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1990

Paleoenvironmental analysis of a late-holocene subfossil Coleopteran fauna from Starks, Maine

Heather A. Hall Colby College

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Recommended Citation Hall, Heather A., "Paleoenvironmental analysis of a late-holocene subfossil Coleopteran fauna from Starks, Maine" (1990). Senior Scholar Papers. Paper 110. https://digitalcommons.colby.edu/seniorscholars/110

This Senior Scholars Paper (Open Access) is brought to you for free and open access by the Student Research at Digital Commons @ Colby. It has been accepted for inclusion in Senior Scholar Papers by an authorized administrator of Digital Commons @ Colby. PALEOENVIRONMENTAL ANALYSIS OF A LATE-HOLOCENE SUBFOSSIL COLEOPrERAN FAUNA FROM STARKS, MAINE

by

Heather A Hall

Submitted in Partial Fulfillment of the Requirements of the Senior Scholars' Program

COLBY COLLEGE 1990 APPROVED:

/--- ~, ---~~:~~------TUTOR Robert E. Nelson

-----~~--~------CHAIR. DEPARTMENT' OF GEOWGY. Donald B. Allen

-----~-~-~~~~------READER Harold R Pestana

------~~ CHAJR. Diane F. Sadoff ABSTRACT

The sandy River in central Maine Is flanked along much of its length by low terraces. Approximately 100 kg of sediment from one terrace in

Starks. Somerset County, Maine was wet-sieved in the field. Over 1100 subfossll Coleoptera were recovered representing 53 individual species of a total of 99 taxa. Wood associated with the fauna is 2000 +/- 80 14C Yr in age (1-16,038). The fauna is dominated by species characteristic of habitats apparent in modem central Maine. The subfossil assemblage is indicative of a wide vartety of environments including open ground (e.g.. Harpalus pensylvanicus), dense forest (e.g., pterostichus honesttLs). aquatic environments (e.g.. Gyrinus, Helophorus), ripartan environments with sand and., gravel substrates (e.g., Bembidion inaequale, Schizogenius lineolatus). and moist. organic-rich terrestrial environments (e.g.. Micropeplus sculptus). The ecological reqUirements for each taxon pennit an environmental reconstruction suggesting an area vegetationally. climatically. and ecologically similar to that of the Sandy River today. The lowest terraces apparently represent the modem-day floodplain of the Sandy River. An average sedimentation rate of l.00 to 1.04 rom per year has been inferred based on radiocarbon dates here and elsewhere on the Sandy River. The Coleopteran fauna suggests that sand and gravel were distinctly abundant, and that the aggradation of point bars. as seen today, contributed to the flood history. Lateral bank erosion of the modem Sandy River accelerated after the State of Maine mandated cessation of bar removal in 1975: flood severity has dramatically increased since that time. Implications suggest that mining of the bars may be necessary to minimize future flooding problems. 2.

INTRODUCTION

Insect fossils make up the most abundant, diverse and identifiable component of Quaternary terrestrial assemblages (Coope. 1978). The fossil record shows evidence of specific constancy for at least the last half million years based on the Cape Deceit Fauna ofAlaska (Matthews. 1974: Coope. 1978). although Repenning (1985) argues constancy for the last two million years. back to at least the late Pliocene. based on a mammalian fauna. Specific identification of the subfossll Coleoptera is therefore based upon the morphological characteristics of modem species. The ecological requirements for each taxon in the fauna pennit an environmental. climatic. and vegetational reconstruction for the area. This research was undertaken with the attempt to contribute to previous paleoenvironmental analyses by Nelson (1987). based on subfossil insects. and Davis et aL (1975). based on pollen studies. who individually concluded that conditions in the central Maine region were essentially modem by the late Holocene (2000-2500 B.P.). The research as well attempted to clarify and defme the environmental factors which have contributed to the long-tenn flood history of the Sandy River. which has increased in severity and frequency in recent decades. as examined by Eastler et aL (1989). 3.

METHODS

Low terraces are common along the Sandy River in its lower reaches from Franklin County to Its confluence with the Kennebec River in Kennebec County, Maine (Figure 1). Terrace sediments were sampled along one river segment in Somerset County on a south-faCing cutbank of the Sandy River, at 44°44.3'N, 69°53.9W (Figure 1). located in the Mercer, Maine 7.5-minute topographic quadrangle (scale 1:24,000). Approximately 5 kg of sediment was collected on September 8. 1989, from a lens with abundant macroscopic plant remains, 2 m below the terrace surface. Additional material was collected three weeks later on September 29, 1989.

The water in the river had risen considerably by this time, and sampling at the same level was impossible. The second sample was taken approximately 30 em above the first, and approximately 100 kg of silt to fme sand was wet­ sieved in the field. Material greater than 0.250 mm was brought back to the laboratory for processing. The samples were wet-sieved in the laboratory using screens with mesh diameters of 2.0 mm. 1.0 mm, and 0.300 nun. Materials greater than 2.0 mm. and between 2.0 nun and 1.0 mm were dried: Coleopteran subfossU parts, along with Trichopteran (caddisfliesl and plant remains, were picked from the dried sediment under a dissecting microscope. The fraction finer than 0.300 mm was discarded. The remaining material, 1.0 mm to 0.300 mm. was processed using standard kerosene flotation techniques as outlined by Coope (1986); the float was similarly picked. The subfossil Coleopteran fragments were taxonomically arranged to family and placed on prepared cardboard microfossil sUdes using gum tragancanth. Identifications were based on comparisons from the modem collection of 4. /.... i/" sandy River----" \ ~ Approximate \ limit of \ ~ terraces STARKS ...... ' .•.....•.•. / ...• I""" ... / Study j NORRIDGEWOCK s1te~ . i ,I \ ! \...... _._..__ : . l~ \\ ...... '. 1 ...... -.._ _ . '.". , o 0.5 1.0 ...i::=::iIII.t:::::iI-t1=_t=:-'-==l1 KM

Study Area

FIG. 1. Map showing the location of the Sandy River site. 5.

R.E. Nelson. selected Coleoptera from tbe Museum of Comparative Zoology, literature descriptions (Lindroth, 1961-1969: Campbell. 1968a.b. 1973. 1982: Dillon and Dillon. 1972; Moore and Legner. 1979: Arnett, 1968: Gordon. 1985; Hilsenhoff. 1973), and expert identification (Schmude. personal communication to R.E. Nelson).

RESULTS

Insect fragments included in this study were restricted to the Order Coleoptera. "Remains of over 1100 individual beetles were recovered, representing 53 species included in 99 taxa and 27 families (Table 1) of a total of 110 families present in North America (Arnett. 1968). Families represented include Carabidae. HaIiplidae. Dytiscidae. Gyrtnidae, Hydrophtlidae. Limnebiidae. Staphylinidae. Pselaphidae. Leodidae. Brathinidae. PtiHidae, Scydmaenidae. Scarabaeidae. Heteroceridae, Elm1dae. Elateridae, Throscidae. Cantharidae, Dermestidae. Nitidulidae, Cucujidae, Cryptophagidae. Coccinellidae. Latbrtdtdae, Chrysomelidae, Curculionidae. and Scolytidae. Pronota and elytra were well represented in the assemblage.

Heads and genitalia were present. but few in number. A log extracted from the same level as the second sample was submitted for analysis. An age of 2000 +/- 80 14C yr B.P. (1-16.038) was determined: a late-Holocene age had been expected based on the modern characteristics of the assemblage. A second piece of the log submitted for radiocarbon dating was submitted for identification. Degradation of the wood led to a lack of the characteristic patterns necessary for positive identification; Carpinus caroliniana Walter (American hombeam) apparently represents the closest match (D. Christensen. oral communication to R.E. 6.

Nelson, 1990).

PALEOENVIRONMENTAL RECONSTRUCTION

The subfossil assemblage is indicative of a wide distribution of ecological requirements (Table 1). and the environments indicated provide a clear picture of the overall character of the lower Sandy River area 2000 years ago. Five major habitat divisions have been inferred based on the characteristics of the assemblage: open ground. dense forest, aquatic. riparian. and moist to wet environments. which mayor may not have been proximal to the surface water. A much wider variety of microhabitats is apparent. The open-ground habitat includes several microenvironments. The fiist is xeric and lightly to moderately vegetated with grasses. as evidenced by the presence of Harpalus pensylvanicus. Tachys granarius and Amara..

Cymindus cribricollis. and T, granarius provide evidence of a microenVironment charactertzed by sand and gravel substrates (Lindroth.

1966.1969). pterostkhus adstri.ctus is a forest species requiring shaded open ground, and thus suggests the existence of a forested area lacking a dense understory (Lindroth. 1966). Hyperaspis binotata is representative of exposed vegetation as are several other taxa: Throscidae live among the flowers. Cantharidae. Apion. Rhynchaenus, Altica and Galeruca among the foliage as adults (Arnett. 1968. Kissinger, 1964). The Scolytidae are associated with trees or wooded vegetation (Arnett. 1968). and NiUdulidae with decaying fruits and fermenting plant Juices (Parsons. 1943). A forest habitat is suggested by the recovery of several species of

Carabidae. Agonwn retractum. Pterostichus honestus. and Sphaeroderus lecontei are forest species usually found in shaded areas with predominantly 7. organic substrates derived from leaf litter (Lindroth. 1961. 1966). P. adstrtctus is one of the most common beetles of the northern coniferous region, and prefers open ground with moderately moist to dry soil (Lindroth. 1966). The subfossil assemblage 1s dOminated by aquatic species represented by the families of Dytiscidae (predacious water beetles), Haliplidae (crawling water beetles), Hydrophilidae (water scavenger beetles), Limnebiidae. Gyrinidae (whirligig beetles). and Elrnidae (riffle beetles). The Dytiscidae. Hydrophilidae. and Limnebiidae occur prinCipally in shallow. slow moving water along an edge typically matted with vegetation. as they rely on self­ contained air stored under the elytra for respiration. and must return to the surface often to replenish the supply; they often prefer back-water pools

OJ and side channels where water movement is slow, so that the chance of being carried away with the current in minimiZed in their relatively frequent trips to the surface (Larson. 1987; Leech and Chandler, 1956; Doyen and

Ulrich. 1978). Gyrtnus lives in sheltered 1entic sites on the surface film of partially open water surfaces (Larson, 1987). As well. HalipLus. Hydraena and Octhebius suggest the presence of large rocks. logs. moss. and other debris in the water (Larson. 1987: Leech and Chandler. 1956: Doyen and Ulrich. 1978). Cryptic behavior Is common among many water beetles; most hide in the substrate or lie concealed among vegetation, rocks or detritus to escape detection (Larson, 1987). Elmidae are found comparatively deeper in lotic sites with well-oxygenated water, where they remain submerged indefinitely due to the presence of retractile gills (Leech and Chandler, 1956: Doyen and Ulrich, 1978), but are typically found as well in shallower water nearshore due to their need to move out of the water to pupate (K. 8.

Schmude, written communication to R.E. Nelson, March 15. 1990).

Dubiraphia can usually be found on submerged roots. stems. and bark, or occasionally on rocks or pebbles encrusted with algae (Brown and White.

1978). Opnoservus is typically found in gravel or coarse sand in fast. shallow water. and Promoresia on roots or moss in fast mountain streams (Brown and White. 1978). The Elmidae are particularly well represented, and include a high specific vanation in the Sandy River fauna.

Species representing the riparian environment (e.g.. Bembidion castor, B. honestum. B. inaequale, SchizogeniUS lineolatus. Tachys saturatus. and Omophron americanum.) suggest open sand and gravel banks. with silty and clay-rich areas (Lindroth. 1961. 1963. 1966). Bembidion quadrimaculatum oppositum prefers a riparian environment. particularly sparsely vegetated with a substrate of fine sand or silt, while B.semicinctum and B. patruele confinn the presence of wet organic substrates (Lindroth.

1963). Large numbers of riparian Carabidae and Oxytelinae (Le.. Bledius and

Carpelimus) further confirm the dominant enVironment as riparian

(Herman, 1986). The Omaliinae Geodromi.cus strictus is as well found on gravel-banked clear steams in Maine (R.E. Nelson, unpublished data). Moist to wet organic-rich substrates are indicated by numerous taxa including Micropeplus sculptus. Oxytelus, Acidota subcarinata. and

Euaesthetus (Campbell. 1968b. 1982: Moore and Legner, 1979). as well as 7 other taxa. This enVironment is typified by moss inhabitants such as

Lesteva pallipes. Rybaxis varicomis and other Pselaphidae. and leaf-litter inhabitants such as Batrisod.es. Trechus crassiscapus. and most PWiidae

(Arnett. 1968: Lindroth. 1961). Arpedium cribratum in Maine has been collected in leaf litter and duff in shaded areas with a moist to wet substrate of silt and very fme sand (R.E. Nelson. unpublished data). The nests of 9. vertebrates are often inhabited by Aphodius, where It feeds on dung (the Scarabaeidae are collectively referred to as the dung beetles) although the genus is often found as well among other organic debris (Arnett. 1968; Borror et at. 1976). Brathinus varicomis in particular is found along the roots of grass groWing near water in cool riparian or swampy-bog habitats (Arnett. 1968; Peck. 1975). It must be noted that these areas are indeed microhabitats and could occur enclosed within the open-ground, forest. or riparian environments.

The ladybird beetle Hyperaspis binotata (Gordon. 1985) and the

Elmid Promoresia elegans (Brown and White, 1978) reach their present northern limits in central Maine. as does Carpinus caroliniana.. if this shrub is truly represented (Little, 1971). suggesting that summer conditions tn this region 2000 years ago were at least as warm as those today.

DISCUSSION AND CONCLUSION

The ecological reqUirements for each taxon pennits an environmental reconstruction suggesting an area vegetationally, climatically. and ecologically similar to the Sandy River today. dominated by aquatic and riparian environments with back-water pools, a sand and gravel bottom and comparatively coarser-grained banks and bars. an adjacent conifer-hardwood forest and at least some marginal open ground. A tremendous diversity of microhabitats would have occurred within these territories. Results from this research agree with the environmental reconstruction of Davis et aL (1975) who suggested a Hardwood-Conifer Period for 2000 b.p. (4700 to 200 years b.p.). This was the modem environment prior to the influences of European settlement (European 10.

Period, 200 years b.p. to present) (Davis et aL, 1975). Further evidence for the modem character of the area 2000 years ago is that many of the same species found in the Sandy River subfossil assemblage have been collected along the modern Sandy River, and in adjacent forest areas (Nelson, unpublished data). Caldwell (1986) and Weddle (1987) in mapping the Sandy River region designated the low terraces as Quaternary alluvium. but did not distinguish between terrace deposits and the modern floodplain. The radiocarbon date of this research suggests that the lowest and broadest surfaces are indeed the modem floodplain of the Sandy River. A 2120 +/­ 80 14C yr B.P. (1-14.818) radiocarbon date from the Sandy River at Strong

(Nelson. 1987) came from 3 m below the ground surface. Based on that date. and the radiocarbon date from this research, as well as the thickness of the sediment, an average sedimentation rate of 1.0 to 1.4 mm per year can be calculated for the Sandy River floodplain.

MODERN ENVIRONMENTAL IMPLICATIONS

The Sandy River has a flood history dating back over 200 years. although discharge records only date back to 1928 (Eastler et aL, 1989).

About 980 cubic feet per second is the average annual discharge at the mouth (Eastler et at. 1989). However, in the last 15 years, flood stage has been reached more qUickly and has become significantly greater in severity.

The April. 1987. flood has ranked first in severity, with a discharge of

51.100 cubic feet per second at Mercer. in comparison to its driest year

With a discharge of 32 -cubic feet per second in 1939 (Bartlett et al.. 1989). The aggradation of sand and gravel bars along the river has contributed 11. to this problem of increasing flood frequency. Gravel had been mined without permit since before 1785. until the State of Maine mandated the cessation of bar removal in 1975 based upon recommendations from the Department of Inland Fisheries and Wildlife. which was concerned with the disruption of biotic environments (Eastler et al.. 1989). Up to 10 m of lateral bank erosion has washed large quantities of sediment annually into the river since that time. derived from land responsible for over 90% of the agricultural productiVity in Franklin County fEastler et al.,1989: York, 1980). This has removed large areas from production while significantly increasing the sediment load of the river. The subfossil assemblage suggests that the area 2000 years ago was similar to unrestricted reaches of the Sandy River today. dominated by broad sandy bars and a gravel bottom. Such features are consistent with rapidly shifting channels due to the meandering and braided path of the river. The radiocarbon dates clearly show that the lowest of the terrace-like surfaces was really the floodplain that prevailed before the European PeIiod of Davis et aL (1975). Removal of sand and gravel bars along the Sandy River encouraged the abandonment of this floodplain by deepening the river channel. thus decreasing the degree of meandering seen in the river; this was also assisted by a series of small dams which helped to halt to flow of sand and gravel within the river. However. because of the aggradation of the bars in recent times, the river is once again being forced outward. thus having the potential of causing severe adverse economic problems. as the lowest terrace surface supports a large part of the agriculture in the county and is coming under increased development pressures. Thus. based on this study and that of Eastler et al.

(1989). it may be suggested that the removal of the gravel from the bar 12. system may be necessary in order to minimize future flooding problems. The State of Maine is currently looking into the proposition: studies are ongoing with regard to the nature of aggradation (Eastler et aI., 1989).

ACKNOWLEDGEMENTS

I am greatly indebted to R.E. Nelson for his outstanding support and dedication, for the loan of his modem collection. and for taxonomic classification. environmental interpretation. field assistance, and manuscript critique. Thanks to D. Furth. Museum of Comparative Zoology, for the loan of valuable comparative specimens. and to K Schmude of the University of Wisconsin for assistance in taxonom1c classification and environmental in'terpretation of Elmidae. to W.E. Doll for assistance in the drafting of figures. and to D.B. Allen and H.R. Pestana for providing constructive suggestions for improvement of the manuscript. This research comprised a senior honors project and was supported by Colby College: the radiocarbon date was funded by the Colby College Natural Science Division Research Grants Committee. Thanks to J. Buckley of Teledyne Isotopes, Westwood. New Jersey for radiocarbon age determination, and to D. Christensen of the Forest Products Lab. U.S.F.S. Madison, Wisconsin for wood identification. 13.

REFERENCES

Arnett. RH. (1968). lhe Beetles of North America". The American Entomological Institute. Ann Arbor. Michigan. Bartlett, W.P.. Higgins, W.B.. and Nichols. W.J. (1989). Water Resources Data Maine. Water Year 1988. USGS Water Data Report ME-88-I. Augusta, Maine.

Borror. n.J.. DeLong, n.M.. and Triplehorn, C.A. (1976). "An Introduction to the Study of Insects (4th ed.r. Holt. Rheinhart. and Winston. New York. Brown. H.P. and White. D.S. (1978). Notes on Separation and Identification of North American Riffle Beetles (Coleoptera: Dryopoidea: Elmidae). Entomological News 89. 1-13. Caldwell. D.W. (1986). Reconnaissance Surficial Geology of the Fannington Quadrangle. Maine. Maine Geological Survey, Open-File Map No. 86-29. Campbell, J.M. (1968a). A New Species of Pycnoglypta Thomson

(Coleoptera: Staphyl1n1dae) from Eastern Canada. Canadian

Entomologist 115, 361-370. Campbell. J.M. (1968b). New World Micropeplinae. Canadian

Entorrwlogist, 100. 225-267, Campbell. J.M. (1973). "A Revision of the Genus Tachtnus (Coleoptera:

Staphylintdae) of North and Central America", Memoirs of the Entomological Society of Canada. No. 90, Ottawa. Campbell. J.M. (1982). A Revision of the North American Omaliinae {Coleoptera: Staphylinidael 3. The Genus Acidota Stephens.

Canadian Entomologist 114. 1003-1029. 14.

Coope, G.R (1978). Constancy of insect species versus inconstancy of Quaternary environments. In "Diversity of Insect Faunas" (Mound. L.A and Waloff, N., eds.). Symposia ofthe Royal Entomological Society of London. No.9, pp.176-187. Blackwell Scientific Publications. London. Coope, G.R (1986). Coleoptera analysis. In "Handbook of Holocene Palaeoecology and Palaeohydrology" (B.E. Berglund, Ed.). pp. 703-713. John Wiley and Sons, New York. Davis. RB., Bradstreet. T.E., Stuckenrath. R, and Borns, H.W. (975). Vegetation and Associated Environments During the Past 14.000 Years Near Moulton Pond. Maine. Quaternary Research 5, 435-465. Dillon. E.S., and Dillon, L.S. (1972). "A Manual of Corrunon Beetles of Eastern North Amenca- Volumes 1,2." Dover Publications. New York. Doyen, J.T. and Ulrich. G. (978). Aquatic Coleoptera. In "An Introduction to the Aquatic Insects of North America". (Merritt. R W. and Cummins, K.W., eds.), pp. 203-232. Kendall/Hunt Publishing. Dubuque, Iowa. Eastler, T., Sproul, J., and Buckland. A. (989). Environmental Geology

Along the Sandy River. In "Guidebook for Field Trips in Southern and West-Central Maine". New England Intercollegiate Geological Conference. 81st Annual Meeting. pp. 200-211. G<>rdon, RD. (1985). The Coccinellidae (Coleoptera) of America North of Mexico. Journal of the New York Entomological Society 93. 1-912. Herman. L.H. (1986). Revision of Bledius. Part IV. Classification of Species Groups, Phylogeny. Natural History, and Catalogue (Coleoptera. Staphylinidae. Oxytelinae). Bulletin of the American Musewn of

Natw-al History 184. 1-367.

Hilsenhoff. W.L. (1973). Notes on Dubiraphia (Coleoptera: Elmidae) with 15.

Descriptions of Five New Species. Annals of the Entomological Society oj America 61. 55-61. Kissinger, D.G. (1964). "Curculionidae of America North of Mexlco. A Key to the Genera." Taxonomic Publications. South Lancaster. Mass.

Larson. D.J. (1987). Aquatic Coleoptera of Peatlands and Marshes in Canada.

Memoirs oj the Entomological Society oj Canada 140, 99-132. Leech. H.B. and Chandler, H.P. (1956). Aquatic Coleoptera. In "Aquatic Insects of California" (Usinger. RL., ed.), pp. 293-371. University of California Press, Berkeley. Undroth, C.H. (l961-1969). 111e Ground-Beetles (Carabldae, excl. Cicindellnae) of Canada and Alaska." Opuscula Entomnlogica, Supplementa 20. 24. 29, 33, 34. Entomologiska Sallskapet. Lund, Sweden. Uttle, E.L. (1971). Atlas of United States Trees- vol. 1: Conifers and Important Hardwoods. U.S. Government Printing Office. Matthews, J.V., Jr. (1974). Quaternary envtrorunents at Cape Deceit (Seward Peninsula, Alaska): Evolution of a tundra ecosystem.

Geological Soctety oj America Bulletin 85, 1353-1384.

Moore. 1. and Legner, E.F. (1979). "An illustrated Guide to the Genera of the Staphylinidae of America North of Mexico. Exclusive of the Aleocharinae (Coleoptera)". University of California Press. Berkeley. Nelson. RE. (1987). A Postglacial flora and subfossil insect fauna from

central Maine. Programme with Abstracts. XII International Union for Quaternary Research (INQUA) (Ottawa), 231. Parsons, C.T. (l943). A Revision of the Nearctic Nitidul1dae (Coleoptera). Bulletin oj the Musewn of Comparative Zoology 42, pp. 121-278. Peck, S.B. (1975). A Review of the Distribution and Habitats of North 16.

American BrathInus (Coleoptera: Staphylintdae: Omallinae). Psyche 82.59-66. Repennmg. C.A (985). Pleistocene mammalian faunas: climate and

evolution. Acta Zoologica Fennica 170. 173-176.

Weddle. T.K. (1987). Reconnaissance Surficial Geology of the Norridgewock Quadrangle. Maine. Maine State Geological Survey. Open-File Map No. 87-23.

York, V. (1980). !he Sandy River and its Valley". Knowlton and McLeary Company. Farmington. Maine. 17.

Table 1 Subfossil beetles recovered from the Sandy River and their modern habitats (families and subfamilies arranged taxonomically according to Arnett (1968); within families. taxa arranged alphabetically)

Taxonomy Sample 1 Sample 2 Habitat ------FAMILY: Carabidae

Agonwn retractum LeConte (2) D,IV

Agonum spp. (2) E,3 Amarasp. (1) C.2 Bembidion castor Lindroth (l) (6) BJ,II B. fronta1e LeConte (3) E,III,rv.3 B. gractliforme Hayward (2 ) E,I1I.3 B. hDnestum Say (1) B,I1 B. inaequale Say (1) BJ,In,l B. patruele Dejean (1) B,lV.l.2

B. quadnmaculatum oppositum Say (1) B.III.l

B. (?) rusticum Casey (2) B.I1 B. semlcinctum Notman ( 1) ( l) B.IV

B. (?) versicolor LeConte (1) E.N.l

Bembidton sp. (3) Cymindis crtbricolUs Kirby (1) C.I.l,2 Harpalus pensylvanicus DeGeer ( 1) C.2 Omophron americanwn Dejean (1) E,I pterosttchus adstrictus Eschscholtz (2 ) (1) C.D P. hDnestus Say (1) D,3 P. luctuosus Dejean (1) E.3 P. pensylvanicus leConte (1) D,II,IV

Pterostichus sp. (2) ( 1) &hizogenius lineolatus Say (1) B,I1,l

Sphaeroderus lecontei Dejean ( 1 ) ( 1) D.IV

Tachys (?) ancilla Casey (l) T. jlavicauda Say (1) D T. granarius Dejean ( 1) C.I,II,l

T. incurous leConte (2) E,I.II,l

T. saturatus Casey (1) (2) B,I1.3

T. scltulus LeConte (5) B,E,I,III Trec1w.s crassiscapus Undroth (1) E.IV Genus indet. (2) FAMILY: Ha11plidae

HaIiplus sp. (2) A

Genus indet. ( 1 ) A FAMILY: Dytiscidae

Agabussp. (1) ( 1) A

Coelambus sp. (1) (1) A

Hydroporus sp. ( 1) (1) A

Neoporus sp. (1) A FAMILY: Gyrinidae

Gyrinus sp. (1) A

Genus indet. (1) A FAMILY: HydrophJlldae

Berosus sp. ( 1 ) A

Cercyon sp. (1) A Enochrus sp. (1) A

Helocombus bifid.us (leConte) ( 1) A Heloplwrus sp. (6) A ? Hydrochara sp. ( 1) A Hydrochus sulxupreus Randall (1) A

Tropistemus sp. (1) A

Genus fidet. (1) (1) FAMILY: Umnebtidae

Hydraena spp. (4) (30) A,E

Octhebius sp. ( 1) A FAMILY: Staphyllnidae SUBFAMILY: Micropeplinae Micropeplus sculptus LeConte (1) E.rv.l,2 SUBFAMILY: Oxytelinae

Bled1us spp. (27) B.I,l CarpeUmus spp. (5) B.E.I,III Oxytelus sp. ( I ) IV Genus indet. ( 1) (2 ) SUBFAMILY: Omaltinae

Acidota subcarinata Erichson (3) E,IV Arpedium cribratum Fauvel (2) B.E.I1,IV Geodromicus strictus Fauvel (2) B.I1 Lesteua pallipes leConte ( 1 ) E.IV Olophrum sp. (1) E Omaliwnsp. ( 1) E

Pycnoglypta l.urida Gyllenhall (2) IV Genus indet. (1) (1) SUBFAMILY: Steninae TRIBE: Stenini

Stenus spp. (3 ) (15) B.E. I.III. 1 SUBFAMILY: Euaesthetmae Euaesthetus sp. (1) E.IV SUBFAMILY: Paedertnae

RugUus spp. (1 ) (4) IV SUBFAMILY: Staphylininae TRIBE: XanthoUnini

Genus indet. (3 ) (42) TRIBE: Quedllni

Genus indet. (1) (11 ) .. SUBFAMILY: Tachyponnae TRIBE: Tachyporin!

Tachinus homi Campbell ( 1 ) IV

T. (?) scrutator Gemminger & Harold (2) IV Tachinus sp. (2) (1) IV

TachypOT11S sp. (2) TRIBE: Bolltoblini Lordithon sp. (1) IV (Fungi) SUBFAMILY: Aleocharinae

MyUaena. spp. (9)

Genus indet. (2) (29) SUBFAMILY fidet. (5) FAMILY: Pselaphidae

BatrisDdes sp. (1) IV Decarthron sp. (1) E 21.

PUopius piceus LeConte (1) IV

Reichenbachia spp. (2 ) E Rybaxis varicomis (Brendel) (1) IV Genus indet. (3) FAMILY: Leodidae

Agathidiwn sp. ( 1) (1) IV Genus indet. (2) FAMILY: Brathinidae

Brathinus varicomis LeConte (3) E,B.2 FAMILY: Ptillidae SUBFAMILY: Ptiliinae

TRIBE: Ptiliini Genus lndet. (1) E,IV TRIBE: Acratrichini

Genus indet. (1) E,IV FAMILY: Scydmaenidae

Genus indet. (2 ) E,IV FAMILY: Scarabaeidae

Aegelia sp. (7) B,E,l Aphodius sp. (1) IV FAMILY: Heteroceridae

Heterocerus sp. (3) B.E,I.III,l FAMILY: Elmidae

Dubiraphia (?) vittata (Melsheimer)(I) (14) A

Dublraphia spp. (11) A

Macronychus glabratus Say (2) A

MicrocyllDepus pusiUus (LeConte) ( 1 ) A Oulimntus latlusculu.s (LeConte) (1) (9) A

OptiDservus trivittatus (Brown) (1 ) A

OptiDservus spp. (2) A Promoresia elegans (LeConte) (4) (9) A

Promoresia spp. (29) A

Stenelmis (7) bi£wtnata LeConte (2) A

S. crenata (Say) (2) A S. mera Sanderson (3) A

S. mirabUis Sanderson (2) (5) A FAMILY: Elatertdae

? Cardiophorus cardisce (Say) (1) Hypolithos sp. ( 1 ) ?B FAMILY: Throscidae Genus indet. (1) 2.3 FAMILY: Canthartdae ? Podabrus sp. (1) 2,3 Genus indet. ( 1 ) 2,3 FAMILY: Dermestidae

Genus indet. (1) IV FAMILY: Nitidulidae

Genus indet. (1) IV FAMILY: CucuJidae

Genus indet. (1) IV FAMILY: ? Cryptophagidae

Genus indet. (1) IV FAMILY: Cocc1nellidae

Hyperaspis bmotata (Say) ( 1) 2-3 Genus lndet. (1) FAMILY: Lathrtdtidae

Genus indet. (1) IV FAMILY: Chrysomelidae

SUBFAMILY: Donaclinae

Donacia a.equaUs Say ( 1 ) E,2,3 D. subtUis Kunze ( 1 ) E,2.3 Donacia sp. (1) (1) E,2,3 Platel1JTlQJis sp. (1) E,2,3 SUBFAMILY: Eumolpinae

Xanthonia decemnotata (Say) (3) D SUBFAMILY: Galerucinae

Galeruca sp. (10) rv,2.3 Genus indet. (1) SUBFAMILY: Altlcinae Altica sp. (1) (1) 2,3

Genus indet. (15)

SUBFAMlLY: Orsodacninae

Orsodacne atra (Ahrens) ( 1) 2,3

SUFAMlLY: indet. (3) (9) FAMILY: Curcu1iontdae

SUBFAMILY: Apioninae ? Apion sp. (2 ) rv.2.3 SUBFAMILY: Curculioninae

Rhynchaenus sp. (8 ) 8.2,3

SUBFAMILY: lndet. (3 ) (27) FAMILY: Scolytidae 24.

Genera lndet. (7) D FAMILY: lndet. (5) (21)

Totals: 1100 Individual beetles 99 taxa 52 identified species

Ecological coding:

Local Habitat Substrate Vegetation Density A=aquatic I=sand 1=bare or sparsely B=ripartan lI=gravel vegetated C=open ground. xeric III=silt and clay 2=moderately vegetated D=forest IV=organics 3=densely vegetated E=hygrophilollS (6) = number refers to the minimum number of individuals based on the most abundant skeletal part. ? =subfossils lack diagnostic characters needed for positive identification, or suitable series of reference speCimens not available.