesper-
oleucas and Rhinichthys. The abdominal vertebra is distinctive from Rhinichthys,
but very similar to G. bicolor as well as G. orcutti.
Three morphologically defined subspecies of stickleback in California are de-
fined by a progressive reduction of bony armor in more southerly populations,
particularly the number of lateral plates (Miller and Hubbs 1969; Bell 1976). The
fossil pelvic and dorsal spines of stickleback from La Brea have the relatively
reduced features of unarmored threespine stickleback (Gross 1978). The complete
pelvic spines have fewer poorly developed serrations along the ventral edge, about
the same as fish (G. a. williamsoni) from the Los Angeles Basin. However, G. a.
williamsoni from the upper Santa Clara River have more, as do G. a. microcephalus
from farther downstream (Table 1). The fossil dorsal spines lack serrations (Fig.
1C), again a condition of Los Angeles Basin G. a. williamsoni only. Only an
occasional adult male of Los Angeles Basin G. a. williamsoni will have up to three
serrations basally on the dorsal spines. Adult fish of both sexes from the upper
and lower Santa Clara River consistently have such serrations. The serrations are
even more numerous and better developed in G. a. aculeatus. The adult size of
the fossil fish is not known. The lengths of the fossil spines do not exceed lengths
of those of G. a. williamsoni and G. a. microcephalus. If they had the same
proportions as recent stickleback from the Los Angeles Basin, they were 35 to 40
mm SL (standard length = 1.63 spine length + 32.2, N = 15, r = .16). The low
r value demonstrates considerable variability in spine length. The evidence in-
dicates the fossil stickleback was as unarmored as the fish present into historical
times in the Los Angeles Basin. The stickleback fossils represent at least 23 separate
fish (the number of left pelvic spines).
The fish fossils come from three areas in Pit 91: 1) stream drift from the southeast
corner, also containing mollusks and abundant macroscopic plant material, 2) a
lens-shaped deposit in the northeastern corner with fragmentary remains of ver-
tebrates equal to or smaller than juvenile bison, and 3) the central-western side
on the western edge of the “central bone mass” (George Jefferson, pers. comm.;
Shaw and Quinn 1986). The lens-shaped deposit shows evidence of subaerial
exposure. The stream drift contained all of the bones of arroyo chub, and all but
two bones of the rainbow trout; the latter came from the lens-shaped deposit.
Threespine stickleback came from all three areas as follows: 1) 13 bones, 2) 31
bones, and 3) 7 bones.
Discussion
Deposits at Pit 91 are a complex of stream and overbank deposits (Maloney
and Akersten 1976). Bones occur in poorly sorted sandy and gravelly portions of
a channel up to 8 m wide that was later invaded by liquid tar. Mammalian long
SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
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LA BREA FOSSIL FISH 99
bones, predominantly from carnivores and scavengers, are generally aligned in
the direction of the channel and show little wear or abrasion. Silt and blue-green
clays lateral to the channel contain most of the plant fossils, but fewer vertebrate
fossils. Plant fossils are from several distinct plant communities including closed
cone pine forest, relict coast redwood, inland dry foothills, and nearby pond or
marsh. Two primary agencies of deposition are believed responsible for the present
geological facies: entrapment of animals in soft asphalt pools during warm weather
and subsequent asphalt invasion of stream collected bone concentrations (Wood-
ard and Marcus 1973; Maloney et al. 1974; Marcus and Berger 1984; Harris and
Jefferson 1985).
Ecological indications from the plant fossils are that the Pit 91 deposits were
accumulated at a time when winters were wetter and summers dry but cooler
than today (Marcus and Berger 1984). Radiocarbon dates for the sites where the
fish fossils were found range from 25,000 + 1,000 to 33,000 + 1,750 yr BP
(Marcus and Berger 1984), and at that time the sea level was somewhat higher,
possibly to within a few kilometers of La Brea (Nardin et al. 1981; Nardin 1983).
Late Pleistocene climates varied widely, but coastal areas (including La Brea)
apparently fluctuated much less because the marine influence maintained Medi-
terranean climatic conditions (Johnson 1977; Miller 1971; Harris 1985).
Seven species of fishes inhabited the inland freshwaters of southern California
into historical time, namely Lampetra tridentata (Pacific lamprey), Lampetra cf.
pacifica (Pacific brook lamprey), Salmo gairdneri, Gila orcutti, Rhinichthys osculus,
Catostomus santaanae, and Gasterosteus aculeatus (Culver and Hubbs 1917; Fol-
lett 1961; Moyle 1976). All seven species occurred in the Los Angeles River,
which flowed westward three to four km south of La Brea until 1851 when its
course was fixed by man to the present southerly route to San Pedro (Shepard
and Wanless 1971). Lampreys lack bone and would not fossilize, and Rhinichthys
and Catostomus have not been found. These last two species exist today in moun-
tainous tributaries of the Los Angeles River in the San Gabriel Mountains, farther
upstream than Gila or Gasterosteus. The lack of Rhinichthys and Catostomus
indicate (tentatively) no long-distance transport of fish to the site. In addition,
although the individual fossils are disarticulated and sometimes broken, they show
little or no abrasion, presumably reflecting local deposition of an in situ fauna.
As with other ectothermic organisms, amphibians and reptiles (Gehlbach 1965)
and insects (Miller 1982), no evidence exists for prehistoric extinctions among
the freshwater fishes at La Brea in the last 32,000 years. However, in the last 45
years the Pacific brook lamprey disappeared (Hubbs 1967), and the unarmored
threespine stickleback became an endangered species (Moyle 1976).
Considerable extinction occurred among the larger birds and mammals. Little
or none is documented for the small birds and mammals, 1.e., passerines, rodents
(Stock 1956; Harris and Jefferson 1985). However, these small endotherms, par-
ticularly passerine birds are difficult to identify and future work may disclose
extinct forms. Otherwise, whatever led to the megafaunal extinctions did not lead
to elimination of small aquatic vertebrates nor of endothermic ones.
The presence of fossils of these three fish species is additional evidence arguing
for permanent, small riparian stream conditions in the area of the La Brea deposits.
All could survive only under the following conditions: 1) water temperature about
<22°C, 2) dissolved oxygen = 8%o, due to large deep pools, adequate ground water
100 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
inflow, and/or shaded conditions, and 3) available hiding places to provide pro-
tection from predators (Moyle 1976). The three fossil taxa present an ecologically
graded series with Sa/mo gairdneri requiring the coldest and most oxygenated
water (Moyle 1976). Gila orcutti requires the warmest and least oxygenated water
(Castleberry and Cech 1986), and Gasterosteus aculeatus has intermediate re-
quirements (Feldmuth and Baskin 1976). The abundance of fossil stickleback and
trout suggest an environment on the cooler side of this range that could be at-
tributed to colder climate than today or local spring-fed conditions. However, by
themselves, the fish remains are also consistent with the fish fauna present into
historical times.
A large literature cited exists on the selection by piscine predators, particularly
trout, for fully, or at least well, armored stickleback (Wootton 1976; Gross 1978;
Bell 1984). Native trout do not occur with the only remaining population of G.
a. willlamsoni in the upper Santa Clara River (Bell 1978). However, unarmored
stickleback were known from the Los Angeles and San Gabriel River systems
which also harbored runs of steelhead trout Sa/mo gairdneri (Hubbs 1946). If
poorly armored stickleback necessitates lack of piscine predators (or at least lack
of trout), perhaps their interaction was limited to a few weeks during midwinter
high flows when adult trout migrated to, and young smolts descended from,
headwater tributaries lacking stickleback. The present evidence indicates all three
species lived in some proximity on the Los Angeles Basin. They could have
occupied separate habitats but still had their remains washed together in the stream
deposits of Pit 91.
The fossils come from three separate places within Pit 91 and were separated
over as much as 10,750 years (including error). Thus dry-warm periods as well
as wet-cool ones could be included. The fact that only small taxa or small spec-
imens of larger ones were taken indicates small, local tributaries rather than a
main drainage.
This is the first fossil record for Gila orcutti, but Salmo gairdneri and, partic-
ularly Gasterosteus aculeatus have extensive fossil records in the western United
States (G. R. Smith 1981; M. L. Smith 1981; Cavender and Miller 1982; Bell
1973; Bell et al. 1985). Recently fossil stickleback have been discovered in Pleis-
tocene beds of Lake Manix in the Mojave drainage (Roeder 1985). Bell (1982)
discovered recent stickleback living in Holcomb Creek, a tributary to the Mojave
River, but subsequent electrophoretic analysis suggests they were introduced with
trout from the Santa Clara River (D. Buth, pers. comm.). These fossils will be of
great interest, since this gap separates distinct evolutionary lines in each taxon
today.
Acknowledgments
The excavation of the site was supported by NSF Grant GB-24819. William
A. Akersten, George L. Jefferson, Shelly Cox, and Christopher Shaw have regularly
provided an access to specimens and information about the “dig” at Rancho La
Brea. R. R. Miller and R. M. Bailey (UMMZ) permitted me to examine and
borrow skeletal material under their care. Helpful comments on the manuscript
have been provided by W. A. Akersten, J. Harris, G. Jefferson, D. Buth, L.
Grande, and M. Bell. James Diana, James Long (trout), and Jonathan Baskin
LA BREA FOSSIL FISH 101
(stickleback) provided recent comparative material, and Mario Ruiz performed
some of the statistical computations.
Literature Cited
Akersten, W. A. 1980. Fossils in asphalt. [Letter to] Science, 208:552.
, C. A. Shaw, and G. T. Jefferson. 1983. Rancho La Brea: Status and future. Paleobiology,
9(3):211-217.
Bell, M. A. 1973. Pleistocene threespine sticklebacks, Gasterosteus aculeatus, (Pisces) from southern
California. J. Paleo., 47(3):479-483.
. 1976. Evolution of phenotypic diversity in Gasterosteus aculeatus superspecies on the Pacific
coast of North America. Syst. Zool., 25(2):211-227.
——. 1982. Melanism in a high elevation population of Gasterosteus aculeatus. Copeia, 1982(4):
829-835.
Bell, M. 1978. Fishes of the Santa Clara River system, southern California. Contrib. Sci., Nat. Hist.
Mus., Los Angeles Co. No. 295, 20 pp.
1984. Evolutionary genetics and phenetics: the threespined stickleback, Gasterosteus acu-
leatus, and related species. Pp. 431-528 in Evolutionary Genetics of Fishes (Bruce Turner, ed.),
Plenum Press, New York.
———,,J. V. Baumgartner, and E.C. Olson. 1985. Patterns of temporal change in single morphological
characters of a Miocene stickleback fish. Paleobiology, 11(3):258-271.
Castleberry, D. T., and J. J. Cech. 1986. Physiological responses of a native and introduced desert
fish to environmental stressors. Ecology, 67(4):912-918.
Cavender, T. M., and R. R. Miller. 1982. Salmo australis, a new species of fossil salmonid from
southwestern Mexico. Contributions from the Museum of Paleontology, The University of
Michigan, 26(1):17 pp.
Culver, G. B., and C. L. Hubbs. 1917. The fishes of the Santa Ana system streams in southern
California. Lorquina, 1(2):82-83.
Feldmuth, C. R., and J. N. Baskin. 1976. Thermal and respiratory studies with reference to tem-
perature and oxygen tolerance for the unarmored stickleback Gasterosteus aculeatus williamsoni
Hubbs. (sic) Bull. S. Calif. Acad. Sci., 75(2):127-131.
Follett, W. I. 1961. The freshwater fishes—their origins and affinities. Pp. 212-232 in Symposium:
The biogeography of Baja California and adjacent seas. Syst. Zool., 9(3/4).
Gehlbach, F.R. 1965. Amphibians and reptiles from the Pliocene and Pleistocene of North America:
A chronological summary and selected bibliography. Texas J. Sci. 27(1):56-70.
Gross, H. P. 1978. Natural selection by predators on the defensive apparatus of the three-spined
stickleback, Gasterosteus aculeatus L. Canad. J. Zool., 56(2):398—413.
Harris, A. H. 1985. Late Pleistocene vertebrate paleoecology of the West. University of Texas Press,
Austin, vii + 293 pp.
Harris, J. M., and G. L. Jefferson (ed.). 1985. Rancho La Brea: Treasures of the tar pits. Nat. Hist.
Mus. Los Angeles Co., Sci. Ser. 31, vil + 87 pp.
Hubbs, C. L. 1946. Wandering of pink salmon and other salmonid fishes into southern California.
Calif. Fish and Game, 32(2):81-86.
—. 1967. Occurrence of the Pacific lamprey, Entosphenus tridentatus, off Baja California and
in streams of southern California, with remarks on its nomenclature. Trans. San Diego Nat.
Hist. Soc., 14(21):301-312.
Johnson, D. L. 1977. The Late Quaternary climate of coastal California: Evidence for an Ice Age
refugium. Quat. Res., 8(2):154-179.
Maloney, N. J.,and W. A. Akersten. 1976. Formation of calcareous sandstone at asphalt-groundwater
contacts in fluvial sediments, Rancho La Brea, California. Geol. Soc. Amer., Cordilleran Section,
Abstracts of Programs for 1976:393.
—., J. K. Warter, and W. A. Akersten. 1974. Probable origin of the fossil deposits in Pit 91,
Rancho La Brea Tar Pits, California. Geol. Soc. Amer., Cordilleran Section, Abstracts of
Programs for 1974:212.
Marcus, L. F., and R. Berger. 1984. The significance of radiocarbon dates for Rancho La Brea. Pp.
159-183 in Quaternary extinctions. A prehistoric revolution. (Paul S. Martin and Richard G.
Klein, eds.), University of Arizona Press, Tucson.
102 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Miller, G. H. 1972. Some new and improved methods for recovering and preparing fossils as
developed on the Rancho La Brea project. Curator, 14(4):293-307.
Miller, R. R. 1965. Quaternary freshwater fishes of North America. Pp. 569-581 in The quaternary
of the United States. (E. H. Wright and D. G. Frey, eds.), Princeton University Press, Princeton.
, and C. L. Hubbs. 1969. Systematics of Gasterosteus aculeatus, with particular reference to
intergradation and introgression along the Pacific Coast of North America: A commentary on
a recent contribution. Copeia, 1969(1):52-69.
Miller, S.E. 1982. Quaternary insects of the California asphalt deposits. Proc. Third North American
Paleo. Con., 2:377—380.
Miller, W. E. 1971. Pleistocene vertebrates of the Los Angeles Basin and vicinity (exclusive of
Rancho La Brea). Bull. Los Angeles Co. Mus. Nat. Hist. No. 10, ii + 124 pp.
Moyle, P. B. 1976. Inland fishes of California. University of California Press, Berkeley, viii + 405 pp.
pp.
Nardin, T. R. 1983. Late Quaternary depositional systems and sea level change—Santa Monica and
San Pedro Basins, California Continental Borderland. Amer. Assoc. Petroleum Geol., Bull.,
67(7):1104-1124.
—., H. Osborne, D. J. Bottjer, and R. C. Scheidemann, Jr. 1981. Holocene sea-level curves for
Santa Monica shelf, California Continental Borderland. Science, 213(4505):33 1-333.
Roeder, M. A. 1985. Late Wisconsin records of Gasterosteus aculeatus (threespine stickleback) and
Gila bicolor mohavensis (Mohave tui chub) from unnamed Mojave River sediment near Daggett,
San Bernardino County, California. Pp. 171-174 in Geological investigations along Interstate
15, Cajon Pass to Manix Lake. (Robert E. Reynolds, compiler.), San Bernardino County Mu-
seum, Riverside, California.
Savage, D. 1951. Late Cenozoic vertebrates of the San Francisco Bay region. Univ. Calif. Publ.,
Dept. Geol. Sci., Bull., 28:215-314.
Shaw, C. A. 1982. Techniques used in excavation, preparation, and curvation of fossils from Rancho
La Brea. Curator, 25(1):63-77.
, and J. Quinn. 1986. Rancho La Brea: A look at coastal southern California’s past. Calif.
Geol., 39(6):123-133.
Shepard, F. P., and H. R. Wanless. 1971. Our changing coastlines. McGraw Hill, New York. 579
pp.
Smith, G. R. 1981. Late Cenozoic freshwater fishes of North America. Ann. Rev. Ecol. Syst., 12:
163-193.
Smith, M.L. 1981. 2. Late Cenozoic fishes in the warm deserts of North America: A reinterpretation
of desert adaptations. Pp. 11-38 im Fishes in North American Deserts. (Robert J. Naiman and
David L. Soltz, eds.), John Wiley and Son, New York.
Stock, C. 1956. Rancho La Brea. A record of Pleistocene life in California. Los Angeles Co. Mus.
Nat. Hist. Sci. Ser. No. 20 (Paleontology No. 11), 81 pp.
Woodard, G. D. and L. F. Marcus. 1973. Rancho La Brea fossil deposits: A re-evaluation from
stratigraphic and geological evidence. J. Paleo., 47(1):54:69.
Wootton, R. J. 1976. The biology of the sticklebacks. Academic Press, New York, x + 387 pp.
Accepted for publication 28 November 1988.
Bull. Southern California Acad. Sci.
88(3), 1989, pp. 103-116
© Southern California Academy of Sciences, 1989
Notes on Marine Algae of San Diego County
Including Merger of Murrayellopsis with Veleroa
Joan G. Stewart
A-002, Scripps Institution of Oceanography, University of California,
La Jolla, California 92093
Abstract.— The history of marine algal studies in San Diego County is reviewed.
Ten taxa of Rhodophyta have ranges that extend into this southernmost part of
California on the basis of recent collections. The distributions of these taxa were
previously reported to terminate to the north either on the offshore California
Channel Islands or on the mainland. San Diego records are summarized for four
species that have been added to the California flora since Abbott and Hollenberg’s
Marine Algae of California was published (1976). Three species earlier considered
rare in southern California have been found to be extremely abundant and wide-
spread in algal turf on rocky beaches of San Diego County. Recent subtidal col-
lections of an undescribed Phycodrys are treated as a major range extension of P.
cerratae, described from central Peru. Abundant material from San Diego County
sites supports the merger of Murrayellopsis and Veleroa (Rhodophyta) both at
the generic and species level; for reasons of priority, Murrayellopsis dawsonii
becomes a synonym of Veleroa subulata.
During early days of exploration in California, ship-based studies focussed
primarily on the coast between Monterey and San Francisco. Vancouver’s ex-
pedition with Menzies as a biologist, however, spent nearly 2 weeks in San Diego
late in 1793 before sailing south to El Rosario in Baja California, thence to Hawaii
(Jepson 1929). The few plants preserved from this voyage were described in
scattered reports; the only alga that might have been collected near San Diego is
the species now known as Egregia menziesii. Work of early collectors of marine
algae throughout California has been described by Papenfuss (1976).
In 1885, Daniel Cleveland, a 60 year resident of San Diego, listed the algal
species he had collected ‘at San Diego” in Orcutt’s checklist of the flowering
plants of southern and Baja California (Cleveland 1885). This was the first pub-
lished information for marine algae on the coast of San Diego County. Both he
and Edward Palmer, another local botanist, sent specimens to Farlow at Harvard
to be identified and, if new to science, to be named. These two early phycologists
were recognized by the names of several species that were first collected locally
(Sargassum palmeri; Ozophora, Platysiphonia, and Pterosiphonia clevelandii).
During the years just before and after 1900, Mary Snyder collected extensively
from San Diego beaches and sent material to herbaria throughout the United
States. Snyder’s specimens were cited by other workers but her collections never
were assembled as the basis for local floristic analyses.
Somewhat later Setchell and Gardner, working in Berkeley, brought together
all available information about Chlorophyta and Phaeophyta along the entire
103
104 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Pacific coast of North America (Setchell and Gardner 1920, 1925). These two
volumes include important new information about algae of southern California
and Baja California, but because the introductory section describing the sources
of the various records was intended to be included in a third volume on the
Rhodophyta that was never prepared, it is often unclear which algae represent
new records for particular areas. The detailed descriptions of habitats, morpho-
logical variation, and local distribution patterns for the green and brown algae
remain useful for San Diego County despite numerous nomenclatural changes
that have taken place in the intervening years.
From Snyder’s time until the arrival of E. Yale Dawson in La Jolla in 1942,
San Diego lacked resident phycologists. Dawson’s three years of intensive work
in San Diego County led to the publication of an “‘annotated list’’ of marine algae
(Dawson 1945) by the San Diego Natural History Museum. This combined earlier
records for the area with information based on his own collections.
With the use of SCUBA in nearshore biological studies after 1956, diving
botanists could directly observe attached organisms and it was no longer necessary
to depend on beach drift to describe subtidal algal populations. At Scripps Insti-
tution of Oceanography, the first diving studies of submerged algal assemblages
focussed on Macrocystis beds, but since the early 1960’s subtidal algae have been
collected systematically from sites along the coast of San Diego County. New
records and detailed information based on these local studies were not available
at the time the first comprehensive attempt at describing the California algal flora
was prepared by Abbott and Hollenberg (1976). To incorporate records of subtidal
collections, data from intertidal studies, and previously published information on
algal distributions into a single document, a checklist of benthic marine plants of
San Diego County has been prepared for the use of marine biologists working in
this part of California (Stewart 1989).
Approximately 362 species of marine algae are now known from the various
habitats in San Diego County. Collections from depths beneath about 13 m, or
beneath the depth of warmer summer water temperatures, disclose an algal flora
rather different from the intertidal flora. Seasonal distributional patterns occur in
subtidal as well as in intertidal habitats. The list (Stewart 1989) includes taxa that
can be assigned to several different biogeographical categories: 1) intertidal species
typical of intertidal floras both north and south of San Diego; 2) subtidal taxa
that are widespread in deep-water sites along the Pacific Coast of North America;
3) intertidal species that occur intertidally or in shallow water in warmer regions
of Pacific Baja California and in the Gulf of California; 4) deep-water taxa that
grow intertidally or in very shallow water in central and northern California.
A group of species found in central and northern California and in northwestern
Baja California is conspicuously absent from the coast of San Diego County. At
least 15 of these taxa are large and could not be overlooked if they were present.
Examples include such easily recognized species as Leathesia nana, Laminaria
setchellii, large Porphyra thalli, and Mastocarpus (Gigartina) papillatus. These
disjunct distributions are usually explained in terms of water temperatures. San
Diego County beaches lie in the southern part of the southern California bight, a
broad embayment where water circulation patterns include wide and changeable
eddies as well as north-south currents. Wind and wave directions along the coast
differ between seasons and affect upwelling of cold deeper water. The particular
MURRAYELLOPSIS, VELEROA; SAN DIEGO MARINE ALGAE
Table 1.
Species
Antithamnion
hubbsii
Callithamnion
catalinense
Callophyllis
thompsonii
Chondria arcu-
ata
Colacodasya
californica
Gigartina tepida
Heterosiphonia
japonica
Pikea robusta
Previous
distribution
in California
Santa Catalina Is-
land
California Chan-
nel Islands
Monterey
to Orange County
to Orange County
to Orange County
to Orange County
to Santa Barbara
County
Habitat in San
Diego County
Subtidal, on bryozoan
Subtidal; epiphytic, in
garibaldi nests, on
sponges, rocks,
shells
drift, Imperial Beach
intertidal algal turf
on intertidal Heterosi-
phonia erecta
Mission Bay Channel;
10-13 m off Pt.
Loma; intertidal, La
Jolla
Subtidal, 10-23 m, on
various substrates
Drift, on Imperial
Beach
105
New Southern Range Extensions: taxa treated in MAC but not previously recorded from
San Diego county.
New records
27 m, La Jolla, JS 1991
17-37 m, numerous collections, all
reproductive phases, throughout
year. Specimens vary and do
not consistently conform to ear-
lier descriptions; some resemble
forms of C. biseriatum, a widely
distributed subtidal species
Several similar thalli; one identi-
fied by I. A. Abbott, JS 1141
abundant, La Jolla (see Abun-
dance in text)
JS 1453, 2853, 3170 (all in No-
vember)
JS 1715 (cystocarpic), det. G. I.
Hollenberg; 8 additional collec-
tions
JS 2432, 2438 cystocarpic; 13 ad-
ditional collections
Dawson (1945b) recorded speci-
mens (as P. pinnata) from the
beach near Coronado, near the
area of our drift collections; JS
1153, 3225
Polysiphonia to Santa Monica, Intertidal, Bird Rock, JS 2294
brodiaei Los Angeles Co. La Jolla
Porphyrella cal- Laguna Beach, Or- On mussels and bar- abundant, high intertidal rocks,
ifornica ange County nacles, La Jolla early spring
localities on the Pacific coast of Baja California where populations of “‘northern”’
algae occur, characteristically are sites strongly influenced by persistent patterns
of upwelling. Seasonal sand deposition on rock surfaces, and the nature and
inclination of rocky substrates, may also be important to help explain the marked
contrasts between the algal vegetation of San Diego County and that to the north
and south.
Unless otherwise cited, voucher specimens are filed with JS collections, pres-
ently on loan to SIO herbarium. Subtidal data are based primarily on collections
made by J. R. Stewart. Individual specimens are cited in the relevant sections.
Southern range extensions. —Ten taxa of Rhodophyta with ranges that extend
into San Diego County on the basis of recent collections are shown in Table 1.
The distributions of these taxa previously were reported (Abbott and Hollenberg
1976) to terminate to the north either on the offshore California Channel Islands
or on the mainland.
106 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Fig. 1. Berkeleya hyalina (Round & Brooks) Cox. Whole plant from San Diego Co.
Species New to California.—San Diego records for species that have been added
to the California flora since Abbott and Hollenberg’s Marine Algae of California
was published (1976) are summarized as follows.
Naviculaceae, Bacillariophyta
Berkeleya hyalina (Round and Brooks) Cox, a colonial benthic diatom, was
identified (Chastain and Stewart 1985) on the basis of culture and field studies
and microscopic examination of cleaned frustules. This species, as well as the
better known B. rutilans, secretes and lives in tubes of mucilage that are aggregated
into attached tufted thalli that resemble filamentous ectocarpoid algae. In the field,
B. hyalina has an easily recognizable morphology (Fig. 1); details of cell and tube
morphology are illustrated in the earlier report (Chastain and Stewart 1985). The
species is presently known from Baja California, mainland southern California
and offshore channel islands (K. A. Miller, pers. comm.) and the type locality in
Togo, West Africa. In San Diego County it is often abundant in mid-intertidal
habitats, April-October, on rocks or mollusc shells. Thalli grow to 3 cm high, are
dark olive-brown, and slippery to the touch.
MURRAYELLOPSIS, VELEROA; SAN DIEGO MARINE ALGAE 107
Ulvaceae, Chlorophyta
Chloropelta caespitosa Tanner. The genus and species were described (Tanner
1980) from collections in Los Angeles and Orange counties. Among specimens
collected earlier from La Jolla by Dawson and originally identified as Ulva cali-
fornica, Tanner recognized Chloropelta thalli. In 1976 he collected specimens
from high intertidal rocks at Tourmaline Surfer Park in Pacific Beach, and in the
summer of 1986 he again visited the area and located Chloropelta on two La Jolla
beaches (Tanner, pers. comm.); I have recently collected the species from these
same sites.
Cutleriaceae, Phaeophyta
Cutleria cylindrica Okam. Thalli that were conspicuously different from any
previously collected San Diego species were found in Point Loma intertidal pools
in the winter months 1984-87, with 1984 being a year of unusually warm water
that followed a winter of unusually strong storms. The thalli were absent the
following winters (1987-89). Subsequent study recognized the specimens as the
same alga collected at Santa Catalina Island in 1973 and identified as Cutleria
cylindrica by Hollenberg (1978), a species described from Japan. He discussed
the similarities of this “Cutleria’’ species to Myriogloia and other taxa in the
family Chordariaceae and questioned its affinities with Cutleriaceae. The same
alga was collected by Terry Klinger subtidally from San Clemente Island several
times in 1986-87, and from shallow subtidal rocks off Bird Rock in La Jolla, also
in 1986.
Delesseriaceae, Rhodophyta
Apoglossum gregarium (Daws.) Wynne. The southern California plants de-
scribed by Stewart (1974) as Phrix gregarium are delicate, minute (less than 1 cm
high and 1—2 mm wide), subtidal blades that grow on sponges, bryozoans, in nests
of the garibaldi fish, or with other algae in various subtidal rocky habitats. They
are not rare, but difficult to recognize in the field. Reproductive as well as vegetative
thalli have been collected from depths of 17—28 m at all times of the year in sites
beneath the summer thermocline between La Jolla Bay and the outer kelp beds
off Point Loma. Elsewhere, specimens have been collected by R. Moe (pers.
comm.) subtidally at the Galapagos Islands and off Palos Verdes in Los Angeles
County, and by J. R. Stewart at San Benitos Islands, Mexico.
The rippled, undulate blades are unlike any other small algal blades Iam familiar
with, and in freshly collected algal material are so conspicuous as to make rec-
ognition of the species obvious. I earlier (Stewart 1974) suggested that this strik-
ingly characteristic appearance could be explained as the consequence of frequent
continuing divisions in blade cells while the central and pericentral cells elongate
but do not divide transversely. Growth to either side of the midline therefore
forms an increasingly ruffled blade, with the “ruffles” extending to the midline,
rather than observed only near the margins.
Phycodrys cerratae Daws., Acleto & Foldv.—In 1970 and again in 1981 attached
Phycodrys blades (Fig. 3) were found in La Jolla Canyon offshore from Scripps
Institution of Oceanography, San Diego County (JS 1744, 3551). The site has
been observed frequently in intervening years but additional plants have not been
108 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
found. These collections could not be attributed to either of the two other species,
P. profunda and P. setchellii, previously known from San Diego County. In 1940,
G. J. Hollenberg collected 15 specimens of an unidentified Phycodrys species
(Figs. 4, 5) from drift at Cabrillo Beach near San Pedro, Los Angeles County (GJH
# 3102). These were later given to me by I. A. Abbott for comparison with the
thalli collected in La Jolla Canyon.
The genus Phycodrys world-wide is large and generally conceded to be in need
of monographic treatment. Pending such a study, I have identified the three
southern California collections cited above as P. cerratae. A review of the mor-
phology and ecology of species attributed to the genus in the eastern Pacific
recognized two that were particularly similar to the unidentified specimens. Both,
P. isabellae and P. cerratae, are subtidal and not often collected. The type specimen
of Phycodrys cerratae was collected in drift material near Lima, Peru in 1960 and
described and figured by Dawson et al. (1964). An isotype (Fig. 2) was deposited
at HAFH, now in LAM. P. isabellae was described from a subtidal plant collected
from the San Juan Islands, Washington (Norris and Wynne 1968). Subsequently
other subtidal plants from Oregon and central California were identified as rep-
resenting this latter species.
Compared with P. cerratae and P. isabellae (Table 2), the southern California
specimens resemble P. cerratae in size, appearance of tetrasporangial sori, and in
structure of the marginal bladelets. Primary blades of Hollenberg’s drift specimens
are to 12 cm high, to 3.5 cm wide; secondary blades are to 4.2 cm long and 1.5
cm wide. San Diego blades are to 9 cm (primary), 7 cm (secondary) high and 2
cm wide. Lateral blades are subtended by distinct stipes closely spaced on margins
of primary and secondary blades. No rhizoids or other proliferations for secondary
attachment from blade margins were found.
Phycodrys isabellae has not been recognized in subtidal collections from San
Diego County. The specimens from the Monterey peninsula, described by Abbott
and Hollenberg (1976), are to 7 cm high but mostly smaller and as figured, the
marginal bladelets tend to grow from the blade edge without a distinct stipe.
Tetrasporangial sori in type material (Fig. 23, Norris and Wynne 1968) are con-
spicuous and discrete; the specimen illustrated as Fig. 587 (in Abbott and Hol-
lenberg 1976) appears to show tetrasporangia in diffuse “‘sori”’ filling the space
between lateral nerves. Tetrasporangia are scattered and inconspicuous on blades
collected in the La Jolla Canyon. Abbott and Hollenberg (1976) describe margins
of P. isabellae as undulate, whereas blades collected in San Diego County are flat.
In support of the assignment of southern California collections to P. cerratae,
a species described and known only from central Peru, there are numerous other
algae with similar distribution patterns. Santelices and Abbott (1978) accepted
bipolar distributions for a number of species in the algal flora of Chile and report
aK
Fig. 2. Phycodrys cerratae Daws., Acleto & Foldv. Isotype, Barranco, Lima, Peru, Jan. 9, 1960,
collected by Emma Cerrate 2997. AHF 75435. Resembles La Jolla canyon collection (Fig. 3) and San
Pedro drift material (Figs. 4, 5) in size, midrib, margins without rhizoids, and numerous lateral
bladelets.
Fig. 3. Phycodrys cerratae. 20 m, La Jolla Canyon, San Diego Co., 4 Sept. 1981. JS 3551.
Figs. 4,5. Phycodrys cerratae. G. J. Hollenberg 3102, “‘cast ashore leeward side of gov. breakwater,
Cabrillo Beach, San Pedro, Calif. 24 July, 1940.” Annotated ““Phycodrys n. sp. I.A.A., 1973.”
MURRAYELLOPSIS, VELEROA; SAN DIEGO MARINE ALGAE 109
TTI TTT
Z
}
OL
|
HCPL HNLP PITT
| i | mln
wale
If! WHA
Bins p ? ps
Der OY Os Ban awCo, Leute
tal
| |
110
SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Table 2. Comparison of published descriptions of two species and undescribed collections.
Blade length
Blade shape
Blade width
Midrib
Lateral veins
Margins
Marginal blade-
lets
Phycodrys isabellae
Norris & Wynne 1968
to 7 cm
oblanceolate
to 2 cm
weakly developed or promi-
nent
“some blades with weakly
developed opposite veins
from midrib”
sometimes denticulate, with
secondary attaching rhi-
zoids
primary blade single or with
few to many bladelets
Southern California
collections, Fig. 3-5
to 12 cm
both
to 3.5 cm
prominent
faint, not distinct
no rhizoids, smooth
to 7 cm in length,
closely spaced
Phycodrys cerratae
Dawson et al. 1964,
and Fig. 2
to 12 cm
lanceolate
to 3 cm
prominent, broad
opposite, distinct (but
see Fig. 2)
(see Fig. 2. No rhizoids
shown)
Margins entire, or with
numerous closely-
spaced pedicillate
bladelets to 20
mm+
Tetrasporangia sori between lateral veins, faint sori between lateral
large and distinct in photo veins
Gametangiathalli described not found not found
(in their Tables 1, 2, and elsewhere in the text) 17 species from Chilean sites that
occur in San Diego County (Stewart 1989). Dawson described the intertidal algal
flora of central Peru as sharing many features in common with southern California
(Dawson et al. 1964). Excluding taxa that are difficult to identify (e.g., crustose
corallines), Dawson et al. (1964) list 29 species from Peru that occur in San Diego
County. Phycodrys cerratae is added to the flora of San Diego County and thus
to the California marine algal flora on the basis of the J. Stewart and G. J.
Hollenberg collections cited above.
Abundance.—Three species described (Abbott and Hollenberg 1976) as occa-
sional, relatively rare, or rare in southern California, have been found to be
extremely abundant and widespread in algal turf on rocky beaches of San Diego
County.
Ceramium flaccidum (Kitz) Ardissone [as C. gracillimum var. byssoideum
(Harv.) Maz. and C. taylorii Daws. in Abbott and Hollenberg 1976]. Thalli cover
coralline anchor taxa with a deep-red layer of filaments in late summer months,
with peak abundance in October at La Jolla sites. The thalli are extremely fine
but easily recognized by the elongated internodes contrasted with narrow nodal
bands and the darker apical regions where central cells have not yet elongated
(Fig. 6). All reproductive phases can be found in San Diego county material.
Womersley (1978) illustrates microscopic detail of nodal cortication and provides
a thorough study of the species, considering it ““‘probably cosmopolitan in cold
temperate to tropical seas.”
Binghamia forkii (Daws.) Silva. Thalli (Fig. 7) are easily found between May
and November on the platform rock beaches between Ocean Beach and Point
Loma where they form a yellowish or greenish epiphytic mat over algal turf. Less
MURRAYELLOPSIS, VELEROA; SAN DIEGO MARINE ALGAE 111
250 um
Fig. 6. Ceramium flaccidum (Kiitz.) Ardissone. Axis with elongated proximal internodes, San
Diego specimen. Long uncorticated internodes below, delicate size distinguish thalli in field.
abundant, it also grows at the same season on beaches north of Mission Bay. All
reproductive phases are easily found in these intertidal habitats.
Chondria arcuata Hollenb. Thalli are dull red when growing in shaded habitats,
but more commonly are pale or almost yellowish in intertidal turf. The species
is illustrated by Fig. 672 in Abbott and Hollenberg (1976). Erect branches are
cylindrical, often somewhat arched or curved as they grow horizontally over other
algae, and secondarily attached at frequent intervals. A tuft of trichoblasts from
the apical pit may be conspicuous in freshly collected or young material but in
older plants or thalli that have been several hours in containers, this feature may
be misleadingly absent. The species is one of the most common and abundant
epiphytes growing on Pterocladia capillacea (Gmel.) Born. and Thur. and Cor-
allina pinnatifolia (Manza) Daws. on La Jolla beaches, less common elsewhere
in the County.
Taxonomic Merger: Veleroa and Murrayellopsis
Type species of these two monotypic genera, V. subulata and M. dawsonii, are
small polysiphonous red algae with four pericentral axial cells and radially ar-
112 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
©:5:em
Fig. 7. Binghamia forkii (Daws.) Silva. Characteristic thalli, with tetrasporangial specimen on left.
Whole epiphytic thallus comprises branches as shown, commonly overgrowing algal turf.
ranged monosiphonous, determinate lateral branchlets. Tetrasporangia occur in
main axes rather than in stichidia. Kylin (1956) placed Veleroa in the Lophothalia
group of Rhodomelaceae, a group characterized by persistent pigmented tricho-
blasts termed monosiphonous branchlets by Dawson (1944, 1963) and ramuli by
Abbott and Hollenberg (1976). Closely allied genera differ in the number of
pericentral cells, in being corticated, or by being bilaterally branched (Kylin 1956).
Murrayellopsis was distinguished (Dawson 1963; Abbott and Hollenberg 1976)
from Veleroa by having paired tetrasporangia in axial segments and mostly branched
““trichoblasts.”’ Post (1962, 1963) described Murrayellopsis dawsonii gen. et spec.
nov. without mention of Vel/eroa which had been described earlier by Dawson
(1944). A subsequent paper (Post 1964) added references to Veleroa in the text
and included this genus in a key where it was distinguished from Murrayellopsis
by the number of tetrasporangia in a segment. She stated her opinion that “in no
case is Murrayellopsis a luxuriant Veleroa.”
Dawson’s treatment (1963) of the two species is based only on the two original
descriptions as no additional collections of either were known at that time. Since
then, thalli distinguishable only by the number of tetrasporangia per segment have
been identified in collections from subtidal habitats along the coast of California.
Material studied:
Veleroa subulata. Holotype LAM 500006, Dawson 281d, dredged in 22 m,
Tepoca Bay, 4 Feb., 1940. LAM 509949, R. Setzer 7721, 3 m, at west end of
breakwater on inside, Los Angeles Harbor, 7 Nov., 1973. LAM 599947, R. Moe
2B, Palos Verdes Point, 7 m, 19 Oct. 1972. LAM 599948, R. Moe 120, Palos
Verdes Point, 7 m, 19 Oct. 1972. K.A. Miller material, collected from San Miguel
Island off the coast of central California, 17 Sept. 1983, 5-10 m depth, 5 slides,
unaccessioned UCB.
MURRAYELLOPSIS, VELEROA; SAN DIEGO MARINE ALGAE 113
Murrayellopsis dawsonii. Holotype LAM 500009, det. Erika Post, from Hyp-
sypops rubicunda (garibaldi fish) nest, 12 m, New Hope Rock off Point Loma,
San Diego Co., 18 April 1961. Paratype (LAM designation) LAM 500007, det.
Erika Post, from vertical surface at 7 m, N.E. side of North Island, Islas Coronados,
2 June 1961. UCB 1462114, col. A. Nonomura, 10 m, sand pool, Mussel Pt.,
Pacific Grove, 8 Aug. 1973. CMMEX (unaccessioned), L. Aguilar 32, 20 m, Isla
Todos Santos, Baja California, Mexico, 19 June 1982.
Veleroa subulata/Murrayellopsis dawsonii. Santa Barbara Co. JS 2215, 10 m,
Naples Reef, May 1972; San Diego Co. JS 832a,b, 30 m, on bowl, La Jolla
Submarine Canyon, 10 Nov. 1967; 906e, 27 m on rock, La Jolla Canyon, 6 Feb.
1968; 1233a-b, 22 m, inshore from Quast Rock, La Jolla, May 1969; 1381, 13
m, New Hope Rock, 5 Sept. 1969; 1809 a-b, 13 m, New Hope Rock, 23 Jan.
1971; 1877, 1878, 1896, 13 m, New Hope Rock, 13 Feb. 1971; 2420, 2434, 2435,
27 m, Quast Rock, La Jolla, 10 April 1973; 2941, 15 m, garibaldi nest, Loma
Sea Cliff, Jan. 1977.
All of the above specimens fit within the original description (Dawson 1944)
of Veleroa subulata with the following amendments.
Genus VELEROA: Irregularly arranged indeterminate branches can replace
pigmented monosiphonous, determinate branchlets; tetrasporangia borne in a
single or double spiral row, one or two per segment in slightly swollen upper
portions of main axes and branches; carpogonial branches developed on next-to-
basal cells of monosiphonous branchlets.
Veleroa subulata: Plants to 20 mm high; pericentral cells of polysiphonous erect
axes and branches to 50-70 um, more often 10-30 um, diameter, as much as 6
diameters long; lateral branchlets to 700 um long, each with up to 16 cells that
are individually mostly less than 60 um long and 10-40 wm diameter at base,
with a basal cell about half the length of succeeding cells; branchlets simple or
with up to 4 second order branchlets on lowermost cells and these infrequently
once-forked; branchlets tapering to a very small sharply-pointed apical cell; tetra-
sporangia borne one or two per segment in up to about 15 successive segments
of main axes and branches, to 60 wm diameter when mature.
Cells of polysiphonous prostrate axes of similar diameter as pericentral cells in
erect axes, but often longer, to 160 um. Polysiphonous, or multiseriate, rhizoids
develop from the prostrate axes, with very long, to 300 wm, mostly narrow cells.
On entangled plants, e.g., growing around bryozoans, several rhizoids grow in
clumps from single segments of an axis and separately or apparently coalesced
attach to substrate in a disc-like pad. In such thalli the distinction between pros-
trate and erect axes is lacking. Monosiphonous branchlets and rhizoids may de-
velop from adjoining segments. The basal cell in each lateral monosiphonous
branchlet is distinctly shorter than, but with the same diameter as, the more distal
cells of the branchlet (Fig. 8). When branchlets detach, the basal cell usually
remains and is seen on older proximal parts of the thalli as a scar cell.
Type: Dawson, 381d, growing on a small hydroid, dredged in 22 m, Tepoca
Bay, Sonora, in the upper Gulf of California, 4 Feb. 1940. Herb. AHF no. 63.
When this type material was examined for the present study, no reproduction
was found, suggesting that the branches photographed as Fig. 2 in Plate 72 of
Dawson (1944) have been lost. The single vial representing the type collection
contains mostly fragmented axes and branchlets. Cell measurements conform to
114 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Figs. 8-11. Veleroa subulata Daws. Thalli collected from nests of the garibaldi on or near New
Hope Rock, Pt. Loma, San Diego Co., the type locality of Murrayellopsis dawsonii Post. Fig. 8. JS
2941. Polysiphonous branch with monosiphonous branchlets. 1 cm scale = 80 um. Fig. 9. JS 1809.
Cystocarps on next-to-basal cell of monosiphonous branchlets. 1 cm scale = 25 um. Fig. 10. JS 2941.
Tetrasporangia in single row, smaller distally. 1 cm scale = 100 um. Fig. 11. JS 2941. Single and
paired tetrasporangia in same row. | cm scale = 100 um.
the above description. In general appearance, the algae described as Veleroa sub-
ulata are indistinguishable from any of the collections referred to Murrayellopsis
dawsonii.
Post (1962, 1963, 1964) stated that pericentral cells in Murrayellopsis dawsonii
were 4—5 in number but all of the material examined in this study consistently
showed four. Several instances of “‘cortication’”? were observed where the peri-
central cells in several adjacent axial segments were apparently replaced by ir-
regularly arranged, irregularly shaped small cells. These portions of the thalli were
mostly basal, lacked branching, and in the habitats where the plants were collected,
were 1n a layer of debris or sand grains close to the substrate.
Polysiphonous branching is infrequent, but in those thalli where the axes branch,
all such branches occur in the upper half or third of the axis, each one replacing
a monosiphonous branchlet insofar as could be seen. A relatively densely branched
axis bears up to 11 branches and each of these may bear 2,3 third-order branches
which are unbranched or once-forked. The distal clustering above the long un-
branched proximal portions of the axes gives a characteristic bushy aspect to
individual axes. Degree of monosiphonous branchlet development parallels the
extent of polysiphonous branching. Many axes are unbranched, with mostly short
unbranched or once-forked branchlets; others with polysiphonous branches also
bear longer, more-branched monosiphonous branchlets. Lower parts of most axes
MURRAYELLOPSIS, VELEROA; SAN DIEGO MARINE ALGAE 115
are often unbranched or bear only short monosiphonous spine-like branchlets.
Branching from the cells of the monosiphonous branchlets is also radial, appar-
ently spiral from sequential “‘corners” of the single cells of the uniseriate filaments.
From the type material for Murrayellopsis that was collected at New Hope
Rock, I segregated 19 tetrasporangial axes from the large amount of vegetative
material. All were densely branched, and contained mostly paired tetrasporangia.
Proximal segments, just below the swollen tetrasporangia-bearing segements, show
very narrow, irregularly aligned and separated pericentral cells, indicating that
tetrasporangia had been shed from these segments. Apically, distal to the swollen
segments, small cells were formed in addition to immature tetrasporangia which
are distinct by differential pigments. In her revision of the privately published
1962 description of Murrayellopsis, Post (1964) changed the number of tetraspo-
rangia per segment from two, to 2, 3, or 4. I carefully squashed several axes to
spread the tetrasporangia and in no case were more than 2 per segment found.
Without this treatment, the size and congestion of the tetrads in the axis did not
allow this determination. The amount of either polysiphonous or monosiphonous
branching is uncorrelated with development of single or double rows of tetraspo-
rangia.
A cystocarpic specimen (Fig. 9) and numerous tetrasporangial thalli have been
collected from nests of the garibaldi on New Hope Rock as well as from other
nearby rocky subtidal habitats (JS specimens). In these thalli many of the tetra-
spores are in paired rows as described for the type material. Some thalli, however,
have only single rows of sporangia (Fig. 10); single tetrasporangia occasionally
are interspersed with pairs in the same axis (Fig. 11). Figures 8—11 illustrate recent
collections from the type locality for Murrayellopsis dawsonii. I suggest that tetra-
sporangia develop in a single row along an axis, in a %4 spiral from successively
arranged pericentral cells. A second row of sporangia can then develop, usually
subsequent to the enlargement of those in the first row. Tetrasporangia in the first
row are mostly retained while the second row develops, resulting in two rows of
large tetrasporangia. The single row, one per segment (as described first for Veleroa
subulata but now found in Murrayellopsis specimens) therefore appears to rep-
resent a developmental stage, rather than to characterize a distinct taxon. For this
reason I propose merging the two taxa both at the generic and species level; for
reasons of priority, Murrayellopsis dawsonii becomes a synonym of Veleroa sub-
ulata, typified by the few scraps now deposited in collections at LAM.
Acknowledgments
I am grateful to Paul Silva for clarifying the correct usage of taxonomic terms,
specifically, the use of ‘“‘merger” for the discussion of Veleroa subulata. I thank
V. Anderson at LAM, P. Silva at UCB, K. Miller at UCB, and R. Aguilar at
CMMEX who have loaned institutional and personal collections of Veleroa and
Murrayellopsis. Figs. 1, 6, and 7 were drawn by Nancy Hurlburt. I am grateful
for I. A. Abbott’s gift of the Hollenberg ““Phycodrys n. sp.”’ collection. Steven
Murray’s review comments greatly improved the presentation of the information.
Literature Cited
Abbott, I. A., and G. J. Hollenberg. 1976. Marine algae of California. Stanford University Press, xii
+ 827 pp.
116 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Chastain, R. A., and J. G. Stewart. 1985. Studies on Berkeleya hyalina (Round & Brooks) Cox, a
marine tube-forming diatom. Phycologia, 24:83-92.
Cleveland, D. 1885. Algae. Species collected at San Diego by Daniel Cleveland. Pp. 12-13 in Flora
of southern and lower California. (C. R. Orcutt, ed.), San Diego.
Dawson, E. Y. 1944. The Marine algae of the Gulf of California. Allan Hancock Pacific Expeditions,
3:189-453.
1945. An annotated list of the marine algae and marine grasses of San Diego County,
California. Occ. Pap. San Diego Natural History Soc. No., 7:1-97 (reprinted with corrections
May 1952).
1963. Marine red algae of Pacific Mexico. Part 7. Ceramiales: Ceramiaceae, Delesseriaceae.
Allan Hancock Pacific Expeditions, 26:1—207.
—., C. Acleto, and N. Foldvik. 1964. The seaweeds of Peru. Beihefte zur Nova Hedwigia, 13:
1-111, 80 pl.
Hollenberg, G. J. 1978. Phycological notes VIII. Two brown algae (Phaeophyta) new to California.
Bull. So. Calif. Acad. Sci., 77:28-35.
Jepson, W. L. 1929. The botanical explorers of California. VI. Madrono, 1:262-270.
Kylin, H. 1956. Die Gattungen der Rhodophyceen. Gleerups, Lund, xv + 673 pp.
Norris, R. E., and M. J. Wynne. 1968. Notes on marine algae of Washington and southern British
Columbia, III. Syesis, 1:133-146.
Papenfuss, G. F. 1976. Landmarks in Pacific North American marine phycology. Pp. 21-45 in
Marine algae of California. (I. A. Abbott and G. J. Hollenberg, eds.), Stanford University Press,
xii + 827 pp.
Post, E. 1962. Murrayellopsis dawsonii gen. et spec. nov. Post aus einem Goldfisch-Nest. 3 pp (no
pagination), 2 figures (privately printed).
1963. Murrayellopsis dawsonii Post gen. et spec. nov. aus einem marinen Goldfischnest.
Naturwissenschaften, 2:49.
—. 1964. Murrayellopsis dawsonii Post gen. et spec. nov. aus einem marinen Goldfisch-Nest.
Hydrobiologia, 23:274—280.
Santelices, B., and I. A. Abbott. 1978. New records of marine algae from Chile and their effect on
phytogeography. Phycologia, 17:213-222.
Setchell, W. A., and N. L. Gardner. 1920. The marine algae of the Pacific Coast of North America.
II. Chlorophyceae. Univ. Calif. Publ. Bot., 8:139-374, 25 pls.
—. 1925. The marine algae of the Pacific Coast of North America. III. Melanophyceae. Univ.
Calif. Publ. Bot., 8:383-739, 73 pls.
Stewart, J.G. 1974. Phrix: a new genus in Delesseriaceae (Rhodophyta). Phycologia, 13:139-147.
. 1989. Benthic marine algae and seagrasses of San Diego County. San Diego Natural History
Museum, Occ. Pap. submitted.
Tanner, C. E. 1980. Chloropelta gen. nov., an ulvaceous green alga with a different type of devel-
opment. J. Phycol., 16:128-137.
Womersley, H. B. S. 1978. Southern Australian species of Ceramium Roth (Rhodophyta). Aust. J.
Mar. Freshwater Res., 29:205—257.
Accepted for publication 25 April 1989.
Bull. Southern California Acad. Sci.
88(3), 1989, pp. 117-122
© Southern California Academy of Sciences, 1989
Research Notes
Late Pleistocene Chipmunk, Tamias (Mammalia: Sciuridae),
from Rancho La Brea, Los Angeles, California
The Rancho La Brea paleontological sites in Hancock Park in the western part
of Los Angeles have produced one of the most impressive and complete records
of late Pleistocene and early Holocene life in the world (Akersten et al. 1983;
Harris and Jefferson 1985; Shaw and Quinn 1986). Early major excavations at
Rancho La Brea between 1905 and 1930 sampled mostly larger organisms. This
was primarily due to collecting methods that tended to overlook smaller fossils.
In 1969, the Natural History Museum of Los Angeles County (LACM) began a
new scientific excavation at Rancho La Brea, using modern collecting techniques,
that has led to the recovery of abundant samples of smaller vertebrate fossils such
as lizards, snakes, and rodents (Shaw 1982; Akersten et al. 1983). An examination
of samples of these smaller fossils has revealed the presence of a chipmunk, a
taxon that was previously unknown from Rancho La Brea.
The collective fossil biota recovered from Rancho La Brea represents a diverse
Late Pleistocene flora and fauna. The majority of plants and animals are still
represented by living forms, but Rancho La Brea is better known for its records
of large, extinct mammals (Stock 1956; Harris and Jefferson 1985).
Hancock Park and vicinity is underlain by approximately 45 m of Late Pleis-
tocene strata that Woodard and Marcus (1973) referred to the Palos Verdes Sand.
Most of the thickness of the Palos Verdes Sand in this area was deposited in a
shallow marine environment, but the upper 9 m to 10 m were deposited on an
alluvial plain that developed between the Santa Monica Mountains and the Pacific
Ocean. Fossil excavations have only penetrated to depths of a little over 8 m, but
drill cores have yielded both vertebrate and marine invertebrate fossils at greater
depths. Although the fine details of depositional history within any single exca-
vation may be complex, radiometric age determinations at varying depths have
shown an approximate age range of between 35,000 and 11,000 years before
present for the fossils from these excavations (Woodard and Marcus 1973; Marcus
and Berger 1984).
The continuing scientific excavation begun by the LACM in 1969 is centered
over one of the 1915 excavations, or “‘Pits,” as they were commonly called. This
site, Pit 91, was abandoned in 1915 before removal of fossils was completed. The
modern excavation, assigned the locality number LACM 6909, is approximately
9 m square, and is quarried in six-inch depth intervals using a horizontal square
yard grid system commonly employed in archeological excavations (Shaw 1982).
As larger fossil specimens are removed, surrounding matrix is retained for pro-
cessing to recover smaller fossil specimens (pollen, seeds, small mollusks, insects
and other arthropods, and small vertebrates). Over 400 tons of fossiliferous matrix
have been recovered to date, but only a small percentage has been fully cleaned
and the fossils removed (Akersten et al. 1983).
A substantial sample of the matrix recovered from a concentration of fossils
encountered early in the new excavation (1969-1970) in the northeast corner of
117
118 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Ze
Pa aA a
Figs. 1-2. Teeth of Tamias cf. T. merriami from Rancho La Brea. Fig. 1, nght M', LACMRLP
52223, occlusal view. Fig. 2, right M,, LACMRLP 52229, occlusal view. Scale line equals 2 mm.
LACM 6909 has been cleaned (field numbers GJM 273, 275, 295, 480, 537, 539,
and 550). This matrix was from grid designations N-3 and M-3 from a depth
between 5 feet and 7.5 feet below datum (see Shaw 1982, for a detailed explanation
of the grid and sub-locality designations). The fossils occur in fine grained silty
asphaltic sandstone with lenses of brown silt, sediments that represent an infilling
of an older stream channel. These stream sediments have yielded a radiometric
date using '*C of 32,600 + 2800 years before present based on one specimen of
a juvenile Smi/odon left femur from this concentration (specimen number R-51285,
from locality GJM 550, grid M-3 at 7-7.5 feet below datum; UCLA 1738-D;
Marcus and Berger 1984).
The small fossils recovered from this northeast deposit contain a diverse as-
semblage of snails, lizards, snakes, and small mammals, including nine isolated
chipmunk teeth that are described below. Dental terminology and measurements
follow Black (1963). Measurements are made on an EPOI Shopscope optical
micrometer. Taxonomy for chipmunks follows Levenson et al. 1985.
Family Sciuridae Gray, 1821
Subfamily Sciurinae Baird, 1857
Tamias cf. T. merriami (Allen, 1889)
Figs. 1 and 2, Table 1
Material. —(Specimen numbers prefixed LACMRLP) Right M! (52223) from
locality GJM 273; left P* (52226), right dP, (52225), left P, (52227), right M,
(52224) from locality GJM 539; right M! (52230), left M? (52231), right P, (52228),
right M, (52229) from locality GJM 550.
Description. —Teeth with tall lophs(ids); upper molars subquadrate in outline,
protoconule and metaconule absent, metaloph only slightly constricted at con-
nection to protocone, anterior cingulum well developed but lacking distinct para-
style, mesostyle absent; P* reduced antero-posteriorly, lacking anterior cingulum;
lower molars narrow (labial-lingual) compared to typical Tamias spp., mesostylids
absent, mesoconid reduced on M,, absent on M,, anterior cingulum well developed
and well separated from protoconid in M,, metalophid (connecting metaconid
RESEARCH NOTES 119
Table 1. Measurements of teeth of Zamias cf. T. merriami from Rancho La Brea. Abbreviations:
A/P = greatest anterior-posterior dimension, TR = greatest labial-lingual dimension. Measurements
in millimeters.
Specimen number
Pt M! M! M? dP, P, P, M, M,
52226 52223 52230 52231 52225 52228 52227 52229 52224
A/P 1.15 1.58 19, ESS 1.10 1.46 1.36 1.47 1.54
TR 1.43 1.75 1.86 1.86 0.90 1.15 1.04 1.46 1.66
and protoconid) incomplete in both M, and M,, particularly in M,; P, relatively
long (compared to M,), cusps subdued, mesostylid absent; dP, a smaller version
of P, but with more reduced ectolophid and complete lingual connection of pos-
terolophid to metaconid.
Comparisons.—The nine isolated fossil teeth were compared to samples of
Tamias merriami, T. sonomae Grinnell, 1915, T. speciosus (Merriam, 1890), and
T. townsendii (Bachman, 1839) from the collections of the Section of Mammalogy,
Natural History Museum of Los Angeles County. These species were selected for
comparison because they are either extant chipmunks which live in habitats
inferred to have been present near Rancho La Brea in the Late Pleistocene (T.
merriami, T. sonomae, and T. townsendii) or are those living closest to Hancock
Park today (7. merriami and T. speciosus).
In overall size, the fossil teeth fall in the upper range of 7. merriami or lower
range of 7. sonomae, and they are clearly larger than those of 7. speciosus. The
reduced P* resembles 7. merriami in contrast to the other species of Tamias which
have a more completely developed P* with a distinct anterior cingulum (=para-
style). In contrast, the relatively long P, more closely resembles 7. sonomae. The
narrowness of the lower molars most closely resembles those of 7. speciosus. The
well developed anterior cingulum and reduced metalophid on the lower molars
resemble the condition in both 7. merriami and T. sonomae.
The sample of fossil teeth is too small to yield information on variability, but
based on the modern species examined, the fossils appear intermediate between
T. merriami and T. sonomae in both size and morphology. If larger samples
become available, it may be possible to demonstrate that this fossil taxon from
Rancho La Brea represents a distinct species near 7. merriami or, alternatively,
it may represent a population of 7. merriami with teeth near the upper size limit
of the living populations. Goodwin and Reynolds (1989) have reported the oc-
currence of a large, undescribed, fossil species of Tamias from Kokoweef Cave
in the eastern Mojave Desert which they compare to large living species of Tamias
including 7. merriami. The fossils from Rancho La Brea may be related to this
large, Kokoweef species, but the sample is too small to draw definite conclusions.
Discussion
There are neither historic records nor previous fossil records (Miller 1971) of
chipmunk in the Los Angeles Basin. Their closest occurrence to Hancock Park is
30 km northeast, across the Los Angeles Basin on the chaparral-covered slopes
of the western San Gabriel Mountains (7. speciosus). The Santa Monica Mountains
form the headwaters of the drainages crossing the Rancho La Brea area both in
120 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
the Late Pleistocene and today (Shaw and Quinn 1986). There are no documented
historic records, and no modern observations of chipmunks in the Santa Monica
Mountains (R. Hannum, personal communication), even though ample suitable
habitat exists to support them.
This new record suggests that chipmunks previously inhabited the Santa Monica
Mountains, the Santa Monica plain, and quite possibly the Los Angeles basin.
Their modern absence from these areas lends further support to the well docu-
mented southward shift of more equable paleoclimates during periods of the
Pleistocene, or, a less likely alternative, a change in the habitat preference of
chipmunks.
It is also noteworthy that the nine 7amias teeth are the only record of chipmunk
derived from more than 75 years of collecting at Rancho La Brea. Considerable
matrix from many horizons within Pit 91 (LACM 6909) has been processed for
small vertebrates, but the chipmunk specimens are restricted to a single deposit
in the northeast corner of the excavation.
The fossils found with the chipmunk teeth do not have a strikingly different
composition from the typical collective assemblage recovered from the Rancho
La Brea deposits, although there are subtle differences. The associated large fossils
are those typical at Rancho La Brea including the carnivores (sabretooth, Smi-
lodon; large lion, Felis atrox; wolf, Canis dirus; and coyote, C. latrans) and large
herbivores (large ground sloth, Glossotherium; horse, Equus occidentalis; Bison
sp.; pronghorn, Capromeryx). Large fossils from the northeast corner deposit are
not as numerous nor as well preserved as are the smaller biota. R. Lamb (personal
communication) provided the following about the fossil mollusks: “‘All but the
rarest species of land and freshwater mollusks identified from Pit 91 are found
in the same deposit as the chipmunk. The land snails associated with the chipmunk
fossils are characteristic of riparian environments. A quantitative analysis of all
mollusks shows an increase in individuals of permanent water and streambank
species in the stratigraphic levels bearing chipmunks.” Only one of the three fish
identified at Rancho La Brea is present (Swift 1989), the threespine stickleback,
Gasterosteus aculeatus. There is a diverse snake assemblage (La Duke 1983) with
the garter snake, Thamnophis sirtalis forming 70% of the snakes; however, this
is also the common snake in other, well-sampled sites. The skink, Eumeces skil-
tonianus Baird and Girard, 1852, and the alligator lizard, Gerrhonotus multicar-
inatus Blainville, 1825, form 76% of the lizard fauna, a slightly larger percentage
than in other areas of Pit 91 examined to date. The small mammals associated
with the chipmunk are all those previously recognized at Rancho La Brea (Harris
and Jefferson 1985) with the exception of the bats, mole, and desert shrew, but
these missing taxa are generally rare.
Considered together, this fossil assemblage does not appear to contain clear
evidence to explain the restricted occurrence of chipmunk in the northeast deposit.
The assemblage, including the chipmunk, suggests a riparian habitat surrounded
by more open and/or brushy terrain. There is ample evidence from other studies
of the fossil plants and animals that Rancho La Brea has preserved several com-
munities including costal sage shrub, chaparral, deep canyon, and riparian (Tem-
pleton 1964; Warter 1976; Shaw and Quinn 1986). Consequently, the preferred
habitat of present-day 7. merriami (chaparral and foothill scrub) was present
during the span of time represented by the Rancho La Brea deposits and is present
RESEARCH NOTES 121
today in the Santa Monica Mountains. The restriction of chipmunks to a single,
small fossil deposit and their present-day absence in ample, suitable habitat in
the adjacent Santa Monica Mountains are thus a bit enigmatic. One possible
reason for this restricted fossil occurrence of Tamias is that the deposit that yielded
the specimens is one of the oldest sampled at Rancho La Brea and conditions
may have been slightly different from those that prevailed during later times.
In conjunction with this study, all sciurid specimens recovered from Pit 91 and
the other Rancho La Brea excavations (over 500 specimens) were also examined.
Notable by its absence is the tree squirrel, Sciurus griseus Ord, 1818 (see also
Shaw and Quinn 1986). The paleoecological reconstructions for the Late Pleis-
tocene at Rancho La Brea certainly support the presence of the oak/foothill wood-
land association favored by this tree squirrel today. Acorns, the preferred food
of S. griseus, are common as fossils at Rancho La Brea (Templeton 1964). Ad-
ditionally, this squirrel is currently found in close association with 7. merriami
in the coast ranges of central California.
Acknowledgments
The author wishes to thank S. Cox and L. Orloff for assistance in analyzing the
fossil samples reported in this study. This study has benefitted from discussions
with S. George, G. Jefferson, R. Lamb, and C. Shaw. R. Hannum provided insights
on the present-day range of chipmunks. L. Barkley provided assistance with
modern comparative material.
Literature Cited
Akersten, W. A., C. A. Shaw, and G. T. Jefferson. 1983. Rancho La Brea: status and future. Paleobiol-
ogy, 9(3):211-217.
Black, C. C. 1963. A review of the North American Tertiary Sciuridae. Bull. Mus. Comparative
Zool., Harvard Univ., 130(3):113-248.
Goodwin, H. T., and R. E. Reynolds. 1989. Late Quaternary Sciuridae from Kokoweef Cave, San
Bernardino County, California. Bull. Southern Calif. Acad. Sci., 88(1):21-32.
Harris, J. M., and G. T. Jefferson. 1985. Rancho La Brea: treasures of the tar pits. Nat. Hist. Mus.
Los Angeles Co., Sci. Ser., 31:1-87.
LaDuke, T. C. 1983. The fossil snake fauna of Pit 91, Rancho La Brea, Los Angeles County,
California. M.S. Thesis, Dept. Zool., Michigan State Univ., 57 pp.
Levenson, H., R. S. Hoffmann, C. F. Nadler, L. Deutsch, and S. D. Freeman. 1985. Systematics of
the Holarctic chipmunks (Tamias). Jour. Mamm., 66(2):219-242.
Marcus, L. F., and R. Berger. 1984. The significance of radiocarbon dates for Rancho La Brea. Pp.
159-183 in Quaternary extinctions, a prehistoric revolution. (P. S. Martin and R. G. Klein,
eds.), Univ. Arizona Press, Tucson.
Miller, W.E. 1971. Pleistocene vertebrates of the Los Angeles basin and vicinity (exclusive of Rancho
La Brea). Bull. Nat. Hist. Mus. Los Angeles Co., 10:1-124.
Shaw, C. A. 1982. Techniques used in excavation, preparation, and curation of fossils from Rancho
La Brea. Curator, 25(1):63-77.
,and J. P. Quinn. 1986. Rancho La Brea: a look at coastal southern California’s past. California
Geol., 39(6):123-133.
Stock, C. 1956. Rancho La Brea. A record of Pleistocene life in California. Sixth edition. Los Angeles
Co. Mus. Nat. Hist., Sci. Ser. 20, Paleont., 11:1-81.
Swift, C. C. 1989. Late Pleistocene freshwater fishes from the Rancho La Brea deposit, Los Angeles,
California. Bull. Southern Calif. Acad. Sci. 88(3):93-102.
Templeton, B.C. 1964. The fruits and seeds of the Rancho La Brea Pleistocene deposits. Unpublished
Ph.D. dissertation, Oregon State Univ., 224 pp.
Warter, J. K. 1976. Late Pleistocene plant communities—evidence from the Rancho La Brea tar
122 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
pits. Proceedings, Symposium on Plant Communities of Southern California, Spec. Publ. Cal-
ifornia Native Plant Soc., 2:32-39.
Woodard, G. D., and L. F. Marcus. 1973. Rancho La Brea fossil deposits: a re-evaluation from
stratigraphic and geological evidence. Jour. Paleont., 47(1):54-69.
Accepted for publication 22 June 1989.
David P. Whistler, Section of Vertebrate Paleontology, Natural History Museum
of Los Angeles County, 900 Exposition Blvd., Los Angeles, California 90007.
Bull. Southern California Acad. Sci.
88(3), 1989, pp. 123-126
© Southern California Academy of Sciences, 1989
A Note on the Ontogenetic Age of the
Rancho La Brea Hominid.
Los Angeles, California
Although human remains from Pit 10, Rancho La Brea (now Hancock Park)
have been known since 1914 (Merriam 1914) they have received little attention
by physical anthropologists or anatomists. Kroeber examined the remains of “La
Brea Woman” (LACM HC 1323) with Merriam in 1914 (Heizer 1962) but did
not publish a report; his notes were eventually published posthumously (Kroeber
1962). In that publication Kroeber assessed the age of the individual as “‘perhaps
25 years old” but did not discuss how that age was determined. In 1981 the
cranium, mandible and a few post cranial bones of “‘La Brea Woman”’ received
a more systematic assessment (Bromage and Shermis 1981) but that study dealt
only superficially and incompletely with the ontogenetic age of the individual. It
is the purpose of this note to reassess that age based on a fuller analysis of the
preserved cranial and post cranial indicators.
Bromage and Shermis (1981) estimated the age of the La Brea hominid at 25-
30 years based largely on dental attrition. They suggested that the dentition showed
“third stage dental wear’ yet gave no reference, discussion or comparative data
to support that estimate. They appear, in fact, to have used Brothwell’s ageing
classification (1972), which contains a “‘third stage dental wear” phase consistent
with the features on LACM HC 1323 that they cite. If they have, in fact, used
Brothwell’s system they should have noted the possible inaccuracies resulting
from use of a system which does not include or consider the often heavily attrited
coastal California materials. Brothwell’s ageing classification was based on British
materials from the Neolithic through Medieval periods. Bromage and Shermis
supported their age assessment by pointing out that all of the adult teeth had
erupted at the time of death (=14+) and that the medial epicondyle of the humerus
had fused (=16+). They suggested, without discussion or any supporting data,
that a lack of fusion of the iliac crest may have been due to “‘dietary deficiencies.”
In the current study, initial analysis of all the hominid skeletal materials from
Pit 10 suggests that only a single individual is represented. All the remains are
from a small individual of very delicate build and there is no duplication of
materials. Features of the innominate, such as a wide sciatic notch and a mod-
erately developed preauricular sulcus, suggest that the individual is female. None
of the preserved long bones are complete enough for a secure stature estimate.
Re-evaluation of the La Brea hominid materials allows a more precise, and
younger, age than that estimated by either Kroeber or Shermis and Bromage.
The dental eruption sequence appears to have been complete although all of
the anterior dentition was lost post mortem. In the maxilla, 3 teeth remain: the
RM!” and the LM!. Alveoli for the missing molar teeth are present and suggest
that these teeth were in full eruption at death. An ectopic right canine may have
been present but this issue does not affect the estimated age of the individual. In
the mandible, only a single tooth remains, the LM; which was impacted; the
123
124 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
occlusal surface of this tooth was rotated mesially with the long axis of the tooth
at approximately 45 degrees to the long axis of the horizontal ramus. Radiograph-
ically the root apices of this tooth appear to be closed. Radiographic and visual
examination suggest that a similarly impacted M, may have been present on the
right but was lost ante mortem. McKern and Stewart (1957) reported that the
third molars in their study (on males only) had generally erupted between 17 and
22 years. Other authors have reported, however, a much longer eruption period;
Romanes (1972), for example, reported that third molars may erupt between 13
and 25 years.
Most authors are now in agreement that closure of the cranial sutures show
such high individual variability that they should not be used as age indicators
(i.e., Hrdlicka 1939; Singer 1953; Brooks 1955; McKern and Stewart 1957; Krog-
man 1962). Closure of the spheno-occipital synchondrosis (the “basilar suture’),
however, 1s an exception and shows high consistency of age at closure (McKern
and Stewart 1957). Moreover, ossification of post cranial bones from secondary
or epiphyseal centers provides an important and reliable source of information
regarding ontogenetic age. Although variability exists in rates of closure between
different individuals and between the sexes, most ossification activity at epiphyseal
centers can be shown to occur during a consistent and restricted span of time.
Variability in epiphyseal closure rates between races seems to be relatively minor:
“Intra-racial variability is much more marked than inter-racial differences” (Krog-
man 1962:32). Several sources are available for assigning age to post cranial
remains (Stevenson 1924; Todd 1930; Stewart 1934; Krogman 1962); however,
the largest and best documented study of epiphyseal closure was published by
McKern and Stewart in 1957. It should be noted that McKern and Stewart’s data
were derived from male Korean war dead and while sexually homogeneous the
sample was racially mixed. In males, growth is slower than in females and occurs
over a longer time period; therefore ontogenetic age estimates derived from male
data would represent maximum values for a female individual, as the La Brea
hominid undoubtedly is (see above).
In the cranium of the La Brea hominid the spheno-occipital synchondrosis 1s
not fused (confirmed by radiographs), although it is in the last phase (Stage 3) of
fusion. McKern and Stewart (1957) indicate that complete fusion of this syn-
chondrosis took place in 97% of their male sample by 19 years and in 100% by
21 years of age. Huschke’s foramina are present bilaterally (contra Bromage and
Shermis); however since these foramina, normally present during ossification of
the tympanic plate, usually close by the 5th year (Clemente 1985) their presence,
while of interest, has no bearing on the age determination of this individual.
The post cranial remains of the La Brea hominid are incomplete and few in
number: a fragmentary scapula, right humerus, radius (side?; destroyed for ra-
diocarbon dating), ulnar shaft (side?), right femur (now lost), left femur (destroyed
for radiocarbon dating but represented by a cast), left innominate, cervical (N =
2) and lumbar (N = 2) vertebrae. Of this material, the scapula and innominate
contain at least two epiphyseal centers which had not yet completely fused. On
the innominate, much of the iliac crest was lost, apparently through post mortem
abrasion, and the condition of the diaphyseal/epiphyseal interface cannot be de-
termined. However, at the anterior superior iliac spine a small portion of the iliac
crest is preserved and is fused. Posterior to this fused area, at the superior surface
RESEARCH NOTES 125
of the iliac diaphysis, a few vertical striations, characteristic of an active, fusing
diaphyseal surface are seen. At 5.1 cm posterior to the anterior superior iliac spine,
at the superior terminus of the acetabulo-cristal buttress, a small area of epiphysis
is adherent to the diaphyseal surface. The closure of the iliac crest at the anterior
superior iliac spine and in the middle third of the crest are consistent with a
“partial fusion”’ stage (Webb and Suchey 1985). In that study of 116 females,
100% of their sample aged 16 and 17 years demonstrated a similar “‘partial fusion”
stage; this stage could continue, however, until 23 years of age. In no individual
was full fusion present before the age of 18 years. Although the scapula contains
a number of secondary ossification centers, only the acromial center on this
specimen is undamaged enough for evaluation. On the lateral margin of the
acromion process there is a small area of fusing epiphysis consistent with a Stage
1 fusion. Again, confirmation of active fusion was confirmed radiographically. In
McKern and Stewart’s male data (1957) 40% of their sample showed complete
fusion by 17 years; 95% of their sample showed complete fusion by 21 years and
all had fused by 23 years.
It is clear from the cited evidence that the La Brea hominid had not yet reached
full adult status at the time of death although a precise determination of her age
is not possible. Difficulties are encountered because although inter-racial vari-
ability in closure of the basi-occipital synchondrosis and the post-cranial epiphyses
in recent populations is relatively minimal we do not know that this was also the
case for pre-contact southern California populations. Moreover, the undoubted
female status of the La Brea hominid means that her age in years would very
likely be somewhat less that the skeletal age of the all-male comparative sample
of McKern and Stewart. Nevertheless, the fully erupted adult dentition suggests
that an age younger than 17 years is unlikely, while the evidence of the unfused
spheno-occipital synchondrosis and the unfused condition of the acromial and
iliac crest epiphyses suggest that an age of more than 18 years is also unlikely. A
reasonable estimate of the age of the La Brea hominid would therefore seem to
be 17-18 years.
Acknowledgments
I would like to thank John Harris, Los Angeles County Museum and George
Jefferson, Page Museum, for generously making the La Brea hominid available
to me for study, and Suellen Gauld for valuable research assistance. Queen of
Angeles-Hollywood Presbyterian Hospital generously provided the radiographs
and Michael Zucker, M.D. provided assistance in the interpretations of those
radiographs. Remaining errors, however, are my own.
Literature Cited
Bromage, T., and S. Shermis. 1981. The LaBrea Woman (HC 1323); descriptive analysis. Soc. Calif.
Arch. Occ. Pap., 3:59-75.
Brooks, S. 1955. Skeletal age at death: reliability of cranial and pubic age indicators. Am. J. Phys.
Anthropol., 13:567-597.
Brothwell, D. 1972. Digging up bones. Brit. Mus. (Nat. Hist.), London, xiii + 194 pp.
Clemente, C. D. 1985. Henry Gray. Anatomy of the human body. 39th American Edition. Lea and
Febiger, Philadelphia, xvii + 1676 pp.
Heizer, R. 1962. The Rancho La Brea skull. Am. Ant., 27:416.
Hrdlicka, A. 1939. Practical anthropometry. Wistar Inst., Philadelphia, xiv + 231 pp.
126 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Kroeber, A. 1962. The Rancho La Brea skull. Am. Ant., 27:416-417.
Krogman, W. 1962. The human skeleton in forensic medicine. Charles Thomas, Springfield, III.,
337 pp.
Merriam, J. 1914. Preliminary report on the discovery of human remains in an asphalt deposit at
Rancho La Brea. Science, NS, 40:198-203.
McKern, T., and T. Stewart. 1957. Skeletal age changes in young American males. Headquarters
Quartermaster Research and Development, Tech. Rep. EP-45. Natick, Mass.
Romanes, G. 1972. Cunningham’s textbook of anatomy, 11th Ed. Oxford U. Press, London, xv +
996 pp.
Singer, R. 1953. Estimation of age from cranial suture closure. J. For. Med., 1:52-59.
Stevenson, P. 1924. Age order of epiphyseal union in man. Am. J. Phys. Anthropol., 7:53-93.
Stewart, T. 1934. Sequence of epiphyseal union, third molar eruption and suture closure in Eskimos
and American Indians. Am. J. Phys. Anthropol., 19:433-452.
Todd, T. 1930. The anatomical features of epiphyseal union. Child Dev., 1:186-194.
Webb, P., and J. Suchey. 1985. Epiphyseal union of the anterior iliac crest and medial clavicle in a
modern multiracial sample of American males and females. Am. J. Phys. Anthropol., 68:457—
466.
Accepted for publication 22 June 1989.
G. E. Kennedy, Department of Anthropology, University of California, Los Angeles,
California 90024.
Bull. Southern California Acad. Sci.
88(3), 1989, pp. 127-130
© Southern California Academy of Sciences, 1989
The Natal Pterylosis of Closed-nest Building
Tyrant Flycatchers (Aves: Tyrannidae)
The tyrant flycatchers (Tyrannidae) are perhaps the most diverse and dominant
family of New World passerine birds (Traylor and Fitzpatrick 1982). Although
the systematics, morphology, behavior, and ecology have received much attention
(Traylor 1977, 1979; Traylor and Fitzpatrick 1982 and references therein) the
natal pterylosis has been described for only a few temperate zone species (Weth-
erbee 1957, 1958) of this largely Neotropical family.
This paper provides detailed information on the natal pterylosis of seven species
in six genera of Neotropical tyrannids (Table 2); field observations of additional
species are also summarized in Table 1. One newly hatched nestling (stage A,
Wetherbee 1957) of the Slaty-capped Flycatcher, Leptopogon superciliaris, was
collected 11 km north of Maracay, Est. Aragua, Venezuela on 26 June 1972 by
C. T. Collins; an additional two specimens (stage A) were collected, from one
nest, 1 km west of Macas, Prov. Morona-Santiago, Ecuador on 13 August 1988
by M. Marin A. Single specimens of the Common Tody Flycatcher, Todirostrum
cinereum, Yellow-breasted Flycatcher, To/momyias flaviventris, Rusty-margined
Flycatcher, Myiozetetes cayanensis, Social Flycatcher, Myiozetetes similis, and
Great Kiskadee, Pitangus sulphuratus and two specimens of the Pied Water-tyrant
Fluvicola pica were collected at Fundo Pecuario Masaguaral, 45 km south of
Calabozo, Est. Guarico, Venezuela between 22 April 1979 and 27 May 1980 by
B. T. Thomas. The specimen of Pitangus sulphuratus had pin feathers emerging
through the skin (Stage B); all of the others were late stage embryos or newly
hatched (Stage A) (Wetherbee 1957). The classification of the Tyrannidae followed
here is that of Traylor (1977, 1979); common names are those of Meyer de
Schauensee (1966).
Specimens were examined under a binocular dissecting microscope and the
number and distribution of natal downs (neossoptiles) in each species recorded
(Table 2). The terminology for neossoptile tracts and regions within tracts follows
Wetherbee (1957). Neossoptiles were present in the coronal, occipital regions and
scapular tract in all nine specimens with downs present; the specimen of TJol-
momyias was completely naked! The spinal and caudal tracts had neossoptiles
present in 8 of these 9 specimens. Alar, ventral, and postauricular neossoptiles
were present only in Fluvicola, Myiozetetes, and Pitangus. Ventral cervical neos-
soptiles, which are only infrequently observed in passerine birds, were present in
only the single specimen of Myiozetetes similis.
Each of the closed-nest building species considered here (Table 1) has chicks
which are either naked at hatching or have a relatively low total number of
neossoptiles in a small number of tracts. By contrast eighteen specimens from ten
genera of open-cup nesting tyrannids averaged 401 neossoptiles (range 154-607)
(Wetherbee 1967; Collins, in prep). Although the number and distribution pattern
of passerine neossoptiles has proven in some cases to reflect taxonomic affinities
(Collins and Kemp 1976), that is unlikely to be the case here as one or more
“groups” of genera (Traylor and Fitzpatrick 1982) in each of three proposed
127
128 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Table 1. Natal down pattern in closed-nest building tyrant flycatchers.
Species Total number of neossoptiles Source
Elainiinae
Paltry Tyrannulet dense long grey down Skutch, 1960:471.
(Zimmerius villissimus)
Ocre-belled Flycatcher ‘sparse but long grey natal down” Skutch, 1960:568.
(Mionectes oleaginea) 52 Collins, in prep.
Slaty-capped Flycatcher 53-64 this study
(Leptopogon superciliaris)
Common Tody Flycatcher 16 this study
(Todirostrum cinereum)
Slaty-headed Tody Flycatcher “tufts of grey down on the crown oc- Skutch, 1960:495.
(Todirostrum sylvia) ciput, shoulders and middle of
back”
Eye-ringed Flycatcher ‘“‘sparse but rather long grey down” Skutch, 1960:511.
(Rhynchocyclus brevirostris)
Yellow-breasted Flycatcher 0 this study
(Tolmomyias flaviventris)
Yellow-olive Flycatcher “devoid of down” Skutch, 1960:502.
(Tolmomyias sulphurescens)
Fluvicolinae
Pied Water-tyrant 154-156 this study
(Fluvicola pica)
Royal Flycatcher “utterly naked” Skutch, 1960:526.
(Onychorhynchus mexicanus)
Ruddy-tailed Flycatcher “devoid of down” Skutch, 1960:535.
(Terenotriccus erythrurus)
Sulphur-rumped flycatcher “devoid of down” Skutch, 1960:545.
(Myiobius sulphureipygius)
Black-tailed Flycatcher “devoid of down” Skutch, 1960:552
(Myiobius atricaudus) “naked” Gross, 1964
Tyranninae
Rusty-margined Flycatcher 185 this study
(Myiozetetes cayanensis)
Social Flycatcher 252 this study
(Myiozetetes similis) “sparse light grey down” Skutch, 1960:441.
Gray-capped Flycatcher “sparse light grey down” Skutch, 1960:418.
(Myiozetetes granadensis)
Great Kiskadee 269 this study
(Pitangus sulphuratus)
subfamilies of the Tyrannidae (Traylor 1977, 1979) are represented by the genera
considered in this study. Of greater significance is an ecological correlation among
all of the genera considered here—they all build covered, domed, or ball-shaped
nests, a habit which is thought to have been evolved independently in all three
subfamilies (Traylor and Fitzpatrick 1982). Although the reason for the absence
or reduction in number of neossoptiles has been considered by a number of
authors, with little agreement (see Wetherbee 1957:353), this pattern does seem
RESEARCH NOTES 129
Table 2. Distribution and counts of neossoptiles.
Species?
Todi- Myioze-
rostrum Myioze- tetes Pitangus
Leptopogon cine- tetes caya- __ sulphura-
Region superciliaris reum Fluvicola pica similis nensis tus
Ocular? — — — — 6/9 4/5 11/11 9/9 18/18
Coronal Wi WSs We 5/5 10/10 8/8 19/19 6/6 16/16
Postauricular — _— — — 2/2 0/0 2/0 — 1/1
Occipital 4/4 4/3 4/3 1/1 5/5 4/3 8/8 3/3 6/6
Cervical — — — — — — 2/2 _ _
Middorsal 7/6 8/8 6/6 _ 7/7 9/9 17/16 6/7 20/21
Pelvic® _ — —_ — 0 3 3 4 5
Scapular 6/6 6/6 4/5 3/1 5/10 8/8 13/12 13/13 15/17
Femoral 0/2 0/0 0/0 — 6/6 7/6 11/10 12/12 15/15
Ventral — — _ — 10/5 8/9 9/16 23/10 10/10
Caudal 6/6 6/6 6/6 — 3/3 6/6 6/6 5/6 6/6
Crural — — _ — 3/6 6/4 8/10 8/7 7/8
Primaries — — — — 1/1 0/0 — — =
Secondaries 2/1 0/0 0/0 — 8/8 0/0 2/2 1/1 —
Greater Secondary Coverts — — 5/6 10/11 6/8 5/4 9/8
Middle Secondary Coverts — — 2/3 7/5 7/5 6/5 Wi
Lesser Secondary Coverts _ _ — — 1/1 — —
Carpal Remix Covert _ — 0/0 1/1 0/1 — —
Patagial Coverts _ _ _ _ — — —
Total 64 60 £53 16 154 156 252 185 269
4 Tolmomyias flaviventris was entirely naked and omitted from this table.
> Number of neossoptiles (right/left).
¢ Single midline row; all other tracts bilaterally paired.
to be common in taxa which utilize enclosed, cavity or domed nests. In the
Tyrannidae an open cup nest is most common, and, has been thought to be
primitive (Traylor and Fitzpatrick 1982). If so, the absence or reduced number
of neossoptiles in some tyrannid genera is most likely to be also derived along
with the closed nesting habit. It is interesting to note that Pitangus and Myiozetetes
build loosely constructed bulky, ball-shaped nests in rather conspicuous open
places in trees or other structures (Traylor and Fitzpatrick 1982: Fig. 3g; Skutch
1960). The remaining genera considered here (Table 1) all build tightly woven,
globular or pyriform shaped nests frequently suspended from an overhanging twig
or root (Traylor and Fitzpatrick 1982: Fig. 31-31). Pitangus sometimes builds an
initially open cup-shaped nest which is gradually covered over during later stages
of incubation. This is perhaps indicative of a more recently derived tendency and
shows an evolutionary pathway from a primitive open to a derived closed, type
of nest construction (Traylor and Fitzpatrick 1982:16). Pitangus and Myiozetetes
also show the least reduction in number of neossoptiles which would be in keeping
with species having only recently derived the habit of building a closed nest. The
Piratic Flycatcher, Legatus leucophaius, which pirates covered nests of other species
and the Sulphur-bellied Flycatcher, Myiodynastes luteiventris, which builds its
cup-shaped nest in a cavity in a dead tree or old woodpecker hole both seem to
have the denser down covering typical of open-nesting tyrannids (Skutch 1960)
130 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
from which they also may be more recently derived. Apparent exceptions to the
above noted correlation are the Golden-crowned Spadebill, Platyrinchus coro-
natus, which builds an open cup nest and yet has nestlings which are entirely
naked (Skutch 1960:335), and the Paltry Tyrannulet, Zimmerius vilissimus which
builds a closed, domed nest with a side opening, and yet has “‘on its crown, back,
sides, long grey down which is dense for a passerine” (Skutch 1960:471).
Information on the natal pterylosis of tyrant flycatchers still is drawn from a
few individuals constituting only a small fraction of the 374 species and 88 genera
of the Tyrannidae. Included are forms representing only part of the broad range
of morphological and geographical diversity which occurs in this dominant New
World family. When detailed information on the natal pterylosis is available for
a greater array of taxa it is possible that some clearer taxonomic as well as ecological
correlates will be found.
Acknowledgments
We are indebted to Manuel Marin Aspillaga and Betsy Trent Thomas for pro-
viding specimens utilized in this study and Stuart L. Warter for comments on the
manuscript.
Literature Cited
Collins, C. T., and M. H. Kemp. 1976. Natal pterylosis of Sporophila finches. Wilson Bull., 88:154—
SWE
Gross, A. O. 1964. Nesting of the Black-tailed Flycatcher on Barro Colorado Island. Wilson Bull.,
76:248-256.
Meyer de Schauensee, R. 1966. The species of birds of South America and their distribution. Acad.
Nat. Sci., Philadelphia, xviii + 577 pp.
Traylor, M. A., Jr. 1977. A classification of the tyrant flycatchers (Tyrannidae). Mus. Comp. Zool.
Bull., 148:129-184.
—. 1979. Tyrannidae. Pp. 1-228 in Peters’ Check-list of birds of the world, vol. 8. (M. A.
Traylor, Jr., ed.), Museum Comparative Zoology, Cambridge, x + 365 pp.
, and J. W. Fitzpatrick. 1982. A survey of the tyrant flycatchers. Living Bird, 19:7-45.
Skutch, A. 1960. Life histories of Central American birds. Pac. Coast Avif., 34:1-593.
Wetherbee, D. K. 1957. Natal plumages and downy pteryloses of passerine birds of North America.
Bull. Am. Mus. Nat. Hist., 113:339-436.
—. 1958. New descriptions of natal pterylosis of various bird species. Bird-Banding, 29:232-
236.
Accepted for publication 12 May 1989.
Charles T. Collins and Kathleen M. McDaniel, Department of Biology, California
State University, Long Beach, California 90840.
Bull. Southern California Acad. Sci.
88(3), 1989, pp. 131-134
© Southern California Academy of Sciences, 1989
A New Species of the Genus Traubella
(Siphonaptera: Ceratophyllidae)
Among fleas found on canyon mice (Peromyscus crinitus) collected in south-
western Utah were several specimens of an undescribed species of Traubella
Prince, Eads and Barnes 1976, the first time this genus had been found in the
state. The present paper describes this new taxon.
Traubella grundmanni, new species
Figs. 4-7
Diagnosis. —Selected generic characteristics shared with Traubella neotomae (I.
Fox, 1940) include: head with frontoclypeal tubercle present but scarcely notice-
able; eye relatively small and lightly pigmented except for outer rim; trabecula
centralis present; row of minute setae (one to three setae in width) extending full
length of dorsal margin of antennal fossae in both sexes. Labial palps five-seg-
mented and about 90 percent as long as forecoxae. Apical spinelets present on
first four abdominal terga. One long and two minute flanking antepygidial setae
in male, and three long, well-developed antepygidial setae, each of a different
length but center one longest, in female. Penis rods not coiled. Ventral anal lobe
of female similar in shape, and number, arrangement and size of setae for both
species.
The description that follows compares the new species with Traubella neotomae
(I. Fox, 1940), the only other known species in the genus.
Modified Abdominal Segments, Male. —Finger longer than fixed process (Fig.
4) rather than equal (Fig. 1), finger almost flat-topped (Fig. 4), not gradually tapered
(Fig. 1), and with about 6—7 marginal, unpigmented or lightly pigmented, mostly
long, tapering setae of unequal lengths; of these the anterior seta longest, and one
(usually the third or fourth seta posterior to apex) much shorter and less developed
than others. Apex of fixed process (Fig. 4) somewhat less rounded than in neotomae
(Fig. 1), tipped with one seta rather than three. Two acetabular setae in about the
same position but longer than the two in neotomae.
Modified Abdominal Segments, Female. — Anal stylet with one long apical seta
and two minute setae along its shaft, as opposed to stylet with one long apical
seta flanked by a shorter ventrolateral subapical seta. Posterior border of sternum
VII variable (Fig. 7) with deep sinus separating upper narrow, well developed
lobe from much broader lower lobe compared to a shallow sinus and broadly
rounded, scarcely differentiated upper and lower subequal lobes (Fig. 2). Differ-
ences in numbers and position of setae on sternum VII as in Figs. 2 and 5.
Bulga of spermatheca (Fig. 6) approximately the same length as neotomae (Fig.
3) but much narrower in diameter and with slightly concave upper and lower
borders, not inflated with well rounded upper border and straight lower border;
hilla strongly bent and about the same length in both species, but narrower and
without sclerotized tip in grundmanni.
Size (total length mm, mounted specimens). —Males (N = 3), range 2.7—3.2,
average 2.9; Females (N = 9), range 2.8-3.7, average 3.4.
131
132 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
7.
Figs. 1-3. Traubella neotomae. Fig. 1, finger and fixed process of male; Fig. 2, sternite VII of
female; Fig. 3, spermatheca.
Figs. 4-7. Traubella grundmanni, n. sp. Fig. 4, finger and fixed process of male; Fig. 5, sternite
VII of female; Fig. 6, spermatheca; Fig. 7, variation in outline of posterior margin, sternite VII of
female.
Types. — Holotype male and Allotype female ex Peromyscus crinitus stephensi
Mearns 1897 (H.J.E. host no. 19135) collected at mouth of Snow Canyon, 3.2
km northeast of Ivins: Washington County, Utah, elev. 945.5 m, 19 November
1988, H. J. Egoscue original numbers 11179 and 11181, respectively.
RESEARCH NOTES 133
Paratypes, one male, one female, same data, H. J. Egoscue; two males, five
females, same host species, location and elevation, 19 November 1988, H. J.
Egoscue; one male, same host and location, elevation 950 m, 17 December 1988,
J. Kucera; one female, same data, 18 December 1988, J. Kucera; one male, one
female, same data, R. E. Elbel.
Additional material.—Two males (very badly damaged), host unknown, col-
lected 28 or 29 December 1961 at the type locality by A. W. Grundmann and
unknown students.
The holotype and allotype will be deposited in the U.S. National Museum,
Washington, D.C. Paratypes collected by me will remain, at least temporarily, in
my collection; those borrowed from R. E. Elbel and J. Kucera are being returned.
This new species is named for Dr. Albert W. Grundmann, Professor Emeritus,
Department of Biology, University of Utah, in recognition of his many years of
teaching and research in the field of parasitology and for providing the first spec-
imens of this flea.
Discussion. — Traubella neotoma (1. Fox 1940) (Figs. 4-7), type host, Neotoma
lepida, was originally described by Fox (1940) as a species of Amphipsylla. Prince
et al. (1976) carefully reviewed the controversial taxonomic history of the taxon
and corrected the situation by erecting the genus 7raubella to hold neotomae.
They also adequately discussed the many similarities between Traubella and
Malaraeus and indicated the closest affinities of Traubella were with what was
then called the telchinus group (eremicus, sinomus and telchinus) with which
Traubella was compared.
Traubella grundmanni is known only from the type locality, where its preferred
or true host appears to be Peromyscus crinitus stephensi. Special efforts to find
grundmanni on other species of small mammals at Snow Canyon were unsuc-
cessful. More intensive year-around collecting at Castle Cliff and environs less
than 32 km distant on the west slope of Beaver Dam Mountains yielded no
specimens of the new flea from either woodrat nests or numerous hosts of several
rodent species which, however, included very few canyon mice.
The type locality of gr'undmanni is located among colorful sandstone formations
in what is often called “‘red rock country.” Dominant shrubs along the dry wash
leading out of Snow Canyon include creosote bush, Larrea tridentata, and sand
sagebrush, Artemisia filifolia. But the lava flows and cliffs favored by canyon mice
there are very sparsely vegetated and lack characteristic shrubs. The climate in
this part of Utah is characterized by low annual rainfall, wide ranges in temper-
atures, long, hot summers and relatively mild winters—conditions probably quite
similar to those in southern California and other parts of the arid southwest where
Traubella neotomae has been reported (Prince et al. 1976). Both species of 7rau-
bella are apparently winter fleas.
Acknowledgments
I am grateful to R. E. Elbel and J. Kucera of the University of Utah for their
help, encouragement and companionship on several field trips and for critically
reading the manuscript. H. E. Stark also reviewed the paper and gave useful
suggestions. R. E. Lewis of Iowa State University kindly took time from his busy
schedule to provide taxonomic advice. The work was supported in part by BRSG
S07 07092 awarded by the Biomedical Research Support Program, Division of
134 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Research Resources, National Institutes of Health to the University of Utah.
Permission to use drawings based on photomicrographs of Traubella neotomae
in The Rothschild Collection of Fleas, The Ceratophyllidae: Key to the Genera
and Host Relationships, by Traub, Rothschild and Haddow (1983) was given by
Academic Press and Meriam Rothschild.
Literature Cited
Fox, I. 1940. Siphonaptera from western United States. J. Washington Acad. Science, 30:272-276.
Prince, F. M., R. B. Eads, and A. M. Barnes. 1976. Traubella, a new genus of flea (Siphonaptera:
Ceratophyllidae). J. Medical Entomol., 13:162-168.
Traub, R., M. Rothschild, and J. Haddow. 1983. The Rothschild collection of fleas: the Cerato-
phyllidae, key to the genera and host relationships. Univ. Press, Cambridge, England, 288 pp.
Accepted for publication 10 May 1989.
Harold J. Egoscue, Post Office Box 787, Grantsville, Utah 84029.
Bull. Southern California Acad. Sci.
88(3), 1989, pp. 135-136
© Southern California Academy of Sciences, 1989
INDEX TO VOLUME 88
Bennett, Tony, see Diane N. Waugh
Collins, Charles T. and Kathleen M. McDaniel: The Natal Pterylosis of Closed-
nest Building Tyrant Flycatchers (Aves: Tyrannidae), 127
Dailey, Murray D. and Loris S. Fallace: Prevalence of Parasites in a Wild Pop-
ulation of the Pacific Harbor Seal (Phoca vitulina richardsi), |
Dugoni, Thomas, see Diane N. Waugh
Egoscue, Harold J.: A New Species of the Genus 7raubella (Siphonaptera: Cer-
atophyllidae), 131
Fallace, Lori S., see Murray D. Dailey
Foley, Christopher J. and Brian N. White: Occurrence of Ephydra hians Say
(Diptera: Ephydridae) in Deep Water in Mono Lake, California, 41
Garthwaite, Ronald L. and Stefano Tairi: Platyharthrus aiasensis Legrand (Isopo-
da: Oniscidea: Platyarthridae) in the Americas, 42
Goodwin, H. Thomas and Robert E. Reynolds: Later Quaternary Sciuridae from
Kokoweef Cave, San Bernardino County, California, 21
Kennedy, G. E.: A Note on the Ontogenetic Age of the Rancho La Brea Hominid.
Los Angeles, California, 123
Littler, Mark, see Steven N. Murray
Martin, Robert A. and Robert H. Prince: A New Species of Early Pleistocene
Cotton Rat from the Anza-Borrego Desert of Southern California, 80
McDaniel, Kathleen M., see Charles T. Collins
Morris, Penny A., Dorothy F. Soule, and John D. Soule: Bryozoans, Hermit Crabs
and Gastropods: Life Strategies Can Affect the Fossil Record, 45
Murray, Steven N. and Mark M. Littler: Seaweeds and Seagrasses of Southern
California: Distributional Lists for Twenty-one Rocky Intertidal Sites, 61
Prince, Robert H., see Robert A. Martin
Reynolds, Robert E., see H. Thomas Goodwin
Shannon, Michael Mishima: Collective Vigilance Enhances Feeding Rates of the
Opaleye Girella nigricans (Girellidae), 88
Sigmodon lindsayi, n. sp., 80
Soule, Dorothy F., see Penny A. Morris
Soule, John D., see Penny A. Morris
Stewart, Joan G.: Notes on Marine Algae of San Diego County Including Merger
of Murray ellopsis with Veleroa, 103
135
136 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Swift, Camm C.: Late Pleistocene Freshwater Fishes from the Rancho La Brea
Deposit, Los Angeles, California, 93
Taiti, Stefano, see Ronald L. Garthwaite
Traubella grundmanni, n. sp., 131
Waugh, Diane N., Tony Bennett, and Thomas Dugoni: The Incidence of the
Cymothoid Isopod Lironeca californica on fishes in Campbell Cove, Sonoma
County, California, 33
Whistler, David P.: Late Pleistocene Chipmunk, Tamias (Mammalia/Sciuridae)
from Rancho La Brea, Los Angeles, California, 117
White, Brian N., see Christopher J. Foley
Wickstein, Mary K.: Key to the Palaemonid Shrimp of the Eastern Pacific Region,
11
Now York Botanical ‘Garden Libra
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3 MExE
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CONTENTS
Late Pleistocene Freshwater Fishes from the Rancho La Brea Deposit,
Southern California By Camm C. Swift
Notes on Marine Algae of San Diego County Including Merger of Murray-
ellopsis with Veleroa By Joan G. Stewart
Research Notes
Late Pleistocene Chipmunk, Tamias, (Mammalia: Sciuridae), from Rancho La Brea, Los An-
geles, California By David P. Whistler
A Note on the Ontogenetic Age of the Rancho La Brea Hominid. Los Angeles, California By
G. E. Kennedy
The Natal Pterylosis of Closed-nest Building Tyrant Flycatchers (Aves: Tyrannidae) By Charles
T. Collins and Kathleen M. McDaniel
A New Species of the Genus Traubella (Siphonaptera: Ceratophyllidae) By Harold J.
Egoscue
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