Quaternary Proceedings No. 5,1997 185-191 
? Quaternary Research Association, London 
Late Quaternary Insects of Rancho La Brea^ 
California^ LISA 
Scott E. Miller 
Scott E. Miller, 1997. Late Late Quaternary insects of Rancho La Brea, California, USA. In Studies in Quaternary Entomology - An Inordinate Fondness for Insects. 
Quaternary Proceedings No. 5, John Wiley & Sons Ltd., Chichester, pp. 185-191. 
Abstract 
Asphalt-impregnated sediments at Rancho La Brea, Los Angeles County, California, provide a rich 
Quaternary insect record. Ages of various sites at Rancho La Brea range from more than 40,000 "C 
yr B.P. to modern. The major palaeoecological insect groups are: (1) ground dwellers, (2) aquatics, 
(3) scavengers, and (4) miscellaneous. Only two apparent terminal Pleistocene extinctions are 
recognized, both dung beetles (Cole?ptera: Scarabaeidae). 
KEYWORDS: Cole?ptera, Quaternary, asphalt, Rancho La Brea, California, Extinction, Palaeoecology. 
Scott E. Miller, Department of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, 
Hawaii 96817, USA & Research Associate in Entomology, Natural History Museum of Los Angeles 
County, USA. 
Quaternary 
Proceedings 
WILEY 
Publisbas Since 1807 
Appreciation 
In the late 1970s, I noticed that while studies by Russell 
Coope and his students in Europe and North America 
showed no recognizable extinction of insects at the end of 
the Pleistocene, many supposedly extinct insect species 
had been described from the asphalt deposits at Rancho La 
Brea. The combination of Professor Coope's 
reinterpretation of the temporal stability of insect species 
with a revolution in understanding of the taphonomy of the 
asphalt deposits catalyzed by Leslie Marcus, stimulated my 
interest in Rancho La Brea insects. Therefore, I am very 
pleased to contribute this review of Rancho La Brea insects 
to this volume honouring Professor Coope. 
Introduction 
The Rancho La Brea asphalt deposits are among the world's 
most renowned Late Pleistocene vertebrate localities (Stock 
& Harris 1992), but the associated abundant invertebrate 
and plant fossils have received little attention. Most of the 
early entomological papers contained significant errors in 
identifications and interpretations. Preliminary studies were 
made on limited samples from the early excavations by the 
University of California Museum of Palaeontology (Grinnell 
1908; Blaisdell 1909; Essig 1931). W.D. Pierce published a 
series of papers from 1946 to 1957 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 palaeoecological indicators, 
but his publications and taxonomic procedures were 
replete with errors (see Miller 1983). Rancho La Brea 
insects were reviewed by Miller (1983), which culminated 
a series of papers evaluating the previously described taxa 
(Moore & Miller 1978; Miller & Peck 1979; Doyen & Miller 
1980; Miller ef a/. 1981; Gagne & Miller 1982; Miller 
1983; Nagano ef al. 1983). The present paper reviews this 
body of knowledge again and suggests directions for future 
research. 
The popular conception of fossil accumulation at 
Rancho 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. However, recent studies (see Marcus 
& Berger 1983; Stock & Harris 1992) 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. The "pits" at Rancho 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 concentration of bones 
in stream channels or ponds. Once buried, the fossils were 
impregnated by asphalt permeating upward and laterally. 
Over eighty species of insects in over thirty-one families 
have been recorded from Rancho La Brea, along with 
largely unidentified millipedes, centipedes, spiders, 
harvestmen, and isopods (Miller 1983; Larew 1987; Stock 
& Harrris 1992). Many more families and species will be 
recognized as material already excavated is analysed in 
detail. Almost all Rancho La Brea insect specimens are 
cpnsidered conspecific with Holocene species and fall 
within reasonable ranges of geographic variation. While 
studies by other workers indicate that almost all Pleistocene 
insect fossils represent extant species (Elias 1994; Coope 
186 QUATERNARY PROCEEDINGS 
1995), two Rancho La Brea species appear to be extinct. 
Localities 
The Rancho La Brea asphalt deposits are located in 
Hancock Park, Los Angeles, Los Angeles County, California 
(Fig. 1). More than one hundred individual excavations or 
"pits" have been made since 1905. Most of these were 
unproductive test holes and fewer than fifteen are 
considered major sources of vertebrate fossils. The most 
important site for insect fossils is Locality (Pit) 91 (=LACM 
Vertebrate Paleontology Locality 6909). No insects were 
available from the original partial excavation of Locality 91 
in 1915, but the renewed excavation starting in 1969 and 
continuing to the present (Shaw 1982), has yielded a vast 
assemblage of insects. Before the reopening of Locality 91, 
emphasis was placed on large vertebrates; insects and other 
small fossils were neglected by early excavators. Pierce's 
specimens were largely salvaged from the miscellaneous 
material associated with the vertebrate collections, 
including matrix inside the skull cavities of large 
vertebrates. 
Almost all the insect fossils from Rancho La Brea are 
housed at the George C. Page Museum, a branch of the 
Natural History Museum of Los Angeles County (LACM). 
An estimated 100,000 insect specimens have been sorted 
and catalogued in some 13,500 catalogue numbers at 
LACM. Of these, over 9,000 catalogue numbers and the 
bulk of the individual specimens are from Locality 91. At 
least another 100,000 insect specimens, predominantly 
from Locality 91, remain uncatalogued (C.A. Shaw, 
personal communication). Only a small percentage of the 
Locality 91 specimens have received detailed analysis and 
identification. The University of California Museum of 
Paleontology (UCMP) houses a small amount of Rancho La 
Brea insect material, most of which represents vouchers for 
the earliest studies by Grinnell (1908), Blaisdell (1909), and 
Essig (1931). 
Pierce (1946a; 1947) mentioned the occurrence of 
insects from the following Localities (Pits) at Rancho La 
Brea: A, B, 2, 3, 4, 9, 10, 13, 16, 28, 29, 36, 37, 51, 60, 
61, 67, n, 81, 101, "Academy", "Bliss 29" and "Pit X". 
Original matrix was available to Pierce only from Localities 
A, 81, 101, and presumably "Bliss 29" and "Pit X". Insects 
from Localities 3, 4, 13, 60, 61, 67, and 77 were recovered 
from the brain cavities of Smilodon skulls. Although 
material from such skulls is probably contemporaneous 
with the skulls, it may not be (e.g. Harington 1980, 197). 
Miller (1983) described the insect fossil localities within 
Rancho La Brea in greater detail (see also Marcus & Berger 
1983; Stock & Harris 1992). It is worth noting that the age 
of some of Pierce's material is questionable, especially his 
"Bliss 29" and "Pit X". 
Palaeoecology 
Interpretation of accumulation and preservation in asphaltic 
matrix is difficult, and much remains to be learned about 
insect palaeoecology in general (Elias 1994). 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 latter appears to have 
been more common at Rancho La Brea. Viscous asphalt 
flows or pools can trap animals by (1) appearing as water, 
(2) having sticky, melted surfaces at high temperatures 
(solid at lower temperatures), and/or (3) being camouflaged 
with a covering of dirt, leaves or debris. 
Insects can be attracted to asphaltic sediments in the 
following three ways, but they can also simply be 
accidentally trapped, without attraction, by crawling or 
falling into asphalt and not being able to free themselves. 
(1 ) Attraction to carrion or other material already trapped or 
otherwise in contact with the asphalt, 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 1946b, fig. 6; 1957; 
Kennedy 1917). Many aquatic insects locate water visually 
and apf)ear to have poor sense of smell. "For insects that 
see in the ultraviolet, oil surfaces closely mimic the 
polarization and reflectivity characteristics of water" 
(Hor?th & Zeil 1996). Some water beetles will "land on the 
shiny tops of cars, mistaking them for water" (Leech & 
Chandler 1956, 309), and even water itself is an effective 
trap for many terrestrial insects (Kenward 1978, 40). (3) 
Attraction to warmth or perhaps chemicals, versus visual 
attraction to the asphalt itself. Some insects appear to be 
attracted to the fresh tar coating on roads (Saylor 1933; 
Borell 1936; Hubbs & Walker 1947), although this 
attraction may result from the warmth of the roads because 
some beetles (e.g., Melanophila spp.) have infrared sense 
organs (Evans 1966). In practice, taphonomy is a product of 
all these processes, each of which occur 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 (Miller & Peck 1979, 101), where 
modern insects and debris appear to be incorporated into 
Pleistocene matrix by accumulating in the bottoms of 
natural cracks and rodent burrows which penetrate into the 
matrix. 
La 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) ground-dwelling beetles and other 
Figure 1. Map showing the location of Rancho La Brea (1) and 
other fossil-bearing asphalt deposits in California: 
McKittrick (2), Carpinteria (3), and Maricopa (4) 
(drawing by L. Meszoly). 
LATE QUATERNARY INSECTS OF RANCHO LA BREA 187 
Table 1.      Beetle families recorded from asphalt deposits around 
the world. See text for r?f?rences. 
UM..U Tilwi ITOM. ?'Mt-r 
*?. UlltonU tallhrtla Nn TrMMM toKuat 
bMM? 
OnbWM X X X X X X 
Hil?M?* X X t 
Oit?cidH X X X X X X 
Grnnnlai X X 
Hi<U'<4i? X T X 
Hy*o?Md?? X X X X X X 
X X X X 
S4>MM X X X X 
Sc>r*t?iKlia X X X X X X 
eup'iKtidit X X X 
(Ul*'i44? X X 
H?t*loc?litf?? X 
0?nn?ilidlt X 7 X 
Cl?r<4>* X 
1trtbnon*u X X X X X X 
ZoptitndM X 
CMCiuilid? X X 
Amtiicidio X 
C??imbyc4ii X t x 
X X. 
ScctytKlM X 
CxciOonHlutil) X x X X 
terreslrlal crawling species, (4) miscellaneous stray 
individuals of herbivorous and other species which are not 
preserved in significant numbers, and (5) fragments of 
insects from faeces or gut contents of vertebrates. Members 
of the last category are minor and cannot now be 
distinguished from the others, but careful studies of 
vertebrate feeding habits {e.g. Dodson & Wexler 1979) 
should eventually allow differentiation. 
Detailed understanding of La Brea insects has been held 
back by the following problems: (1) Conclusions of early 
authors clouded understanding of ecological and extinction 
patterns. These conclusions have been largely reevalualcd 
{e.g. Miller 190?), providing a new foundation for research. 
(2) More data needs to be accumulated from sorting and 
identification of fossils, esfXicially from Locality 91. This is 
hampered by the tremendous diversity of the modern 
California fauna, as well as the paucity of identification aids 
in the literature (e.g., keys and illustrations). The modern 
California insect fauna includes over 30,000 species, many 
of which are poorly understood or characterized (Powell & 
Hogue 1979). Studies of Quaternary insects of northern 
North America have prospered in recent years in large part 
because the modem fauna is less diverse and better known. 
In contrast to works like Lindroth (1961-1969), no mo<lern 
review exists for Carabidae or Tenebrionidae of California. 
Over 90 years have passed since the last general review of 
southern California beetles (Fall 1901), and it contained no 
keys or illustrations! (3) More data needs to t)e accumulated 
on the ecology of living insects. Progress is being made, 
however, especially in areas such as aquatic insects (e.g. 
Merrill & Cummins 1996) and forensic entomology (Smith 
1986). 
Palaeoecological conclusions that can be drawn from 
Rancho La Brea insects are limited at present, but further 
study of the modern and fossil faunas promises to yield 
significant contributions to palaeoecoiogy. La 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 (e.g. FALINMAP 1996). 
Insects are also known from other Quaternary asphalt 
deposits in California (Fig. 1): Carpinteria, Santa Barbara 
County (Miller 1978; Moore & Miller 1978; Miller & Peck 
1979; Doyen & Miller 1980; Miller, unpublished data); 
Maricopa, Kern County (Doyen & Miller 1980); and 
McKitlrIck, Kern County (Miller 1983). Similar deposits also 
occur near Talara, Peru (Churcher 1966), Fyzabad, Trinidad 
(Blair 1927), and Binagady, in the Caucasus Region 
(Bogachev 1948; Burchak-Abramovich & Dzhafarov 1955; 
Vereschagin 1967). Although the faunas of most of these 
remain incompletely known, comparison of the beetle 
families recorded from them shows distinct similarity (Table 
1). The major paleoecological groups are: (1) ground 
dwellers (Carabidae, Tenebrionidae); (2) aquatics 
(especially Dytiscidae, Hydrophilidae, with other families); 
(3) scavengers (especially Scarabaeidae), and (4) 
miscellaneous families, A detailed analysis of these deposits 
is beyond the scope of the present paper, but there are 
certainly similarities between these deposits in their Late 
Quaternary ages, taphonomy, palaeoecoiogy, and fauna! 
composition (see also Akersten 1980). Churcher (1966) also 
questionably listed Silphidae from Talara, but S.B. Peck 
(personal communication) was unable to locate any 
Silphidae in Churcher's samples at the Royal Ontario 
Museum. Zopheridae has only recently been recognized as 
separate from Tenebrionidae, so may yel be recorded from 
the other deposits. 
Extinction 
The causes of extinctions among large vertebrates at the 
end of the Pleistocene remain a topic of debate (Beck 1996 
and references therein). Axelrod (1967) and Martin and 
Neuner   (1978)   suggested   that   low   seasonalily   of 
Figure 2. Holotype head of iJnihophjgui everesMe from 
Locality 81, Rancho La Broa (width about 2 mm; 
LACM IP 3057; scanning electron micrograph by 
L.E.C. Ling, Carleton University). 
188 QUATERNARY PROCEEDINGS 
Pleistocene environmenis permitted the establishment of 
very complex communities that lack modern analogs, and 
that the inception of Holocene seasonal environments was 
a major cause o? terminal Pleistocene extinctions. Despite 
the potential data at Rancho La Brea and elsewhere, 
relatively little remains known about the Quaternary 
climatic and vegelational history o? southern coastal 
Caliiornia (Johnson 1977; Cole & Liu 1994). Many birds 
and mammals recorded irom Rancho La Brea are now 
extinct (Stock & Harris 1992). 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 (Warier 1976; Miller 1979). 
A( Rancho La Brea, the extinction picture for insects 
was clouded by Piercc's many supposedly extinct taxa. 
Most of these have now bc>en placed in synonymy of extant 
taxa, two have been recognized as being extinct, and 
others are impossible to identify because of the nature o? 
the type specimens and the taxonomic condition of the 
groups concerne?) (Miller 1983). 
The scarab beetles Copris prisiinus Pierce and 
Onthophaaoiis everestae Pierce (Figure 2) 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 availal>ility 
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 fauna 
had survived into the Holocene, the scarabaeinc fauna of 
California might not have survived the present seasonal dry 
periods, since their successful reproduction requires 
adequate moisture (Miller ef al. 1981). Of course, possibly 
one or both of these species still lives in inadequately 
collected regions of Mexico and remain to be rediscovered. 
Several Pierce laxa remain problematic, but it is best to 
assume they are extant until contrary evidence is found 
(Miller 1983). It is presently impossible to identify properly 
S?rica kanakoffi Pierce, but there is no good reason to 
assume that il is not conspecific with an cxiant (but 
unidentified) species. S?rica is a large, poorly underst(K)d 
genus of plant feeding beetles (Miller ei al. 1981). 
Likewise, the specific identity of the scarab Phanaeus 
labreae (Pierce) also cannot presently be determined due to 
Figure 3. Hololype of Spirobolui' ?ustralis from Rancho la 
Brea (width about 7 mm; UCMP Invertebralc 
specimen 10008). 
the broken and distorted condition of the holotype, 
although it probably is assignable to an extant species 
(Miller ef al. 1981; Edmonds 1994, 91). The millipede 
'Spirobolus' australis Grinnell (Figure 3) is another 
unidentifiable laxon. known only from the fragmentary 
ly(x>s. It is probably a Hiltonius or Tylobolus (Miller 1983, 
9(j). The tenebrionid be<?tle Conionlin ronmaf}S Pierce is not 
conspecific with any described spwcies (Doyen & Miller 
1980), but this does not mean it is extinct. Conioniii is a 
genus of numerous pKwrly understood S(>ecies, probably 
including at least several undescribed species. C. remnant 
probably still lives in southern California, but has yet to be 
recognized (most Coniontii in museum collections are 
unidentified or misideniified). 
Directions for future research 
The high quality of preservation, both in physical 
characteristics and geochemistry (e.g. McMenamin cf al. 
1982), as well as the large numbers of individuals available 
for study, make Rancho La Brea fossils excellent subjects 
for many kinds of research. This presents the possibility of 
evaluating 40.000 years of change in nwrplioiogy, genetics, 
and isolopic composition in individual insect species! 
The high quality of data associated with specimens from 
Locality 91 will allow palacoccological studies. These 
could be especially illuminating if undertaken as part of 
muliidisciplinary research including analysis of plants, 
vertebrates, and sediments. The availability of computer 
data processing and storage now allows correlation of data 
from many sources. Any further research will require a 
major program of sorting and identification. For some taxa, 
this will require r??valuation o? the modern California 
fauna to place the fossils in a context {e.g. Miller & Peck 
1979). Plant fossils should also be examined for signs of 
insect damage, such as the gall illustrated in Figure 4, and 
other phytopathological data for comparison to modern 
studies such as Gagne (1989), Keifer ef al. (1982), and 
Opler(1974). 
Recent developments in isolopic and molecular analysis 
techniques open new horizons on the information that can 
be extracted from Rancho La Brea insect sj)ecimens. These 
insects have already been used in carbon isotope studies of 
climatic change (QC-1049, QC-IOSO; IF. Marcus & S.E. 
Miller, unpublished data). La Brea specimens should be 
suitable for other isolopic studies (e.g. R. Miller 1991; R. 
Miller ef al. 1993). Accelerator radiocarbon dating 
techniques now being applied to bone can be extended to 
insects as well (Elias & Toolin 1990; Stafford ef al. 1991). 
Likewise, some of the techniques of nucleic acid extraclior? 
and analysis now being applied to population genetics and 
syslematics of insects (Hillis ef al. 1996) should be 
applicable to Rancho La Brea specimens (see also 
Janczewski et al. 1992). 
Conclusion 
Asphalt-impregnated sediments at Rancho La Brea. Los 
Angeles County, California, provide a rich fossil record 
spanning the last 40,000 years. The abundance of insect 
specimens and their excellent preservation, especially at 
Locality 91, provide tremendous but untapped 
opportunities for palaeoecological and climate change 
studies. The resolution of basic questions about apparent 
terminal Pleistocene extinctions provides a foundation for 
LATE QUATERNARY INSECTS OF RANCHO LA BREA 189 
Figur? 4. Unidentified cynipid gall (Hymenoptera; Cynlpldae) 
ofi Quercus (?) ip. (Fagaceael from Pit A, Rdncho La 
Bfea (lengih about 20 mm; specimen LACM RLP 
4308). 
further work on this and other asphalt deposits. 
Acknowledgements 
The Natural Museum of Los Angeles County (W.A. 
Akersten, C.L. Hogue, G.T. Jefferson, R.L. Reynolds, C.A. 
Shaw, E.C. Wilson), Santa Barbara Museum of Natural 
History (R.S. Gray, PC. Hochberg), Queens College, 
C.U.N.Y. (L.F. Marcus), Smithsonian Institution and 
University of California al Santa Barbara supported this 
research. This research svould not have been possible 
without the collaboration of many laxonomisls 
acknowledged elsewhere (Miller 1983) . O. Kukul, D. 
McMenamin, and V.V. Zherichin helped me understand 
the literature about Binagady. The manuscript was 
reviewed by P. Buckland, N.L. Evenhuls, C.L. Hogue, G.T. 
lefferson and C.A. Shaw. 
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