Late quaternary insects of Rancho La Brea and McKittrick, California

QUATERNARY

RESEARCH

20, 90-104 (1983)

Late Quaternary

Insects of Ranch0 McKittrick, California

La Brea and

SCOTT E. MILLER Museum

of Comparative

Zoology.

Harvard

University.

Cambridge.

Massachusetts

02138

Received June 7. 1982 Asphalt-impregnated sediments at Ranch0 La Brea (Los Angeles County) and McKittrick t Kern County) in California provide a rich Quaternary insect record. Ages of various sites at Rancho La Brea range from more than 40,000 1JCyr B.P. to modern. McKittrick insects studied by W. D. Pierce are not contemporaneous with the late Pleistocene vertebrate fauna. but are only about 7000 1JC yr old. The major paleoecological groupings are: (I) ground dwellers, (2) aquatics. (3) scavengers, and (4) miscellaneous. Contrary to conclusions of earlier authors. most specimens represent modern species. Only two apparent terminal Pleistocene extinctions are recognized. bdth dung beetles (Scarabaeidae).

taxa to be considered “identified”. In addition, some of his drawings were inaccurate (Matthews and Halffter, 1968). His taxonomic procedures and lack of adequate modern ecological data also led him to radical paleoecological conclusions (Miller and Peck. 1979. p. 99). Insects are also known from asphalt deposits near Carpinteria, Santa Barbara County (Lance, 1946; Miller, 1978; Moore and Miller, 1978; Miller and Peck. 1979; Doyen and Miller, 1980), and Maricopa. Kern County (Doyen and Miller. 1980). These are not included here because they were not studied by Pierce and have been only partially studied in recent years. Late Quaternary insect-bearing asphalt deposits which are similar in many respects of fauna1 composition, taphonomy, and age to the California deposits also occur near Talara, Peru (Churcher. 1966). Fyzabad, Trinidad (Blair, 1927), and Binagady, in the Caucasus Region of the USSR (Bogachev, 1948). In order to provide a base for future research, this paper compiles and, where possible, reevaluates all published and some unpublished records of Quaternary insects from Ranch0 La Brea and McKittrick, and discusses paleoecology and extinction. This paper summarizes studies since 1976 by the

INTRODUCTION The Ranch0 La Brea and McKittrick asphalt deposits are among the world’s most renowned late Pleistocene vertebrate localities, but the associated abundant invertebrate and plant fossils have received little attention (Fig. 1). A few entomologists have published on Ranch0 La Brea and McKittrick insects, but most early papers contain significant errors in identifications and interpretations. Preliminary studies were made on limited samples from the early excavations by the University of California Museum of Paleontology (Grinnell, 1908; Blaisdell, 1909; Essig, 193 I ). From 1944 to 1954, W. D. Pierce (1945a1961) worked in detail on insects from excavations by the Natural History Museum of Los Angeles County. Pierce was among the first workers in North America to appreciate the significance of Quaternary insects and their potential as paleoecological indicators (Pierce, 1961; Ashworth, 1979), but his publications and taxonomic procedures were replete with errors. Serious identification problems result from his erection of taxa based on fragmentary specimens (Miller and Peck, 1979). Pierce felt that fragments he could not associate with known species should be described as new 90 0033-.5894/83 $3.00 Copyright

@I) 1983 by the Universtty

of Washington

FOSSIL INSECTS

author and collaborators (see also Moore and Miller, 1978; Miller and Peck, 1979; Doyen and Miller, 1980; Miller et al., 1981; Gagne and Miller, 1982; Miller, 1982; Nagano et al., 1983). Because of the great number of specimens available for study and the elementary state of taxonomic knowledge of many of the groups involved, this paper has been primarily limited to previously published records and no attempt has been made to make specific identifications on many of the generic or family identitications in the literature. Thus, the lists of taxa recorded from these sites (Appendixes 1 and 2) are incomplete and not representative of fauna1 composition, since many more taxa will eventually be added, especially as a result of recent excavation at Ranch0 La Brea Pit 91. The earlier collections, which almost entirely lack detailed stratigraphic data, are best used only as a supplement to the more significant material from Pit 91, Almost all Ranch0 La Brea and McKittrick insect specimens are considered conspecific with Holocene species and fall within reasonable ranges of geographic variation. The use of subspecitiic names is inappropriate, slince the fossils do not represent geographic races and there is no morphological basis upon which to found chronosubspecies. While studies by other workers indicate that almost all Pleistocene insect fossils represent extant species (Ash-

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worth, 1979; Coope, 1978; Morgan and Morgan, 1980b), several Ranch0 La Brea species do appear to be extinct. LOCALITIES

AND DATA

Ranch0 La Brea The Ranch0 La Brea asphalt deposits are located in Hancock Park, Los Angeles, Los Angeles County, California (Fig. 1). More than 100 individual excavations or “pits” have been made since 1905. Most of these were unproductive test holes and fewer than 15 are considered major sources of vertebrate fossils (Howard, 1962; Stock, 1956). Before the reopening of Pit 91 in 1969, emphasis was placed on large vertebrates; insects and other small fossils were neglected by the early excavators. Pierce’s material was largely salvaged from the miscellaneous material associated with the vertebrate collections at LACM.] The specimens reported by Grinnell (1908) were taken from an unstated locality or localities in 1906, probably during excavation of UMCP lot. 1059. Likewise, the speci1 CIT, California Institute of Technology (VP collection now at LACM); LACM, Natural History Museum of Los Angeles County; LACMIP, LACM Invertebrate Paleontology Collection; lot., locality; RLP. Ranch0 La Brea Project, George C. Page Museum, LACM (catalogue number prefix); UCMP. University of California Museum of Paleotology, Berkeley; VP, Vertebrate Paleontology.

FIG. 1. Map showing fossil-bearing asphalt deposits in California: 1, Ranch0 La Brea. 2. McKittrick, 3. Carpinteria, 4. Maricopa.

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mens reported by Essig t 193 1) were taken from an unstated locality or localities. probably UCMP vertebrate excavations (UCMP lot. 2051 (Essig, 1931, Fig. 3) and probably others also). Pierce (1946a, 1947a) mentioned that occurrence of insects from the following pits at Ranch0 La Brea: A, B, 2. 3, 4, 9, 10, 13, 16,28,29,36, 37,.51, 60,61.67,77,81, 101, “Academy”, “Bliss 29”. and “Pit X”. Original matrix was available to Pierce only from Pits A, 81, 101. and presumably Bliss 29 and Pit X. Insects from Pits 3, 4, 13. 60, 61, 67, and 77 were recovered from the brain cavities of Smilodon ccdifornicrts Bovard skulls. Although material from such skulls is probably contemporaneous with the skulls, it may not be (e.g., Harington, 1980, p. 197). Pierce never published data on insects from Pits 2, 10, 16, 29, 36, 5 1, 60, 61, 77, and Academy, so these excavations will receive no further attention in this paper. Details on the other localities follow (for additional details see Howard, 1962; Woodard and Marcus, 1973; Miller and Peck, 1979; Kurten and Anderson, 1980; Marcus and Berger, 1983). Pits A, B, and C were excavated in 1929. and Pierce’s Bliss 29 specimens were collected in 1929 by W. Bliss from these sites after official excavation ended. The excavators had considerable difficulty with caveins and flooding during the excavations (unpublished field notes in LACM), so contamination is quite possible. The age of Bliss 29 insects is even more questionable due to unknown origins and possible contamination. The Bliss 29 material included “burr clover and modern goat bones,” indicative of contamination (J. Everest, unpublished notes in LACMIP, 1942). “A very few examples of burr clover” were also found in Pit B material (Everest. l.c.). Pits 3, 4, 9, 13, 28, 37, and 67 (61 and 67 were actually the same deposit) were among the more important vertebrate fossil-bearing sites excavated by LACM between 1913 and 1915 (Howard, 1962; Woodard and Marcus, 1973). Very few insects are available from

any of these. since they were not saved during the excavations, but some insects were salvaged by Pierce from skull cavities. The bulk of the fossiliferous matrix from Pit 81 was excavated as a block in 1915 and stored intact. In 1945, matrix removed from the block during preparation for exhibit yielded many insects. Pit 81 vertebrates were never studied, but this site was an important source of insects for Pierce. Bone collagen of Equus from Pit 81 (lacking stratigraphic data) yielded a radiocarbon date of 10,940 ? 510 yr B.P. (QC-405. Marcus and Berger, 1983). Pit 91 ( = LACM VP lot. 6909). partially excavated in 1915, was reopened in 1969 by LACM (Warter, 1976; Miller and Peck, 1979). No insects were available before 1969. but the excavation, undertaken with great care (Shaw, 1982), has yielded huge quantities of insects, of which only some 12,000 have been sorted so far. A few new identifications from Pit 91 are reported here: comprehensive coverage must await further study.

Pit 101 was excavated in 1947 and matrix was collected for Pierce from 1.4-m (4.5-ft) and 1.8-m (6-ft) depths. The age is unknown. Pierce’s Pit X, or “Pit t?),” refer to “mixed material lacking data” (Pierce, 1954a), and may not be fossil. The popular conception of fossil accumulation at Ranch0 La Brea presents a picture of great pools of continuously active liquid asphalt (“tar pits”) that trapped unwary animals, which in turn attracted scavengers that also became trapped (e.g., Stock, 1956). However, recent studies (Woodard and Marcus, 1973; Maloney rt nl., 1974) indicate that such “death traps” had only a minor role in the accumulation of fossils. Stratigraphic reinterpretation and radiocarbon dating indicate that the fossil deposits were formed at the sites of discontinuously active asphaltic seeps during the accumulation of alluvium from the late Pleistocene to the present. Most deposits are stratified and can be correlated with faties of surrounding sediments which are not

FOSSIL

INSECTS

impregnated with oil. The pits at Ranch0 La Brea were artifacts of excavation, and did not represent naturally occurring deep pools of liquid asphalt. While the larger and more continuous fossil accumulations probably represent areas where animals were mired in flows or small pools of asphalt, many of the smaller accumulations probably represent localized fluvial concentrations of bones in stream channels or ponds. Once buried, the fossils were impregnated by asphalt permeating upward and laterally. McKittrick

The McKittrick asphalt deposits lie about 0.8 km south of McKittrick, Kern County, in the southern San Joaquin Valley of California (Fig. I). The original 1921 excavation for vertebrates (UCMP lot. 4096) was on the southeast side of the present northern fork of State Highway 58, but the 1925-1927 excavations (UCMP lot. 7139 and CIT VP lot. 138 = LACMIP lot. 5103) lay across that road on the northwest side. The CIT VP lot. 138 “comprises essentially the same areaas UCMPRocality7139” (Schultz, 1938, p. 130) and was about 260 m from the intersection of Highways 58 and 33. The biota is considered late Pleistocene in age, although there is some “admixture of a later (Recent, but not present day) assemblage” (DeMay, 1941, p. 59). Plant material from a UCMP excavation (but lacking further data) yielded a date of 38,000 + 2500 yr B.P. (UCLA-728, Berger and Libby, 1966). Schultz t 1938), Sternburg (1932), and Blum (undated field notes at LACM) all mention the occurrencle of young layers of better stratified sediments with asphaltic matrix containing insects and small vertebrates lying above the Pleistocene vertebratebearing layers at these original localities. Specimens reported by Essig (193 1) were taken from an unstated locality or localities by E. L. Furlong of UCMP in 1926, probably from surf&e layers. Judging from the fauna1 composition reported by Essig, the material was similar in age to that from Pierce’s sites 3 and 4.

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Pierce’s sites 3 and 4 (LACMIP lot. 260) lay southeast of the original localities, on the east side of the present State Highway 33 about 1.2 km south of McKittrick. In 1945, Pierce excavated matrix from a depth of 0.6 m (2 ft) below the surface in a fracture in the recently exposed bank, designating this site 3 (Pierce, 1947b, and unpublished notes). In 1947, Leonard Bessom of LACM collected from a depth of 1.2 m (4 ft) near site 3. Pierce designated Bessom’s locality site 4, and wrote that the 0.6m (2-ft) depth at site 3 would correspond to the 0.6 to 0.76-m (24 to 30-in) layer at site 4. Pierce believed site 3 was younger than site 4, and that the insects from site 3 indicated drier conditions than did those from site 4. Stratigraphic relationships of Pierce’s localities to the CIT and UCMP localities have not been determined, but Pierce’s localities appear to be much younger. Woody stem fragments from Pierce’s excavation of site 4 yielded an age of 7645 ? 250 yr B.P. (QC-782). Plant material and insect fragments that I collected in 1979 from apparently the same stratigraphic layer very near site 4 (the locality has undergone significant erosion) yielded ages of 5255 t 250 and 12,200 ? 250 yr B.P. (QC-995A and QC995B). QC-995B was based on a small sample, which may account for its anomalously high age by increasing the effects of any contaminates (e.g., asphalt). A Cybister elytron from l-m depth in this roadcut (horizontal relationship to Pierce’s sites unknown) provided an experimental 14C accelerator mass spectrometer date of 7975 ? 470 yr B.P. by the Chalk River Nuclear Laboratory, Ontario, Canada (Nagano et al., 1983; A. V. Morgan, personal communication, 1982). Pierce (1957) suggested that site 4 represented a moister environment than site 3 (i.e., the 1.2 m (4 ft) versus 0.6 m (2 ft) depths, respectively); this is probably correct. Taxa recorded from site 3 include the aquatic (Dytiscidae and Hydrophilidae), scavenging (Silphidae), and ground-dwelling (Tenebrionidae) groups, while taxa re-

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corded from site 4 include aquatic (Odonata, Belostomatidae. Nepidae, Notonectidae. Corixidae, Gerridae. Hydrophilidae, Dytiscidae, Haliplidae, Heteroceridae, Cicindelidae, and Carabidae), scavenging (Silphidae and Staphylinidae), ground-dwelling (Tenebrionidae, Carabidae), and miscellaneous groups (Orthoptera, Hymenoptera and Araneida). The diversity, especially of aquatic taxa, at site 4 suggests the presence of at least a semipermanent stream or pool (e.g., presence of damselfly nymphs). The differing composition at site 3 is probably due to a drier depositional environment, but may be partially due to the site being less well studied than site 4 and to unidentified factors that may have allowed better preservation at site 4. Aquatic insects commonly occur in modern asphalt flows at McKittrick, despite considerable distance from bodies of water. Age differences between sites 3 and 4 cannot presently be determined. So few insects are available from the original McKittrick vertebrate fossil excavations that meaningful fauna1 analyses cannot be made. The known taxa (Nicrophoms, Eleodes, etc.) do fit into previous paleoecological reconstructions for these localities. The presence of Eleodes contrasts with their absence at Pierce’s sites 3 and 4, but the paleoecological implications are not clear. The depositional environment at McKittrick was discussed by Schultz (1938) ..

oil reached the surface [from numerous small discontinuously active petroleum seeps] and spread out in sheets of a fraction of an inch or so in thickness [and] it became intercalated with clay. sand, gravel, and windblown material. The resulting product is a rudely stratified material consisting of fine and coarse sediments more or less uniformly saturated with petroleum. The upper layers which contain a Recent vertebrate fauna seem to be somewhat better stratified than the lower levels which contain the Pleistocene vertebrates. VanderHoff (1934) contends that it was mainly during the summer months that the oil became fluid enough to spread over large areas; while the winter rains carried in most of the elastic material.”

The result is a brea’ belt representing a complicated sequence of events. Pierce (1957, p. 945) suggested that “the depth of one inch represents a minimum of ten years, a maximum of more than 25 years.” but data are not adequate to support this suggestion. PALEOECOLOGY Interpretation of accumulation and preservation in asphaltic matrix is difficult, especially in light of our present poor knowledge of insect paleoecology (Kenward, 1978). Two distinct processes appear to be involved: direct preservation (entrapment in viscous asphalt) and indirect preservation (impregnation with asphalt subsequent to death and burial). Accumulation of asphalt deposits occurs in two major ways (Campbell, 1979): the horizontal flow (forming thin, radiating layers) and the filling of natural depressions (forming lense-like deposits). The former accumulation is more common at McKittrick and the latter more common at Ranch0 La Brea, but both seem to occur at each locality. Viscous asphalt flows or pools can trap animals by ( 1) appearing as water, (2) having sticky, melted surfaces at high temperatures which are solid at lower temperatures, and/or (3) being camouflaged with a covering of dirt, leaves. or debris. Entrapment of insects occurs in four ways. (1) Attraction to carrion or other material already trapped or otherwise in contact with asphalt, and is expected among the scavenging insects. (2) Attraction. especially of aquatic species. to pools of oil and water which appear to be water (Hanna. 1924; Miller, 1930; Pierce, 1949b (Fig. 6). 1957: Kennedy, 1917); many aquatic insects locate water by sight and have poor sense of smell. Even water is an effective trap for many terrestrial insects (Kenward, 1978, p. 40). (3) Attraction to the asphalt itself. Some insects are attracted to the fresh tar coating 1 Brea is the viscous of volatile components

asphalt formed by evaporation from oil in seepages.

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INSECTS

on roads (Saylor, 1933; Borell, 1936; Hubbs and Walker, 1947), and some are attracted to burning oil (Linsley, 1943). This attraction may be due, however, to the warmth of the roads and fires. (4) Accidental trapping, without attraction, when insects crawl or fall into asphalt and are not able to free themselves. In practice, taphonomy is a product of all these processes, each of which has occurred at modern asphalt seeps, but the relative importance of each process is not known. The physical characteristics of the asphalt seep and the circumjacent microhabitat(s) are also important; some species may have been more attracted than others to the: particular microhabitat(s) present. A special case of preservation has been observed at asphalt deposits near Maricopa, Kern County, which may be important to evaluation of Pierce’s McKittrick fossils (Miller and Peck, 1979, p. 101). It seems that modern insects and debris can become incorporated into Pleistocene matrix by accumulating in the bottoms of natural cracks and rodent burrows which penetrate into the matrix. Brea fossil insects can be crudely classified as: (1) aquatic and semiaquatic species that lived in water overlying the tar or were attracted to tar appearing as water, (2) scavenging species that lived on or were attracted to carrion or dung, (3) grounddwelling beetles and other terrestrial crawling species, (4) miscellaneous stray individuals of herbivorous and other species which are not preserved in significant numbers, and (5) fragments of insects from feces or gut contents of vertebrates. Members of the minor last category cannot now be distinguishled from the others, but careful studies of vertebrate feeding habits (e.g., Dodson and Wexlar, 1979) should eventuepally allow differentiation. An accurate paleoecological understanding of brea insects requires: (1) reevaluation of previous conclusions, which heretofore prevented an understanding of ecological and extinction patterns, (2) iden-

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tification of more fossils, and (3) accumulation of ecological data on living insects. This paper addresses the first problem, but the others require additional research on the systematics and ecology of the California entomofauna. Paleoecological conclusions that can be drawn from California brea insects are limited at present, but further study of the modern and fossil faunas promises to yield significant contributions to paleoecology (such as those presently possible in Europe, where the modern fauna is well known). Brea insects offer data not only on Quaternary climatic changes, but also on changes in community composition, which may allow evaluation of postulated terminal Pleistocene changes in community ecology. EXTINCTION Extinctions among large vertebrates at the end of the Pleistocene were probably due largely to climatic and resultant ecological changes, with perhaps some influence from human hunting (Axelrod, 1967; Johnson, 1977; Martin and Neuner, 1978). Axelrod (1967) and Martin and Neuner (1978) have suggested that low seasonality of Pleistocene environments permitted the establishment of very complex communities that lack modern analogs, and that the inception of Holocene seasonal environments was a major cause of terminal Pleistocene extinctions. Many birds and mammals recorded from Ranch0 La Brea and McKittrick are now extinct. Terrestrial and aquatic invertebrates other than insects have not been adequately studied, but none have been recognized as extinct. No extinct plants are known from any of the California asphalt deposits (Warter, 1976; Miller, 1979). Studies elsewhere in North America (Ashworth, 1979; Morgan and Morgan, 1980b) and in Europe (Coope, 1978) indicate that no extinction of insects took place at the end of the Pleistocene. At Ranch0 La Brea, the extinction picture has been clouded by Pierce’s many supposedly extinct taxa. Most of these have now been

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placed in synonymy of extant taxa, others have been recognized as being extinct, and others are impossible to identify because of the nature of the types and the taxonomic condition of the groups concerned. For example, it is presently impossible to identify properly Serica kanakoffi, but there is no good reason to assume that it is not conspecific with an extant (but unidentified) species. Serica is a large, poorly understood genus of plant feeding beetles; it is best to assume that this species is extant until contrary evidence is found. The specific indentity of the scarab Phanacus lahreae also cannot presently be determined due to the broken and distorted condition of the holotype. The millipede “Spiroholus” australis is another unidentifiable taxon, known only from the fragmentary types (Fig. 2). The genus Spiroholus is now restricted to the Old World (Keeton, 1960), and australis is probably a Hilton& or Ty-

MILLER

loholus (R. L. Hoffman. personal communication 198 I ). The tenebrionid beetle Coniontis rernnans is not conspecific with any described species (Doyen and Miller, 1980), but this does not mean it is extinct. Corziontis is a genus of numerous poorly understood species, probably including at least several undescribed species. C. retnnans probably still lives in southern California, but has not yet been recognized (most Cotsiontis in museum collections are unidentified or misidentified). The scarab beetles Copris pristinus and Onthophugous everestae appear to be extinct. The most closely related species occur today in Mexico (and in Texas for Copris) and are associated with mammal dung. Probably the reduction of dung availability due to terminal Pleistocene mammal extinctions, as well as direct effects of the changing climate, caused the demise of these scarabs. However, even if the large mammal

FIG. 2. Holotype of Spirobolus nustralis Grinnell, University Invertebrate Specimen 10008. Ranch0 La Brea. Ventral view

of California Museum of Paleontology on left, dorsal view on right.

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fauna had survived into the Holocene, the scarabaeine fauna of California might not have survived the present seasonal dry periods, since their successful reproduction requires adequate moisture (Miller et al., 1981). Of course, possibly one or both of these species still lives in inadequately collected regions of Mexico and remain to be rediscovered.

APPENDIX 1: INSECTS RECORDED FROM RANCH0 LA BREA

This list contains all previously published records for insects and other terrestrial arthropods, as well as a few new records denoted by *. Order

Orthoptera

(grasshoppers and relatives) Family Acrididae (grasshoppers) undetermined species (Pierce, 1947a) Family Stenopelmatidae Cjerusalem crickets) Stenopelma,tus species (Pierce, 1947a) Order

lsoptera

(termites) Family undetermined undetermined species (frass only) (Pierce, 1947a) Order

Hemiptera

(true bugs) Family Corixidae (water boatmen) undetermined species (Pierce, 1947a) Family Notonectidae (backswimmers) Notonecta species (Pierce, 1947a, 1948a) Family Belostomatidae (giant water bugs) Lethocerus americanus (Leidy) (Usinger, 1956; Lauck and Menke, 1961; Menke, 1963, 1979a,b) Family Gerridae (water striders) *Gerris species, partial thorax RLP 6966E, Pit A (det. A. S. Menke and S. E. Miller) Family Reduviidae (assassin bugs) “Rasahus species cf. R. guttatus (Say), thorax RLP 6913E, Pit A (det. R. C. Froeschner)

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Order

Coleoptera

(beetles) Family Cicindelidae (tiger beetles) (Pierce, 1947a; Nagano et al., 1983) Cicindela haemorrhagica LeConte Cicindela oregona LeConte Family Carabidae (ground beetles) Amara insignis Dejean (Grinnell, 1908) Calosoma semilaeve LeConte (Grinnell, 1908) *Elaphrus species cf. E. jmitimus Casey, partial elytron C127a, Pit A (det. H. Goulet) = Elaphrus ruscarius foveatus Pierce, 1948a (misidentification) *Elaphrus californicus Mannerheim, right elytron RLP 2356E, Pit 91, GJM 8961 L-4 (det. H. Goulet) Platynus species cf. P. funebris LeConte (Grinnell, 1908) Pterostichus species (Grinnell, 1908; Essig, 193 1) unidentified species Family Haliplidae (crawling water beetles) undetermined species (Pierce, 1947a) Family Dytiscidae (predaceous diving beetles) Dytiscus marginicollis LeConte (Grinnell, 1908) undetermined species (Pierce, 1947a) Family Histeridae (hister beetles) (Pierce, 1947a) Saprinus species undetermined species Family Hydrophilidae (water scavenger beetles) undetermined species (Pierce, 1947a) Family Silphidae (carrion beetles) (Pierce, 1949a; Miller and Peck, 1979) Heterosilpha ramosa (Say) Nicrophorus guttula Motschoulsky = Nicrophorus

guttulus

punctostriatus

Pierce, 1949a Nicrophorus maginatus Fabricius = Nicrophorus guttulus labreae Pierce,

1949a = Nicrophorus = Nicrophorus

1949a

mckittricki Pierce, 1949a obtusiscutellum Pierce,

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Nicrophorus nigrita Mannerheim = Nicrophorus investigator alpha Pierce, 1949a Thanatophifus lapponicus (Herbst) Family Staphylinidae (rove beetles) undetermined species (Moore and Miller, 1978) Family Dermestidae (dermestid beetles) Dermestes species (Pierce, 1947a, 1949a) Family Tenebrionidae (darkling ground beetles) (Doyen and Miller, 1980) Apsena laticornis Casey (Essig. 193 1; Pierce, 1954b) = Apsena labreae Pierce, 1954b Apsena pubescens (LeConte) (Pierce, 1954b) Apsena rufipes (Eschscholtz) (Pierce, 1954b) Coniontis abdominalis LeConte (Grinnell, 1908; Pierce, 1954~) = Coniontis abdominalis caseyi Pierce, 1954c = Coniontis abdominalis fragmans Pierce, 1954~ = Coniontis abdominalis labreae Pierce, 1954~ = Coniontis blissi Pierce, 1954~ = Coniontis pectoralis paraelliptica Pierce, 1954~ = Coniontis pectoralis interrupta Pierce, 19.54~ = Coniontis tristis alpha Pierce, 1954~ = Coniontis tristis asphalti Pierce, 1954c = Coniontis tristis latigula Pierce, 1954c Coniontis elliptica Casey (Grinnell, 1908; Pierce, 1954~) Coniontis remnans Pierce, 1954c Coniontis lamentabilis Blaisdell (Pierce, 1954c) Eleodes acuticaudus LeConte (Grinnell, 1908; Essig, 1931) Eleodes laticollis LeConte (Grinnell. 1908; Essig, 1931) Eleodes grandicollis LeConte (Grinnell, 1908; Essig, 1931) = Eleodes elongatus Grinnell, 1908 Eleodes osculuns (LeConte) (Essig, 193 I)

= Eleodes behri Grinnell. 1908 = Eleodes intermedius Grinnell, 1908 Eleodes omissrrs LeConte (Essig, 1931) Nyctoporis carinata LeConte (Essig. 1931; Pierce, 1954a) Family Zopheridae Noserus plicatus LeConte (Doyen and Miller, 1980) = Noserus corrosus of Pierce, 1954a (misidentification) Phloeodes pustulosus LeConte (Pierce, 1954a; Doyen and Miller, 1980) Family Scarabaeidae (scarab beetles) (Miller et al., 1981). Canthon (Boreocanthon) simplex LeConte = Canthon simplex antiquus Pierce, 1946b Canthon (Boreocanthon) praticola LeConte = Canthon praticola vetustus Pierce, 1946b Copris pristinus Pierce, 1946b (Matthews and Halffter, 1968) Onthophagus everestae Pierce, 1946b Phanaeus labreae (Pierce, 1946b) = Palaeocopris labreae Pierce, 1946b Phileurus illatus LeConte (Saylor, 1948) Serica kanakoffi Pierce, 1946~ Family Elateridae (click beetles) (det. E. C. Becker) “Acolus species, partial prothorax RLP 898E. Pit 91, GJM 575/K-5 *Anchastus cinercipennis (Eschscholtz), prothorax RLP 1253E. Pit 91, GJM 7871 L-5 “Aplastus species, prothorax RLP 7506E. Pit A *Cardiophorus species, partial prothorax RLP 3957E, Pit 91, GLM 408/L-4; 4 partial prothoraces, 1 prosternum RLP 3316E, Pit 91, GJM 295/M-3 *Dalopius species 1, pronotum RLP 4043E, Pit 91, GJM 295/M-3 *Dalopius species 2, prothorax RLP 3408E, Pit 91, GJM 355/K-5; prothorax RLP 4078. Pit 91, GJM 914/F-7 “Dalopius species 3. prothorax RLP 40378, Pit 91, GJM 914/F-7

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species, prothorax RLP 7507E, Pit A “Melanotus species, partial prothorax RLP 2218E, Pit 91, GJM 874/I-6 *Melanotus (?) species, prothorax RLP 7508E, Pit A *undetermined species, left elytron RLP 3243E, GJM 273/N-3 Family Coccinellidae (ladybird beetles) *Coccinella californica Mannerheim, RLP 4048E ( = C 109a) from Pit 16 dump; probably this species, head and pronotum RLP 6965E ( = C 164a), from Pit A (det. R. D. Gordon) Family Anthicidae (antlike flower beetles) “Notoxus s,pecies in sparsus species group, two pronota RLP 3281E, Pit 91, GJM 295/M-3; RLP 391 lE, Pit 91, GJM 401/L-3 (met. D. Chandler) Family Cerambycidae (long-horned beetles) undeterminetd species (Pierce, 1947a) Family Curculionidae (weevils) *Apion species, elytron RLP 4044E from Smilodon skull LACM 2001-546, Pit 3, C-4, 2 m (6.5 ft); elytron RLP 4045E from Pit 101, C-5, 1.4 m (4.5 ft); 2 elytra RLP 4046E-4047E from Pit 101, E-5, 1.4 m (4.5 ft) (det. D. G. Kissinger) undetermined species (Pierce, 1947a) Family Scolytidae (bark beetles) *Gnathotrichus species, probably G. retussus (LeConte), maybe G. sulcatus (LeConte), prothorax RLP 4049E from Smilodon skull LACM 2001-546, Pit 3, C-4, 2 m (6.5 ft) *Limonius

Order Diptera

(flies) Family Bibionidae (march flies) *Dilophus species, female head and thorax RLP 3420E, Pit 91, GJM 355/K5 (det. G.. W. Byers) Family Calliphoridae (blow flies) Cochliomyia macellaria (Fabricius) (Gagne and Miller, 1982) = Protochrysomyia howardae Pierce, 1945b

99

IN CALIFORNIA

Order Hymenoptera

(ants, wasps, and bees) Family Ichneumonidae (ichneumon wasps) *undetermined species, 2 heads RLP 4072E, Pit 91, GJM 914/F-7 (det. A. S. Menke and R. W. Carlson) Family Formicidae (ants) Veromessor andrei Mayr (Pierce, 1954b) undetermined species Family Vespidae (vespid wasps) undetermined species (Pierce, 1947a) CLASS

DIPLOPODA

(millipedes) Order Julida

Family Julidae undetermined

species (Pierce

1947a)

Order Spirobolida

Family Spirobolidae “Spirobolus”

australis Grinnell,

1908

CLASS ARACHNIDA

(arachnids) Order Araneida

undetermined

(spiders) species (Pierce,

1947a)

Order Phalangida

undetermined

(harvestmen) species (Pierce, 1947a)

APPENDIX 2: INSECTS RECORDED FROM McKlTTRlCK

This list contains all previously published records for insects, including a few reidentifications (denoted by *; see Appendix 3 for details). Important citations are noted. Order Odonata

(dragonflies and damselflies) Suborder Anisoptera (dragonflies) undetermined species (Essig, 193 1; Pierce, 1947b) Suborder Zygoptera (damselflies) undetermined species (including nymphs) (Pierce, 1947b)

SCOT-I' E. MILLER

100 Order Orthoptera

(grasshoppers and relatives) Family Acrididae (grasshoppers) undetermined species (Pierce, 1947b) Family Stenopelmatidae (Jerusalem crickets) Stenopelmatus species (Pierce, 1947b) Order Hemiptera

(true bugs) Family Corixidae (water boatmen) undetermined species (Pierce, 1947b) Family Notonectidae (backswimmers) *Notonecta (Paranecta) unifasciata Guerin-Meneville = Notonecta (Paranecta) badia Pierce, 1948a Family Nepidae (waterscorpions) *Ranatra bessomi Pierce, 1948a *Ranatra asphalti Pierce, 1948a Family Belostomatidae (giant water bugs) “Belostoma bakeri Montandon = Belostoma species (Pierce, 1947b) *Lethocerus americanus (Leidy) = Lethocerus species (Pierce, 1947b) Family Gerridae (water striders) Gerris species cf. G. incognitus Drake and Hottes or G. gillettei Lethierry and Severin (Pierce, 1947b; Calabrese, 1978) Order Coleoptera

Colymbetrs species (Pierce. 1947b) Agabus species (Pierce, 1947b) Family Hydrophilidae (water scavenger beetles) Hydrophilus triungularis Say (Essig. I93 1) Hydrophilus species (Essig, 193 I ; Pierce, 1947b) Tropisternus species (Pierce, 1947b) Family Silphidae (carrion beetles) (Pierce, 1949a; Miller and Peck, 1979) Nicrophorus guttula Motschoulsky Nicrophorus marginatus Fabricius = Nicrophorus mckittricki Pierce, 1949a Family Staphylinidae (rove beetles) Philonthus species (2 species) (Moore and Miller, 1978) Family Buprestidae (metallic wood-boring beetles) (Pierce. 1947b) *Acmaeodera nigrovittata Van Dyke Family Heteroceridae (variegated mudloving beetles) Heterocerus species (Pierce, 1947b; Morgan and Morgan, 1980a) Family Tenebrionidae (darkling ground beetles) (Doyen and Miller, 1980) Eleodes species (several species) Stenomorpha mckittricki (Pierce, 1954a) = Parasida mckittricki Pierce, 1954a Family Scarabaeidae (scarab beetles) undetermined species (Pierce. 1947b) Order Lepidoptera

(beetles) (butterflies and moths) Family Cicindelidae (tiger beetles) undetermined species (Pierce, 1947b) Cicindela haemorrhagica LeConte (Nagano et al., 1983) = Cicindela species (Pierce, 1947b) Order Diptera Family Carabidae (ground beetles) (flies) *Elaphrus finitimus Casey undetermined species (Pierce, 1947b) = Elaphrus ruscarius foveatus Pierce, 1948b undetermined species Order Hymenoptera Family Haliplidae (crawling water beetles) (ants, wasps, and bees) Haliplus species (Pierce, 1947b) Family Chrysididae (cuckoo wasps) Family Dytiscidae (predaceous diving beeundetermined species (Pierce, 1947b) tles) Cybister expfanatus LeConte (Essig, Family Vespidae (vespid wasps) undetermined species (Pierce, 1947b) 193 1; Pierce, 1947b)

FOSSIL

INSECTS

APPENDIX 3: NEW IDENTIFICATIONS AND SYNONYMIES OF McKlTTRlCK INSECTS HEMIPTERA:

NOTONECTIDAE

Notonecta (Paranecta) unifasciata Guerin-Meneville Notonecta (Paranecta) badia Pierce 1948a, p. 24-25, Figs. 4-9, N. SYN. The almost complete holotype (LACMIP 2527 = McK 7a) and the paratypes (LACMIP 25282562, 5661-5687) of Notonecta badia, all from site 4 depth 1.2 m (4 ft) at McKittrick, are Notonecta unifasciata (F. S. Truxal, personal communication, 1982). N. unifasciata is widespread in the western United States, western Canada, and Mexico and exhibits a wide range of tolerance to its aquatic medium (Truxal, 1979). HEMIPTERA:

NEPIDAE

Ranatra asphalti Pierce

IN

101

CALIFORNIA

communication, 1980). The paratypes (LACMIP 2566 = McK llf, 2567 = McK lle, 2568 = McK lld, 2569 = McK lib, and 2570 = McK 1 la; Sphon (1973) assigned two numbers (2570 and 2571) to paratype 2570) lack diagnostic characters necessary for identification. The comments above under R. asphalti apply here also; positive identification must await revision of Ranatra. HEMIPTERA:

BELOSTOMATIDAE

Belostoma bakeri Montandon

Three individuals (1 head, 1 head and pronotum, and 1 disarticulated head, thorax, and abdomen) from site 4, depth 1.2 m (4 ft) (LACMIP 6447 = McK 3 1) are apparently this species, although the pronota are suggestive of Belostoma JZumineum Say (A. S. Menke, personal communication, 1980). An additional almost complete, but broken, specimen (LACMIP 6446) from site 4, depth unknown, is probably B. bakeri.

Ranatra asphalti Pierce 1948a, p. 29-30, Figs. 13-15 and 19. The holotype of RanLethocerus americanus (Leidy) atra asphalti, which consists of head, Specimen LACMIP 6448 (= McK 35) is thorax, anterior abdomen, and basal left an almost complete female of this species middle femur from site 4, depth 1.2 m (4 ft) from site 4, depth 1.1 m (3.5 ft) (A. S. (LACMIP 2564 = McK 12a), is probaMenke, personal communication, 1980). bly Ranatra brevicollis Montandon (A. S. Menke, personal communication, 1980). COLEOPTERA: CARABIDAE However, the holotype is not complete Elaphrus finitimus Casey enough to positively assign it to R. brevicollis. The ranges of R. brevicollis and RanElaphrus ruscarius foveatus Pierce 1948b, atra fusca Palisot de Beauvois presently p. 53-55, Figs. 1-3, N. SYN. The holotype overlap in the California central valley and of Eluphrus ruscarius fovea&s (LACMIP intergrades occur which are not presently 2949 = McK 13a), a left elytron horn site 4, understood (Menke, 1979a); positive iden- depth 1.2 m (4 ft) at McKittrick is Elaphrus tification of the McKittrick Ranatra must finitimus (H. Goulet, personal communiawait needed revision of the genus. cation, 1979). The species is not known to occur presently in the southern Central Ranatra bessomi Pierce Valley of California (Goulet and Baum, Ranatra bevsomi Pierce 1948a, p. 27-29, 1982). Figs. 11, 12, and 16-18. The male holotype of Ranatra bessomi, which consists of COLEOPTERA: BUPRESTIDAE thorax, broken abdomen, and basal front Acmaeodera nigrovittata Van Dyke femora from site 4, depth 1.2 m (4 ft) This species is represented by three (LACMIP 2565 = McK llc), is probably McKittrick specimens: 1 head, thorax, and Ranatra f14sca (A. S. Menke, personal

102

ScoT’r E. MII,LEK

anterior abdomen with anterior elytra from site 4, depth 1.2 m (4 ft) (LACMIP 6445): 1 thorax and anterior abdomen with anterior elytra from site 4, depth 1.2 m (4 ft) (LACMIP 6444): and 1 abdomen with elytra from matrix collected for radiocarbon dating QC-995 (equivalent to site 4. depth 1.2) (R. L. Westcott, personal communication, 1981). ACKNOWLEDGMENTS The Santa Barbara Museum of Natural History (especially R. S. Gray and F. G. Hochberg). the Natural History Museum of Los Angeles County (W. A. Akersten, C. L. Hogue, G. T. Jefferson. P. G. Owen. R. L. Reynolds, C. A. Shaw, and E. C. Wilson). Queens College, CUNY (L. F. Marcus and R. R. Pardi), the Smithsonian Institution. and a University of California. Santa Barbara. President’s Research Fellowship supported this research. My studies were made possible by the generous help of many taxonomists including E. C. Becker, D. E. Bright. G. W. Byers. D. M. Calabrese. R. C. Froeschner. H. Goulet. A. R. Hardy, R. L. Hoffman, D. H. Kavanaugh, D. G. Kissinger, H. B. Leech, I. Moore. C. D. Nagano, T. N. Seeno. F. S. Truxal. R. L. Westcott. and especially J. T. Doyen, R. J. Gagne. R. D. Gordon. H. F. Howden. A. S. Menke, and S. B. Peck. For providing information and assistance, I also thank A. C. Ashworth. D. I. Axelrod. H. A. Bronstein. F. M. Carpenter, K. R. Cheesman, C. R. Churcher. C. R. Harington, F. G. Hochberg. D. L. Johnson. 0. Kukul. R. J. McGinley. H. M. Miller. P. M. Miller. A. V. Morgan. E. Nesbit, J. H. Peck, D. Pierce. D. L. Schulte. A. W. Skalski. D. W. Steadman. J. K. Warter. D. P. Whistler, E. Wing. R. F. Yopp, and V. V. Zherichin. L. Meszoly drew the map. 1 am especially indebted to W. A. Akersten. C. L. Hogue, L. F. Marcus. and E. C. Wilson for their continued support and encouragement of my brea studies.

REFERENCES Ashworth, A. C. (1979). Quatemary Coleoptera studies in North America: Past and present. In “Carabid Beetles: Their Evolution, Natural History. and Classification.” (T. L. Erwin. G. E. Ball and D. R. Whitehead. Eds.). pp. 395-406. Dr. W. Junk B.V.. The Hague. Axelrod. D. I. (1967). Quaternary extinctions of large mammals. Uni\aersity of Ca/$ornia Publications it2 Geological Science 74, l-42. Berger. R., and Libby. W. F. (1966). UCLA radiocarbon dates V. Radiocarbon 8, 467-497. Blair, K. G. (1927). Insect remains from oil sand in Trinidad. Transactions of the Entomological Society ofLondon 75, 137-141.

Blaisdell. F. E.. Sr. (1909). A monographic revision of the Coleoptera belonging to the tenebrionid tribe Eleodiini inhabiting the United State\. Lower California, and adjacent islands. fhtlieri~7 C$I/IC United Slairs Nationcrt Muserrm 63, I-524. Bogachev. A. (1948). Fauna of the Binagady asphalt deposits: Coleoptera. Trady Esfes\senno-Istoricheskii Muzei, Buklc 1-2, 137-160. [In Russian] Borell. A. E. (1936). A modern La Brea tar pit. Auk 53, 298-300. Calabrese, D. M. (1978). Identity of Gerri.c (Hemiptera: Heteroptera) species from Pleistocene asphalt deposits at McKittrick, California. Entomolo~icul News 89, 59-60. Campbell. K. E.. Jr. (1979). The non-passerine Pleistocene avifauna of the Talara tar seeps. Northwestern Peru. Royal Onturio Mfrserrm, Life Scicnws Contributions 118. l-203. Churcher. C. S. (1966). The insect fauna from the Talara tar-seeps, Peru. Canadian Journal qf Zoology 44. 985-993 Coope. G. R. (1978). Constancy of insect species versus inconstancy of Quaternary environments. Symposia of‘ the Royal Entomologit ul Society of‘ London 9, 176-187. DeMay. I. S. (1941). Quaternary bird life of the McKittrick asphalt, California. Carne,qie In.stitation of Washington Publication 530, 35-60. Dodson. P., and Wexlar. D. (1979). Taphonomic investigations of owl pellets. Puleobiology 5, 275-284. Doyen, J. T.. and Miller. S. E. (1980). Review of Pleistocene darkling ground beetles of the California asphalt deposits (Coleoptera: Tenebrionidae, Zopheridae). Pan-Paci’c Eniomologist 56, I-IO. Essig. E. 0. (1931). “A History of Entomology.” MacMillan, New York. Gagne. R. J., and Miller. S. E. (1982). PrrjtochtT.somyia houurdae from Ranch0 La Brea. California, Pleistocene. new junior synonym of Cochliomyiu muce~luria (Diptera: Calliphoridae). Bulletin of the Southern Calijiornia Ac,ademy of Sciences 80, 95-96 Goulet. H., and Baum. B. R. (1982). Analysis of variation in the Elaphrus finitimas complex of North America (Coleoptera: Carabidae). Canadian Joarm~l of Zoology 60, 2424-2433. Grinnell. E (1908). Quaternary myriapods and insects of California. Urksersity i7f Calijknitr Pah/ication.c in Geology 5, 207-2 15. Hanna. G. D. (1924). Insects in the California tar traps. Science 59. 5.55. Harington, C. R. (1980). Pleistocene mammals from Lost Chicken Creek. Alaska. Canadian Journal of Earth Sciei7(.es 17, 168-198. Howard, H. (1962). A comparison of avian assemblages from individuals pits at Ranch0 La Brea, California. Natural History Museum of Los Angeles County Contributions in Science 58, l-24.

FOSSIL INSECTS Hubbs, C. L., and Walker, B. W. (1947). Abundance of desert mammals indicated by capture in fresh road tar. Ecology 28., 464-466. Johnson, D. L. (1!)77). The California ice-age refugium and the Rancholabrean extinction problem. Qltuternury Research 8, 149-153. Keeton, W. T. (1960). A taxonomic study of the milliped family Spirobolidae (Diplopoda: Spirobolida). Memoirs of the American Entomological Society 17, 1-146. Kennedy, C. H. (1917). Notes on the life history and ecology of the dragonflies (Odonata) of Central California and Nevada. Proceedings of the United States National Museum 52, 483-635. Kenward, H. K. (1978). The analysis of archaeological insect assemblages: A new approach. The Archaeology of York 119, l-68. Kurt&. B., and Anderson, E. (1980). “Pleistocene Mammals of North America.” Columbia Univ. Press, New York. Lance, J. F. (1946). Fossil arthropods of California. 9. Evidence of termites in the Pleistocene asphalt of Carpinteria, California. Bulletin of the Southern California Academy of Sciences 45, 21-27. Lauck, D. R., and Menke, A. S. (1961). The higher classification of the Belostomatidae (Hemiptera). Annals sf the Entomological Society of America 54, 644-657. Linsley, E. G. (1943). Attraction of Melnnophila beetles by fire and smoke. Journul of Economic Entomology 36, 341-342. Maloney, N. J., Warter, J. K.. and Akersten, W. A. (1974). Probable origin of the fossil deposits in Pit 91. Ranch0 La\ Brea tar pits, California. Geological Society of America, Abstracts with Programs 6, 212. Marcus. L. F., and Berger, R. (1983). The significance of radiocarbon dates for Ranch0 La Brea. In “Quaternary Extinctions: A Prehistoric Revolution.” (P. S. Martin and R. G. Klein, Eds.), in press. Univ. of Arizona Press., Tucson. Martin, L. D., and Neuner, A. M. (1978). The end of the Pleistocense in North America. Transactions of the Nebraska Academy of Sciences 6, 117-126. Matthews, E. G., and Halffter, G. (1968). New data on American Corpis with discussion of a fossil species (Coleopt., Scarab.). Ciencia (Mexico City) 26, 147-162. Menke, A. S., (1963). A review of the genus Lethocerus in North and Central America, including the West Indies. Annuls qf the Entomological Society of America 56, 261-267. Menke. A. S. (1979a). Family Nepidae. In “The Semiaquatic and Aquatic Hemiptera of California (Heteroptera: Helmiptera)” (A. S. Menke, Ed.). California Insect Survey Bullrrin 21, 70-75. Menke, A. S. (1979b). Family Belostomatidae. In “The Semiaquatic
IN CALIFORNIA

103

Miller, L. (1930). Dragon fly psychology. Pomona College Journal of Entomology and Zoology 22, 45-46. Miller, S. E. (1978). A fossil of Scaphinotus interruptus from the Pleistocene Carpinteria asphalt deposit, Santa Barbara County, California (Coleoptera. Carabidae). Pun-Pacific Entomologist 54, 74-75. Miller, S. E. (1979). Reevaluation of Pyrus hoffmannii (Rosaceae) from the Pleistocene Carpinteria asphalt deposit, California. Madrono 26, 190-191. Miller, S. E. (1982). Quaternary insects of the California asphalt deposits. Third North Americun Paleontological Convention, Proceedings 2, 377-380. Miller, S. E.. Gordon, R. D.. and Howden. H. F. (1981). Reevaluation of Pleistocene scarab beetles from Ranch0 La Brea, California (Coleoptera: Scarabaeidae). Proceedings of the Entomological Society of Washington 83, 625-630. Miller, S. E., and Peck, S. B. (1979). Fossil carrion beetles of Pleistocene California asphalt deposits, with a synopsis of Holocene California Silphidae (Insecta: Coleoptera: Silphidae). Transactions of the San Diego Society of Natural History 19, 85-106. Moore, I., and Miller, S. E. (1978). Fossil rove beetles from Pleistocene California asphalt deposits (Coleoptera: Staphyiinidae). Coleopterists’ Bulletin 32, 37-39. Morgan, A. V., and Morgan, A. (1980a). Beetle bitsThe science of paleoentomology. Geoscience Canada 7, 22-29. Morgan, A. V., and Morgan, A. (1980b). Fauna1 assemblages and distributional shifts of Coleoptera during the late Pleistocene in Canada and the northern United States. Cunadian Entomologist 112, 1105-l 128. Nagano, C. D., Miller, S. E., and Morgan, A. V. (1983) Fossil tiger beetles (Coleoptera: Cicindelidae): review and new Quaternary records. Psyche 89, 339-346. Pierce, W. D. (1945a). Fossil arthropods of California. 1. Introductory statement. Bulletin of the Southern California Academy of Sciences 43, 1-3. Pierce. W. D. (1945b). Fossil arthropods of California. 8. Pleistocene myiasis from the La Brea pits. Bu/letin of the Southern Colifornin Academy of Sciences 44, 8-9. Pierce, W. D. (1946a). Fossil arthropods of California. 10. Exploring the minute world of the California asphalt deposits. Bullefin of the Southern C’alifornitr Academy of Sciences 45, 113-I 18. Pierce, W. D. (1946b). Fossil arthropods of California. I I. Descriptions of the dung beetles (Scarabaeidae) of the tar pits. Bulletin of the Soufhern Califr,rniu Academy ofsciences 45, 119-131. Pierce, W. D. (1946~). Fossil arthropods of California. 12. Description of a sericine beetle from the tar pits. Bulletin of the Southern California Academy of Sciences 45, 131-132. Pierce, W. D. (1947a). Fossil arthropods of California.

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13. A progress report on the Ranch0 La Brea asphaltum studies. Bullefin ofrhe Southern Cal~%rnict Academy of Sciences 46, 136-138. Pierce, W. D. (3947b). Fossil arthropods of California. 14. A progress report on the McKittrick asphalt field. Bulletin

qf the Southern

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ences 46, 138-143. Pierce, W. D. (194Sa). Fossil arthropods of California. IS. Some Hemiptera from the McKittrick asphalt field. Bulletin of the Souther-n California Acudeq of Sciences 47, ‘l-33. Pierce. W. D. (1948b). Fossil arthropods of California. 16. The carabid genus Elaphtxs in the asphalt deposits. Bulletin of the Southern California Acadelny of Sciences

47, 53-54.

Pierce, W. D. (1949a). Fossil arthropods of California. 17. The silphid burying beetles in the asphalt deposits. Bulletin of the Southern California Academy of Sciences

48, 55-70.

Pierce, W. D. (1949b). A modern asphalt seep tells a story. Los Angeles County Museum Quarterly 7(3). 11-17. Pierce, W. D. (1954a). Fossil arthropods of California. 18. The Tenebrionidae-Tentyriinae of the asphalt deposits. Bulletin of the Southern California Academy of Sciences

53, 35-45.

Pierce, W. D. (1954b). Fossil arthropods of California. 19. The Tenebrionidae-Scaurinae of the asphalt deposits. Bulletin of the Southern California Academy of Sciences

53, 93-98.

Pierce, W. D. (1954~). Fossil arthropods of California. 20. The Tenebrionidae-Coniontinae of the asphalt deposits. Bulletin of the Southern California Academy of Sciences 53, 142-156. Pierce. W. D. (1957). Insects. GeoloXicul Society of America

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67, 943-952.

Saylor. L. W. t 1948). Synoptic revision of the United States scarab beetles of the subfamily Dynastinae. 4: Tribes Oryctini (part). Dynastini. and Phileurmi. Journrcl

qf‘rhe

Washington

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of Scicrrc,cr

38.

176-183. Schultz. J. R. (1938). A Late Quaternary mammal fauna from the tar seeps of McKittrick. California. CNTnegie Institution of Washington Puhlic~nrbm 487, I I I215. Shaw. C. A. (1982). Techniques used in excavation. preparation. and curation of fossils from Ranch0 La Brea. Curator 25, 63-77. Sphon, G. G. (1973). Additional type specimen5 of fossil Invertebrata in the collections of the Natural History Museum of Los Angeles County. Natural Hi.rtar? Museum of Los Angeles in Science 250, l-75.

County

Contributions

Sternberg, C. H. (1932). “Hunting Dinosaurs on Red Deer River. Alberta, Canada.“ C. H. Stemberg, San Diego. Stock. C. (1956). Ranch0 La Brea: A record of Pleistocene life in California. Natural History Mu~eom of Los Angeles County Science Bulletin 20, l-81, Truxal. F. S. (1979). Family Notonectidae. in “The Semiaquatic and Aquatic Hemiptera of California (Heteroptera: Hemiptera)” (A. S. Menke, Ed.). Ca&rnia Insect Surl,ey Bulletin 21, 139-147. Vander Hoof. V. L. (1934). Seasonal banding in an asphalt deposit at McKittrick. Proceedings of the Geological Society of America. p. 332. Warter. J. K. (1976). Late Pleistocene plant communities-Evidence from the Ranch0 La Brea tar pits. In “Plant Communities for Southern Caiifornia” tJ. Latting, Ed.), pp. 32-39. California Native Plant Society Special Publication No. 2, Berkeley. Woodard, G. D., and Marcus, L. F. ( 1973). Ranch0 La Brea fossil deposits: A re-evaluation from stratigraphic and geological evidence. Journrrl r>fPa/eon-

Pierce, W. D. (1961). The growing importance of paleoentomology. Proceedings of the Entomological Society of Washington 63, 21 l-217. tolo,qy 47, 54-69. Saylor. L. W. (19331. Attraction of beetles to tar. Pcrn- Usinger. R. L. (1956). “Aquatic insects of CaliPacific Entomologist 9, 182. fornia.” Univ. of California Press. Berkeley