Christopher D. Nagano, Scott E. Miller, and Alan V. Morgan.
Fossil Tiger Beetles (Coleoptera: Cicindelidae): Review and New Quaternary Records.
Psyche 89:339-346, 1982.

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FOSSIL TIGER BEETLES (COLEOPTERA: CICINDELIDAE): REVIEW AND NEW QUATERNARY RECORDS
BY CHRISTOPHER D. NAGANO~, SCOTT E. MILLER^ AND ALAN V. MORGAN
Fossil Cicindelidae are extremely rare in the stratigraphic record, probably due to the fragile nature of their exoskeleton. In this paper we summarize previous records and comment on new finds, as well as describe cicindelid specimens found in the southern California asphalt deposits which were noted, but not identified, by Pierce (1947a, 1947b).
Both the well known McKittrick asphalt deposit in Kern County, California and the Rancho La Brea sequence, Los Angeles County, California, have produced identifiable cicindelid specimens. Pierce's specimens are deposited in the Natural History Museum of Los Angeles County (LACM). A fairly well-preserved specimen of Cicin- dela haemorrhagica LeConte (LACM Invert. Paleo. hypotype 4944) from the McKittrick asphalt deposit, retains complete markings on the elytra, and the elytral pleura still show a blue coloration similar to that of populations found along the sea coast of central San Diego County, California. A mandible (LACM Invert. Paleo. hypo- type 4945) also from McKittrick, is probably a cicindelid but family placement is uncertain due to the poor condition of the specimen. Both McKittrick specimens were collected by L. Bessom from W. D. Pierce's "site 4" (LACM Invert. Paleo. Loc. 260), at a depth of 4 feet (1.3m) (Pierce 1947b, Miller and Peck, 1979). This site has not yet been radiocarbon dated, but a Cybister elytron taken from a depth of 3 feet (1 m) in the road cut at McKittrick has provided an Natural History Museum of Los Angeles County, Los Angeles, California 90007. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138.
'Department of Earth Sciences, University of Waterloo, Waterloo, Ontario, Canada N2L 3G 1.
Manuscript received by the editor May I, 1982.



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340 Psyche [vo~. 89
experimental sample for a 14C mass spectrometer run at Chalk River Nuclear Laboratory, Ontario, Canada. The resultant age of ca 8,000 yr B.P. suggests that the samples are probably of Holocene age, (Miller and Peck 1979, Morgan and Morgan 1980a) or of very late Pleistocene age as stated by Pierce (1947b). Three Cicindela specimens are known from Rancho La Brea; two thoraces of C. haemorrhagica (RLP 8779E and RLP 9014E) and a metasternurn of C. oregona LeConte (RLP 9465E). RLP 8779E and 9465E are from Pierce's "Bliss 29" material, which was collected in 1929 by W. Bliss from the sites of pits A, B, and C. The samples are probably late Pleistocene in age, but this is questionable due to unknown locality and possible contamination (Miller and Peck, 1979). RLP 9014E is from Pierce's "Pit X", which refers to mixed material, lacking data, and of questionable age. The Quaternary presence of C. haemorrhagica and C. oregona in southern California is not unexpected. Both are presently widely distributed in fresh and marine littoral habitats in western North America. Although these two species are not narrowly restricted to specific microhabitats, they are always found near permanent sour- ces of water.
Beside the above mentioned localities, a rekindled interest in the examination of coleopterous faunas has recently revealed fragments of cicindelids in a number of sites in the United States and Canada (Fig. 1). All but one of these records are more recent than the last review of North American fossil insects (Morgan and Morgan 1980b). The oldest specimen which is stratigraphically interpreted as pre last interglacial (pre-Sangamon) is a partial elytral fragment which is probably of the genus Omus from the Mountain View Dump site near Palo Alto, California (D. Adam pers. comm. 1978, Morgan unpublished). In the Pacific northwest, a recently examined site (Nelson and Coope, 1982) from Discovery Park, Fort Lawton, Seattle, has produced the remains of Cicindela oregona LeConte. A large and varied assemblage accompanies this find which is from sediments previously radiocarbon dated at between 23,000 and 18,000 yr. B.P. The site pre-dates the last major (Vashon) ice advance in the area and suggests a cooler climate with more open



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19821 Nagano, Miller & Morgan- Fossil Tiger Beetles 34 1 Figure 1.
Location of North American sites mentioned in the text. vegetation, an interpretation similar to that made from another 18,000 year old coleopterous assemblage from Port Moody in southern British Columbia (Miller, Morgan and Hicock, 1982). In the central eastern section of the continent three sites post- dating the retreat of Laurentide ice have produced cicindelid remains. The Norwood site in Minnesota (Ashworth et al., 198 1) is a late-glacial kettle which has a sequence of silts overlain by peat dated at 12,400k60 yr, B. P. (QL- 1083). A left elytron of Cicindela cf. C. sexgutiata Fabricius was recovered from the upper silt, an horizon characterized by a number of open ground beetle species. In the Canadian province of Ontario two sites at Kitchener and Brampton contain specifically identified cicindelids. The sites are approximately equivalent in age to the Norwood locality (ca. 12,400 to 12,000 yr. B.P.) and both slightly post-date the last major (Port Huron) ice readvance in the region. The Gage Street site, Kitchener,



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342 Psyche [vo~. 89
produced a solitary, well-preserved mandible of Cicindela repanda Dejean from the basal level of a marl deposit (Schwert, 1978) found in association with open-ground but largely boreal species. The Brampton site near Toronto, is a kettle deposit from which a soli- tary well-preserved right elytron and mandible fragment of Cicin- dela limbalis Klug was recovered (Morgan and Freitag, 1982). Once again the cicindelid fragments were associated with a fauna resident today in open ground regions within the boreal zone (Morgan, Morgan and Motz, 1982).
The presence of cicindelids in these early deposits of late Wiscon- sinan sequences is not surprising. In all cases, with the exception of the California examples, the tiger beetle remains are associated with species which inhabit open ground situations. Undoubtedly the ice merely forced many cicindelid populations southward at the time of maximum advance and they remained there to successfully recolo- nise sandy terrain after ice retreat. Tiger beetle remains also are present due to the nature of the sediments; the very fine silts and clays which are typical of most of these sequences is ideal for the preservation of the extremely thin elytral chitin found in cicindelids. In coarse sediments, or in sequences which are organic-rich, the detritus would abrade, distort and fragment the remains to a degree where most skeletal parts would become unrecognizable. In Table 1, we have attempted to compile known fossil Cicindeli- dae records including those described in this paper, and we have also commented, where appropriate, on some of the early identifi- cations.




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Table I: Known Fossil Cicindelidael
Age Deposit Locality Species Reference
Eocene
Green River USA: Colorado Cicindelopsis eophilus Cockerel1 1920 Cockerel12
Cf. Collyris sp. Larsson 1978
Odontochila sp. Horn 1907
Megacephala Carolina Horn 1906
Linnaeus
C. lunulata Fabricius, Bogachev 1948
C. campestris pontica Fisch.,
C. germanica Linnaeus
Megacephala cf. Blair 1927
M. femoralis Perty4
C. haemorrhagica H orn 1 876
LeConteb; C.sp.
Omus sp? this paper
Cicindela oregona LeConte this paper
C. haemorrhagica LeConte
Cicindela oregona LeConte
Coope and Nelson,
I982
Cicindela campestris Linnaeus Coope 1959 Cicindela s p. Coope and Sands
I966
C. campestris Linnaeus Coope 1977
Oligocene
Oligocene
Olig~cene(?)~
Baltic Amber
Baltic Amber
Baltic Amber
Quaternary Binagady brea
USSR: Caucasus
Region
Quaternary Fyzabad brea Trinidad
Port Kennedy USA: Pennsylvania
Pleistocene
Late Pleistocene
Mountain View Dump
Rancho La Brea
USA: California
USA: California
Late Pleistocene Seattle
USA: Washington
Late Pleistocene
Late Pleistocene
U. K: Cheshire
U.K: Warwickshire'
Late Pleistocene Low Wray Bay,
Windermerelo
Glanllynnau 1 1
U. K: Westmoreland
Late Pleistocene
C. campestris Linnaeus
Coope and Brophy
1972
C. campestris Linnaeus Osborne 1972
Cicindela sp, Osborne 1980
Cicindela sylvatica Linnaeus Coope 1979
Late Pleistocene
Late Pleistocene
Late Pleistocene
Church Stretton12
U.K: Shropshire
U, K: Staffordshire
U.K.
West Bromwichl3




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Age ~e~osit Locality Species Reference
Late Pleistocene N orwood USA: Minnesota Cicindela cf. sexguttata Fabricius Ashworth et al., 1981
Late Pleistocene Gage St., Kitchener Canada: Ontario Cicindela repanda Dejean Schwert 1978 Late Pleistocene Brampton Canada: Ontario Cicindela limbalis Klug Morgan and Freitag, 1982
Holocene McKittrick brea USA: California C. haemorrhagica LeConte this paper NOTES
'Matthews' (1976 and 1977) record of a cicindelid mandible from the late Tertiary Beaufort Formation on Meighan Island has been reidentified as Sialis sp. (Neuroptera: Sialidae) larva (Matthews, pers. comm.). 2As suggested by Cockerel1 (1920), this may not be a cicindelid. 3Thought by Cockerel1 (1920) to be a fake. doriginally reported as 'prope viridis' Tatumy now a synonym. The Port Kennedy mammal fauna is considered middle Pleistocene (Kurten & Andersen 1980)å but stratigraphic relation- ships of the insects are unknown.
6Willis (1967) suggests this may be a misidentification of C. rujiventris Dejean. 'Because of county boundary revisions in many parts of England and Wales former counties (in text) have disappeared ie. (Caernarvonshire in North Wales is now included in the new county of Gwynedd). Refer to original papers for national grid coordinates.
gca. 70,00040,000 yrs. B.P.
9ca. 32,000 yrs. B.P.
%a. 13,000 yrs. B.P.
lka. 12,500 yrs. B.P.
I2ca, 12,50H1,500 yrs, B,P,
%a. 12,100 yrs. B.P.




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19821
Nagano, Miller & Morgan- Fossil Tiger Beetles We would like to thank C. L. Hogue, P. C. Owen, C. A. Shaw and E. C. Wilson (all of LACM), J. V. Matthews Jr. (Geological Survey of Canada), A. Morgan (University of Waterloo), R. E. Nelson (University of Seattle) and D. P. Schwert (North Dakota State University) for providing specimens and data. ASHWORTH, A. C., Schwert, D. P,, Watts, W, A. and H. E. Wright, Jr. 1981. Plant
and Insect fossils at Norwood in south-central Minnesota: A record of Late- glacial succession. Quaternary Res. 16: 66-79. BLAIR, K. G. 1927. Insect remains from oil sand in Trinidad. Trans. Ent. SOC. Lond. 75: 137-141.
BOGACHEV, A. 1948. Fauna of the Binagady asphalt deposit: beetles-Coleoptera. Trudy Estestvenno-Istoricheskii Muzei, Baku 1-2: 137-160. [in Russian]. COCKERELL, T. D. A. 1920.
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A Late Pleistocene insect fauna from Chelford, Cheshire. Proc. Roy. SOC. (London) Ser. B. 151: 70-86. COOPE, G. R.
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COOPE, G. R. 1979. The Carabidae of the glacial refuge in the British Isles and their contribution to the post glacial colonization of Scandinavia and the North Atlantic Islands. pp. 407-424 in Erwin, T. L. et al. (ed.), Carabid beetles: their evolution, natural history, and class~jkation. Junk, The Hague. COOPE, G. R. AND J. A. BROPHY.
1972. Late Glacial environmental changes indi- cated by a coleopteran succession from North Wales. Boreas l(2): 97-142. COOPE, G. R. AND C. H. S. SANDS. 1966. Insect faunas of the last glaciation from the Tame Valley, Warwickshire. Proc. Roy. SOC. (London) Ser. B. 165: 38M 12.
HORN, G. H. 1876. Notes on some coleopterous remains from the bone cave at Port Kennedy, Pennsylvania. Trans. Amer. Ent. SOC. 5: 241-245. HORN, W.
1906. Ueber das verkommen von Tetracha carolina L. im preufsischen bernstein und die phylogenie der Cicindela-arten. Deut. Ent. Zeit. pp. 329-336. HORN, W. 1907. Brulle's bbOdontochila aus dem baltischen Bernstein" und die Phylogenie der Cicindeliden. (Col.). Deut, Ent. Zek pp. 461-466. KURTEN, B. AND E. ANDERSON.
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346 Psyche [VOI. 89
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Klug (Coleoptera: Cicindelidae) in a late-glacial site at Brampton, Ontario. Coleop. Bull. 36: 105-108.
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the base of a late-glacial sequence near Brampton, Ontario. Prog. with Abs. Geol. Assoc. Can. Annual Meeting, Winnipeg 67. NELSON, R. E. AND G. R. COOPE. 1982. A pre-Vashon (Late Pleistocene) insect fauna from Seattle, Washington. Abs. Vol. VII, AMQUA Conf. Seattle, Wash., 146.
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