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1061 Correlation of Grassy Lake and Cedar Lake ambers using infrared spectroscopy, stable isotopes, and palaeoentomology Ryan C. McKellar, Alexander P. Wolfe, Ralf Tappert, and Karlis Muehlenbachs Abstract:The Late Cretaceous GrassyLake andCedarLake amber depositsofwestern Canada are amongNorthAmeri- ca’smost famousamber-producing localities. Althoughit hasbeen suggestedforoveracentury that CedarLake amber fromwestern Manitoba maybe a secondarydeposit havingoriginated fromstrata inAlberta, thishypothesishasnotbeen tested explicitly usinggeochemical fingerprinting coupled tocomparative analyses ofarthropod faunal content. Although there are many amber-containing horizons associated withCretaceous coalsthroughout Alberta, most are thermally mature andbrittle, thuslacking the resilience tosurvivelongdistance transportwhile preserving intact biotic inclusions.Oneof the few exceptions isthe amber foundin situat GrassyLake. We present asuite ofnewanalyses fromthese andother Late Cretaceous ambers fromwestern Canada, including stable isotopes(HandC),Fouriertransforminfrared (FTIR) spectra, andan updated faunal compendium forthe GrassyandCedarlakes arthropodassemblages. Whencombined with amber’s physicalproperties andstratigraphic constraints, the resultsoftheseanalyses confirm that Cedar Lake amber is derived directly fromthe GrassyLake amber deposit oran immediate correlative equivalent. Thisenables the palaeoenvir- onmental context ofGrassyLake amber to beextended tothe Cedar Lake deposit,making possiblea moreinclusive sur- veyofCretaceous arthropodfaunas. Re´sume´ : Lesgisements ambrife`resduCre´tace´ supe´rieur de GrassyLake et CedarLake, dansl’Ouestcanadien, figurent parmi lessitesde production d’ambre lesplusce´le`bresd’Ame´riqueduNord.Bien que,pendant plusd’un sie`cle, onait soutenuquele gisement de CedarLake, de l’OuestduManitoba,pourrait eˆtre unde´poˆtsecondaire ayant sonorigine dans desstrates albertaines, cette hypothe`sen’apase´te´ mise explicitement a` l’e´preuve a` l’aide de me´thodesge´ochimiques combine´esa` desanalyses comparatives de la faune d’arthropodesqu’il contient. S’ilexiste, danstoute l’Alberta, denom- breuxhorizonsambrife`resassocie´sauxcharbonscre´tace´s,danslaplupart descas,l’ambre qu’ilsrenferment estthermique- ment mature et cassant, ne pouvantdoncpasre´sistera` untransport surdelongues distances tout enpre´servant des inclusions biotiquesintactes. Unedesrares exceptions estl’ambre trouve´ en place a` GrassyLake. Nouspre´sentonsune se´rie de nouvelles analyses de spe´cimens provenant de cesgisements et d’autresgisements ambrife`resduCre´tace´ tardif de l’Ouestcanadien, dontdesanalyses desisotopesstables(Het C)et desspectresinfrarouges par transforme´ede Fourier (IRTF),ainsi qu’uncompendium faunique a` jour desassemblages d’arthropodes de GrassyLake et CedarLake. Jumele´s auxdonne´essurlesproprie´te´sphysiqueset auxcontraintes stratigraphiques desambres,lesre´sultats de cesanalyses con- firmentquel’ambredeCedarLakeestdirectement de´rive´ dugisementambrife`redeGrassyLakeoud’une´quivalent corre´- latif imme´diat. Ceci permet d’e´largir le contexte pale´oenvironnemental de GrassyLake au gisement de CedarLake, rendant ainsi possibleunrecensement pluscomplet desfaunesd’arthropodes cre´tace´es. [Traduit par la Re´daction] Historical background tant because they were formed during a crucial time for understanding the adaptive radiations of modern insect The Canadian amber deposits at Grassy Lake (southern groups and the co-evolutionary processes associated with Alberta) and Cedar Lake (western Manitoba) are renowned the rise of angiosperms. Furthermore, these faunas closely internationally because of the richness of their arthropod in- predate the K–T (Cretaceous–Tertiary) extinction event, pro- clusions (Pike 1994). These ambers are particularly impor- viding an important baseline with which to assess its conse- quences for terrestrial arthropods. Received 8February 2008.Accepted 16September 2008. Publishedonthe NRCResearch PressWeb siteat cjes.nrc.ca on 3November 2008. Cedar Lake amber ThestudyofCedarLakeambercanbetracedasfarbackas Paperhandled byAssociateEditor C.Hillaire-Marcel. 1890 and includes such famous personages as the geologist R.C.McKellar,1A.P.Wolfe, andK.Muehlenbachs. Joseph Burr Tyrrell and the palaeoentomologist Frank Car- Department ofEarth andAtmospheric Sciences, 1-26Earth penter(Grimaldi1996).Amberfromthislocalityhasbeenre- Sciences Building, UniversityofAlberta, Edmonton, ferred to as chemawinite or cedarite (Aber and Kosmowska- ABT6G2E3,Canada. Ceranowicz 2001),andthe former name hasbeen incorrectly R.Tappert.School ofEarth andEnvironmental Sciences, extendedbysomeauthorstoencompassthebulkofCanadian UniversityofAdelaide, SouthAustralia, 5005,Australia. amber. The most extensive reviews of this deposit are those 1Correspondingauthor (e-mail: [email protected]). of Carpenter et al. (1937) and subsequently McAlpine and Can.J.EarthSci.45:1061–1082(2008) doi:10.1139/E08-049 #2008NRCCanada 1062 Can. J.EarthSci.Vol. 45,2008 Martin (1969). Whereas both provide general overviews of dent that the Cedar Lake and Grassy Lake ambers might be thearthropodfauna,andtheformerincludesdetaileddescrip- related in at least a general sense. It was obvious to J.B. tions for some taxa, an exhaustive taxonomic survey of the Tyrrell that Cedar Lake amber was a secondary deposit that CedarLakefaunahasnotpreviouslybeenattempted. must have originated somewhere within the Cretaceous of Amber along the beaches of Cedar Lake was once so the prairies (Carpenter 1937). By following the course of plentiful that the Hudson’s Bay company collected it for the Saskatchewan River drainage upstream, and identifying manufacturing varnish, and early collecting trips were able potential source rocks that it incises, Carpenter delineated a to collect >100 kg of raw amber with relative ease. Later wide range of Cretaceous strata that may have originally collections sawdiminishingreturns, but still foundfaunal in- contained the amber (Fig. 1). The matter is complicated by clusions in 2% of the raw amber collected (McAlpine and the areal extent of lithologically similar marginal to fully Martin 1969). Construction of the Grand Rapids dam, com- marine sediments, as well as flood plain deposits, that may pleted in 1965, inundated the amber-rich beach. This ren- host amber-rich coal measures. dered the collections at the Harvard University Museum of Fortunately, the arthropod assemblage contained in Cedar Comparative Zoology (Cambridge, Massachusetts), the Lake amber suggests a Campanian age (McAlpine and Mar- Royal Ontario Museum (Toronto, Ontario), and the Cana- tin 1969), which greatly restricts the extent of potential dian National Collection of Insects and Arthropods (Ottawa, source rocks (shaded area of Fig. 1). Even if the arthropods Ontario) the bases for most published work associated with are assigned to a broader time interval (e.g., Santonian to the deposit. Given the importance of Cedar Lake amber, it Maastrichtian), the putative source formations are limited by has been characterized geochemically using both infrared the conditions required for amber accumulation. Large am- spectroscopy (Langenheim and Beck 1965; Kosmowska- ber concentration deposits are generally found in association Ceranowicz et al. 2001; Aber and Kosmowska-Ceranowicz with coal and lignified plant tissues, which are often associ- 2001) and stable isotopes (Nissenbaum and Yakir 1995). In ated withdeltaic andcoal-producing terrestrial andnearshore recent syntheses on amber (e.g., Rice 1980; Poinar 1992; environments (Grimaldi 1996). This focuses attention on the Grimaldi 1996), the provenance of Cedar Lake amber is Foremost Formation, and to a limited extent upon the given merely as ‘‘near Medicine Hat’’, and the deposit has younger Oldman, Dinosaur Park,and Horseshoe Canyon for- been assigned cautiously a Late Cretaceous age. mations, and the older Milk River Formation. Amber has a specific gravity close to that of seawater (*1.025 g/mL), so Grassy Lake amber a concentration deposit within distal, fully marine sediments Amber from Grassy Lake was first mentioned as a deposit is unlikely. This effectively excludes the marine Bearpaw associated with a coal mine ‘‘near Medicine Hat, Alberta’’ Formation overlying the Foremost Formation, and the ma- discovered by P. Boston in 1963 (McAlpine and Martin rine Pakowki Formation that underlies it (Fig. 2). 1966, p. 531). Aside from isolated references in the entomo- Within the Foremost Formation, there are six coal seams logical literature, GrassyLake amber waslargely unexplored that are also referred to as the Taber coal zone. However, it until Pike’s (1995) dissertation on the locality. Pike (1995) is unclear which of these seams outcrop at the Grassy Lake provided a provisional taxonomy for many of the arthropod locality(Pike1995).Inadetaileddescriptionofthedepositio- inclusions, as well as a detailed account of taphonomy and nal environments of the Foremost Formation (Ogunyomi and collection biases. Subsequent collaborations described por- Hills1977),therewasnomentionofamberconcentrationsin tions of the fauna in detail (e.g., Heie and Pike 1992, 1996; any of the six coal seams from the sections logged. Because O’Keefe et al. 1997; Poinar et al. 1997), while others have thesecoal seamslikely representlocalized cyclesoflagoonal reported on additional arthropod groups represented in the and salt marsh deposition in the area (Pike 1995), the possi- deposit (e.g., Yoshimoto 1975; Gagne´ 1977; Wilson 1985; bility exists that amber is not widespread within these units Penney and Selden 2006). Thus, the palaeoentomology of andthatitsoccurrencemaybemildlydiachronous. Grassy Lake amber is somewhat better understood than that of the Cedar Lake deposit. Objectives of study Grassy Lake amber is associated with a thin (*70 cm Despite the longstanding suggestion that the Grassy Lake thick) coal seam and two 30 cm thick overlying shale beds. and Cedar Lake amber deposits are coeval, detailed compar- Largeportionsofthiscoalseam(belowtheambercontaining isons of the amber from these two localities have not been interval) havebeenstrip-mined intheregionsouthofthevil- attempted using multiple lines of geochemical and palaeo- lage of Grassy Lake, Alberta, and the resulting tailings piles biological evidence. The methodologies employed in this arethemainsourceofamber.Theamberformsresistantnod- paper aim to establish new guidelines for comparisons be- ule-shaped masses that weather out of the disturbed sedi- tween in situ and reworked amber deposits and explore the ments and are easily surface-collected, but the distribution of potential effects of long-distance transport on the character- amberwithinthesesedimentsisvariable.Themajorityofcu- istics of amber. The present study also contributes the larg- rated specimens from Grassy Lake are located at the Royal est stable isotopic data set generated from amber to date, Tyrrell Museum in Drumheller, Alberta, Canada. Due to the obtained with sufficient stratigraphic control to address the relatively recent interest in Grassy Lake amber, basic analy- source and fate of these important amber deposits. ses of physical properties, stable isotopic composition, and infrared spectra have been limited todate (e.g.,Zobel 1999). Materials and methods Stratigraphic constraints Materials From these pioneering investigations, it has become evi- SpecimensofGrassyLakeamber werecompared withCe- #2008NRCCanada McKellar etal. 1063 Fig.1.Inset showinglocation ofstudyarea. Subcropdiagram ofthe Oldman andForemostformations inthe prairie provinces, withthe SaskatchewanRiver system superimposed.Light greyshadingindicates the extent ofthe outcrop andsubcropofthe OldmanandForemost formations andthe Belly River Group.Finelines anddarkgreyshadingrepresent the courseofriver drainage andthe position oflakes, respectively. Thisdiagram ismodified from fig. 24.1ofDawsonet al. (1994);Energy,MinesandResourcesCanada’s (1985)Canada— DrainageBasinsmap. darLakeamber,aswellaswithotherCretaceousambersfrom thick) were taken from both freshly fractured and weathered the Horseshoe Canyon Formation (early Maastrichtian) col- surfaces of Grassy Lake amber pieces. Samples were taken lected at localities near Drumheller and Edmonton, Alberta. from a wide range of amber morphological types (isolated Additional comparisons were made with samples with less drips,schlauben,runnels,anddisks)andamberclaritygrades stratigraphicconstraintfromthevicinityofMedicineHat,Al- (opaquetobone,clear,andclearwithavarietyofinclusions). berta (Campanian). Cretaceous amber used in stable isotope These flakes were placed on a NaCl disk and their infrared analyses was collected from the surface of two coal seams absorption was determined in transmitted mode, at wave- separated by 10 m stratigraphically, at the Morrin Bridge lengths ranging from 2.5 to 15 mm (wavenumbers from 650 crossing north of Drumheller (base of section at to4000cm–1),usingaThermo-NicoletNexus470FTIRspec- 51838’50.3@N, 112854’30.8@W); from a single coal seam in a trometer equipped with a Nicolet Continuum IR microscope. road cut near the Royal Tyrrell Museum of Palaeontology, For each spectrum 200 individual interferograms were aver- west of Drumheller (51828’41@N, 112847’20@W); and from a aged. To minimize the continuum and to avoid the necessity singlecoalseamatthebottomofEdmonton’sNorthSaskatch- of baseline corrections, only samples with a consistent thick- ewan River valley (53831’38@N 113829’27@W). In all cases, nesswereanalyzed. theamber sampleswerecollected fromwithincoal seams,or To objectively compare the FTIR spectroscopic results from shales directly adjacent to a coal. Additional samples obtained from western Canadian amber samples, hierarchical within the FTIR spectral library at the University of Alberta, (agglomerative) cluster analysis was undertaken (e.g., Nau- Edmonton, Alberta, came from deposits within Horsethief mann et al. 1991). Fourteen FTIR spectra were included in Canyon, Alberta (near 51832’21.9@N, 112852’06.8@W). Pur- this analysis, representing different weathering states of chased specimens from the Raritan Formation (Turonian) at Grassy Lake amber (n = 8), Cedar Lake amber (n = 2), and Sayerville,NewJersey,USA,werealsoanalyzed,butthereis single exemplars for each of Horsethief Canyon, Medicine littlecontrolovertheircollection. Hat, Edmonton, and Drumheller Valley ambers. The finger- print regions of each spectrum (800–1100 cm–1) were first FTIR spectroscopy normalized to unity, to offset the potential influence of vari- Fourier transform infrared (FTIR) spectroscopy was used able sample thickness and angle of incidence. The clustering to determine the infrared (IR) absorption characteristics of employs Ward’s (1963) minimum variance algorithm, which ambersfromeachdeposit.Numerousthinflakes(*5–10mm was applied to a dissimilarity matrix of squared Euclidean #2008NRCCanada 1064 Can. J.EarthSci.Vol. 45,2008 Fig.2.Stratigraphic diagram forAlberta from Santonian to Maas- air dried, but were not chemically or thermally pre-treated. trichtian Stages,modified fromtext-fig. 1ofBraman (2001)and Amber samples of 2–7 mg were combusted at 800 8C in fig.24.3ofDawsonet al. (1994).Fm.,Formation; Gp.,Group; vacuum-sealed quartz glass tubes, to which CuO (2 g), Cu Mbr.,Member; Maast.,Maastrichtian. (100 mg), and Ag (100 mg) had been added. Values of d13C and dD were obtained from the CO and H O evolved 2 2 during combustion using a Finnigan MAT-252 isotope-ratio mass spectrometer. The results are expressed in d notation relative to Vienna standard mean ocean water (VSMOW) for dD, and the Vienna Pee Dee Belemnite (VPDB) for d13C. The precision for the lab procedures just outlined are ±3% for dD and ±0.1% for d13C. Palaeoentomology A faunal comparison was made between the arthropods from the Grassy Lake and Cedar Lake ambers, using pub- lished records dedicated to these sites (e.g., Pike 1995; Skid- more 1999), as well as the compilation of all isolated taxonomic works that refer to material from either locality. Published images of taxa from these localities are somewhat limited, soin most cases the published identifications are ac- cepted as definitive. Published identifications were aug- mented by firsthand observations of specimens within the Royal Tyrrell Museum collection of Grassy Lake amber made using primary literature and keys (e.g., Goulet and Huber 1993). Selected inclusions were photographed through a Wild M400 Photomakroskop with a Nikon D100 digital camera, and individual images were compiled using Auto-Montage software. The compiled taxonomic list comprising primary and sec- ondary identifications is presented in Table 1. In all cases where the determining authority is unknown or the identifi- cation was uncertain, the taxa are presented with question marks. Comparisons of the Grassy Lake and Cedar Lake amber faunas are made at high levels of taxonomy (order and family), but emphasis is placed upon lower taxonomic ranks (genus and species). Results FTIR spectra Comparisons of the FTIR spectra of ambers from Cedar Lake and Grassy Lake indicate only minor differences (Fig. 3). There is a similar degree of variability within the spectra generated from different forms of Grassy Lake am- ber, as well as from weathered versus fresh surfaces. None- theless, the same absorption peaks can be identified distances to create the resulting dendrogram. Cluster analy- throughout the 8–10 mm (1000–1250 cm–1) range in all sis was performed with MVSP version 3.1 software (Kovach Grassy Lake and Cedar Lake spectra, although they are 1999). slightly more subdued in the latter. This window is consid- ered to represent the primary fingerprinting region for the Stable isotopes spectral characterization of ambers (Langenheim and Beck The carbon (d13C) and hydrogen (dD) stable isotopic 1968). Compared with all of the FTIR spectra we have gen- compositions were measured from portions of the same am- erated from Cretaceous amber localities in Alberta, Cedar ber specimens used for FTIR spectroscopy, as well as from Lake amber is the most similar to Grassy Lake amber additional samples of similar morphology and grade. Speci- (Fig. 3), while amber from Edmonton, Medicine Hat, Horse- mens from Cedar Lake, Edmonton, Drumheller, Morrin thief Canyon, and Drumheller Valley all differ to varying Bridge, and the Raritan Formation were also analyzed iso- degrees. However, because the variability observed within topically, but these did not encompass as wide a range of Grassy Lake FTIR spectra is sufficient to encompass the amber grades and flow morphologies as were recognized at spectra of certain other Late Cretaceous deposits, as well as Grassy Lake. All specimens were fragments from freshly that of Cedar Lake amber, FTIR does not provide an exclu- broken surfaces that were cleaned with distilled water and sive match. For example,amber collected in Horsethief Can- #2008NRCCanada McKellar etal. 1065 Table1.Palaeoentomological comparison (see table noteforexplanation ofsymbols). Grassy Cedar Taxon Lake Lake Source ORDERHYMENOPTERA Superfamily Ceraphronoidea Family Megaspilidae xx (Skidmore 1999) Lygocerusdubitatus Brues,1937 ? (Brues1937) Family Stigmaphronidae xx (Skidmore 1999) Family Ceraphronidae xx (McAlpine andMartin1969) Superfamily Chalcidoidea Family Eulophidae xx (McAlpine andMartin1969) Family Eupelmidae x (Pike1995) Family Mymaridae xx xx (Skidmore 1999) Carpenterianatumida Yoshimoto,1975 xx (Yoshimoto1975) Macalpinia canadensisYoshimoto,1975 xx (Yoshimoto1975) ProtooctonusmasneriYoshimoto,1975 xx (Yoshimoto1975) TriadomerusbulbosusYoshimoto,1975 xx xx (Yoshimoto1975) Family Tetracampidae xx xx (Skidmore 1999) Baeomorphadistincta Yoshimoto,1975 xx xx (Yoshimoto1975) Baeomorphadubitata Brues,1937 xx xx (Yoshimoto1975) Baeomorphaelongata Yoshimoto,1975 xx xx (Yoshimoto1975) Baeomorphaovatata Yoshimoto,1975 xx xx (Yoshimoto1975) Bouceklytus arcuodensYoshimoto,1975 xx (Yoshimoto1975) Distylopusbisegmentus Yoshimoto,1975 xx (Yoshimoto1975) Family Torymidae x (Pike1995) Family Trichogrammatidae xx xx (Skidmore 1999 EnneagmuspristinusYoshimoto,1975 xx (Yoshimoto1975) Superfamily Proctotrupoidea Family Diapriidae xx (Skidmore 1999) Family Proctotrupidae (Serphidae) xx (Skidmore 1999) Superfamily Evanioidea Family Gasteruptiidae xx (Skidmore 1999) Superfamily Ichneumonoidea Family Braconidae xx xx (Skidmore 1999) Diospilusallani Brues, 1937 x (Brues1937) Neoblacusfacialis Brues,1937 x (Brues1937) Pygostolus patriarchicusBrues,1937 x (Brues1937) Family Ichneumonidae xx (Skidmore 1999) Family Paxylommatidae xx (Skidmore 1999) Superfamily Platygastroidea Family Platygastridae xx (Skidmore 1999) Family Scelionidae xx xx (Skidmore 1999) Baryconusfulleri Brues,1937 x (Brues1937) Proteroscelio antennalis Brues,1937 x (Brues1937) Superfamily Chrysidoidea Family Bethylidae xx (Skidmore 1999) Family Chrysididae xx xx (Skidmore 1999) Procleptes carpenteri Evans,1969 x (Evans 1969) Family Dryinidae xx (Skidmore 1999) DryinuscanadensisOlmi, 1995 x (Olmi 1995) Superfamily Cynipoidea Family Protimaspidae xx xx (Skidmore 1999) ProtimaspiscostalisKinsey,1937 x (Kinsey1937) Superfamily Mymarommatoidea Family Mymarommatidae xx xx (Skidmore 1999) Archaerommaminutissima(Brues),1975 xx xx (Yoshimoto1975) Archaerommanearctica Yoshimoto,1975 xx xx (Yoshimoto1975) Superfamily Serphitoidea Family Serphitidae Brues,1937 xx xx (Skidmore 1999) #2008NRCCanada 1066 Can. J.EarthSci.Vol. 45,2008 Table1(continued). Grassy Cedar Taxon Lake Lake Source Serphites paradoxusBrues,1937 x (Brues1937) Superfamily Apoidea Family Sphecidae x x (Pike1995;Evans 1969) LisponemasingularisEvans,1969 x (Evans 1969) Superfamily Vespoidea Family Formicidae xx (Skidmore 1999) Cananeuretusoccidentalis Engel andGrimaldi, 2005 x (Engel andGrimaldi 2005) Canaponedentata Dlussky,1999 x (Dlussky1999) Eotapinomamacalpini Dlussky,1999 x (Dlussky1999) SphecomyrmacanadensisWilson,1985 x (Wilson 1985) ORDERDIPTERA ‘‘Nematocerans’’ Family Anisopodidae xx (Skidmore 1999)nodet. Family Bibionidae xx (McAlpine andMartin1969) Plecia myersiPeterson,1975 x (Peterson 1975) Family Canthyloscelididae (Synneuridae) x (Pike1995) Family Cecidomyiidae xx xx (Skidmore 1999) Cretocatocha mcalpinei Gagne´,1977 xx (Gagne´ 1977) Cretocordylomyia quadriseriesGagne´,1977 xx (Gagne´ 1977) Cretomiastorferejunctus Gagne´, 1977 xx xx (Gagne´ 1977) Cretowinnertzia angustalaGagne´,1977 xx (Gagne´ 1977) Family Ceratopogonidae xx xx (Skidmore 1999) Adelohelea glabraBorkent,1995 xx xx (Borkent 1995) Culicoides canadensis(Boesel), 1937 xx x (Borkent 1995) Culicoides agamusBorkent,1995 xx (Borkent 1995) Culicoides annosusBorkent,1995 xx xx (Borkent 1995) Culicoides bullusBorkent,1995 xx (Borkent 1995) Culicoides filipalpis Borkent,1995 xx xx (Borkent 1995) Culicoides obuncusBorkent,1995 xx xx (Borkent 1995) Culicoides tyrrelli (Boesel), 1937 xx x (Borkent 1995) Dasyheleasp. xx (Skidmore 1999) HeleageronarenatusBorkent,1995 xx xx (Borkent 1995) Atriculicoides globosus(Boesel), 1937 xx x (Borkent 1995) Atriculicoides sp.Borkent,1995 xx xx (Borkent 1995) LeptoconopsprimaevusBorkent,1995 xx xx (Borkent 1995) Minyohelea pumilisBorkent,1995 xx (Borkent 1995) Palaeobrachypogon aquilonius(Boesel), 1937 xx xx (Borkent 1995) Palaeobrachypogon vetus Borkent,1995 xx xx (Borkent 1995) Palaeobrachypogon remmiBorkent,1995 xx xx (Borkent 1995) Peronehelea chrimikalydia Borkent,1995 xx xx (Borkent 1995) Protoculicoides depressusBoesel, 1937 xx x (Borkent 1995) Family Chironomidae xx xx (Skidmore 1999) Metriocnemussp.Poinaret al., 1997 x (Poinaret al. 1997) Metriocnemuscretatus Boesel, 1937 x (Boesel 1937) SpaniotomaconservataBoesel, 1937 x (Boesel 1937) Spaniotoma(Smittia)veta Boesel, 1937 x (Boesel 1937) Family Culicidae x (Poinaret al. 2000) Paleoculicis minutusPoinaret al., 2000 x (Poinaret al. 2000) Family Lygistorrhinidae xx (Blagoderov andGrimaldi 2004) Plesiognoriste carpenteri Blagoderov andGrimaldi, 2004 xx (Blagoderov andGrimaldi 2004) Family Mycetophilidae xx xx (Blagoderov andGrimaldi 2004) SyntemnafissurataBlagoderov andGrimaldi, 2004 xx (Blagoderov andGrimaldi 2004) Saigusaiapikei Blagoderov andGrimaldi, 2004 xx (Blagoderov andGrimaldi 2004) Synaphalongistyla Blagoderov andGrimaldi, 2004 xx (Blagoderov andGrimaldi 2004) NedocosiacanadensisBlagoderov andGrimaldi, 2004 xx (Blagoderov andGrimaldi 2004) Zeliinia occidentalis Blagoderov andGrimaldi, 2004 xx (Blagoderov andGrimaldi 2004) #2008NRCCanada McKellar etal. 1067 Table1(continued). Grassy Cedar Taxon Lake Lake Source Lecadonileia parvistyla Blagoderov andGrimaldi, 2004 xx (Blagoderov andGrimaldi 2004) Family Psychodidae xx (Skidmore 1999) Sycorax sp. xx (Skidmore 1999) Family Scatopsidae xx xx (Skidmore 1999) Family Sciaridae xx xx (Skidmore 1999) Family Tipulidae (Limoniidae) xx xx (Skidmore 1999) Dicranomyiaalbertensis (Krzemin´skiandTeskey), 1987 x xx (Brookset al. 2007) Macalpina incomparabilisKrzemin´skiand Teskey,1987 x xx (Krzemin´skiandTeskey1987) TrichoneuracanadensisKrzemin´skiand Teskey,1987 x (Krzemin´skiandTeskey1987) Brachycera Family Atelestidae xx xx (Grimaldi andCumming1999) Apalocnemis canadambrisGrimaldi andCumming, 1999 x (Grimaldi andCumming1999) ArchichrysotusmanitobusGrimaldi andCumming, 1999 x (Grimaldi andCumming1999) CretodromiaglaesaGrimaldi andCumming, 1999 x x (Grimaldi andCumming1999) Cretoplatypalpus americanusGrimaldi andCumming, 1999 x x (Grimaldi andCumming1999) Mesoplatypalpus carpenteri Grimaldi andCumming, 1999 x (Grimaldi andCumming1999) NemedromiacampaniaGrimaldi andCumming, 1999 x (Grimaldi andCumming1999) Nemedromiatelescopica Grimaldi andCumming, 1999 x (Grimaldi andCumming1999) Family Bombyliidae xx (Skidmore 1999) Family Dolichopodidae ? (Skidmore 1999) Family Empididae xx xx (Skidmore 1999) Family Ironomyiidae x (McAlpine 1973) Cretonomyia pristinaMcAlpine, 1973 x (McAlpine 1973) Family Platypezidae xx (Skidmore 1999) Family Rhagionidae ? (Skidmore 1999) Family Stratiomyidae xx (McAlpine andMartin1969) Cretaceogaster pygmaeusTeskey,1971 x (Teskey 1971) Cyclorrhapha Family Phoridae xx xx (Skidmore, 1999) PrioriphoracanadambraMcAlpineandMartin, 1966 xx (McAlpine andMartin1966) Priophoraintermedia BrownandPike, 1990 xx xx (BrownandPike1990) Priophoralongicostalis BrownandPike, 1990 xx xx (BrownandPike1990) Priophorasetifemoralis BrownandPike, 1990 xx (BrownandPike1990) Family Sciadoceridae xx (Skidmore 1999) SciadophorabostoniMcAlpineandMartin, 1966 xx xx (McAlpine andMartin1966) ORDERHEMIPTERA SuborderAuchenorrhyncha Superfamily Cicadoidea Family Cercopidae xx xx (Skidmore 1999) Family Cicadellidae xx (Skidmore 1999) Family Jascopidae xx xx (Skidmore 1999) Jascopusnotabilis Hamilton, 1971 x (Hamilton 1971) Family Membracidae xx (Skidmore 1999) Superfamily Fulgoroidea Family Caliscelidae xx (Skidmore 1999) Family Cixiidae xx (Skidmore 1999) Family Fulgoridae xx (Skidmore 1999) Family Issidae ? (Skidmore 1999) SuborderHeteroptera Superfamily Cimicoidea Family Nabidae xx (Skidmore 1999) Superfamily Coreoidea Family Anthocoridae x xx (Skidmore1999;McAlpineandMartin1969) Superfamily Reduvioidea Family Reduviidae xx (Skidmore 1999) #2008NRCCanada 1068 Can. J.EarthSci.Vol. 45,2008 Table1(continued). Grassy Cedar Taxon Lake Lake Source SuborderSternorrhyncha Superfamily Aphidoidea Family Adelgidae xx xx (Skidmore 1999) Family Canadaphidae xx xx (Skidmore 1999) Alloambriacaudata Richards, 1966 xx (Richards 1966) Alloambriainfelicis KaniaandWegierek, 2005 x (Kania andWegierek 2005) Canadaphiscarpenteri Essig,1937 x x (Pike1995;Essig1937) PseudambrialongirostrisRichards, 1966 xx (Richards 1966) Family Cretamyzidae Heie andPike,1992 x (Heie andPike1992) Cretamyzuspikei Heie andPike,1992 x (Heie andPike1992) Family Drepanosiphidae xx (Richards 1966) Aniferella bostoniRichards, 1966 xx (Richards 1966) Family Palaeoaphidae xx xx (Kania andWegierek 2005) AmbaraphiscostalisRichards, 1966 xx (Richards 1966) Ambaraphiskotejai Kania andWegierek, 2005 x (Kania andWegierek 2005) Longiradiusfoottitti Heie, 2006 x (Heie 2006) Palaeoaphis archimediaRichards, 1966 xx (Richards 1966) Palaeoaphidiella abdominalis Heie,1996 x (Heie 1996) Family Tajmyraphididae Grassyaphispikei Heie,1996 x (Heie, 1996) Family indet. Canaphisalbertensis Heie, 2006 x (Heie 2006) Superfamily Coccoidea Electrococcus canadensisBeardsley, 1969 x (Beardsley 1969) Neutrococcus albertaensisPike, 1995 x (Pike1995)nom.nudem Superfamily Phylloxeroidea Family Mesozoicaphididae Heie andPike,1992 x (Heie andPike1992) Albertaphis longirostrisHeie andPike,1992 x (Heie andPike1992) CalgariaphisunguiferaHeie andPike, 1992 x (Heie andPike1992) CampaniaphisalbertaeHeie andPike, 1992 x (Heie andPike1992) MesozoicaphuscanadensisHeie andPike, 1992 x (Heie andPike1992) Mesozoicaphuselectri Heie andPike, 1992 x (Heie andPike1992) MesozoicaphusparvaHeie andPike, 1992 x (Heie andPike1992) Mesozoicaphustuberculata Heie and Pike,1992 x (Heie andPike1992) Superfamily Psylloidea Family Psyllidae xx xx (Skidmore 1999) ORDERCOLEOPTERA Family Bruchidae x (Poinar2005) MesopachymerusantiquaPoinar,2005 x (Poinar2005) Family Carabidae (larva) ? (Skidmore 1999) Family Chrysomelidae xx (Skidmore 1999) Family Cleridae xx (Skidmore 1999) Family Curculionidae xx (Skidmore 1999) Family Dascillidae xx (Skidmore 1999) Family Eucnemidae x (Pike1995) Family Helodidae ? (Skidmore 1999) Family Mordellidae xx (Skidmore 1999) Family Orthoperidae ? (Skidmore 1999) Family Scydmaenidae xx xx (Skidmore 1999) Palaeoleptochromus schaufussiO’Keefe,1997 x (O’Keefe et al.1997) Family Staphylinidae xx xx (Skidmore 1999) Family Trogositidae ? (Skidmore 1999) 1 (Skidmore 1999) ORDERSTREPSIPTERA ORDERNEUROPTERA Family Berothidae ? (Skidmore 1999) #2008NRCCanada McKellar etal. 1069 Table1(continued). Grassy Cedar Taxon Lake Lake Source Family Coniopterygidae xx xx (Skidmore 1999) Family Hemerobiidae x (Klimaszewski andKevan1986) PlesiorobiusCanadensisKlimaszewski andKevan,1986 x (Klimaszewski andKevan1986) Family Raphididae xx (Skidmore 1999) Family Sisyridae xx ? (Skidmore 1999) ORDERPSOCOPTERA Family Liposcelidae xx (Skidmore 1999) Family Sphaeropsocidae xx xx (Skidmore 1999) SphaeropsocitescanadensisGrimaldi andEngel, 2006 x (Grimaldi andEngel 2006) Sphaeropsocussp. xx (Skidmore 1999) ORDERTHYSANOPTERA SuborderTerebrantia Family Aeolothripidae xx (Skidmore 1999) Family Thripidae xx (Skidmore 1999) x (Pike1995) SuborderTubulifera ‘‘StemGroupThysanoptera’’ Family Lophioneuridae xx (Skidmore 1999) ORDERTRICHOPTERA Family Electralbertidae x (Pike1995) Electralberta cretacica Botosaneanu andWichard, 1983 x (Botosaneanu andWichard 1983) Family Psychomyiidae ? (Skidmore 1999) ORDERLEPIDOPTERA Superfamily Incurvaroidea 1 (Skidmore 1999) 4 (Pike1995) ORDERPHASMIDA 1 (Pike1995) ORDERDERMAPTERA 23 (Skidmore 1999) ORDERBLATTARIA 1 (Skidmore 1999) ORDERZORAPTERA 4 (Skidmore 1999) ORDERISOPTERA 1 (Skidmore 1999) ORDERPLECOPTERA ORDERCOLLEMBOLA Family Brachystomellidae xx (Christiansen andPike2002) Bellingeria cornuaChristiansen andPike, 2002 xx (Christiansen andPike2002) Family Hypogastruridae x (Pike1995) Family Isotomidae xx (Christiansen andPike2002) ProtoisotomamicromucraChristiansen andPike xx (Christiansen andPike2002) Family Neanuridae x (Pike1995);below Campanuridaelectra Pike, 1995 x (Pike1995)nom.nudem Pseudoxenylla fovealis Christiansen andPike, 2002 xx (Christiansen andPike2002) Family Oncobryidae Christiansen andPike, 2002 xx (Christiansen andPike2002) Oncobryadecepta Christiansen andPike, 2002 xx (Christiansen andPike2002) Family Poduridae xx (Skidmore 1999) Family Protentomobryidae x x (Pike1995) Protentomobryapalliseri Pike, 1995 x (Pike1995)nom.nudem Protentomobryawalkeri Folsom1937 x (Folsom1937) Family Sminthuridae xx (Skidmore 1999) BrevimucronusanomalusChristiansen andPike xx (Christiansen andPike2002) KeratopygosmegalosChristiansen andPike,2002 xx (Christiansen andPike2002) Family Tomoceridae xx (Skidmore 1999) EntomocerusmirusChristiansen andPike,2002 xx (Christiansen andPike2002) ORDERTHYSANURA Family Lepismatidae xx (Skidmore 1999) #2008NRCCanada 1070 Can. J.EarthSci.Vol. 45,2008 Table1(concluded). Grassy Cedar Taxon Lake Lake Source 1 (Skidmore 1999) ORDERPROTURA ORDERACARI Acariformes SuborderOribatida Family Oribatidae xx xx (Skidmore 1999) Superfamily Damaeoidea xx (Skidmore 1999) Superfamily Gymnodamaeoidea Family Gymnodamaeidae xx (Skidmore 1999) Superfamily Hermannielloidea xx (Skidmore 1999) Superfamily Hermannioidea Family Hermanniidae xx (Skidmore 1999) Superfamily Nothroidea Family Camisiidae xx (Skidmore 1999) Superfamily Oppioidea xx (Skidmore 1999) SuborderProstigmata Superfamily Bdelloidea Family Bdellidae xx xx (Skidmore 1999) Bdella vetusta Ewing,1937 x (Ewing1937) Family Cunaxidae xx (Skidmore 1999) Superfamily Erythroidea xx (Skidmore 1999) Family Erythraeidae xx xx (Skidmore 1999) Leptussp. xx (Skidmore 1999) Superfamily Eupodoidea Family Eupodidae xx (Skidmore 1999) Parasitiformes SuborderMesostigmata Superfamily Parasitoidea Family Parasitidae xx (Skidmore 1999) ORDERARANEAE Family Araneidae ? xx (Skidmore 1999) Family Ctenidae xx (Skidmore 1999) Family Erigonidae ? (Skidmore 1999) Family Huttoniidae xx xx (Skidmore 1999;PenneyandSelden2006) Family Lagonomegopidae x (Penney2004) Grandoculuschemahawinensis Penney,2004 x (Penney2004) Family Linyphiidae xx (Skidmore 1999) Family Oonopidae x (Penney2006) OrchestinaalbertensisPenney, 2006 x (Penney2006) Family Theridiidae ? xx (Skidmore 1999) 4 (Skidmore 1999) ORDERPSEUDOSCORPIONIDA Note: ‘‘x’’markersindicatethespecimenavailabilitywithinstudiesoneachtaxon:‘‘x’’indicatesthatauthorsonlyobservedmaterialfromonelocality, whereas‘‘xx’’indicatesspecimensfrombothlocalitieswereobserved.Abold‘‘x’’denotesataxonthatwasconfirmedbyMcKellarinapreliminarystudy ofaportionoftheRoyalTyrrellMuseum’sGrassyLakeambercollection.Rareordersaremarkedwiththenumberofknownspecimens,andallcommon orderswereobservedbyMcKellar.TheworksofMcAlpineandMartin,aswellasmanyotherdeterminingauthorities,arecitedinSkidmore(1999).The mostrecenttaxonomicsourceavailableislistedpreferentiallyhere,forthesakeofbrevity. yon produces a fingerprint quite similar to that of Grassy Stable isotopes Lake amber, but lacks the absorption peak near 1240 cm–1 The stable isotopic ratios obtained from North American (8.06 mm) seen in all examples of Grassy Lake material. Cretaceous ambers are presented in Fig. 4. The data have The FTIR spectra of Cedar Lake and Grassy Lake am- also been compiled as a co-isotopic plot (Fig. 5) that in- bers presented here are generally comparable to previous cludes previous measurements from Nissenbaum and Yakir reports (Langenheim and Beck 1968; Aber and Kosmow- (1995). The new raw isotopic data are contained in Appen- ska-Ceranowicz 2001; Kosmowska-Ceranowicz et al. dix A. Both Grassy Lake and Cedar Lake ambers have dD 2001). However, our results are the first to include com- values that exhibit a wide range. In the Grassy Lake parisons with additional Cretaceous ambers from western samples, dD ranges from –351.9% to –265.2%, with a Canada. mean value of –301.8%. The Cedar Lake material pro- #2008NRCCanada

Description:
Values of. d13C and dD were obtained from the CO2 and H2O evolved during combustion and family), but emphasis is placed upon lower taxonomic . Family Bibionidae xx .. This may compromise the utility of d13C values for
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