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Journal of Archaeological Science (1999) 26, 83–94 Article No. jasc.1998.0305, available online at http://www.idealibrary.com on The Fatty Acid Composition of Native Food Plants and Animals of Western Canada M. E. Malainey Department of Anthropology, University of Manitoba, Winnipeg, MB R3T 5V5, Canada R. Przybylski Department of Foods and Nutrition, University of Manitoba, Winnipeg, MB R3T 2N2, Canada B. L. SherriDepartment of Geological Sciences, University of Manitoba, Winnipeg, MB R3T 2N2, Canada (Received 4 November 1997, revised manuscript accepted 10 April 1998) In order to facilitate the identification of residues absorbed into the walls of Late Precontact vessels, a collection of more than 130 native food plants and animals from Western Canada was established. The fatty acid composition of uncooked plant and animal samples were determined using gas chromatography. Hierarchical cluster and principal component analyses show similarities in fatty acid composition of these foods which generally correspond to meaningful categories. Large mammal fat, large herbivore meat, fish, plant greens, roots and berries/seeds can be dierentiated. The fatty acid composition of small mammals was more similar to plants than to large mammals. ? 1999 Academic Press Keywords: NATIVE FOOD, FATTY ACIDS, COMPOSITION, WESTERN CANADA, GAS CHROMATOGRAPHY. Introduction P ottery was commonly manufactured during the Late Precontact Period (2000 to European contact) by the hunting and gathering peoples of Western Canada. Although hundreds of vessels are found in sites dating to this period, the introduction of the European copper kettle in the 18th century led to the rapid disappearance of the Native pottery technology. Relatively little is known about how these conoidal and globular vessels were used, although the presence of carbonized residues suggest many func- tioned as cooking pots (Malainey, 1995). The identifi- cation of residues absorbed in the walls of these vessels may provide information about pottery use and expand our understanding of cooking practices, diet and general subsistence strategies beyond that derivable from traditional sources, such as faunal remains and tool assemblages. Certain aspects of the analysis of absorbed vessel residues remain unchanged since it was first accom- plished by Condamin et al. (1976); in particular, the use of gas chromatography to characterize the lipid component of the residue. In the past 10 years, vessel residue analysts have turned from GC alone to gas chromatography combined with mass spectroscopy (GC/MS) (Deal & Silk, 1988; Deal, 1990; Heron & Pollard, 1988; Heron, 1989; Evershed, Heron & Goad, 1990, 1991; Gerhardt, Searles & Biers, 1990; Heron, Evershed & Goad, 1991; Evershed et al., 1992; Skibo, 1992; Heron & Evershed, 1993; Charters et al., 1995). The higher cost of GC/MS and more limited access to these instruments are serious disadvantages. Therefore, a method of using GC alone to identify vessel residues would be beneficial. Lipids used in residue characterization include sterols, waxes and fatty acids. Sterols are biological regulators which can be used to discriminate animal, plant and animal/plant combinations in the vessel residues (Evershed et al., 1992; Evershed, 1993). The presence of a particular wax, a long-chain alkyl com- pound, enabled the very precise identification of the former contents of a vessel as plants of the Brassica family (Evershed, Heron & Goad, 1991). One major 83 0305–4403/99/010083+12 $30.00/0 ? 1999 Academic Press

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Page 1: Fatty Acids Composition in Canada

Journal of Archaeological Science (1999) 26, 83–94Article No. jasc.1998.0305, available online at http://www.idealibrary.com on

The Fatty Acid Composition of Native Food Plants andAnimals of Western Canada

M. E. Malainey

Department of Anthropology, University of Manitoba, Winnipeg, MB R3T 5V5, Canada

R. Przybylski

Department of Foods and Nutrition, University of Manitoba, Winnipeg, MB R3T 2N2, Canada

B. L. Sherriff

Department of Geological Sciences, University of Manitoba, Winnipeg, MB R3T 2N2, Canada

(Received 4 November 1997, revised manuscript accepted 10 April 1998)

In order to facilitate the identification of residues absorbed into the walls of Late Precontact vessels, a collection ofmore than 130 native food plants and animals from Western Canada was established. The fatty acid composition ofuncooked plant and animal samples were determined using gas chromatography. Hierarchical cluster and principalcomponent analyses show similarities in fatty acid composition of these foods which generally correspond tomeaningful categories. Large mammal fat, large herbivore meat, fish, plant greens, roots and berries/seeds can bedifferentiated. The fatty acid composition of small mammals was more similar to plants than to large mammals.

? 1999 Academic Press

Keywords: NATIVE FOOD, FATTY ACIDS, COMPOSITION, WESTERN CANADA, GASCHROMATOGRAPHY.

Introduction

P ottery was commonly manufactured during theLate Precontact Period (2000 to Europeancontact) by the hunting and gathering peoples

of Western Canada. Although hundreds of vessels arefound in sites dating to this period, the introductionof the European copper kettle in the 18th century ledto the rapid disappearance of the Native potterytechnology. Relatively little is known about how theseconoidal and globular vessels were used, although thepresence of carbonized residues suggest many func-tioned as cooking pots (Malainey, 1995). The identifi-cation of residues absorbed in the walls of thesevessels may provide information about pottery useand expand our understanding of cooking practices,diet and general subsistence strategies beyond thatderivable from traditional sources, such as faunalremains and tool assemblages.

Certain aspects of the analysis of absorbed vesselresidues remain unchanged since it was first accom-plished by Condamin et al. (1976); in particular, the

830305–4403/99/010083+12 $30.00/0

use of gas chromatography to characterize the lipidcomponent of the residue. In the past 10 years, vesselresidue analysts have turned from GC alone to gaschromatography combined with mass spectroscopy(GC/MS) (Deal & Silk, 1988; Deal, 1990; Heron &Pollard, 1988; Heron, 1989; Evershed, Heron & Goad,1990, 1991; Gerhardt, Searles & Biers, 1990; Heron,Evershed & Goad, 1991; Evershed et al., 1992; Skibo,1992; Heron & Evershed, 1993; Charters et al., 1995).The higher cost of GC/MS and more limited access tothese instruments are serious disadvantages. Therefore,a method of using GC alone to identify vessel residueswould be beneficial.

Lipids used in residue characterization includesterols, waxes and fatty acids. Sterols are biologicalregulators which can be used to discriminate animal,plant and animal/plant combinations in the vesselresidues (Evershed et al., 1992; Evershed, 1993). Thepresence of a particular wax, a long-chain alkyl com-pound, enabled the very precise identification of theformer contents of a vessel as plants of the Brassicafamily (Evershed, Heron & Goad, 1991). One major

? 1999 Academic Press

Page 2: Fatty Acids Composition in Canada

84 M. E. Malainey et al.

advantage of employing GC/MS for vessel residueanalysis is its capability to detect and identify traces ofsterols and waxes.

Fatty acids are the major constituents of fats and oilsand occur in nature as triacylglycerols, consisting ofthree fatty acids attached to a glycerol molecule byester-linkages. Unsaturated fatty acids decomposemore readily than saturated fatty acids, sterolsand waxes. In the course of decomposition, simpleaddition reactions may occur at points of unsaturation(Solomons, 1980) or peroxidation may lead to theformation of a variety of volatile and non-volatile endproducts which continue to degrade (Frankel, 1991).Determining the composition of uncooked plants andanimals is an important first step in the identificationof archaeological residues; but because of fatty aciddecomposition, direct comparisons between uncookedplants and animals and archaeological residues are notpossible.

In this study, food plants and animals native toWestern Canada were collected and their fatty acidcompositions were determined using GC.

Experimental Methods

Selection and processing of food samples

The compilation of Manitoba food plants by Shay(1980) and ethnographic accounts of cooking practicesdetermined which samples to include in this study. Theedible plants identified are consistent with thoserecognized in other sources (Gilmore, 1919; Gaertner,1967; Kindscher, 1987; Young, 1993). Over 130 plants,mammals, fish and birds used by natives as food weregathered, mostly from the plains, parkland andsouthern boreal forest of Manitoba. For each plantvariety collected, one specimen was pressed andretained for identification; the others were cleaned withdistilled water and then air dried at room temperature.In cases where there were seasonal differences in theexploitation of the greens, seeds and roots of a plant,the fatty acid composition of each part was analysedseparately. The collection includes berry samples fromWood Mountain, Saskatchewan. Native varieties ofcorn, squash, sunflower and red bean were obtainedfrom the University of North Dakota. Samples ofblack bear, deer, small mammals, grouse and catfishwere donated to the study; whereas samples of bisonmeat, bone marrow and all other fish were purchasedcommercially.

Researchers have found that the fatty acid com-position of some domestic oilseeds are affected byvariety, planting conditions and climatic conditionsduring the course of the growing season (Sheppard,Iverson & Weihrauch, 1978). Most wild plantspecimens were collected in 1994; adequate levels ofrain and sun produced favourable growing conditions.Plant specimens were collected as close as possible totheir optimal time of utilization.

Lipid extraction of food samplesImmediately prior to lipid extraction, dried plantsamples were ground with an electric coffee grinderwhile flesh samples were finely chopped. Lipids wereextracted by homogenization (2#2 min) of 10 g of thesample in a 2:1 v/v mixture of chloroform and metha-nol (2#50 ml) (Folch, Lees & Stanley, 1957). Solidsand the non-lipid component were removed by filteringthe solvent/lipid mixture into a separatory funnel fol-lowed by a distilled water wash (33 ml). After separ-ation, the lower lipid-chloroform layer was collectedand solvents were removed by evaporation. Aftertransferring the sample to a small, pre-weighed vial, thelipid was dried with benzene under nitrogen until itsweight remained constant. Lipid samples were storedin chloroform-methanol (2:1 v/v), flushed with nitrogenin a vial sealed with a teflon-lined cap and placed in a"20)C freezer.

Fatty acids were transesterified by treating 200 mg ofthe total lipid extract dissolved in 1 ml of petroleumether with 12 ml of 0·5 N anhydrous hydrochloric acidin methanol (65–70)C, 60 min). Fatty acids whichoccur in the sample as di- or triglycerides are detachedfrom the glycerol molecule and converted to methylesters. After cooling to room temperature, 5 ml ofiso-octane was added, followed by 6 ml of distilledwater and the sample was mixed. When the upper layerhad clearly separated, 2·5 ml of the fatty acid methylester solution was placed in a vial and crimped with analuminum cap with a teflon lining.

Gas chromatography analysis parametersThe separation was performed on a Hewlett-Packard5890 gas chromatograph fitted with a flame ionizationdetector connected to a Hewlett-Packard 3390 com-puting integrator. Samples were separated using aSupelcowax 10 fused silica capillary column(30 m#0·25 mm I.D., Supelco, Oakville, ON). Anautosampler injected a 1 ìl sample using a split injec-tion system with ratio set at 1:80. Hydrogen was usedas the carrier gas at a linear velocity of 40 cm/s. Thecolumn temperature was programmed from 190 to235)C at 2)C per min; lower and upper temperatureswere held for 3 and 10 min, respectively. Peaks wereidentified through comparisons with several externalqualitative standards (NuCheck Prep, Elysian, MN).Using this procedure, fatty acids are detectable to thenanogram level.

Statistical analysisIn this study, the total fatty acid compositions ofmodern food samples were characterized by hierarchi-cal cluster and principal component analyses. Thesemethods are able to process the high number ofvariables resulting from fatty acid analysis. Further-more, the results of the hierarchical cluster analysis canbe independently verified by the principal component

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Native Food Plants and Animals of Western Canada 85

analysis. The SAS hierarchical cluster analysis pro-gram, PROC CLUSTER, set to determine averagelinkage, was used to identify relationships among thefatty acid compositions of the modern food samples.The clusters generated provided a framework for pre-senting the compositional data. The data was alsoanalysed with principal component analysis, a multi-variate technique, using the SAS program, PROCPRIN.

Results

1.3Average root-mean-square distance

1.2 1.1 1.0 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0

Roots

Mixed

Greens

Berries

Roots

Seeds + berries

Mixed

Berries + nuts

Fish

GreensC

B

A

Large mammal

V

VI

VII

VIII

IX

X

XI

XIII

XII

XIV

IIIDeer fatIIIIV

Squirrel

XV—RootsGrouseCattail seed

Figure 1. Dendrogram of the hierarchical cluster analysis of the fatty acid compositions with subclusters identified.

Hierarchical cluster analysis

Hierarchical cluster analysis of the fatty acid com-position of food samples results in the formation ofthree major clusters, A, B and C, each consisting of anumber of subclusters (Figure 1). The three largeclusters reflect broad similarities in the fatty acidcomposition of its individual members. Cluster Acontains all the large mammal and fish samples.Cluster B contains primarily plant seeds and berries,but some plant roots and three small mammals are also

included. The samples in cluster C are mainly plantgreens and roots, but rosehips and bearberries appearas well. Four samples, deer fat, cattail seeds, grouseand squirrel, did not readily cluster with any othersamples. Tight clusters indicate strong similarities inthe fatty acid composition of the samples; loose link-ages signify lower degrees of correspondence. Thediscriminating characteristics of the food sampleclusters are summarized in Table 1. The fatty acidcomposition of the members of the major subclusters, Ito XV, and the four samples which did not readily linkwith other samples, deer fat, squirrel, grouse andcattail seed, are described below. The fatty acidcompositions of the food plant and animals samplesare presented in Tables 3–9. In cases where more thanone specimen was collected at the same stage ofmaturation, an average composition is given.

Cluster A, the large mammal and fish cluster,includes 11 samples. It consists of four subclusters, I toIV, and the deer fat sample which does not readilycluster. The samples in subcluster I, bear fat and cowbone marrow, have levels of C18:1 isomers averaging57%. The different location of the single point of

Page 4: Fatty Acids Composition in Canada

86 M. E. Malainey et al.

Table 1. Summary of average fatty acid compositions of food sample clusters

Cluster A B C

Subcluster I II III IV V VI VII VIII IX X XI XII XIII XIV XV

Type

Mammalfat andmarrow

Largeherbivore

meat Fish Fish

Berriesandnuts Mixed

Seedsand

berries Roots Seeds Mixed Greens Berries Roots Greens Roots

C16:0 19·90 19·39 16·07 14·10 3·73 12·06 7·48 19·98 7·52 10·33 18·71 3·47 22·68 24·19 18·71C18:0 7·06 20·35 3·87 2·78 1·73 2·36 2·58 2·59 3·55 2·43 2·48 1·34 3·15 3·66 5·94C18:1 56·77 35·79 18·28 31·96 54·00 35·29 29·12 6·55 10·02 15·62 5·03 14·95 12·12 4·05 3·34C18:2 7·01 8·93 2·91 4·04 37·85 35·83 54·69 48·74 64·14 39·24 18·82 29·08 26·24 16·15 15·61C18:3 0·68 2·61 4·39 3·83 0·93 3·66 1·51 7·24 5·49 19·77 35·08 39·75 9·64 17·88 3·42VLCS 0·16 0·32 0·23 0·15 0·71 4·46 2·98 8·50 5·19 3·73 6·77 9·10 15·32 18·68 43·36VLCP 0·18 4·13 39·27 23·24 0·02 0·64 0·42 0·39 0·54 0·88 0·22 0·14 0·30 0·18 0·40

VLCS – Very Long Chain (C20, C22 and C24) Saturated Fatty Acids.VLCP – Very Long Chain (C20, C22 and C24) Polyunsaturated Fatty Acids.

Table 2. Variation in modern references explained by principal components 1 to 11

Principalcomponent Eigenvalue

Fatty acids with highcomponent loadings

Percentagevariation

Cumulativevariation

PRIN1 8·01 Polyunsaturates 22·24 22·24PRIN2 4·25 Medium chain; C18:1; C18:2 11·80 34·04PRIN3 3·12 Medium chain 8·66 42·71PRIN4 2·50 C17:0; C19:0 6·93 49·64PRIN5 1·87 Unsaturates 5·19 54·84PRIN6 1·72 C16:0; C17:0; C18:0 4·78 59·61PRIN7 1·45 C18:3ù6, C18:4 4·03 63·64PRIN8 1·44 monounsaturates 3·99 67·63PRIN9 1·15 C17:1; C20:2; C20:4ù6 3·19 70·82PRIN10 1·06 C20:1; C22:1 2·93 73·75PRIN11 1·03 C14:1; C18:1ù11; C20:4ù6 2·86 76·61

unsaturation in the two C18:1 fatty acid isomers,C18:1ù9 and C18:1ù11, results in slightly differentretention times and peak separation; the value pre-sented represents the sum of the two peaks. Theaverage levels of C16:0 (20%), as well as, C18:0 andC18:2 (both 7%) are also significant. Subcluster IIincludes the meat of two herbivores: bison and deer.The average fatty acid composition of these samples ischaracterized by high levels of C18:1 isomers (36%),C18:0 (20%), C16:0 (19%) and C18:2 (9%). SubclustersI and II link and are then joined by deer fat. Deer fatdiffers from the other samples in terms of its higherlevel of C18:0 (39%) and lower level of C18:1 isomers(24%).

While diet can alter the fatty acid composition ofanimal tissue, these changes tend to impact poly-unsaturated fatty acids and absolute levels are affectedmore than relative values. There are broad similaritiesin the composition of large herbivore products. Thefatty acid composition of bison and deer meat aresimilar to values reported for the meat of moose,caribou, domestic oxen, Cape buffalo and several

different species of African antelope (Crawford et al.,1970; Appavoo, Kubow & Kuhnlein, 1991). The fattyacid composition of the subcutaneous fat of white taildeer from southeast Manitoba is very similar to that ofthe wild African eland (Crawford et al., 1970).

All of the fish samples appear in subclusters III andIV. Subcluster III includes pickerel, perch, catfishand smoked trout; subcluster IV contains whitefish andsmoked goldeye. The division between the two sub-clusters reflects higher levels of C18:1 isomers andlower levels of C22:6 in whitefish and smoked goldeye,as compared to the other fish samples. The averagefatty acid composition of the fish samples is character-ized by high levels of both very long chain (20 or morecarbons in length) polyunsaturated fatty acids (VLCP)— 39% for subcluster III and 23% for subcluster IV —and C18:1 isomers — 18% for subcluster III and 32%for subcluster IV. The average level of C16:0 in thesubclusters is about 15%; the average level of C16:1isomers is 8% for subcluster III and 12% for subclusterIV. Smoking is a method of preservation which doesnot affect the fatty acid composition of the fish

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Native Food Plants and Animals of Western Canada 87

Table 3. The fatty acid composition of mammal and bird samples

Sample bear cow bison deer deer beaver muskrat squirrel grouse

Type fat marrow meat meat fat meat meat meat meat

Subcluster I I II II N/A VI X N/A N/A

C14:0 1·12 3·77 2·64 1·41 2·32 0·83 0·87 0·22 0·55C16:0 18·16 21·65 18·80 19·98 22·60 17·43 16·86 9·77 15·77C16:1 2·20 4·89 4·90 1·82 1·34 4·57 1·48 0·67 3·11C17:0 0·35 0·51 1·20 1·07 1·93 0·26 1·04 0·32 0·21C18:0 5·89 8·22 22·34 18·35 38·78 6·41 6·08 7·98 13·25C18:1 60·57 52·98 38·07 33·51 23·70 38·31 17·16 18·71 19·28C18:2 8·54 5·47 6·50 11·37 3·46 24·07 34·83 43·66 17·95C18:3ù3 0·89 0·48 0·60 4·63 2·64 2·87 18·24 0·75 6·86C20:4ù6 0·04 0·05 1·65 3·51 0·10 0·78 0·51 8·26 9·92C20:5 0·00 0·00 0·00 1·75 0·00 1·18 0·11 0·40 6·59Others 2·24 1·98 3·29 2·61 0·88 3·29 2·28 1·70 3·07

Table 4. The fatty acid composition of fish samples

Sample catfish pickerel perch trout whitefish goldeye

Type fish fish fish smk fish fish smk fish

Subcluster III III III III IV IV

C14:0 2·29 1·40 1·02 3·39 3·19 2·51C16:0 16·00 17·12 18·38 12·77 13·86 14·34C16:1 8·12 7·63 7·34 6·95 13·96 10·40C18:0 4·52 3·67 3·70 3·58 2·95 2·60C18:1 20·27 17·10 15·33 20·42 28·58 35·34C18:2 3·45 2·02 2·09 4·07 2·26 5·81C18:3ù3 6·39 4·19 3·00 3·98 1·98 5·68C18:4 1·33 1·19 0·67 3·18 1·57 1·71C20:1 0·79 0·54 0·35 0·81 1·22 0·28C20:4ù6 6·01 3·64 5·39 3·12 1·95 2·23C20:4ù3 0·00 1·51 0·90 3·93 0·97 2·96C20:5 9·30 8·69 8·08 5·56 9·38 6·16C22:3 1·36 1·59 1·57 1·54 0·54 0·42C22:4 0·00 1·64 1·73 2·47 0·69 0·12C22:5 3·50 3·03 3·15 4·12 3·52 2·78C22:6 8·30 20·86 23·72 15·10 9·18 3·24Others 8·38 4·19 3·57 5·03 4·21 3·41

samples. For a complete review of the factors whichinfluence the lipid composition of freshwater fish,readers should consult Henderson & Tocher (1987).

Cluster B consists of 46 samples divided into sixsubclusters, V-X, of seeds, berries, roots and smallmammals. Squirrel meat also appears in cluster B, butit is only loosely linked with the other samples. Sub-cluster V is a small cluster (N=6) of chokecherry,pincherry and wild hazelnuts in two minor clusters.The average fatty acid composition of these samplesfeatures very high levels of C18:1 isomers (54%) andC18:2 (38%).

Subcluster VI contains samples of dock (N=3) andknotweed (N=1) seeds, two varieties of mushroom,

acorn and beaver meat in two minor clusters. Theaverage fatty acid composition of the samples in sub-cluster VI contain high levels of C18:2 (36%) and C18:1isomers (35%). The average level of C16:0 is 12%. Thecompositions of the mushrooms and beaver differ fromother samples with respect to their higher levels ofC16:1 isomers. The alpha linolenic fatty acid, C18:3ù3is virtually absent from mushrooms. The level of C18:2in beaver meat is more than 10% lower than the meanfor this subcluster.

Subcluster VII consists of a tight cluster of eightseeds and berries, including most of the Native culti-gens: Mandan Sweet Corn, Assiniboin Flint Corn,Arikara winter squash and Mandan sunflower. Wild

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88 M. E. Malainey et al.

Table 5. The fatty acid composition of berry samples

Sample chokecherry pincherry gooseberry saskatoon rosehips bearberry blueberry juniper silverberryhaw

thornred

osier

Subcluster V (N=4) V&VII (N=2) X (N=2) VII (N=2) XII (N=4) XII (N=2) X X XIV VII XII

C16:0 4·05&0·50 2·15&0·22 8·42&0·23 7·61&0·23 4·05&1·73 2·32&2·69 6·88 8·68 20·88 7·16 26·60C18:0 2·00&0·71 0·83&0·16 1·79&0·33 1·92&0·18 1·52&0·64 0·98&0·20 1·71 3·39 3·24 1·96 1·36C18:1 52·93&3·95 43·10&7·46 12·89&2·21 25·43&1·92 16·16&2·36 12·52&1·99 17·19 14·84 7·81 23·45 16·47C18:2 38·25&4·88 52·19&8·05 36·44&1·10 52·36&0·34 33·13&4·63 20·99&6·33 41·70 36·43 15·81 56·87 38·09C18:3ù3 1·05&0·08 0·65&0·31 22·41&5·28 2·64&0·52 39·33&1·12 40·60&6·40 28·76 20·66 18·38 2·13 14·76C20:0 0·24&0·04 0·13&0·04 1·26&0·40 2·69&0·33 1·32&0·91 0·40&0·11 1·32 0·57 4·08 1·63 0·16C20:1 0·18&0·02 0·18&0·03 0·17&0·01 1·03&0·08 0·66&0·09 0·47&0·02 0·25 1·14 0·12 0·69 0·24C22:0 0·45&0·58 0·19&0·26 0·68&0·96 3·57&0·69 1·07&1·03 6·64&3·72 0·46 2·03 17·70 0·65 0·14C24:0 0·29&0·10 0·04&0·01 1·97&0·09 1·30&0·24 0·84&0·87 13·85&7·58 0·56 0·95 7·53 1·00 0·44Others 0·56&0·17 0·56&0·09 13·99&2·381 1·48&0·05 1·94&1·25 1·24&0·42 1·05 9·992 4·46 4·45 1·74

1 C18:3ù6=8·51&1·68%; C18:4=3·43&0·33%.2 C12:0=3·86%; C20:2=3·35%; C22:0=2·03%.

Table 6. The fatty acid composition of nut, bean and mushroom samples

Sample acorn hazelnutNative

redbean vetchling Armillaria Leccinum

Type nut nut bean pods mushroom mushroom

Subcluster VI V XI VIII VI VI

C16:0 9·13 3·87 13·71 14·69 10·01 12·85C16:1 0·38 0·37 0·08 2·71 6·61 1·80C17:0 0·30 0·07 0·19 0·42 0·03 0·11C18:0 1·82 1·46 2·32 2·23 2·31 0·90C18:1 37·74 65·23 10·88 9·48 45·50 39·76C18:2 41·10 27·62 34·74 54·09 30·71 40·65C18:3ù3 5·79 0·53 35·35 9·54 0·00 0·14C20:0 0·59 0·24 0·45 1·07 0·30 0·14C22:0 0·74 0·08 0·74 1·34 0·26 0·21C24:0 0·85 0·17 0·93 2·18 0·57 0·18Others 1·54 0·36 0·61 2·25 3·60 3·24

varieties in this subcluster include bulrush, saskatoonand hawthorn, which later links with a pincherrysample. The average fatty acid composition of thesesamples is characterized by very high levels of C18:2(55%) and C18:1 isomers (29%), which is typical formany vegetable oils. The pincherry sample differs fromthe others in terms of its very low level of C16:0 andC18:1 isomer level, almost 9% higher than the sub-cluster mean. Most berries and the bulrush seed havehigher levels of C18:3ù3 and C24:0 than other samplesin this cluster, saskatoon and bulrush seed have higherlevels of C22:0. The level of C18:0 in sunflower andwinter squash is somewhat higher than in the othersamples. The fatty acid composition of Native corndiffered from commercial corn oil (Sheppard, Iverson& Weihrauch, 1978) in that the average level of C18:1isomers was 5% higher and C18:2 was 5% lower in theNative cultigens. The values obtained for Mandansunflower corresponded well to average values for

commercial sunflower oil (Sheppard, Iverson &Weihrauch, 1978).

Subcluster VIII consists of 10 plant roots, includingfalse solomon’s seal, cow parsnip, Jerusalem artichoke,wild onion, prairie turnip, tiger lily, water parsnip andwild calla, as well as a sample of vetchling pods in twominor clusters. The average fatty acid composition ofthe samples is characterized by very high levels ofC18:2 (49%) and C16:0 (20%). Some roots containlevels of medium chain or very long chain saturated(VLCS) fatty acids which are significantly higher thanthe mean for the entire cluster. The level of C12:0 inJerusalem artichoke is almost six times higher than themean; C15:0 levels are significantly higher in bothJerusalem artichoke and cow parsnip. The level ofC22:0 is 6% in prairie turnip, double the mean of 3%.The level of C24:0 is over 12% in tiger lily root andapproaches 8% in prairie turnip, whereas the subclustermean is 4%.

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Native Food Plants and Animals of Western Canada 89

Table 7. The fatty acid composition of plant seed samples

Sample marshelder arrowgrass dock1 knotweed goosefoot2Nativecorn3

Nativesunflower

Nativesquash bulrush cattail

Subcluster IX IX (N=2) VI(N=3), X, XIII IX XIII(N=2), X VII (N=2) VII VII VII N/A

C16:0 9·01 6·78&0·94 12·56&2·14 11·56 13·37&4·76 10·61&0·16 5·72 10·15 5·88 9·16C16:1 1·32 1·13&0·53 1·25&0·99 0·81 1·94&1·73 0·04&0·01 0·02 0·11 0·20 0·44C17:0 0·29 0·32&0·16 0·22&0·06 0·34 0·26&0·06 0·09&0·01 0·09 0·15 0·20 20·25C18:0 6·29 2·18&0·08 1·82&0·33 1·61 1·77&0·26 2·17&0·30 4·46 7·41 0·54 3·34C18:1 5·95 12·06&1·53 24·31&5·87 35·57 15·37&1·69 32·16&0·49 31·60 28·97 25·01 4·58C18:2 59·70 66·36&6·08 34·85&6·88 36·70 38·72&12·66 52·40&0·25 55·71 51·81 60·40 11·13C19:0 0·12 0·21&0·27 0·00 0·00 0·02&0·03 0·00 0·00 0·00 0·00 22·21C18:3ù3 5·13 5·67&3·49 10·30&7·58 5·88 16·13&5·28 1·16&0·04 0·13 0·18 3·16 3·11C20:0 3·42 1·66&0·65 1·64&0·62 0·70 1·39&0·48 0·54&0·05 0·41 0·53 0·71 2·96C20:1 0·47 0·17&0·02 0·94&0·38 1·41 1·07&0·45 0·32&0·01 0·21 0·08 0·49 0·27C22:1 0·00 0·00 1·24&1·12 0·92 1·09&0·75 0·00 0·00 0·00 0·07 0·63C22:0 2·36 1·68&0·81 2·19&0·75 1·34 2·51&0·93 0·18&0·01 1·24 0·14 1·52 13·55C22:2 1·06 0·00 0·02&0·04 0·00 0·10&0·09 0·00 0·00 0·00 0·00 0·28C24:0 0·95 1·07&0·35 6·13&4·20 1·22 2·25&1·89 0·23&0·02 0·30 0·12 1·05 6·11Others 3·92 0·74&0·35 2·11&0·81 1·95 3·23&0·59 0·13&0·01 0·12 0·34 0·83 1·99

1 N=5.2 N=3.3 Mandan flint corn and Assiniboin sweet corn.

Subcluster IX is a small cluster of marsh-elder (N=1)and arrow-grass (N=2) seeds. The seeds have anaverage fatty acid composition which is extremely highin C18:2 (64%), with lower levels of C18:1 isomers(10%) and C16:0 (8%). The levels of other fatty acidsvary widely between these three samples.

Subcluster X contains mainly plant material, dockand lamb’s-quarters seeds and gooseberry, blueberryand juniper berries, but muskrat meat also joinedthis subcluster. This subcluster contains a number ofminor clusters and the average fatty acid compositionsare quite variable. In general, the average level ofC18:2 is high (39%) and significant amounts ofC18:3ù3 (20%), C18:1 isomers (16%) and C16:0 (10%)are present. Juniper berries differ from the othersamples on the basis of its higher levels of C12:0 andC14:0. The level of C18:0 is higher in muskrat while thetwo gooseberry samples have high levels of C18:3ù6and C18:4. The levels of very long chain saturated andmono-unsaturated fatty acids vary widely betweensamples.

Squirrel meat likely appears in cluster B because ofits high level of C18:2 (44%), but it does not closelyresemble any of the samples.

Cluster C consists of five subclusters of plants, XI toXV, and two samples that did not readily cluster(grouse flesh and cattail seed). Subcluster XI is domi-nated by plant greens, specifically false solomon’s seal,fireweed, golden rod, violet, dock and goosefoot whichform two minor clusters. In total, there are 24 plantsamples, 22 of which are plant greens, one plant seedand one plant root. The average fatty acid com-positions are characterized by high levels of C18:3ù3(35%) and C16:0 (19%). The two minor clusters withinsubcluster XI may reflect differences in the levels of

branched C16:0 (brC16:0), C16:1 isomers and C18:2 inthe samples. False solomon’s seal greens, fireweed rootand the Native domesticate, Hidatsa red beans, allhave relatively low levels of C16:1 isomers; false solo-mon’s seal and violet greens both have relatively highlevels of C14:1. Stinging nettle and one sample ofsarsaparilla greens were the last to join establishedclusters. The high C24:0 level in the stinging nettle andthe levels of C18:2, C18:3ù3 and C24:1 in the sarsapa-rilla greens distinguish them from the other samples.

Subcluster XII consists exclusively of rosehips andbearberries. The average fatty acid composition ofthese samples is characterized by high levels ofC18:3ù3 (40%), C18:2 (29%) and C18:1 isomers (15%).Both bearberry samples have higher levels of C22:0and C24:0. One of these has lower levels of C18:1isomers and C18:2 than the other samples and is thelast to join the cluster.

Subcluster XIII is dominated by plant root samples,including bulrush, cattail, water parsnip, giant reed,fireweed, sarsaparilla, water-hore hound, wound wort,ostrich fern and Jerusalem artichoke. Nineteen of the24 samples in subcluster XIII are roots; red osierberries, mare’s tail and chickweed greens, and lamb’s-quarter and dock seeds also appear. The members ofthis subcluster are rather loosely linked, consisting of anumber of minor clusters and the fatty acid com-position of samples is variable. Average levels of C18:2(26%) and C16:0 (23%) are quite high, but there aresignificant amounts of C18:1 (12%) and C18:3ù3(10%), as well. The levels of medium chain fatty acids(C12:0, C14:0 and C15:0) are much higher thanaverage in water parsnip and Jerusalem artichoke.Levels of VLCS (C20:0, C22:0 and C24:0) are very highin some samples, but there is considerable variability.

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90 M. E. Malainey et al.

Subcluster XIV is dominated by plant greens,including cow parsnip, cattail, bulrush and goldenrod. Ten of the 13 samples in this cluster are plantgreens; the other samples are sarsaparilla and arrow-head roots and silverberry. The average fatty acidcomposition is characterized by a high level of C16:0(25%) with moderate levels of VLCS (19%), C18:3ù3(18%) and C18:2 (16%). Branched C16:0 is commonin this subcluster, appearing in all samples exceptsilverberry.

Subcluster XV contains two samples of cattail root.The average fatty acid composition of these samples ischaracterized by very high levels of VLCS; together,C24:0 (19%), C22:0 (13%) and C20:0 (12%) account for

almost 45% of all fatty acids. Significant levels of C16:0(19%) and C18:2 (16%) are also present.

Grouse is likely included in cluster C because itsfatty acid composition is characterized by a number offatty acids (C16:0, C18:0, C18:1 isomers and C18:2)occurring at levels between 13 and 18%, rather thanone or two dominant fatty acids. Cattail seed is onlyloosely linked to cluster C; it differs from the othersamples on the basis of its high level of 17:0, 20%.

Table 8. The fatty acid composition of plant root samples

Samplefalse

solomon’s seal fireweed1water

parsnip2cow

parsnipJerusalemartichoke3 cattail4 bulrush giant reed

Subcluster VIII (N=2) VIII (N=3), XI XIII(N=2), VIII VIII (N=2) VIII, XIII XV (N=2), XIII XIII (N=4) XIII (N=2)

C12:0 0·23&0·14 0·02&0·04 3·60&2·95 0·35&0·01 3·11&1·10 0·13&0·04 0·11&0·13 0·00C15:0 0·69&0·07 0·36&0·10 4·49&4·28 1·20&0·15 4·03&2·27 0·97&0·14 1·09&0·82 0·34&0·25C16:0 23·24&1·15 26·87&5·85 19·22&3·09 18·50&3·00 18·45&3·54 18·15&3·94 25·79&3·27 20·19&4·57C16:1 1·52&1·00 1·45&0·74 1·17&1·54 0·29&0·01 0·69&0·07 1·84&0·90 2·33&1·51 0·40&0·12C16:2 0·00 0·40&0·44 1·10&1·50 0·04&0·06 3·15&2·76 0·00 0·15&0·29 0·14&0·19C18:0 2·79&0·32 3·61&0·68 4·37&0·60 1·84&0·13 1·72&0·42 4·91&1·79 1·90&1·43 2·68&0·25C18:1 5·16&2·20 14·75&10·40 6·96&2·51 4·89&1·55 4·96&0·88 3·97&2·64 13·64&0·99 7·86&3·18C18:2 47·41&6·79 21·37&0·73 37·07&9·31 56·36&0·21 41·77&12·74 21·51&10·26 21·26&7·23 31·57&6·40C18:3ù3 6·02&1·70 16·70&8·54 5·72&0·85 6·14&0·15 8·26&1·31 4·65&2·26 8·21&1·96 6·82&0·91C20:0 1·64&0·50 3·42&2·31 3·08&0·58 1·91&0·92 1·02&0·28 10·03&2·59 2·04&0·53 12·83&2·21C20:1 3·47&2·45 0·64&0·42 0·84&0·44 0·92&0·23 0·62&0·06 0·74&0·21 0·63&0·47 7·08&2·77C22:0 2·61&1·61 1·97&0·39 4·27&1·71 2·09&0·41 3·82&1·73 11·30&2·63 6·76&1·98 3·36&0·62C24:0 3·01&2·18 1·57&0·60 2·29&1·45 2·68&1·23 2·20&0·33 16·81&4·77 12·99&4·78 4·39&0·27C24:1 0·25&0·25 0·85&1·23 0·00 0·51&0·71 1·66&0·52 0·05&0·08 0·44&0·42 0·10&0·13Others 1·97&0·54 6·00&3·24 5·81&3·71 2·32&0·76 4·36&2·06 4·78&2·11 2·66&0·76 2·28&0·33

Sample sarsaparillawild

onion tiger lilywildcalla

prairietumip burreed arrow head

woundwort

ostrichfern

water-horehound

Subcluster XIII, XIV (N=2) VIII VIII VIII VIII XIII XIV XIII XIII XIII

C12:0 0·41&0·31 0·58 0·00 0·00 0·00 0·27 0·12 0·00 0·10 0·00C15:0 0·37&0·09 0·44 0·12 0·34 0·00 0·69 0·07 0·20 0·87 0·41C16:0 22·40&1·09 19·75 20·33 25·10 18·37 19·74 20·30 21·20 32·26 19·09C16:1 3·11&2·28 2·90 1·39 2·53 0·00 3·68 3·66 6·32 2·32 0·87C16:2 1·93&0·10 0·17 0·28 1·69 0·00 0·00 0·48 0·00 3·81 6·14C18:0 2·64&0·81 4·03 2·46 1·19 3·83 3·03 7·17 2·81 2·07 4·74C18:1 7·10&6·06 11·27 7·76 9·44 4·26 19·98 2·99 10·63 10·00 11·98C18:2 26·42&4·84 40·04 45·26 42·46 48·13 20·11 18·41 23·16 34·79 26·39C18:3ù3 13·68&4·94 5·33 3·08 12·85 8·81 3·73 13·22 12·17 6·50 11·25C20:0 2·85&0·08 2·83 1·38 0·30 2·03 4·33 1·35 2·73 0·81 3·06C20:1 0·35&0·09 0·58 0·48 0·55 0·00 1·11 0·17 0·37 0·18 0·35C22:0 5·73&1·56 3·56 3·46 0·97 6·04 5·23 2·59 7·44 1·65 6·55C24:0 7·52&3·96 3·54 12·62 0·90 7·62 14·64 24·59 8·83 2·80 5·17C24:1 0·00 1·59 0·00 0·28 0·00 0·00 0·27 0·85 0·00 1·23Others 5·53&0·86 3·39 1·39 1·41 0·91 3·48 2·75 3·28 1·75 2·76

1 N=4.2 N=3.3 N=2.4 N=3.

Principal component analysisWhen all 37 fatty acids present in the data set areincluded in the analysis, each principal component

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Native Food Plants and Animals of Western Canada 91

explains a relatively small part of the variation. Thefirst 11 principal components have eigenvalues greaterthan one and are considered significant. Together theyaccount for more than 76% of the total variation in themodern food plants and animals (Table 2). Principalcomponent 1, which has high positive componentloadings on polyunsaturated fatty acids with chain-lengths of 20 or more carbons, accounts for 22% of thevariation. Principal component 2, with high positiveloadings on fatty acids with chain-lengths of 16 orfewer carbons and negative loadings on oleic andlinoleic acids, explains almost 12% of the variation.Each of principal components 3 to 11 explain between9 and 3% of the variation. The high loadings onpolyunsaturated fatty acids in the first principal com-ponent results in the separation of fish and grouse in

the plot of PRIN2 versus PRIN1 (Figure 2). Greens,roots, as well as berries, seeds and nuts, each formfairly tight clusters in the plot of PRIN2 versus PRIN3(Figure 3). Reducing the number of variables underconsideration would simplify the principal componentanalysis, but this action may introduce an unacceptablelevel of bias.

Table 9. The fatty acid composition of plant greens samples

Sample false solomon’s seal golden rod1 fireweed cow parsnip bulrush cattail violet

Subcluster XI (N=5) XI (N=3), XIV XI (N=4) XIV (N=3) XIV (N=3) XIV (N=3) XI (N=2)

C14:0 0·53&0·21 0·23&0·09 1·81&0·84 1·54&1·37 0·97&0·40 1·20&0·79 0·62&0·27C14:1 0·89&0·92 0·00 0·00 1·20&0·81 0·00 0·00 2·54&0·14brC16:0 0·82&0·43 1·51&0·64 1·03&0·40 1·44&0·99 1·14&0·22 1·01&0·33 0·53&0·75C16:0 20·55&2·84 21·15&4·98 18·23&1·89 18·25&3·97 30·51&2·99 24·96&3·35 18·84&1·51C16:1 4·22&1·76 8·52&2·58 7·79&2·10 5·49&3·59 8·44&0·97 7·35&1·99 6·24&1·07C16:2 0·87&0·23 2·00&0·32 2·25&0·18 0·97&0·92 1·10&0·12 1·06&0·42 1·97&0·21C17:1 0·36&0·23 0·00 0·06&0·13 3·32&2·52 0·12&0·21 0·18&0·31 0·09&0·13C18:0 2·36&0·42 2·69&0·55 2·86&0·72 2·79&0·47 2·81&0·29 5·13&1·33 2·13&0·30C18:1 3·38&1·65 2·02&0·68 2·95&0·79 3·70&1·82 5·07&0·39 4·26&4·05 6·64&0·05C18:2 25·23&3·93 14·30&3·81 13·84&3·62 24·17&3·40 12·24&2·97 11·91&3·42 23·31&4·38C18:3ù3 32·48&5·14 36·30&6·25 38·10&2·17 19·51&3·45 16·13&4·55 14·94&5·93 31·19&0·78C20:0 1·12&0·22 2·42&0·73 2·99&0·91 2·21&0·59 2·79&0·29 4·87&1·66 0·58&0·40C22:0 2·21&0·28 1·85&0·34 2·19&0·77 3·36&0·66 5·27&1·31 8·20&2·06 1·30&0·97C24:0 2·48&0·48 2·90&1·69 3·13&0·98 7·20&1·40 9·74&1·81 11·48&3·56 2·84&3·39C24:1 0·07&0·07 0·97&0·21 0·56&0·55 0·07&0·12 0·04&0·08 0·00 0·00Others 2·43&0·51 1·89&1·14 2·21&0·60 4·84&4·12 3·67&0·79 3·45&0·87 1·19&0·42

Sample sarsaparilla goosefoot dockwater-hore

houndstingingnettle

chickweed

mare’stail

Subcluster XI (N=2) XI (N=2) XI (N=2) XI XI XIII XIII

C14:0 1·13&0·90 0·85&0·50 0·50&0·21 0·64 0·41 1·52 0·85C14:1 0·00 0·00 0·02&0·02 0·00 0·00 0·00 0·00brC16:0 1·78&0·14 1·90&0·39 1·81&0·33 0·00 2·45 0·00 1·74C16:0 18·13&2·86 18·50&0·75 18·15&1·56 18·53 14·13 19·13 29·79C16:1 9·29&2·41 10·72&2·53 8·15&0·93 8·40 10·14 3·83 10·03C16:2 1·85&0·14 2·27&0·61 1·82&0·88 1·65 1·71 0·86 1·07C17:1 0·06&0·08 1·89&0·45 0·17&0·23 0·14 0·00 0·00 0·00C18:0 2·40&0·92 1·41&0·37 1·77&0·35 5·11 2·14 1·87 4·39C18:1 4·72&0·45 6·43&0·84 9·88&1·27 5·58 2·97 11·02 10·90C18:2 11·34&4·96 13·35&0·54 14·08&2·45 10·35 18·74 31·96 21·26C18:3ù3 41·57&8·42 34·89&0·83 37·61&6·32 30·71 30·85 15·05 9·29C20:0 1·38&1·03 1·90&0·37 0·85&0·22 3·61 2·98 1·02 2·50C22:0 1·90&1·40 2·62&0·16 1·19&0·12 4·24 3·19 1·92 3·28C24:0 2·45&1·48 2·40&0·30 2·64&0·40 3·50 9·51 4·18 3·28C24:1 1·48&1·80 0·00 0·00 1·07 0·00 0·11 0·00Others 0·76&0·23 0·91&0·80 1·43&0·50 6·47 0·79 1·23 0·34

1 N=4.

Discussion and ConclusionsWhen the fatty acid compositions of modern foodplants and animals are subject to principal componentand hierarchical cluster analysis, the resulting group-ings generally correspond to divisions which exist innature. Clear differences in the fatty acid composition

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92 M. E. Malainey et al.

18

6

PRIN1

PR

IN2

6

1

5

4

3

2

0

–1

–2

–3

–2 0 2 4 8 10 12 14–4 16

FatLarge herbivore meatFishBerry/seed/nutGreensRootsSmall mammal meatMushroomBird

Figure 2. Plot of PRIN2 scores versus PRIN1 scores for the modern references.

of large mammal fat, large herbivore meat, fish, plantroots, greens and berries/seeds/nuts can be detected,but the fatty acid composition of medium-sizedmammals meat closely resembles berries/seeds/nuts;their main dietary components.

Samples in cluster A, the large mammal and fishcluster, have elevated levels of C16:0 and C18:1. Fatsamples are high in C18:1 isomers, bison and deer meathave higher levels of C18:0 while fish have high levelsof VLCP. The fatty acid composition of bison and deerproducts resemble those of other large herbivores from

North America (Appavoo, Kubow & Kuhnlein, 1991)and Africa (Crawford et al., 1970).

Statistically significant differences in the fatty acidcomposition of plant roots, greens and berries/seeds/nuts were based on differences in the amounts of C18:2and C18:3ù3 in the samples. The berry, seed, nut andsmall mammal meat samples appearing in cluster Bhave high to extremely high levels of C18:2. Samples insubclusters VIII, IX and X have high to extremely highlevels of C18:1 isomers, as well; squirrel meat hashigher levels of VLCP than other samples in cluster B.

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Native Food Plants and Animals of Western Canada 93

Modern food samples in cluster C are mainly plantroots and greens, but include rosehips and bearberries.These samples all have higher levels of C18:2 exceptfor the berries, all samples have elevated levels ofC16:0. Higher levels of C18:3ù3 and/or VLCS are alsocommon. Grouse meat differs from the other samplesin cluster C because of its higher levels of C18:1isomers and VLCP; cattail seeds have high levels ofC17:0 and VLCS.

The determination of the fatty acid composition ofuncooked food plant and animal samples is an import-ant first step in the identification of vessel residuesbecause it enables cooking experiments to be con-ducted on a representative sample of foods of similar

fatty acid compositions, rather than on all samples inthe collection. The effects of thermal and oxidativedegradation on meat, fish and plants, prepared aloneor in combination, are reported in Malainey,Przybylski & Sherriff (1999).

10

6

PRIN3

PR

IN2

6

1

5

4

3

2

0

–1

–2

–3

–2 0 2 4 8–4

FatLarge herbivore meatFishBerry/seed/nutGreensRootsSmall mammal meatMushroomBird

Figure 3. Plot of PRIN2 scores versus PRIN3 scores for the modern references.

AcknowledgementsThis paper is extracted from Malainey (1997). Theresearch was carried out while MEM held a SocialScience and Humanities Research Council of Canadadoctoral fellowship. This research was supported by aNatural Science and Engineering Research Council

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94 M. E. Malainey et al.

University Research Fellowship and research grant toBLS and a Manitoba Heritage Grant to MEM. Thisproject benefited from the advice and support of M.Latta, R. Zambiazi, G. Monks, B. Watts, E.L. Syms,M. Deal and two anonymous reviewers. The efforts ofall those who aided in the collection of food plant andanimal samples are gratefully acknowledged.

ReferencesAppavoo, D. M., Kubow, S. & Kuhnlein, H. V. (1991). Lipid

composition of indigenous foods eaten by the Sahtu (Hareskin)Dene-Metis of the Northwest Territories. Journal of FoodComposition and Analysis 4, 107–119.

Charters, S., Evershed, R. P., Blinkhorn, P. W. & Denham, V.(1995). Evidence for the mixing of fats and waxes in archaeologicalceramics. Archaeometry 37, 113–127.

Condamin, J., Formenti, F., Metais, M. O., Michel, M. & Blond P.(1976). The application of gas chromatography to the tracing ofoil in ancient amphorae. Archaeometry 18, 195–201.

Crawford, M. A., Gale, M. M, Woodford, M. H. & Casped, N. M.(1970). Comparative studies on fatty acid composition of wild anddomestic meats. International Journal of Biochemistry I, 295–305.

Deal, M. (1990). Exploratory analyses of food residues fromprehistoric pottery and other artifacts from Eastern Canada. SASBulletin 13, 6–12. Houghton: Society for Archaeological Sciences.

Deal, M. & Silk, P. (1988). Absorption residues and vessel function:a case study from the Maine-Maritime region. In (C. C. Kolb &L. M. Lackey, Eds) A Pot For All Reasons: Ceramic EcologyRevisited. Philadelphia: Laboratory of Anthropology, TempleUniversity, pp. 105–125.

Evershed, R. P. (1993). Biomolecular archaeology and lipids. WorldArchaeology 25, 74–93.

Evershed, R. P., Heron, C. & Goad, L. J. (1990). Analysis of organicresidues of archaeological origin by high temperature gas chroma-tography and gas chromatography-mass spectroscopy. Analyst115, 1339–1342.

Evershed, R. P., Heron, C. & Goad, L. J. (1991). Epicular waxcomponents preserved in potsherds as chemical indicators of leafyvegetables in ancient diets. Antiquity 65, 540–544.

Evershed, R. P., Heron, C., Charters, S. & Goad, L. J. (1992). Thesurvivial of food residues: new methods of analysis, interpretationand application. Proceedings of the British Academy 77, 187–208.

Folch, J., Lees, M. & Stanley, G. H. S. (1957). A simple method forthe isolation and purification of total lipids from animal tissures.Journal of Biological Chemistry 266, 497.

Frankel, E. N. (1991). Recent advances in lipid oxidation. Journal ofthe Science of Food and Agriculture 54, 465–511.

Gaertner, Erika E. (1967). Harvest without Planting. Chalk River:self-published.

Gerhardt, K. O., Searles, S. & Biers, W. R. (1990). Corinthian figurevases: non-destructive extraction and gas chromatography-mass

spectrometry. In (W. R. Biers & P. E. McGovern, Eds) OrganicContents of Ancient Vessels: Materials Analysis and ArchaeologicalInvestigation Vol. 7. Philadelphia: MASCA, The UniversityMuseum of Archaeology and Anthropology, University ofPennsylvania, pp. 41–50.

Gilmore, M. (1919) Uses of plants by the Indians of the MissouriRiver region. Thirty-third Annual Report of the Bureau of AmericanEthnology. Washington: Government Printing Office; reprinted1991, University of Nebraska Press, Lincoln.

Henderson, R. J. & Tocher, D. R. (1987). The lipid composition andbiochemistry of freshwater fish. Progress in Lipid Research 26,281–347.

Heron, C. P. (1989). The analysis of organic residues from archaeo-logical ceramics. Ph.D. Dissertation, University of Wales.

Heron, C., Evershed, R. P. & Goad, L.J. (1991). Effects of migrationof soil lipids on organic residues associated with buried potsherds.Journal of Archaeological Science 18, 641–659.

Heron, C. & Evershed, R. P. (1993). The analysis of organic residuesand the study of pottery use. In (M. B. Schiffer, Ed.) Archaeologi-cal Method and Theory 5. Tucson: University of Arizona Press,pp. 247–284.

Heron, C. & Pollard, A. M. (1988). The analysis of natural resinousmaterials from Roman amphoras. In (E. A. Slater & J. O. Tate,Eds) Science and Archaeology Glasgow 1987. Proceedings of aConference on the Application of Scientific Techniques toArchaeology, Glasgow, 1987. Oxford: BAR British Series 196 (ii),pp. 429–447.

Kindscher, K. (1987) Edible Plants of the Prairie. Lawrence: Univer-sity Press of Kansas.

Malainey, M. E. (1995). Functional analysis of precontact potteryfrom West-central Canada. Manitoba Archaeological Journal 5,60–86.

Malainey, M. E. (1997). The reconstruction and testing of subsistenceand settlement strategies for the plains, parkland and southernboreal forest. Ph.D. Thesis, University of Manitoba.

Malainey, M. E., Przybylski, R. & Sherriff, B. L. (1999). The effectsof thermal and oxidative degradation on the fatty acid com-position of food plants and animals of Western Canada: implica-tions for the identification of archaeological vessel residues.Journal of Archaeological Science (this issue).

Shay, C. T. (1980). Food plants of Manitoba. In (L. Pettipas, Ed.)Directions in Manitoba Prehistory: Papers in Honour of ChrisVickers. Winipeg: Association of Manitoba Archaeologists andManitoba Archaeological Society, pp. 233–290.

Sheppard, A. J., Iverson, J. L. & Weihrauch, J. L. (1978). Compo-sition of selected dietary fats, oils, margarines, and butter. In(A. Kuksis, Ed.) Handbook of Lipid Research 1: Fatty Acids andGlycerides. New York: Plenum Press, pp. 341–379.

Skibo, J. M. (1992). Pottery Function: A Use-Alteration Perspective.New York: Plenum Press.

Solomons, T. W. G. (1980). Organic Chemistry. Toronto: John Wiley& Sons.

Young, K. (1993). Wild Seasons: Gathering and Cooking Wild Plantsof the Great Plains. Lincoln: University of Nebraska Press.