Correlation of Grassy Lake and Cedar Lake Ambers Using Infrared Spectroscopy, Stable Isotopes, and Palaeoentomology

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 Grassy Lake and Cedar Lake amber deposits of western Canada are among North Ameri- ca’s most famous amber-producing localities. Although it has been suggested for over a century that Cedar Lake amber from western Manitoba may be a secondary deposit having originated from strata in Alberta, this hypothesis has not been tested explicitly using geochemical fingerprinting coupled to comparative analyses of arthropod faunal content. Although there are many amber-containing horizons associated with Cretaceous coals throughout Alberta, most are thermally mature and brittle, thus lacking the resilience to survive long distance transport while preserving intact biotic inclusions. One of the few exceptions is the amber found in situ at Grassy Lake. We present a suite of new analyses from these and other Late Cretaceous ambers from western Canada, including stable isotopes (H and C), Fourier transform infrared (FTIR) spectra, and an updated faunal compendium for the Grassy and Cedar lakes arthropod assemblages. When combined with amber’s physical properties and stratigraphic constraints, the results of these analyses confirm that Cedar Lake amber is derived directly from the Grassy Lake amber deposit or an immediate correlative equivalent. This enables the palaeoenvir- onmental context of Grassy Lake amber to be extended to the Cedar Lake deposit, making possible a more inclusive survey of Cretaceous arthropod faunas. Re´sume´ : Les gisements ambrife`res du Cre´tace´ supe´rieur de Grassy Lake et Cedar Lake, dans l’Ouest canadien, figurent parmi les sites de production d’ambre les plus ce´le`bres d’Ame´rique du Nord. Bien que, pendant plus d’un sie`cle, on ait soutenu que le gisement de Cedar Lake, de l’Ouest du Manitoba, pourrait eˆtre un de´poˆt secondaire ayant son origine dans des strates albertaines, cette hypothe`se n’a pas e´te´ mise explicitement a` l’e´preuve a` l’aide de me´thodes geóchimiques combineés a` des analyses comparatives de la faune d’arthropodes qu’il contient. S’il existe, dans toute l’Alberta, de nom- breux horizons ambrife`res associe´s aux charbons cre´tace´s, dans la plupart des cas, l’ambre qu’ils renferment est thermique- ment mature et cassant, ne pouvant donc pas re´sister a` un transport sur de longues distances tout en pre´servant des inclusions biotiques intactes. Une des rares exceptions est l’ambre trouve´ en place a` Grassy Lake. Nous pre´sentons une se´rie de nouvelles analyses de spećimens provenant de ces gisements et d’autres gisements ambrife`res du Cre´tace´ tardif de l’Ouest canadien, dont des analyses des isotopes stables (H et C) et des spectres infrarouges par transformeé de Fourier (IRTF), ainsi qu’un compendium faunique a` jour des assemblages d’arthropodes de Grassy Lake et Cedar Lake. Jumele´s aux donneés sur les proprie´te´s physiques et aux contraintes stratigraphiques des ambres, les re´sultats de ces analyses con- firment que l’ambre de Cedar Lake est directement de´rive´ du gisement ambrife`re de Grassy Lake ou d’un e´quivalent corre´- latif imme´diat. Ceci permet d’e´largir le contexte paleóenvironnemental de Grassy Lake au gisement de Cedar Lake, rendant ainsi possible un recensement plus complet des faunes d’arthropodes cre´taceés. [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- Received 8 February 2008. Accepted 16 September 2008. quences for terrestrial arthropods. Published on the NRC Research Press Web site at cjes.nrc.ca on 3 November 2008. Cedar Lake amber The study of Cedar Lake amber can be traced as far back as Paper handled by Associate Editor C. Hillaire-Marcel. 1890 and includes such famous personages as the geologist R.C. McKellar,1 A.P. Wolfe, and K. Muehlenbachs. Joseph Burr Tyrrell and the palaeoentomologist Frank Car- Department of Earth and Atmospheric Sciences, 1-26 Earth penter (Grimaldi 1996). Amber from this locality has been re- Sciences Building, University of Alberta, Edmonton, ferred to as chemawinite or cedarite (Aber and Kosmowska- AB T6G 2E3, Canada. Ceranowicz 2001), and the former name has been incorrectly R. Tappert. School of Earth and Environmental Sciences, extended by some authors to encompass the bulk of Canadian University of Adelaide, South Australia, 5005, Australia. amber. The most extensive reviews of this deposit are those 1Corresponding author (e-mail: [email protected]). of Carpenter et al. (1937) and subsequently McAlpine and

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Martin (1969). Whereas both provide general overviews of dent that the Cedar Lake and Grassy Lake ambers might be the arthropod fauna, and the former includes detailed descrip- 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 Cedar Lake fauna has not previously been attempted. 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 saw diminishing returns, but still found faunal 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 with deltaic and coal-producing terrestrial and nearshore 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, Grassy Lake amber was largely 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 (Pike 1995). In a detailed description of the depositio- inclusions, as well as a detailed account of taphonomy and nal environments of the Foremost Formation (Ogunyomi and collection biases. Subsequent collaborations described por- Hills 1977), there was no mention of amber concentrations in 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 these coal seams likely represent localized cycles of lagoonal 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 and that its occurrence may be mildly diachronous. 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- Large portions of this coal seam (below the amber containing isons of the amber from these two localities have not been interval) have been strip-mined in the region south of the vil- 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 are the main source of amber. The amber forms resistant nod- 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- amber within these sediments is variable. The majority of cu- 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 to date (e.g., Zobel 1999). Materials and methods Stratigraphic constraints Materials From these pioneering investigations, it has become evi- Specimens of Grassy Lake amber were compared with Ce-

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Fig. 1. Inset showing location of study area. Subcrop diagram of the Oldman and Foremost formations in the prairie provinces, with the Saskatchewan River system superimposed. Light grey shading indicates the extent of the outcrop and subcrop of the Oldman and Foremost formations and the Belly River Group. Fine lines and dark grey shading represent the course of river drainage and the position of lakes, respectively. This diagram is modified from fig. 24.1 of Dawson et al. (1994); Energy, Mines and Resources Canada’s (1985) Canada— Drainage Basins map.

dar Lake amber, as well as with other Cretaceous ambers from 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, and disks) and amber clarity grades stratigraphic constraint from the vicinity of Medicine Hat, Al- (opaque to bone, clear, and clear with a variety of inclusions). 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 to 4000 cm–1), using a Thermo-Nicolet Nexus 470 FTIR spec- 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 single coal seam at the bottom of Edmonton’s North Saskatch- of baseline corrections, only samples with a consistent thick- ewan River valley (53831’38@N 113829’27@W). In all cases, ness were analyzed. the amber samples were collected from within coal 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, New Jersey, USA, were also analyzed, but there is single exemplars for each of Horsethief Canyon, Medicine little control over their collection. Hat, Edmonton, and Drumheller Valley ambers. The fingerprint 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 ambers from each deposit. Numerous thin flakes (*5–10 mm was applied to a dissimilarity matrix of squared Euclidean

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Fig. 2. Stratigraphic diagram for Alberta from Santonian to Maas- air dried, but were not chemically or thermally pre-treated. trichtian Stages, modified from text-fig. 1 of Braman (2001) and Amber samples of 2–7 mg were combusted at 800 8Cin fig. 24.3 of Dawson et 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 13 d C and dD were obtained from the CO2 and H2O evolved 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 published 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, so in most cases the published identifications are accepted as definitive. Published identifications were augmented 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 secondary identifications is presented in Table 1. In all cases where the determining authority is unknown or the identification 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 amber, 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 considered 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-

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Table 1. Palaeoentomological comparison (see table note for explanation of symbols).

Grassy Cedar Taxon Lake Lake Source ORDER HYMENOPTERA Superfamily Ceraphronoidea Family Megaspilidae xx (Skidmore 1999) Lygocerus dubitatus Brues, 1937 ? (Brues 1937) Family Stigmaphronidae xx (Skidmore 1999) Family Ceraphronidae xx (McAlpine and Martin 1969) Superfamily Chalcidoidea Family Eulophidae xx (McAlpine and Martin 1969) Family Eupelmidae x (Pike 1995) Family Mymaridae xx xx (Skidmore 1999) Carpenteriana tumida Yoshimoto, 1975 xx (Yoshimoto 1975) Macalpinia canadensis Yoshimoto, 1975 xx (Yoshimoto 1975) Protooctonus masneri Yoshimoto, 1975 xx (Yoshimoto 1975) Triadomerus bulbosus Yoshimoto, 1975 xx xx (Yoshimoto 1975) Family Tetracampidae xx xx (Skidmore 1999) Baeomorpha distincta Yoshimoto, 1975 xx xx (Yoshimoto 1975) Baeomorpha dubitata Brues, 1937 xx xx (Yoshimoto 1975) Baeomorpha elongata Yoshimoto, 1975 xx xx (Yoshimoto 1975) Baeomorpha ovatata Yoshimoto, 1975 xx xx (Yoshimoto 1975) Bouceklytus arcuodens Yoshimoto, 1975 xx (Yoshimoto 1975) Distylopus bisegmentus Yoshimoto, 1975 xx (Yoshimoto 1975) Family Torymidae x (Pike 1995) Family Trichogrammatidae xx xx (Skidmore 1999 Enneagmus pristinus Yoshimoto, 1975 xx (Yoshimoto 1975) 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) Diospilus allani Brues, 1937 x (Brues 1937) Neoblacus facialis Brues, 1937 x (Brues 1937) Pygostolus patriarchicus Brues, 1937 x (Brues 1937) Family Ichneumonidae xx (Skidmore 1999) Family Paxylommatidae xx (Skidmore 1999) Superfamily Platygastroidea Family Platygastridae xx (Skidmore 1999) Family Scelionidae xx xx (Skidmore 1999) Baryconus fulleri Brues, 1937 x (Brues 1937) Proteroscelio antennalis Brues, 1937 x (Brues 1937) 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) Dryinus canadensis Olmi, 1995 x (Olmi 1995) Superfamily Cynipoidea Family Protimaspidae xx xx (Skidmore 1999) Protimaspis costalis Kinsey, 1937 x (Kinsey 1937) Superfamily Mymarommatoidea Family Mymarommatidae xx xx (Skidmore 1999) Archaeromma minutissima (Brues), 1975 xx xx (Yoshimoto 1975) Archaeromma nearctica Yoshimoto, 1975 xx xx (Yoshimoto 1975) Superfamily Serphitoidea Family Serphitidae Brues, 1937 xx xx (Skidmore 1999)

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Table 1 (continued).

Grassy Cedar Taxon Lake Lake Source Serphites paradoxus Brues, 1937 x (Brues 1937) Superfamily Apoidea Family Sphecidae x x (Pike 1995; Evans 1969) Lisponema singularis Evans, 1969 x (Evans 1969) Superfamily Vespoidea Family Formicidae xx (Skidmore 1999) Cananeuretus occidentalis Engel and Grimaldi, 2005 x (Engel and Grimaldi 2005) Canapone dentata Dlussky, 1999 x (Dlussky 1999) Eotapinoma macalpini Dlussky, 1999 x (Dlussky 1999) Sphecomyrma canadensis Wilson, 1985 x (Wilson 1985) ORDER DIPTERA ‘‘Nematocerans’’ Family Anisopodidae xx (Skidmore 1999) no det. Family Bibionidae xx (McAlpine and Martin 1969) Plecia myersi Peterson, 1975 x (Peterson 1975) Family Canthyloscelididae (Synneuridae) x (Pike 1995) Family Cecidomyiidae xx xx (Skidmore 1999) Cretocatocha mcalpinei Gagne´, 1977 xx (Gagne´ 1977) Cretocordylomyia quadriseries Gagne´, 1977 xx (Gagne´ 1977) Cretomiastor ferejunctus Gagne´, 1977 xx xx (Gagne´ 1977) Cretowinnertzia angustala Gagne´, 1977 xx (Gagne´ 1977) Family Ceratopogonidae xx xx (Skidmore 1999) Adelohelea glabra Borkent, 1995 xx xx (Borkent 1995) Culicoides canadensis (Boesel), 1937 xx x (Borkent 1995) Culicoides agamus Borkent, 1995 xx (Borkent 1995) Culicoides annosus Borkent, 1995 xx xx (Borkent 1995) Culicoides bullus Borkent, 1995 xx (Borkent 1995) Culicoides filipalpis Borkent, 1995 xx xx (Borkent 1995) Culicoides obuncus Borkent, 1995 xx xx (Borkent 1995) Culicoides tyrrelli (Boesel), 1937 xx x (Borkent 1995) Dasyhelea sp. xx (Skidmore 1999) Heleageron arenatus Borkent, 1995 xx xx (Borkent 1995) Atriculicoides globosus (Boesel), 1937 xx x (Borkent 1995) Atriculicoides sp. Borkent, 1995 xx xx (Borkent 1995) Leptoconops primaevus Borkent, 1995 xx xx (Borkent 1995) Minyohelea pumilis Borkent, 1995 xx (Borkent 1995) Palaeobrachypogon aquilonius (Boesel), 1937 xx xx (Borkent 1995) Palaeobrachypogon vetus Borkent, 1995 xx xx (Borkent 1995) Palaeobrachypogon remmi Borkent, 1995 xx xx (Borkent 1995) Peronehelea chrimikalydia Borkent, 1995 xx xx (Borkent 1995) Protoculicoides depressus Boesel, 1937 xx x (Borkent 1995) Family Chironomidae xx xx (Skidmore 1999) Metriocnemus sp. Poinar et al., 1997 x (Poinar et al. 1997) Metriocnemus cretatus Boesel, 1937 x (Boesel 1937) Spaniotoma conservata Boesel, 1937 x (Boesel 1937) Spaniotoma (Smittia) veta Boesel, 1937 x (Boesel 1937) Family Culicidae x (Poinar et al. 2000) Paleoculicis minutus Poinar et al., 2000 x (Poinar et al. 2000) Family Lygistorrhinidae xx (Blagoderov and Grimaldi 2004) Plesiognoriste carpenteri Blagoderov and Grimaldi, 2004 xx (Blagoderov and Grimaldi 2004) Family Mycetophilidae xx xx (Blagoderov and Grimaldi 2004) Syntemna fissurata Blagoderov and Grimaldi, 2004 xx (Blagoderov and Grimaldi 2004) Saigusaia pikei Blagoderov and Grimaldi, 2004 xx (Blagoderov and Grimaldi 2004) Synapha longistyla Blagoderov and Grimaldi, 2004 xx (Blagoderov and Grimaldi 2004) Nedocosia canadensis Blagoderov and Grimaldi, 2004 xx (Blagoderov and Grimaldi 2004) Zeliinia occidentalis Blagoderov and Grimaldi, 2004 xx (Blagoderov and Grimaldi 2004)

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Table 1 (continued).

Grassy Cedar Taxon Lake Lake Source Lecadonileia parvistyla Blagoderov and Grimaldi, 2004 xx (Blagoderov and Grimaldi 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) Dicranomyia albertensis (Krzemin´ski and Teskey), 1987 x xx (Brooks et al. 2007) Macalpina incomparabilis Krzemin´ski and Teskey, 1987 x xx (Krzemin´ski and Teskey 1987) Trichoneura canadensis Krzemin´ski and Teskey, 1987 x (Krzemin´ski and Teskey 1987) Brachycera Family Atelestidae xx xx (Grimaldi and Cumming 1999) Apalocnemis canadambris Grimaldi and Cumming, 1999 x (Grimaldi and Cumming 1999) Archichrysotus manitobus Grimaldi and Cumming, 1999 x (Grimaldi and Cumming 1999) Cretodromia glaesa Grimaldi and Cumming, 1999 x x (Grimaldi and Cumming 1999) Cretoplatypalpus americanus Grimaldi and Cumming, 1999 x x (Grimaldi and Cumming 1999) Mesoplatypalpus carpenteri Grimaldi and Cumming, 1999 x (Grimaldi and Cumming 1999) Nemedromia campania Grimaldi and Cumming, 1999 x (Grimaldi and Cumming 1999) Nemedromia telescopica Grimaldi and Cumming, 1999 x (Grimaldi and Cumming 1999) Family Bombyliidae xx (Skidmore 1999) Family Dolichopodidae ? (Skidmore 1999) Family Empididae xx xx (Skidmore 1999) Family Ironomyiidae x (McAlpine 1973) Cretonomyia pristina McAlpine, 1973 x (McAlpine 1973) Family Platypezidae xx (Skidmore 1999) Family Rhagionidae ? (Skidmore 1999) Family Stratiomyidae xx (McAlpine and Martin 1969) Cretaceogaster pygmaeus Teskey, 1971 x (Teskey 1971) Cyclorrhapha Family Phoridae xx xx (Skidmore, 1999) Prioriphora canadambra McAlpine and Martin, 1966 xx (McAlpine and Martin 1966) Priophora intermedia Brown and Pike, 1990 xx xx (Brown and Pike 1990) Priophora longicostalis Brown and Pike, 1990 xx xx (Brown and Pike 1990) Priophora setifemoralis Brown and Pike, 1990 xx (Brown and Pike 1990) Family Sciadoceridae xx (Skidmore 1999) Sciadophora bostoni McAlpine and Martin, 1966 xx xx (McAlpine and Martin 1966) ORDER HEMIPTERA Suborder Auchenorrhyncha Superfamily Cicadoidea Family Cercopidae xx xx (Skidmore 1999) Family Cicadellidae xx (Skidmore 1999) Family Jascopidae xx xx (Skidmore 1999) Jascopus notabilis 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) Suborder Heteroptera Superfamily Cimicoidea Family Nabidae xx (Skidmore 1999) Superfamily Coreoidea Family Anthocoridae x xx (Skidmore 1999; McAlpine and Martin 1969) Superfamily Reduvioidea Family Reduviidae xx (Skidmore 1999)

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Table 1 (continued).

Grassy Cedar Taxon Lake Lake Source Suborder Sternorrhyncha Superfamily Aphidoidea Family Adelgidae xx xx (Skidmore 1999) Family Canadaphidae xx xx (Skidmore 1999) Alloambria caudata Richards, 1966 xx (Richards 1966) Alloambria infelicis Kania and Wegierek, 2005 x (Kania and Wegierek 2005) Canadaphis carpenteri Essig, 1937 x x (Pike 1995; Essig 1937) Pseudambria longirostris Richards, 1966 xx (Richards 1966) Family Cretamyzidae Heie and Pike, 1992 x (Heie and Pike 1992) Cretamyzus pikei Heie and Pike, 1992 x (Heie and Pike 1992) Family Drepanosiphidae xx (Richards 1966) Aniferella bostoni Richards, 1966 xx (Richards 1966) Family Palaeoaphidae xx xx (Kania and Wegierek 2005) Ambaraphis costalis Richards, 1966 xx (Richards 1966) Ambaraphis kotejai Kania and Wegierek, 2005 x (Kania and Wegierek 2005) Longiradius foottitti Heie, 2006 x (Heie 2006) Palaeoaphis archimedia Richards, 1966 xx (Richards 1966) Palaeoaphidiella abdominalis Heie, 1996 x (Heie 1996) Family Tajmyraphididae Grassyaphis pikei Heie, 1996 x (Heie, 1996) Family indet. Canaphis albertensis Heie, 2006 x (Heie 2006) Superfamily Coccoidea Electrococcus canadensis Beardsley, 1969 x (Beardsley 1969) Neutrococcus albertaensis Pike, 1995 x (Pike 1995) nom. nudem Superfamily Phylloxeroidea Family Mesozoicaphididae Heie and Pike, 1992 x (Heie and Pike 1992) Albertaphis longirostris Heie and Pike, 1992 x (Heie and Pike 1992) Calgariaphis unguifera Heie and Pike, 1992 x (Heie and Pike 1992) Campaniaphis albertae Heie and Pike, 1992 x (Heie and Pike 1992) Mesozoicaphus canadensis Heie and Pike, 1992 x (Heie and Pike 1992) Mesozoicaphus electri Heie and Pike, 1992 x (Heie and Pike 1992) Mesozoicaphus parva Heie and Pike, 1992 x (Heie and Pike 1992) Mesozoicaphus tuberculata Heie and Pike, 1992 x (Heie and Pike 1992) Superfamily Psylloidea Family Psyllidae xx xx (Skidmore 1999) ORDER COLEOPTERA Family Bruchidae x (Poinar 2005) Mesopachymerus antiqua Poinar, 2005 x (Poinar 2005) 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 (Pike 1995) Family Helodidae ? (Skidmore 1999) Family Mordellidae xx (Skidmore 1999) Family Orthoperidae ? (Skidmore 1999) Family Scydmaenidae xx xx (Skidmore 1999) Palaeoleptochromus schaufussi O’Keefe, 1997 x (O’Keefe et al. 1997) Family Staphylinidae xx xx (Skidmore 1999) Family Trogositidae ? (Skidmore 1999) ORDER STREPSIPTERA 1 (Skidmore 1999) ORDER NEUROPTERA Family Berothidae ? (Skidmore 1999)

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Table 1 (continued).

Grassy Cedar Taxon Lake Lake Source Family Coniopterygidae xx xx (Skidmore 1999) Family Hemerobiidae x (Klimaszewski and Kevan1986) Plesiorobius Canadensis Klimaszewski and Kevan, 1986 x (Klimaszewski and Kevan 1986) Family Raphididae xx (Skidmore 1999) Family Sisyridae xx ? (Skidmore 1999) ORDER PSOCOPTERA Family Liposcelidae xx (Skidmore 1999) Family Sphaeropsocidae xx xx (Skidmore 1999) Sphaeropsocites canadensis Grimaldi and Engel, 2006 x (Grimaldi and Engel 2006) Sphaeropsocus sp. xx (Skidmore 1999) ORDER THYSANOPTERA Suborder Terebrantia Family Aeolothripidae xx (Skidmore 1999) Family Thripidae xx (Skidmore 1999) Suborder Tubulifera x (Pike 1995) ‘‘Stem Group Thysanoptera’’ Family Lophioneuridae xx (Skidmore 1999) ORDER TRICHOPTERA Family Electralbertidae x (Pike 1995) Electralberta cretacica Botosaneanu and Wichard, 1983 x (Botosaneanu and Wichard 1983) Family Psychomyiidae ? (Skidmore 1999) ORDER LEPIDOPTERA Superfamily Incurvaroidea 1 (Skidmore 1999) ORDER PHASMIDA 4 (Pike 1995) ORDER DERMAPTERA 1 (Pike 1995) ORDER BLATTARIA 23 (Skidmore 1999) ORDER ZORAPTERA 1 (Skidmore 1999) ORDER ISOPTERA 4 (Skidmore 1999) ORDER PLECOPTERA 1 (Skidmore 1999) ORDER COLLEMBOLA Family Brachystomellidae xx (Christiansen and Pike 2002) Bellingeria cornua Christiansen and Pike, 2002 xx (Christiansen and Pike 2002) Family Hypogastruridae x (Pike 1995) Family Isotomidae xx (Christiansen and Pike 2002) Protoisotoma micromucra Christiansen and Pike xx (Christiansen and Pike 2002) Family Neanuridae x (Pike 1995); below Campanurida electra Pike, 1995 x (Pike 1995) nom. nudem Pseudoxenylla fovealis Christiansen and Pike, 2002 xx (Christiansen and Pike 2002) Family Oncobryidae Christiansen and Pike, 2002 xx (Christiansen and Pike 2002) Oncobrya decepta Christiansen and Pike, 2002 xx (Christiansen and Pike 2002) Family Poduridae xx (Skidmore 1999) Family Protentomobryidae x x (Pike 1995) Protentomobrya palliseri Pike, 1995 x (Pike 1995) nom. nudem Protentomobrya walkeri Folsom 1937 x (Folsom 1937) Family Sminthuridae xx (Skidmore 1999) Brevimucronus anomalus Christiansen and Pike xx (Christiansen and Pike 2002) Keratopygos megalos Christiansen and Pike, 2002 xx (Christiansen and Pike 2002) Family Tomoceridae xx (Skidmore 1999) Entomocerus mirus Christiansen and Pike, 2002 xx (Christiansen and Pike 2002) ORDER THYSANURA Family Lepismatidae xx (Skidmore 1999)

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Table 1 (concluded).

Grassy Cedar Taxon Lake Lake Source ORDER PROTURA 1 (Skidmore 1999) ORDER ACARI Acariformes Suborder Oribatida 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) Suborder Prostigmata Superfamily Bdelloidea Family Bdellidae xx xx (Skidmore 1999) Bdella vetusta Ewing, 1937 x (Ewing 1937) Family Cunaxidae xx (Skidmore 1999) Superfamily Erythroidea xx (Skidmore 1999) Family Erythraeidae xx xx (Skidmore 1999) Leptus sp. xx (Skidmore 1999) Superfamily Eupodoidea Family Eupodidae xx (Skidmore 1999) Parasitiformes Suborder Mesostigmata Superfamily Parasitoidea Family Parasitidae xx (Skidmore 1999) ORDER ARANEAE Family Araneidae ? xx (Skidmore 1999) Family Ctenidae xx (Skidmore 1999) Family Erigonidae ? (Skidmore 1999) Family Huttoniidae xx xx (Skidmore 1999; Penney and Selden 2006) Family Lagonomegopidae x (Penney 2004) Grandoculus chemahawinensis Penney, 2004 x (Penney 2004) Family Linyphiidae xx (Skidmore 1999) Family Oonopidae x (Penney 2006) Orchestina albertensis Penney, 2006 x (Penney 2006) Family Theridiidae ? xx (Skidmore 1999) ORDER PSEUDOSCORPIONIDA 4 (Skidmore 1999) Note: ‘‘x’’ markers indicate the specimen availability within studies on each taxon: ‘‘x’’ indicates that authors only observed material from one locality, whereas ‘‘xx’’ indicates specimens from both localities were observed. A bold ‘‘x’’ denotes a taxon that was confirmed by McKellar in a preliminary study of a portion of the Royal Tyrrell Museum’s Grassy Lake amber collection. Rare orders are marked with the number of known specimens, and all common orders were observed by McKellar. The works of McAlpine and Martin, as well as many other determining authorities, are cited in Skidmore (1999). The most recent taxonomic source available is listed preferentially here, for the sake of brevity. 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-

# 2008 NRC Canada McKellar et al. 1071

Fig. 3. FTIR spectra. (A) Inset depicting complete Grassy Lake 15 spectrum at wavelengths from 2.5 to 15 mm. Shading highlights the most informative region and fine lines highlight salient peaks. (B) Blow-ups of absorption between 5 and 15 mm, including the fingerprint region (brackets). From top to bottom: Horsethief Canyon, slightly oxidized Maastrichtian sample producing similar spectrum to that seen within Grassy Lake material; Grassy Lake (G.L.) 10, 13, 15, and 5, a range of amber morphologies and grades, showing minor variations within their spectra, and many peaks lost or reduced in G.L. 5 due to weathering effects; Cedar Lake 1, typical spectrum from this locality, showing extreme weathering effects; Medicine Hat, Edmonton, and Drumheller Val- ley samples produced strikingly different spectra from that seen at G.L., whereas the latter two localities have nearly identical spectra and are represented here by a single line. duced dD values of –353.3% to –252.5%, with a mean value of –291.5%. All other North American ambers have much narrower ranges of dD: Edmonton material ranged from –349.2% to –338.6%, with a mean value of –343.4%; amber collected in the Drumheller Valley ranged from –328.2% to –307.8%, with a mean of –320.7%;amber from the ‘‘upper coal seam’’ at the Morrin Bridge locality ranged from –311.8% to –297.2%, with a mean of –303.8%, whereas that from the ‘‘lower coal seam’’ was distinct, ranging from –280.6% to –270.0%, with a mean of –275.9%. Finally, amber from New Jersey’s Raritan Formation displayed values from –248.8% to –227.6%, with a mean of –236.6%. The new dD data for Cedar Lake amber are consistent with the prior results of Nissen- baum and Yakir (1995), who published a single dD value of –272%, well within the range of new measurements presented here. However, their reported range of dD values in amber (–270% to –160%) needs to be significantly extended, given the common occurrence of lower dD values reported here (Fig. 4). With respect to d13C, Grassy Lake amber ranged from –27.1% to –20.9%, with most samples falling between –24.6% and –20.9%. The mean d13C value for Grassy Lake amber was –23.8%. Cedar Lake material sampled here displayed a range of d13C values from –24.8% to –20.0%, with a mean value of –23.3%. Other North American ambers displayed similar degrees of variability in d13C, but slightly different mean values (Fig. 4). d13C of amber from Edmonton ranged from –25.9% to –23.6% (mean: –24.6%); samples from the Drumheller Valley ranged from –26.5% to –20.7% (mean: –22.8%); Morrin Bridge upper and lower coal seams were not distinct and produced values from –27.6% to –23.1% (mean: – 24.8%); and New Jersey amber ranged from –23.2% to – 20.2% (mean: –21.8%). In the case of Cedar Lake amber, our measurements overlap the range of d13C values reported previously (Nissenbaum and Yakir 1995), and furthermore suggest that the –23.9% to –21.4% range in d13C values they reported should be extended to a range of –24.8% to –20.0%. The greater ranges of d13C values observed here (4.8% for Cedar Lake material and 6.2% for Grassy Lake amber) appear to reflect the greater number of analyses reported herein. This may compromise the utility of d13C values for fingerprinting the provenance of amber deposits,

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Fig. 4. Stable isotope measurements for d13C and dD; value ranges, mean values, and sampling intensity within each locality. VPDB, Vienna Pee Dee Belemnite: VSMOW, Vienna standard mean ocean water.

Fig. 5. Stable isotope comparison and context. Stable isotope compositions measured for Grassy Lake and Cedar Lake ambers, as compared with other North American Mesozoic ambers and a wide range of published values. Ellipses outline the range of values seen at the two focal localities. U. Cret., Upper Cretaceous; Eoc., Eocene; Olig., Oligocene; Mio, Miocene.

# 2008 NRC Canada McKellar et al. 1073 since the value ranges for individual deposits appear much Medicine Hat (Campanian, Belly River Group), is physically greater than originally portrayed (Nissenbaum and Yakir different from the material found at Grassy Lake and Cedar 1995). Lake. In all of these additional localities, the amber col- In the co-isotopic plot (Fig. 5), data points from the two lected was typically very small (1–5 mm), drop-shaped or deposits that form the focal point of this study have been cylindrical, and medium to dark in colour (ranging from yel- circumscribed with ellipses to indicate the degree to which low to a deep, rich red, with the bulk of material dark or- their isotopic values overlap, and their relation to the iso- ange or red in colour). Material from all three of these topic composition of ambers from other localities. The first localities tends to be very brittle, so much so that it is diffi- observation is that the dD and d13C values of amber are in cult to collect it in any great quantity. These amber samples general uncorrelated with each other. With respect to the re- do not appear to represent massive flows, but instead iso- sults from Cedar Lake and Grassy Lake ambers, these two lated drips and perhaps the cylindrical fillings of internal populations have by far the greatest dispersion in both d13C resin ducts: samples 1 cm or larger are rare. Pike (1993) in- and dD values. Yet these broad isotopic ranges overlap al- dicated that some of the Maastrichtian amber collected near most completely, with the exception of two extremely nega- Edmonton and Grande Prairie had been observed to contain tive d13C values from Grassy Lake amber, and that the dD insect inclusions, but amber collected from Edmonton dif- values of this deposit are on average about 10% lower, rela- fers substantially in morphology and inclusion abundance, tive to Cedar Lake amber. All other Mesozoic ambers pro- and material collected near Grande Prairie falls within a dif- duced comparatively narrow ranges of dD values and more ferent drainage system from that of Cedar Lake. highly variable ranges of d13C. This does not create an unequivocal match, but strongly suggests that Grassy Lake ma- Palaeoentomology terial is the only single source capable of producing the Taxonomic studies of included arthropods have suggested range of values seen in Cedar Lake amber. Other localities that the Cedar Lake and Grassy Lake amber are similar, usu- (e.g., Drumheller Valley material) may develop broader ally portraying this relatedness as a result of comparable en- ranges in values with future analyses, but the differences in vironmental conditions during their formation (e.g., Penney mean values seen here suggests that the overlap in values and Selden 2006). Unfortunately, the majority of the low- will not be as extensive as it is between Grassy Lake and level taxonomic work has not yet been completed for both Cedar Lake amber. localities, and few studies make direct comparisons between the taxa found within the different deposits. Of the available Amber morphology and physical properties published data, family-level taxonomy is usually the lowest Amber from Grassy Lake and Cedar Lake possess a dis- level achieved. The family-level taxonomy of the Grassy tinctive set of shared physical characteristics. The size of Lake amber and Cedar Lake amber faunas are contrasted in amber masses from Grassy Lake is usually <1 cm, with rare Table 1 and supplemented with any lower-level data avail- specimens reaching up to 3.5 cm in length (see Fig. 6A, and able. At the order level, 10 out of 23 taxa are shared be- compare to fig. 15 of McAlpine and Martin 1969), while tween the two deposits (the 13 that are present only in pieces recovered from Cedar Lake are ‘‘for the most part Grassy Lake material are exceedingly rare taxa within this smaller than a pea [while some] were found as large as a deposit and were likely found due to sampling intensity). At robin’s egg’’ (Tyrrell 1891, p. 14). The majority of amber the level of families or superfamilies, 41 taxa are in com- from both deposits represents portions of massive flows mon, while 72 are unique to one deposit. Comparisons (schlauben) and bizarre disc-shaped morphologies (often in- made above the species level are not very informative be- terpreted as resin pockets within the tree, but seen to include cause most Cretaceous amber faunas are very similar; thus palynomorphs and fungal hyphae within our material). Iso- species are emphasized here. Of the species described from lated droplets and runnels make up the remainder of the these localities, 24 can be found in both localities, while 96 samples (see Fig. 6B). In terms of colour, amber from each are only present within one deposit or the other (see exem- locality ranges from pale yellow to deep orange-red, with plars in Figs. 6C–6F). the most samples a rich orange-yellow. Clarities or grades within each deposit range from very clear to rare examples Discussion of milky or ‘‘bone’’ amber, clouded by minute bubbles. Grassy Lake amber commonly has a thin rind of carbonized A synthesis of the spectroscopic, stable isotopic and ento- material, which is replaced by a thin weathering rind mological data generated from western Canadian Late Creta- (<1 mm thick) in surface exposed specimens. Prolonged ex- ceous ambers is necessary to assess their potential posure creates a granular texture and slightly crumbly amber relationships and, most importantly, the provenance of sec- masses; this is pervasive in small specimens (<5 mm). Cedar ondary deposits such as the Cedar Lake amber. We empha- Lake amber consistently lacks a carbonized rind, but often size that any single fingerprinting methodology is unlikely displays a surface patina or granular texture. Very dark to provide unequivocal correlations, necessitating the use of flow lines are evident within specimens from both localities, multiple approaches. and the incidence of arthropod inclusions is approximately one in every 50 sampled amber pieces (McAlpine and Mar- Amber spectroscopy, weathering, and botanical source tin 1969; Pike 1995). Although compelling similarities exist between the Grassy Amber collected from other Albertan sources, such as the Lake and Cedar Lake FTIR spectra (Fig. 3), these are insuf- vicinity of Drumheller and Edmonton (early Maastrichtian, ficient for unambiguous correlation. In a general sense, this Horseshoe Canyon Fm. in both cases), and the vicinity of is because the FTIR spectra of most Cretaceous ambers have

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Fig. 6. (A) A sample of Grassy Lake amber (surface collected; note colour and size predominance). Scale is in centimetres. (B) Range of amber morphologies and clarity grades: lower half of circle arrangement composed of runnels containing bulk of insect inclusions (polished specimen at 3 o’clock position), as well as stalactites (9 o’clock position) and schlauben (6 o’clock position); disk-shaped masses in upper half of circle, with partially removed carbonaceous rinds (10–11 o’clock), milky weathering rind (1 o’clock), and results of tumble polishing (2 o’clock); inner circlet displays range of colours and shapes seen within isolated droplets; centerpiece runnel displays near opacity due to dark-coloured drying surfaces. Scale is in centimetres. (C) Archaeromma minutissima (Brues; Hymenoptera: Mymarommatidae) female specimen 0.5 mm long, RTMP 96.9.178: this species is found at both Grassy Lake (G.L.) and Cedar Lake (G.L.). (D) A. nearctica Yoshimoto, 1975, male specimen 0.32 mm long, from G.L., RTMP 96.9.176: this species is known from both G.L. and C.L. localities. (E) Paxylomma- tine hymenopteran, 1.3 mm long, RTMP 96.9.192: this subfamily found only at G.L. to date. (F) Serphitid hymenopterans Serphites n. sp. A (R. McKellar, in preparation), each approximately 1.6 mm long, from G.L., RTMP 96.9.183: this genus is restricted to the Cretaceous and is known from both G.L. and C.L. localities. pronounced overarching similarities. This sentiment echoes Fig. 7. Dendrogram of cluster analysis results obtained from the those of Langenheim (2003, p. 155), who suggested that FTIR spectra of western Canadian Cretaceous ambers. Two chemi- ‘‘the older the amber, the more similar the spectra usually cally distinct populations are evident: one containing Grassy Lake, become’’, and furthermore that ‘‘different spectra may well Cedar Lake, and Horsethief Canyon ambers, the other represented indicate different plant sources ... but very similar spectra by samples from Edmonton, Drumheller Valley, and Medicine Hat. do not necessarily mean the same source’’. Cluster analysis of FTIR spectra from western Canadian ambers (n = 14) confirms some of these difficulties, while reducing the un- certainty inherent in purely visual assessments of FTIR spectroscopic results. The resulting dendrogram (Fig. 7) indicates that two distinct populations exist among the analyzed samples: one containing Horsethief Canyon, Cedar Lake, and variably weathered specimens of Grassy Lake amber (n = 11), and the other containing ambers from Drumh- eller Valley, Medicine Hat, and Edmonton (n = 3). The dissimilarities expressed within the smaller cluster are far greater than those between representatives of the larger ‘‘Grassy Lake’’ cluster. These results have several implications. First, spectral differences associated with different weathering states of Grassy Lake amber (Fig. 3) exert only minor effects, given that all specimens, as well as those obtained from Cedar Lake amber, cluster together closely. Second, the two major clusters defined in Fig. 7 are entirely compat- ible with the following assignment to discrete source rocks: allows discrete samples <1 mg to be routinely measured, the Horseshoe Canyon Formation for ambers from Drumhel- typically from freshly fractured surfaces, and without need ler, Medicine Hat, and Edmonton, and the slightly older to embed the amber in KBr prior to analysis. Accordingly, Foremost Formation for the larger ‘‘Grassy Lake’’ cluster we consider the progressive diminution of absorption peaks (Fig. 2). Third, the FTIR results verify that the primary within the ‘‘Grassy Lake’’ amber series (Fig. 3) and corre- spectral impact of subaerial weathering is the progressive sponding cluster analysis results (Fig. 7) to capture differen- muting of absorption peaks, as first proposed for reworked tial weathering among ambers of similar, and possibly Baltic amber (Beck 1986). Cedar Lake amber, which is pre- identical, original provenance. sumed to have been transported fluvially at least 500 km The inferences just drawn are corroborated by the physi- from its source, produces FTIR spectra that are comparable cal properties of the amber in question. Samples obtained to weathered specimens of Grassy Lake amber, both of from Drumheller, Medicine Hat, and Edmonton are repre- which have subdued peaks relative to fresher specimens sentative of the majority of coal-associated Cretaceous am- (Fig. 3). From this assessment, it appears that the variability ber within Alberta: they are highly brittle and typically of FTIR spectra previously reported for Cedar Lake amber darker in colour; whole specimens >1 cm3 are rare. Grassy (Beck 1986; Aber and Kosmowska-Ceranowicz 2001; Kos- Lake amber from the Foremost Formation has both greater mowska-Ceranowicz et al. 2001) merely reflects differences strength and a lighter colour relative to other Alberta am- in weathering states and, potentially, development in FTIR bers, which is likely attributable to a lower degree of ther- spectroscopic technology, rather than multiple sources as mal maturation and oxidation (Poinar and Mastalerz 2000). suggested by some authors (e.g., Christiansen and Pike Much of the Grassy Lake amber is hosted in shale and not 2002). Conventional FTIR analyses involved samples coal, which may diminish the extent to which the amber is >100 mg, typically represented by a composite of small subjected to compaction, heating, and diagenetic alteration fragments embedded in KBr pellets, and resulting in loss of in situ. The implication of these arguments is that weather- control with respect to the weathering states of the materials ing of this amber occurred primarily during secondary trans- being considered. The use of micro-FTIR spectroscopy here port following exhumation. The quality of entomological

# 2008 NRC Canada McKellar et al. 1075

fossil records from both Grassy Lake and Cedar Lake am- resource is unmatched elsewhere in the Cretaceous of west- bers is potentially an important consequence of these collec- ern Canada. tive attributes, all the more since this palaeontological Unfortunately, the chemical changes that occur during

# 2008 NRC Canada 1076 Can. J. Earth Sci. Vol. 45, 2008 amber diagenesis are not completely understood, particularly ever, discrepancies arise between maximum and mean dD for ambers of this age (Grimaldi et al. 2000). This renders values obtained from the two ambers. The *10% increase difficult any meaningful use of the FTIR data in an effort to of mean dD values seen in Cedar Lake amber, relative to suggest a botanical source for these ambers through compar- Grassy Lake specimens, is of the magnitude and direction isons to modern conifer resins. Although a source tree predicted from weathering effects. A relative dD increase of within the Araucariaceae has been tentatively advanced for *19% has been attributed to weathering effects in previous the Grassy Lake and Cedar Lake ambers (Kosmowska- studies (Nissenbaum and Yakir 1995). If Cedar Lake dD Ceranowicz et al. 2001), none of the botanical inclusions or values are corrected for this effect, the resulting values fall palynomorphs observed to date confirm an araucarian source directly within the range of measured Grassy Lake amber tree. More comprehensive accounts of botanical inclusions dD values. and spectroscopic data may shed light on the composition However, such effects are relatively minor in the context of source forests, but no firm conclusions can be drawn at of either the ranges of dD values obtained from Grassy Lake present. and Cedar Lake specimens (almost 70%) or rare extremely negative dD values (e.g., –353.3%; Fig. 4). As with d13C Inferences from the stable isotopic composition of amber values, a potential explanation for the broad range of dD The analysis of amber stable isotopic signatures was initi- values may reside in distinct flow characteristics among the ated by Nissenbaum and Yakir (1995). These authors sur- analyzed samples. Isolated drops and bubble-rich (bone) am- mised that the modest range of d13C values determined ber consistently produced the lowest dD values, while speci- from a range of different ambers makes this characteristic a mens from massive and multilayered flows (schlauben and relatively useful tool for accurate correlation. Our results runnels) anchor the isotopically enriched portion of the dD from western Canadian Cretaceous ambers produced d13C spectrum. The former group is likely to represent constitu- ranges of 6.2% and 4.8% from Grassy Lake and Cedar tive resins that solidified rapidly at the site of discrete inju- Lake, respectively, proving that variability within a single ries, while the latter population may be attributable to more deposit may be considerably larger than previously envis- extensively induced resinosis. As defensive resins contain a aged from a limited number of analyses. Although these re- greater proportion of volatile (mono- and sesqui-) terpenoids sults might be viewed as diminishing the utility of d13Casa relative to constitutive counterparts (Trapp and Croteau fingerprinting tool, the opposite conclusion may prove 2001), the possibility exists that differential degrees of vola- equally tenable. This is because the ranges of d13C values tilization exert a first-order control on amber dD, with from Grassy Lake and Cedar Lake ambers produce overlap greater volatile losses resulting in increased dD values. and produce nearly identical mean values (Fig. 4), despite Finally, the highly negative dD values reported here from the large amplitude of values. The exceptional range of am- western Canadian Cretaceous ambers are noteworthy from a ber d13C values can be reconciled by variable plant palaeo- palaeoclimate perspective. These samples are unlikely to be physiological factors, including growth rate and carbon overprinted by contamination, because most post-depositio- partitioning between different components of the source tree nal isotopic artifacts lead to an increase in dD values. Frac- (Murray et al. 1998). Attendant factors, such as forest can- tionation of dD between modern conifer resins and their opy structure, the presence of pests and pathogens, and pa- local meteoric waters is in the order of –170% to –200% laeoclimate, may also be implicated in modulating amber (Nissenbaum and Yakir 1995), which is directly comparable d13C values. to the dD fractionation reported from many plant lipids In the case of Grassy Lake amber, variability in d13C val- (Sauer et al. 2001). This similarity is unlikely to be fortui- ues appears to correlate with the morphology of the amber tous, given that considerable biochemical commonalities ex- (Appendix A). Resin flow morphologies relating to limb ist among pathways of isoprenoid synthesis (Bouvier et al. breakage and bark wounding have been identified using di- 2005), whether the end product is a plant lipid or, in the rect analogy to modern conifers (Pinaeceae, Cupressaceae, case of amber, a diverse suite of terpenoid moieties. On and Araucariaceae); these morphologies correspond to the these grounds, our results suggest that the water source that most highly negative d13C values from the range observed. supported the Grassy Lake amber-producing forest had on These samples appear representative of relatively small vol- occasion dD values as low as –150% to –180%, with a umes of released resin. On the other hand, the more massive more central tendency in the –60% to –120% range. This flow morphologies that form the majority of recovered am- broad range is difficult to explain, as it is roughly equivalent ber samples can be ascribed to traumatic responses, or in- to modern precipitation dD values spanning southern On- duced resinosis (Grimaldi 1996). These samples are tario (–65%) to the central Northwest Territories (–165%) associated with relatively high d13C values. Although more (Birks et al. 2004). Thus, we suggest that local water sour- analyses are required to firmly establish these relationships, ces may have been highly variable during intervals of amber they remain consistent with the notion that trees experienc- production in the palaeo-forest. This variability could poten- ing the traumatic stresses that engender rapid and copious tially involve dramatic changes in water sources, relative hu- resin production have a reduced ability to metabolically midity, ambient temperatures, or some combination of these fractionate against 13C (Nguyen Tu et al. 1999). Accord- factors, but our data do not provide any clear indication of ingly, the ranges of d13C values reported here may be attrib- cause. uted to variable forest conditions and tree health. As with d13C, the dD values from Cedar Lake and Grassy Palaeoentomological and stratigraphic considerations Lake ambers are highly variable, with substantial but not ex- Because Grassy Lake amber is represented in published clusive overlap (Fig. 5). In contrast to the d13C results, how- works by as many as ten times the number of studied speci-

# 2008 NRC Canada McKellar et al. 1077 mens documented from Cedar Lake material, there is an in- shale that contains the Taber coal zone produce radiometric herent sampling bias in any palaeoentomological correlation ages between 79.0 and 78.2 Ma (Goodwin and Deino 1989; between these deposits. Very few deliberate comparative Pike 1995), constraining the formation of Grassy Lake am- studies of these faunas exist (Table 1), and the present study ber to this interval. is the first to summarize exhaustively the information avail- The Milk River Formation is the nearest underlying non- able to date. Although a number of taxa have been reinvesti- marine unit, which is overlain uncomformably by marine gated, it remains impossible to ascertain to what extent shales of the Pawkowki Formation that separate it from the further examinations of the fauna from each locality will in- lower Foremost Formation (Stelck et al. 1976). The Milk fluence the overall picture of their assemblages. With this in River Formation has been dated at *83.5 Ma (Braman mind, the absence of several rare orders from Cedar Lake 2001). Above the Foremost Formation, the nonmarine Old- amber that occur sporadically in material from Grassy Lake man and Dinosaur Park formations appear in close succes- cannot be viewed as a criterion for differentiation: in all sion (Dodson 1971), but only in the uppermost portion of likelihood sampling intensity can be invoked. Similarly, just the Dinosaur Park Formation is another coal-rich interval over one third of the superfamilies and families from the encountered: the Lethbridge coal zone, dated *74.9 Ma two ambers have shared representation. However, taxonomic (Eberth and Hamblin 1993). Bearpaw Formation marine works at these levels have typically been restricted to one sediments overlie the Lethbridge coal zone (Fig. 2). deposit or the other. If only studies considering both depos- Within this stratigraphic framework, the closest nonmar- its are included, 41% of superfamilies and families occur in ine units to the known source of Grassy Lake amber in the both deposits. These differences provide a good reflection of Taber coal zone are *3 Ma younger (Lethbridge coal zone) the amount of work dedicated to either deposit: of the 72 or *5 Ma older (Milk River Formation). Given a maximum taxa at this rank that have been recorded, only eight are estimated duration for insect species in the order of 3– documented from Cedar Lake amber alone (Table 1). Simi- 10 Ma (Grimaldi and Engel 2005), it becomes highly im- lar one-sidedness is evident from studies at the species level: probable that 24 species would simultaneously bridge either the shared proportion of species is 20% when studies re- of these temporal gaps. Furthermore, the Lethbridge coal stricted to one or the other deposit are considered, but rises zone is not known to produce appreciable quantities of am- to 41% in investigations addressing both Grassy Lake and ber at the type section of the Dinosaur Park Formation Cedar Lake material simultaneously. (Eberth and Hamblin 1993), and the Milk River Formation The two most comprehensive taxonomic inventories to contains neither substantial coals nor concentrations of plant date address portions of the orders Hymenoptera (Yoshimoto remains. The probability that the Cedar Lake fauna origi- 1975) and Diptera (Borkent 1995), providing some indica- nated from anywhere other than the six coal measures of tion of the trend that accompanies increasingly detailed the upper Foremost Formation is significantly reduced by taxonomic scrutiny. These studies portray the Grassy Lake the outlined stratigraphic constraints placed on the palaeoen- and Cedar Lake amber faunas as remarkably similar, sharing tomological record. 7 of 13 hymenopteran species, and 15 of 18 dipteran species. These similarities prompted Borkent (1995) to postu- Conclusions late that both amber deposits formed in similar ecological situations relative to surrounding habitats, given nearly iden- Although Cedar Lake amber cannot be assigned to the tical species and sex ratios observed among representatives Grassy Lake deposit with absolute certainty without charac- of the biting midges (Diptera: Ceratopogonidae). Indeed, terizing each and every potential in situ source, the balance the dipterans in general, and the diminutive ceratopogonids of evidence pointing to a common source is overwhelming. in particular, provide a strong basis for the palaeoentomo- At the very least these two deposits should be treated as logical correlation of the Grassy Lake and Cedar Lake am- equivalent, if not identical. Faunal inclusions show that the bers. Borkent’s (1995) results imply that both ambers two ambers were formed at a similar time and in similar pa- formed in immediate proximity to the mixture of aquatic laeoenvironments. Grassy Lake amber is one of few Alberta habitats required by specific taxa of larval ceratopogonids ambers of sufficient strength to withstand fluvial transport in (e.g., moist sands, wetland pools, salt marshes). Although excess of 500 km while retaining exquisitely preserved ar- this group of insects is known from other Cretaceous ambers thropod inclusions. The physical properties of Cedar Lake (e.g., Grogan and Szadziewski 1988; Pen˜alver et al. 2007), amber, augmented by FTIR spectroscopic and stable isotopic the abundance of shared species in Grassy Lake and Cedar fingerprints, support this correlation. From a purely strati- Lake ambers is noteworthy. This is because traps capable of graphic perspective, coal seams and associated shales from capturing a nearly identical suite of ceratopogonid taxa, in the upper Foremost Formation become the most viable similar sex ratios, are unlikely to have been widespread. source for the Cedar Lake amber deposit and its associated The documentation of at least 24 shared arthropod species fauna. In turn, the Grassy Lake outcrop is the only known in the Grassy Lake and Cedar Lake deposits can be placed upstream seam from the Foremost Formation that produces in a stratigraphic context to further constrain the possible a faunal record of this quality. age of amber formation, as well as the distribution of suit- The Grassy Lake and Cedar Lake amber deposits also able palaeoenvironments. It is difficult to envisage signifi- contribute to evaluations of the effects of long-distance cant production of arthropod-rich amber outside of the transport on the physical and chemical properties of amber. Taber coal zone at the top of the Foremost Formation, as In a general sense, it appears that peaks within FTIR spectra this is one of few lithologies associated with marginal ma- become more subdued as weathering progresses, while dD rine deposition in the Campanian. Bentonites within the values become increasingly greater, and amber specimen

# 2008 NRC Canada 1078 Can. J. Earth Sci. Vol. 45, 2008 size is reduced by comminution during transport. Beyond Braman, D.R. 2001. Terrestrial palynomorphs of the upper Santo- the minor differences between these deposits, their internal nian – ?lowest Campanian Milk River Formation, southern Al- variability is likely an excellent source of information con- berta, Canada. Palynology, 25: 57–107. doi:10.2113/0250057. cerning the conditions that triggered the formation of amber Brooks, S.E., Cumming, J.M., O’Hara, J.E., Skevington, J.H., and within Late Cretaceous forests. If the relationships between Cooper, B.E. 2007. Diptera types in the Canadian National Col- amber morphology and stable isotope composition proposed lection of Insects. Supplement 3rd edition. Research Branch here are verified by studies of additional amber deposits, the Agriculture and Agri-Food Canada electronic publication. Avail- potential exists to specify the underlying environmental cues able from http://www.nadsdiptera.org/Catalogs/CNCtypes/Suppl. that led to massive resin releases in ancient forests. htm [cited 10 December 2007]. Brown, B.V., and Pike, E.M. 1990. Three new fossil phorid flies Acknowledgements (Diptera: Phoridae) from Canadian Late Cretaceous amber. Ca- nadian Journal of Earth Sciences, 27: 845–848. The authors would like to thank Dr. Brian Chatterton (for Brues, C.T. 1937. Superfamilies Ichneumonoidea, Serphoidea, and comments on a draft of this paper, funding, and amber col- Chalcidoidea. In Insects and arachnids from Canadian amber. lecting); Dr. James Gardner of the Royal Tyrrell Museum University of Toronto Studies, Geological Series, 40, pp. 27–44. and Dr. Philip Perkins of Harvard’s Museum of Compara- Carpenter, F.M. 1937. Introduction. In Insects and arachnids from tive Zoology (for access to museum collections); Shane and Canadian amber. University of Toronto Studies, Geological Ser- Vicki Leuck (for literature and field assistance); Allan Lin- ies, 40, pp. 7–13. doe (for amber preparation help); Dr. Thomas Stachel (for Carpenter, F.M., Folsom, J.W., Essig, E.O., Kinsey, A.C., Brues, FTIR equipment use); Amber Garrett (for FTIR assistance); C.T., Boesel, M.W., and Ewing, H. 1937. Insects and arachnids Robert Skidmore and Jim Troubridge of Agriculture Canada from Canadian amber. University of Toronto Studies, Geological (for information on the Canadian National Collection of In- Series, 40, pp. 7–62. sects and Arthropods); and Dr. Enrique Pen˜alver and Dr. Christiansen, K., and Pike, E. 2002. Cretaceous Collembola (Ar- Claude Hillaire-Marcel (for valuable comments as re- thropoda, Hexapoda) from the Upper Cretaceous of Canada. viewers). Funding for this research was provided by the Nat- Cretaceous Research, 23: 165–188. doi:10.1006/cres.2002.0313. ural Sciences and Engineering Research Council of Canada Dawson, F.M., Evans, C.G., Marsh, R., and Richardson, R. 1994. and the Alberta Ingenuity Fund (to McKellar), and Natural Geological history of the Peace River Arch. In Geological atlas of the Western Canada Sedimentary Basin, G.D. Compiled by Sciences and Engineering Research Council of Canada Dis- Mossop and I. Shetson. Canadian Society of Petroleum Geolo- covery Grants to Chatterton (#7949), Muehlenbachs gists and Alberta Research Council, Calgary, Alberta [online]. (#9165), and Wolfe (#250070). Available from www.ags.gov.ab.ca/publications/ ATLAS_WWW/ATLAS.shtml [cited 2 May 2007]. References Dlussky, G.M. 1999. New ants (Hymenoptera, Formicidae) from Aber, S.W., and Kosmowska-Ceranowicz, B. 2001. Bursztyn i inne Canadian amber. Paleontological Journal, 33(4): 409–412. zywice kopalne swiata. 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Pike, E.M. 1993. Amber taphonomy and collecting biases. Palaios, the Canadian National Collection of Insects. Research Branch 8: 411–419. doi:10.2307/3515016. Agriculture and Agri-Food Canada electronic publication. No Pike, E.M. 1994. Historical changes in insect community structure longer available from http://sci.agr.ca/ecorc/amb/amb_e.htm as indicated by hexapods of Upper Cretaceous Alberta (Grassy [cited 26 October 1999]. Lake) amber. Canadian Entomologist, 126: 695–702. Stelck, C.R., Wall, J.H., and Sutherland, G. 1976. Field trip A-5 Pike, E.M. 1995. Amber taphonomy and the Grassy Lake, Alberta, Guidebook: Mesozoic stratigraphy in central Alberta foothills amber fauna. Ph.D. thesis. Department of Biological Sciences, near Drumheller, May 16–18, 1976. Geological Association of The University of Calgary, Calgary, Alta. Canada – Mineralogical Association of Canada Joint Annual Poinar, G.O., Jr. 1992. Life in amber. Stanford University Press, Meeting 1976, Edmonton, Alta., 67 p. Stanford, Calif. Teskey, H.J. 1971. A new soldier fly from Canadian amber (Dip- Poinar, G.O., Jr. 2005. A Cretaceous palm bruchid, Mesopachy- tera: Stratiomyidae). Canadian Entomologist, 103: 1659–1661. merus antiqua, n. gen., n. sp. (Coleoptera: Bruchidae: Pachymer- Trapp, S., and Croteau, R. 2001. Defensive resinosis biosynthesis ini) and biogeographical implications. Proceedings of the in conifers. Annual Review of Plant Physiology and Plant Mole- Entomological Society of Washington, 107(2): 392–397. cular Biology, 52: 689–724. doi:10.1146/annurev.arplant.52.1. Poinar, G.O., Jr., and Mastalerz, M. 2000. Taphonomy of fossilized 689. resins: determining the biostratinomy of amber. Acta Geologica Tyrrell, J.B. 1891. Fossil resin (‘‘amber’’). Summary reports on the Hispanica, 35(1–2): 171–182. operations of the Geological Survey for 1891 Annual Report, 5: Poinar, G.O., Jr., Krantz, G.W., Boucout, A.J., and Pike, T.M. 14–15. [Annual Report.] 1997. A unique Mesozoic parasitic association. Naturwis- Ward, J.H. 1963. Hierarchical grouping to optimize an objective senschaften, 84: 321–322. doi:10.1007/s001140050405. function. Journal of the American Statistical Association, 58: Poinar, G.O., Jr., Zavortink, T.J., Pike, T., and Johnston, P.A. 2000. 236–244. doi:10.2307/2282967. Paleoculicis minutus (Diptera: Culicidae) n. gen., n. sp., from Wilson, E.O. 1985. Ants from the Cretaceous and Eocene amber of Cretaceous Canadian amber, with a summary of described fossil North America. Psyche, 92: 205–216. doi:10.1155/1985/57604. mosquitoes. Acta Geologica Hispanica, 35(1–2): 119–128. Yoshimoto, C.M. 1975. Cretaceous chalcidoid fossils from Cana- Rice, P.C. 1980. Amber: the golden gem of the ages. Van Nostrand dian amber. Canadian Entomologist, 107: 499–532. Reinhold Company. New York, N.Y. Zobel, A.M. 1999. Cedarite and other fossil resins in Canada. In Richards, W.R. 1966. Systematics of fossil aphids from Canadian Investigations into amber: proceedings of the international inter- amber (Homoptera: Aphididae). Canadian Entomologist, 98: disciplinary symposium Baltic amber and other fossil resins, 746–760. Gdansk, Poland, pp. 241–245. Sauer, P.E., Eglinton, T.I., Hayes, J.M., Schimmelmann, A., and Sessions, A.L. 2001. Compound-specific D/H ratios of lipid bio- markers from sediments as a proxy for environmental and cli- Appendix A matic conditions. Geochimica et Cosmochimica Acta, 65: 213– 222. doi:10.1016/S0016-7037(00)00520-2. Table A1 appears on the following page. Skidmore, R.E. 1999. Checklist of Canadian amber inclusions in

# 2008 NRC Canada McKellar et al. 1081

Table A1. Stable isotope measurements from North American Cretaceous ambers.

Sample Location Age d13C(%) dD(%) Amber observation (colour and grade; morphology; notes) Cedar Lake amber (from Harvard’s Museum of Comparative Zoology, all specimens <3 mm in size) CL2 Cedar Lake, Manitoba Campanian –24.5 –353 Deep red; isolated drip; minor weathering CL3 Cedar Lake, Manitoba Campanian –22.7 –288 Medium orange; drip; minor patina CL4 Cedar Lake, Manitoba Campanian –22.8 –253 Medium orange, milky; flow fragment; highly weathered CL5 Cedar Lake, Manitoba Campanian –22.6 –263 Pale yellow, clear; flow fragment; minor patina CL6 Cedar Lake, Manitoba Campanian –20.0 –280 Deep orange; large flow fragment; minor patina CL7 Cedar Lake, Manitoba Campanian –23.6 –309 Orange-red; flow fragment; partly weathered CL8 Cedar Lake, Manitoba Campanian –23.7 –308 Pale orange; isolated drip; minor patina CL9 Cedar Lake, Manitoba Campanian –23.7 –303 Medium orange, clear; flow fragment; blocky and unweathered CL10 Cedar Lake, Manitoba Campanian –24.2 –320 Medium orange; drip fragment; minor patina CL11 Cedar Lake, Manitoba Campanian –24.8 –282 Yellow, clear; tiny isolated drip CL12 Cedar Lake, Manitoba Campanian –24.7 –260 Orange, milky; flow fragment; pervasively weathered CL pub. Cedar Lake, Manitoba Campanian –23.9 –272 (Nissenbaum and Yakir, 1995, table 1) Grassy Lake amber GL1 Grassy Lake, Alberta Campanian –21.3 –285 Deep orange; flattened disk, ‘‘internal’’; thin oxidative rind GL5 Grassy Lake, Alberta Campanian –25.7 –352 Pale orange; isolated drop GL6 Grassy Lake, Alberta Campanian –22.9 –305 Medium orange, clear; massive flow; milky stringers GL7 Grassy Lake, Alberta Campanian –27.1 –310 Bone grade; massive flow; dark orange rind of clear amber GL8 Grassy Lake, Alberta Campanian –20.9 –295 Golden yellow, clear; massive flow; botanical inclusion GL10 Grassy Lake, Alberta Campanian –23.2 –282 Deep orange, clear; massive flow; carbon-rich GL13 Grassy Lake, Alberta Campanian –23.3 –292 Pale yellow, clear; massive flow; palynomorph inclusions GL14 Grassy Lake, Alberta Campanian –23.0 –295 Dark yellow; massive flow; stringer stains, oxidation surfaces GL15 Grassy Lake, Alberta Campanian –24.6 –284 Weathered surface of amber mass (medium orange; flattened disk) GL16 Grassy Lake, Alberta Campanian –23.3 –300 Pale yellow, clear; massive flow; botanical inclusions GL17 Grassy Lake, Alberta Campanian –24.5 –314 Pale yellow, clear; massive flow; botanical inclusions GL19 Grassy Lake, Alberta Campanian –23.4 –265 Yellow; cylindrical drip; weathered GL20 Grassy Lake, Alberta Campanian –24.0 –280 Pale yellow; isolated drip; unweathered GL21 Grassy Lake, Alberta Campanian –25.3 –304 Bone grade; subspherical mass; dark oxidative crust GL22 Grassy Lake, Alberta Campanian –24.6 –290 Mix of bone grade and dark orange; subspherical; carbon rind GL23 Grassy Lake, Alberta Campanian –22.6 –298 Medium orange; cylindrical drip; weathered to granular GL24 Grassy Lake, Alberta Campanian –24.8 –320 Medium orange; isolated drip; weathered to granular GL25 Grassy Lake, Alberta Campanian –23.9 –320 Medium orange; isolated drip; solid, unweathered GL26 Grassy Lake, Alberta Campanian –23.7 –316 Medium orange; isolated drip; solid, unweathered GL27 Grassy Lake, Alberta Campanian –23.4 –302 Medium orange; isolated drip; solid, unweathered GL30 Grassy Lake, Alberta Campanian –25.4 –318 Dark orange; multi-flow runnel; dark flow lines GL31 Grassy Lake, Alberta Campanian –25.7 –305 Dark orange; multi-flow runnel; dark flow lines GL32 Grassy Lake, Alberta Campanian –22.5 –291 Medium orange; flattened ‘‘internal’’ disk GL33 Grassy Lake, Alberta Campanian –22.9 –281 Mottled bone/medium orange; nodular; heavily weathered, soft GL34 Grassy Lake, Alberta Campanian –22.6 –284 Mottled bone/medium orange; nodular; heavily weathered, soft GL35 Grassy Lake, Alberta Campanian –24.1 –317 Dark orange; multi-flow; inclusions: aphids; scelionid; chironomid GL36 Grassy Lake, Alberta Campanian –23.2 –304 Dark orange; multi-flow; inclusions: aphids; indet. dipteran GL37 Grassy Lake, Alberta Campanian –23.8 –312 Medium orange; flattened ‘‘internal’’ disk Edmonton (river valley) amber ERV1 Edmonton, Alberta Maastrichtian –25.9 –349 Medium orange, cloudy; weathered to granular, crumbling ERV2 Edmonton, Alberta Maastrichtian –25.0 –339 Pale yellow; isolated drop; solid ERV3 Edmonton, Alberta Maastrichtian –24.2 –344 Medium orange; cylindrical fragment; crumbling ERV4 Edmonton, Alberta Maastrichtian –25.0 –343 Medium orange; drip end; crumbling

# 2008 NRC Canada 1082 Can. J. Earth Sci. Vol. 45, 2008

Table A1 (concluded).

Sample Location Age d13C(%) dD(%) Amber observation (colour and grade; morphology; notes) ERV5 Edmonton, Alberta Maastrichtian –23.6 –347 Medium orange; sheet flow fragment; solid ERV6 Edmonton, Alberta Maastrichtian –23.8 –339 Medium orange; spherical drop; granular Morrin Bridge Crossing amber (upper and lower coal seams) MUC1 Morrin Bridge, Alberta Maastrichtian –27.6 –297 Reddish-orange; droplet in lignitic shale; crumbly MUC2 Morrin Bridge, Alberta Maastrichtian –23.7 –312 Reddish-orange; droplet in lignitic shale; crumbly MUC3 Morrin Bridge, Alberta Maastrichtian –23.7 –302 Reddish-orange; droplet in lignitic shale; crumbly MLC1 Morrin Bridge, Alberta Maastrichtian –25.4 –270 Reddish-orange; droplet in lignitic shale; crumbly MLC2 Morrin Bridge, Alberta Maastrichtian –25.1 –281 Reddish-orange; droplet in lignitic shale; crumbly MLC3 Morrin Bridge, Alberta Maastrichtian –23.1 –277 Dark orange; droplet in lignitic shale; pervasively weathered Drumheller Valley amber DV1 Drumheller, Alberta Maastrichtian –20.7 –321 Pale orange; cylindrical flow fragment DV2 Drumheller, Alberta Maastrichtian –20.7 –328 Medium orange; cylindrical flow fragment DV3 Drumheller, Alberta Maastrichtian –22.6 –328 Orange; drop; weathered granular with carbon rind DV4 Drumheller, Alberta Maastrichtian –22.1 –315 Yellow; cylindrical flow fragment DV5 Drumheller, Alberta Maastrichtian –26.5 –325 Medium orange; spherical mass, solid DV6 Drumheller, Alberta Maastrichtian –24.1 –308 Dark orange; isolated drop; weathered granular New Jersey Raritan Formation amber (purchased) NJ1 Sayerville, New Jersey Turonian –21.9 –249 Butterscotch; multiple flows; inclusions: erythraeid, ceratopogonid, wood debris NJ2 Sayerville, New Jersey Turonian –23.7 sample Butterscotch; multiple flows; inclusions: indet. Hymenop- leaked tera, bubble-clouding, wood debris NJ3 Sayerville, New Jersey Turonian –20.2 –248 Butterscotch; clouded with bubbles and wood debris NJ4 Sayerville, New Jersey Turonian –23.2 –229 Golden yellow, clear; large flow fragment; unweathered NJ5 Sayerville, New Jersey Turonian –21.3 –228 Golden yellow, clear; large flow fragment; surface patina NJ6 Sayerville, New Jersey Turonian –20.5 –230 Golden yellow, clear; flow tip fragment, subspherical

# 2008 NRC Canada