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On the other hand, sequence G not shown cannot be identified beyond the class Eudicotyledons level. The results of the phylogenetic analyses are summarized in Table 1. The tree is based on the families Restianaceae Restio tetraphyllus and Elegia sp.

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The last of these is further subdivided into the Panicoideae Z. The outgroup is represented by Xerophyllum tenax Liliales and Asplectrum hyemale Asparagales. The tree is based on the Cupressaceae Juniperus chinensis , Cupressus macrocarpa , Calocedrus formosana , Metasequoia glyptostroboides , Cryptomeria japonica , and Tauwania cryptomerioides , Taxaceae Taxus mairei , Sciadopityaceae Sciadopytis verticillata , Cephalotaxaceae Cephalotaxus wilsoniana , Araucariaceae Araucaria excelsa , Agathis robusta , and Pinaceae Larix leptolepis , Abies lasiocarpa , Picea mariana , Pinus luchuensis , P.

The outgroup is represented by Ginkgo biloba Ginkoales and Ephedra torreyana Ephedrales. The tree is based on the Ophioglossales Botrychium virginianum , Hydropteridales Marsilea schelpeana , Equisetales Equisetum hyemale , Marattiales Danaea cuspidata , Psilotales Tmesipteris obliqua , and Filicales Adiantum raddiatum , Histiopteris incisa , and Pteridium aquilinum orders. The outgroup is represented by Selaginella apoda Lycopodophyta. Normally, plant remains consist of so-called macroremains remains of food and microremains, the latter being represented by pollen.

One way in which pollen is ingested is by adherence to the swallowed food plants. The use of the 18S rDNA system does not allow one to determine whether the plant DNA comes from pollen or from macroremains, as this system is directed to a nuclear gene that is present in both kinds of residues. The colon amplicons grouped into two clusters I, J. Phylogenetic analysis of the consensus sequences shows that the first cluster corresponds to unidentified members of the Eudicotyledon class, whereas the second cluster is identified at the order Poales level.

The ileum clones all grouped in a single cluster H of sequences phylogenetically identified at the family Poaceae level. A phylogenetic analysis allows one to identify three sequences B, F, and I18S at the class level Table 1 ; the first two are Urediniomycetes, and the third is Heterobasidiomycetes. Observation under a light microscope of the food residues from both the ileum and colon of the mummy allows one to easily spot minute fragments of well-preserved muscle fibers Fig. The presence of muscle fibers in the colon, on the other hand, had been previously reported 3 , 4.

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This system is designed to bind to an approximately bp-long fragment of the mitochondrial 12S ribosomal RNA gene 12S rDNA of a wide range of mammals. Both specimens produced amplification bands of the expected size. Amplification products were then cloned and sequenced. For this reason, after all of the ancient DNA analyses had been completed, we sequenced a corresponding portion of the 12S rRNA gene of ibex Capra ibex and found that it perfectly matched the Iceman's sequence.

The screening of the ileum library 17 amplicons , on the other hand, showed that 8 clones were of human origin, and the other 9 clones consensus K corresponded to the database sequence GenBank accession no. However, when we checked a specimen of modern red deer from central Europe, we found that its sequence was identical to the ancient one. On the other hand, a careful comparison of the AF Bootstrapped neighbor-joining 12S rDNA tree of the Pecora infraorder inclusive of the animal consensus K and L sequences from the Iceman's ileum and colon, respectively.

Scientists Have Finally Figured Out What ÖTzi The Iceman Wore When He Died

The tree is based on the Bovidae C. The outgroup is represented by Okapia johnstoni Giraffoidea.


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We can now compare Table 2 the results of our DNA analysis with those of previous ones performed by using light and electron microscopy 3 , 4. The most likely explanation is that pinus and fern sequences reflect the presence of background pollen and spores in the intestines. This conclusion is supported by the lack of pinus and fern sequences in the rbcL libraries. One may wonder how pollen sequences are obtained, as the pollen coat offers a strong obstacle to DNA extraction.

We can only observe that conifer 18S rDNA sequences have also been reported in a previous study on fossil ice cores from northern Greenland The most plausible interpretation of this result is that the DNA comes from ice-trapped pollen. It seems therefore that a prolonged period at low temperature, and possibly the effect of the pressure, may loosen the resistance of the pollen coat, thus making it possible for DNA to be extracted. The vegetation in the Alps correlates closely with climatic variations.

After the end of the last Glacial period, the altitudinal zonation of the forests became progressively established: i mixed deciduous forest oak, elm, lime, ash, maple, and hornbeam in the valleys; ii mixed deciduous and coniferous forest spruce, fir, beech, hazel, birch, and other deciduous species at intermediate altitudes; and iii subalpine coniferous forest Arolla pine, dwarf mountain pine, Scots pine, larch, and spruce up to the tree line With regard to the ferns, these plants grow in moist habitats of the Alps in a wide range of low and intermediate altitudes.

On the other hand, the herb-rich alpine meadows and grasslands were ideal pastures for farm stock. If we now consider the arboreal 18S rDNA spectrum of the colon pines and the pollen spectrum, as determined by microscopy, we can observe that both fit with the distribution of the above-described vegetation on the Neolithic Alps, though they apparently correspond to different altitudinal zones of vegetation. Among the causes of discrepancy, one could consider, for example, variation from one sampling to another because of differences in the content composition inside the colon.

The presence of the macroremains of Poaceae, possibly cereals, in the ileum is suggested by the rbc L DNA analysis cluster H. The same type of analysis indicated the presence of Poales cluster J and Eudicotyledons cluster I macroremains in the colon. As the 18S rDNA analysis of the colon content indicated the presence of Triticeae, we may presume that the Poales fraction of the colon macroremains is composed by cereals.

If we compare the result of the molecular and of the microscopic analysis of the colon Table 2 , it can be observed that although both show the presence of cereals and dicotiledons, the relative proportions of the two types of remains vary considerably from one analysis to another. In this case, the different results could, possibly, be explained by the fact that cereal residues possibly keep morphological details better than dicots, thus leading to an underestimation of the latter when the examination was performed by microscopy.

Identification of fungi by use of the 18S rDNA sequence is less accurate than that of plants. However, the fact that both the Urediniomycetes and the Heterobasidiomycetes classes include psicrophilic genera such as Leucosporidium and Mrakia and that, in addition, the consensus sequence F is very close to a fungal sequence T44NS-7 previously obtained in a study on the DNA of the man's grass clothing 20 , suggest that the fungi present in the intestines are most probably representatives of the cold-adapted microflora that developed after the corpse and the equipment became entrapped in the ice Stable carbon and nitrogen isotope analysis performed on the Iceman's hair led Macko and colleagues 19 to conclude that the man was a vegetarian, or even a vegan.

This conclusion was rejected by Dickson and colleagues 4 on the basis of two kinds of evidence: i paleobotanical analyses showing the presence of unidentified muscle fibers in the food residue from the Iceman's colon together with a prevailing proportion of plant fragments mainly cereals of the wheat and wheat—rye type ; and ii reinterpretation of the Macko and colleagues stable isotope analysis showing that the data were indicative of an omnivorous rather than a vegetarian diet. Of the utmost interest, therefore, are the results of the DNA analysis of the animal component of the intestinal content, although they are obviously indicative only of the composition of the last meals and not of the diet.


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  • In the course of the first archaeological exploration of the Iceman site, two small bone splinters were discovered at the base of the rock shelf, in addition to the other remains. Anatomical and zoological investigations performed by von den Driesch and Peters 23 led them to conclude that the pieces were lateral apophyses of a fourth and a fifth cervical vertebra of a male Alpine ibex C.

    This discovery gave rise to the hypothesis that this meat had represented the Iceman's ultimate food reserve. Such a hypothesis has resisted unchallenged for many years 1 — 4. We are now able to demonstrate that although the use of ibex meat is confirmed by the result of the analyses of the colon content, the last meal of the Tyrolean Iceman was actually composed of red deer meat. The use of red deer meat as a food supply is perfectly consistent with the paleozoological identification of the materials used by the iceman to manufacture his equipment: domestic goat and red deer in the leggings and the punch of a retoucher an implement used to sharpen the edge of stone tools , a long curved spike, and a bundle of four points were all made from red deer antler 1 , 2.

    He lived during the 4th millennium B.C.

    According to some authors 24 , 25 , the deforestation in European Mesolithic was mainly aimed at favoring the growth of red deer herds, which were maintained in a state of semidomestication by means of selective hunting. The strict connection between Neolithic man and red deer is witnessed by the wealth of artistic representation of this animal among the archaeological finds in the central and eastern Alps; the polished and carved stones of the Val Camonica, in particular, evidence how the red deer was central to the interests of the prehistoric populations along the Alpine arc All of the more so because it has been recently proposed that the Iceman might have been the victim of a ritual sacrifice Analyses performed through the years have shown that there is great variation in the plants eaten, and this has given rise to the view that some of the last meals may have had ritual significance, though it is still matter of speculation whether the above-mentioned individuals were actually victims of sacrifices or, rather, of executions.

    Tollund Man, for instance, had eaten many species, including barley, willow-herb, linseed, and gold of pleasure. Grauballe Man and Lindow II had wheat and barley as major cereals in their intestines, together with some weed seeds, whereas a Borre Fen individual had no cereal component, just wild species. On the other hand, corn spurrey Spergula arvensis is present in the Huldremose, Grauballe, and Borre Fen individuals. We can observe that, regarding the presence of cereals, the last meals of the Iceman seem close to those of Grauballe and Lindow II. On the other hand, virtually nothing can be said about the animal component in the intestinal content of the bog bodies, because of the poor state of histological preservation of the remains in the case of Grauballe Man, the consumption of meat has been inferred from the finding of ova of the red squirrel parasite Eimeria mira and of the impossibility of performing molecular analyses because of complete DNA degradation Forensic literature reports that the stomach starts emptying within 10 min from the first food ingestion.

    It is known, however, that the speed of stomach emptying can vary widely according to meal composition whether, for example, the meal is richer in carbohydrates rather than in meat or lipids.

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    In addition, this parameter can vary according to the state of health of the individual, being substantially slower in gastropathic subjects and, what is even more important, in case of protracted emotional stress The latter condition, in particular, seems to apply to the Iceman, because, as reported in the Introduction, he had been hit by an arrow that did not kill him immediately We can finally observe that although the composition of his last meals offers no obvious contribution to the hypothesis on whether the Iceman was or was not the victim of a ritual sacrifice, the finding of ibex and deer meat certainly strengthens the one that, among other possible social roles, he covered that of hunter.

    Almost 30 years ago, Nanna Noe-Nygaard 32 showed that Mesolithic deer and wild boar hunters used to aim their arrows and spears at the left shoulder blade as this gave them the best chance of killing the prey at the first shot.

    We also thank Luca Pedrotti, Parco Nazionale dello Stelvio, for providing an ibex specimen, Rita Lorenzini, Istituto Zooprofilattico, Teramo, for the red deer DNA, Patrizia Ballarini for the microphotograph, and three anonymous reviewers for their constructive criticism. E-mail: francougo. Data deposition: The sequences reported in this paper have been deposited in the GenBank database accession nos.

    NOTE: We only request your email address so that the person you are recommending the page to knows that you wanted them to see it, and that it is not junk mail. We do not capture any email address. Skip to main content. Franco Rollo. Materials and Methods Sample Collection. DNA Extraction. PCR Amplification and Sequencing. Sequence Data Analysis. Aspartic Acid Racemization Analysis. The plants were from at least 75 different species, only 23 of which live in that precise area today. This moss is typically found in wetlands and probably came from the bottom of the Vinschgau valley in South Tyrol, Italy.

    The moss had long been used for staunching wounds because of its mild antiseptic properties. It may have been to treat the deep wound he received to his right palm possibly 48 hours or less before his death, says Dickson. Dickson was also surprised to find fragments of the moss Neckera complanata in his intestines. In his digestive tract, the researchers also found several other species of moss that appear to be from an area north-west of the Schnalstal valley called Kurzras. This is around 15 kilometres from Vinschgau, at an elevation of around metres.