Veget Hist Archaeobot (2009) 18:1322 DOI 10.

1007/s00334-007-0141-7

ORIGINAL ARTICLE

Six mosses from the Tyrolean Icemans alimentary tract and their signicance for his ethnobotany and the events of his last days
James H. Dickson Wolfgang Hofbauer Ronald Porley Alexandra Schmidl Werner Koer Klaus Oeggl

Received: 4 July 2006 / Accepted: 19 March 2007 / Published online: 10 January 2008 Springer-Verlag 2008

Abstract Six different mosses have been recognised in samples taken from the intestinal contents of the 5,200year-old Iceman from the Eastern Alps. Four of the species are important in understanding the lifestyle of the man and/ or bear on the events during the last few days of his life: Anomodon viticulosus, Hymenostylium recurvirostrum, Neckera complanata and Sphagnum imbricatum. The past and present chorology and habitats of the Hymenostylium are discussed in detail, as is the ethnobotany of the day Da n Sphagnum concerning both the Iceman and Kwa `nch , the rst ancient glacier body from North America. Ts Keywords Iceman Neolithic Archaeology Mosses Intestinal contents Austria Italy Alps

Introduction The senior author began studying the numerous remains of tzi) in bryophytes found with the Iceman (often called O 1994. As more and more samples have become available, so the importance of bryophytes, especially mosses, for the understanding of the lifestyle and last days of this 5,200-

J. H. Dickson (&) Graham Kerr Building, Institute of Biomedical and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK e-mail: j.dickson@bio.gla.ac.uk W. Hofbauer A. Schmidl W. Koer K. Oeggl r Botanik, 15 Sternwartestrasse, Institut fu A6020 Innsbruck, Austria R. Porley English Nature, Foxhold House, Crookham Common, Thatcham RG19 8EL, UK

year-old man has increased (Dickson 1997, 2000, 2003; Dickson et al. 1996, 2005a, b). In all, over 100 samples of mineral sediment and ice, and 35 taken from the clothes or extracted from the intestines have been examined for bryophytes and more than 80 species have been recognised, mostly mosses but also some ten or more liverworts. In 1996 the only intestinal sample then being examined by Klaus Oeggl (2000) and colleagues at Innsbruck contained a single moss leaf, which was identied as Neckera complanata by JHD (1997). Because it was already known that the Iceman had been carrying that species (Dickson et al. 1996), the interpretation was that the moss had been used as wrapping for his provisions. In 2000 several more samples of ingesta, from three different organs of the gut, were studied for pollen and macroscopic remains by WK and AS, both of whom were working under the direction of KO. When the body was roughly recovered from the ice, a cordless drill was used to loosen the ice. Very unfortunately, this led to substantial destruction of the left buttock, even the rectum being damaged and so the possibility that the faecal material was contaminated needs to be considered (Egg et al. 1993). Such a consideration does not apply to the ileum and colon (and other internal organs), which were totally unaffected and so remained intact, as they had for over 5,000 years. Though not toxic, mosses are neither palatable nor nutritious. There are no records of mosses eaten as a staple anywhere at any time. There are only very sparse reports of mosses eaten as famine food or as very minor components of meals (Dickson et al. 2000). Similarly, there are exceedingly few records of mosses used for internal medical treatment and none apply to the species discussed in this paper. The Iceman is the rst glacier mummy to have fragments of mosses recovered from the alimentary tract but now there `nch (Long Ago Person Found), the rst day Da n Ts is Kwa

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ancient human body from a North American glacier to be investigated scientically (Dickson et al. 2004; Mudie et al. 2005). This corpse was that of an aboriginal man some aged 1720 years who may have lived about 550 years ago but perhaps more recently than that. He was found melting from a glacier in north-westernmost British Columbia, Canada. Mostly as very small pieces, in the samples taken from the duodenum to the rectum, there are not less than 12 different mosses still under study, double the number previously reportedongoing work by one of us (Dickson) has identied at least six different mosses (Dickson et al. 2005b, p 11). This new study makes an informative comparison with the Iceman mosses, especially in the case of Sphagnum (Bogmosses). Remains of mosses are often recovered from archaeological sites and very frequently they are large, richly branched species, which have shapes which bryologists call wefts, rough mats and fans (Bates 1998). They are soft but springy and do not readily crumble or fall apart when collected. They are highly suitable for such purposes as caulking, stufng, wiping and wrapping. Materials and methods The moss remains from the intestines of the Iceman were discovered on the microscope slides of preparations of the ingesta made for pollen analysis (WK) and analysis of macroscopic remains (AS). In order to interpret best the numerous bryophytes found with the Iceman, the valleys and mountains to the north and south of the Iceman site have been recorded on a 1 km square basis. Several bryologists have been involved, mainly JHD and WH but also RP, Gordon Rothero and Andrew McMullen (Dickson 2003). The results of this survey are shown on Figs. 1, 2 and 3. The habitats and altitudes of the especially important species were noted. Apart from the more than 80 species of bryophytes found with the Iceman other species have been recorded. In all about 500 different taxa have been found growing in the region up till now (summer 2005).

Unidentied fragments The unidentied species consists of two very badly preserved, tiny, similar fragments, which the senior author considers cannot be recognised even to the genus.

Pogonatum/Polytrichum sp.: Haircap The fragment assigned to Pogonatum/Polytrichum is a badly preserved lamina showing only one tooth (singlecelled). Because there have been thousands of fragments of Polytrichum and Pogonatum spp recovered from the sediments and ice samples, it is probable that the fragment can be considered a contaminant and so needs little further discussion. The various species of these two genera are nowadays widespread in the whole recorded ll 1991; Buchner et al. area and indeed in all Tyrol (Du 1993) and an exact identication would probably give little or no information concerning the last days of the Icemans life.

Neckera complanata Hedw., Flat Neckera, life-form: fan (Bates 1998) It is very noteworthy that only Neckera complanata was found in all ve samples whereas every other taxon occurred only once (Table 1). JHD has already extensively discussed the importance of this species for the understanding of the Icemans provenance and way of life (Dickson 2000). Figure 1 reveals that the species occurs today in various localities very nearby to the south of the tzi site, whereas, to the north, it occurs 30 and more O kilometre away; this is a distribution pattern which is true also for Anomodon viticulosus (Fig. 2). It is certain that the intestinal samples represent more than one meal and not less than three. The explanation that this particular moss had been used as food wrapping is strengthened by its occurrence in every one of the ve samples. Any other explanation such as unintentional swallowing in drinking water now seems very much less likely.

Results and discussion The mosses found in the intestines of the Iceman are diverse in ecology and life-form (Table 1). Each species needs to be explained separately and no one explanation accounts for them all collectively. The questions, which arise immediately are 1. 2. Where did the Iceman come in contact with each species? How did each come to enter his alimentary tract? Anomodon viticulosus (Hedw.) Hook. & Tayl., Rambling tail-moss, life-form: rough mat Like Neckera complanata, this moss often grows on limerich, shady rock in woodland and these two species can be found growing side by side (Fig. 4). Both occur in Schnalstal though the Anomodon does not reach the 1,750 m a.s.l., which is the altitudinal limit of the Neckera there (Figs. 1, 2). That the Anomodon had been an accidental

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Fig. 1 Map showing the present distribution of Neckera complanata to the south of the site. This and the two other maps are based on the authors eldwork as mentioned in the text

admixture with the Neckera when the Neckera was gathered is a plausible explanation.

Hymenostylium recurvirostrum (Hedw.) Dix., Hook-beak tufa-moss, life-form: tall turf and large cushion There are eight leaves on one microscope slide and they could well have derived from one very short piece of leafy stem (Fig. 6). The preservation is very good with the cells showing numerous, well developed papillae and so the leaves can be referred to var. scabrum (Linb.) Limpr. [=var. latifolium (J.E. Zetterst.) Wijk. & Margad.]. Four of the 25 Tyrolean and Bavarian specimens in the University of Innsbruck herbarium are that variety, but its taxonomic

value is questionable and its genetic status is unknown. Regarding the species as highly plastic, Zander (1977) doubts the value of infraspecic taxa in this species, as do Crum and Anderson (1981). With regard to var. cataractacum (Schimp.) Podp., Smith (2004, p. 303) states that it appears to be a dustbin taxon in which aberrant forms of H. recurvirostrum are placed and cannot be maintained. In decades of studying mosses recovered from Quaternary deposits and from archaeological layers, the senior author has never before found this species (Dickson 1973, 1986; numerous later papers) nor does he know of discoveries by other palaeoecological and archaeobotanical investigators. Not just because it is a highly unusual subfossil, ASs discovery of the Hymenostylium was not predictable

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Fig. 2 Map of the present distribution of Anomodon viticulosus south of the site

because it is unrepresented in any other of the very many Iceman samples previously examined for plant remains and already carefully searched both quantitatively and qualitatively for bryophytes. The only sample that has produced the Hymenostylium is the rectal one. Despite the damaged rectum, it seems virtually impossible that this particular moss fragment is a contaminant. If that were so, then the fragment had been preserved in the ice immediately beside the corpse or on top of the sediment under it and somehow the fragment became mixed with the faeces as a result of the use of the drill by one of the party of retrievers of the corpse. H. recurvirostrum is very widely spread in Eurasia and the Americas and even reaches 83N in Peary Land, Greenland, the northernmost land in the world (Holmen 1960). However, the concern of this paper is the Tyrol, a region, which is partly western Austria and partly

northernmost Italy (Fig. 5). This geographical restriction arises from the botanical and other archaeological and scientic data which rmly suggest that the Icemans home area was that region and not further aeld, and, more precisely, that he had lived to the south and/or east of where the body was found (Dickson et al. 2005b). In life-form it is a tall turf or large cushion. Herbarium specimens are often about 45 cm tall but 1012 cm can be reached. It is a moss strongly connected with lime-rich rock and numerous European oras and papers mention that it is often encrusted with tufa, a precipitate of calcium carbonate, which forms on the moss when hard water ows or trickles through it, as is often the case (Cortini Pedrotti ll 1991; Du ll and Meinunger 2001; Dierssen 2001; Du 1989; Grims 1999; Hill et al. 1992; Porley and Hodgetts 2005). Dealing with Europe as a whole, Dierssen (2001, p. 116) states often encrusted with a calcareous matrix,

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Fig. 3 Map of the present distribution of Hymenostylium recurvirostrum

Table 1 List of the species, the organ or organs in which each species was found and the number of fragments on each slide Intestinal organ Weight of sample (g) Vegetation zone (from pollen in gut) Anomodon viticulosus Hymenostylium recurvirostrum Neckera complanata Pogonatum/Polytrichum sp. Sphagnum imbricatum s.l. Unidentied sp. Ileum 0.11 Subalpine 7 Colon 1 0.10 Montane 3 1 1 Colon 2 0.04 Montane 1 Colon 3 0.05 Subalpine 2 Rectum 0.39 Subalpine 8 10 1 2

It also shows the vegetation zone in which the Iceman had been when ingestion took place; these deductions were made from pollen analysis compared with pollen surface samples (Oeggl et al. 2007)

below wet cliffs, on damp calcareous rock-ledges. More or less similar statements are found in many bryological texts concerning North America and elsewhere.

rtensson While listing the usual calcareous habitats, Ma (1956, p. 102), dealing with Swedish Lappland states It grows in small scanty specimens almost everywhere if the

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Fig. 4 One of the fragments of Anomodon viticulosus (part of the base of a leaf) from the sample of food residue from the colon (colon 1 sample)

substratum is calcareous enough and the habitat is not too wet (shady small crevices in erratic rocks or boulders, soliuction terraces, frostheaving soil etc.). JHD and WH are not tztal Alps, nor with it familiar with such habitats in the O growing on metalliferous waste, as in Britain (Blockeel 1992), or in Eriophorum-Carex meadows, as in Greenland (Holmen 1960) or growing ... occasionally on the ground or tree bases in forests; also on rocks in glaciers., as in China (Xing-jiang and Crosby 2001, p. 189) or rarely on trees as in Mexico (Zander 1977, p. 255). This ecological preference for lime-rich rock is well shown by the specimens from Tyrol and Bavaria in the Innsbruck herbarium. Of the 25 specimens examined, 18 had obvious limy encrustations or effervesced immediately on the application of 10% HCl, whereas seven did not exhibit either feature. So the preference is clear but the species cannot be said to be exclusive to lime-rich substrata. Nonetheless, its limitation is mainly edaphic and microtopographic rather than macroclimatic. Though it does not reach the highest altitudes in Tyrol, which are well over 3,000 m, in Austria it ascends on occasion well above the timberline, which is around 2,2002,300 m, depending on aspect. The Austrian altitudinal limit is 3,000 m (in the nival zone), according to Grims (1999) who states that it is mainly high montane to alpine in distribution (c. 1,300 to well over 2,000 m). This principal altitudinal range does not appear to be the case in Tyrol, at any rate those parts of Tyrol which were the Icemans territory. Hymenostylium was not recor tztal Alps by Reisigl and ded in the nival zone of the O Pitschmann (1958) or Pitschmann and Reisigl (1954).

tzi. Fig. 5 Map showing the area of the discovery of the Iceman O Places mentioned in the text are shown, as are the international tzi site lies at frontier, rivers and land above 2,000 m a.s.l. The O 3,210 m a.s.l.

In only one of the 125 1 km2 bryologically surveyed as yet (to July 2005) has Hymenostylium been found (Fig. 3). So it has not been found in the very near vicinity of the Iceman site. While a total absence from the squares immediately around that site cannot ever be proved, it seems very probable that this moss is absent or, at any rate, very sparse there where no large outcrops of limestone nor of any other strongly calcareous rock. Because neither the topography nor bedrock geology has changed in any substantial way likely to have been important for the arguments made here, H. recurvirostrum may well have been similarly sparse or absent during the Icemans time, even if the climate was somewhat different, perhaps somewhat warmer or, as some now think, colder (Magny and Haas 2004; Magny et al. 2005). Sparse or absent as it may or may not be close to the site, we have found it, or others have recorded it (Dalla Torre and Sarnthein 1904), not too far off to the south in Vinschgau and nearby areas at several places as follows: near Schlanders (submontane, 900 m, Fig. 5), Valdaunbach near Laas (submontane/lower montane, 900 1,000 m), Partschins Waterfall (submontane; Strassemariern ober dem Partschinser Wasserfalle, c. 500 m; Schiffner ll (submontane/lower montane), Meran (too 1899, above To

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Fig. 6 One of the leaves of Hymenostylium recurvirostrum from the sample of food residue taken from the rectum

vague to designate), Marlingerberg (submontane/montane), llan (submontane, 700 m). hl near Vo Oberthalmu Some 15 km to the northeast, it has been found at Hohe Mut near Obergurgl (subalpine/alpine, 2,500 m) and at the nearby Granatenwand where there are marble outcrops. In his papers on the mosses of Alto Adige (South Tyrol) and Trentino, Venturi (1879, 1899) describes it as ubique (everywhere) and frequente; this is an assessment which does not square with our recording in South Tyrol. In Cortini Pedrottis ora (2001) it is called common from lowland to alpine regions and var. scabrum is mentioned as very rare. How did Hymenostylium come to enter the Icemans alimentary canal? The life-form and hard, limy encrustations of Hymenostylium militate against it being used as wrapping, stufng or wiping. Could the Hymenostylium have been in the rectal sample for the same reason as the Neckera and the Anomodon? However, while these three species can grow in close proximity they are never a part of precisely the same moss community. The Neckera and Anomodon grow in humid, often shady places (e.g. on bark) but not on wet substrata whereas the Hymenostylium is very often on wet substrata, indeed exclusively so in South Tyrol, as far as JHD, WH and RP have observed. A more plausible explanation is that it was swallowed when the Iceman drank water during the last few days of his life. On the assumption of normal throughput rates of ingesta, because the Hymenostylium leaves were recovered

from a faecal sample, it is likely that they were swallowed more than 1 day before the Iceman died. Unpublished pollen analysis indicates that the Iceman was in the subalpine zone at the time represented by the faecal material. However, if the distribution pattern of the Hymenostylium was fundamentally the same 5,200 years ago as known now, one might argue that the accidental ingestion took place when the Iceman was at low or moderate altitude, that is to say the submontane or lower montane zones, but probably not in Schnalstal. However, during our recording we may as yet have missed the Hymenostylium in the subalpine and alpine zones, especially, for instance, at the eastern end of Pfossental where there are extensive outcrops of marble well above the timberline below both the eastern and chl, a peak reaching 2,895 m a.s.l. western slopes of Eisjo (Fig. 6).

Sphagnum imbricatum Hornsch. ex Russ., Imbricate bogmoss, life-form: tall turf Dickson et al. (2005a) considered in detail the discovery of a tiny branch leaf fragment of S. imbricatum s.l. by WK on a pollen slide made from the colon 1 sample (see Oeggl et al. 2008, this volume). On grounds of present biogeography, the fragment is more likely to have derived from S. afne (Ren. & Cardot) Flatberg rather than S. austinii

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(Sull.) Flatberg. It was a very unexpected nd because the species is not known at present in South Tyrol and in North Tyrol it occurs only in the far northeast, in the vicinity of Kufstein, a long way from the Iceman site. We argued that, though the species does not grow now in South Tyrol, it probably did so some 5,000 and more years ago in Vinschgau, the broad, low-lying valley south of the Iceman site and, additionally, we claimed that it could have been ingested accidentally in drinking water. That the colon 1 sample contains cysts of the testate amoeba (Arcella catinus-type) supports the suggestion that the Sphagnum was tzi having drunk water from boggy ground, ingested by O the habitat of the amoeba. However, we did not mention the ethnobotany of Sphagnum which deserves some detailed comments. As discussed by Dickson and Dickson (2000), bogmosses have very detailed Holocene and archaeobotanical histories and a well-documented ethnobotany, especially from North America (Johnson and Vitt 1996). Alone among mosses, species of Sphagnum have been used for external medical treatment as wound dressings, a use that continued on a large scale well into the 20th century during both world wars. Dried Sphagnum is highly absorbent and, much more than that, there are the wound-healing properties of Sphagnum holocellulose (sphagnan, a kind of pectin), discussed by Painter (2003). This acts by immobilizing bacterial cells and depriving them of their nutrients. The native peoples of North America have used bogmosses as diapers (nappies) for their babies and found these plants highly efcient for that purpose. Dry bogmosses will soak up much liquid and their acidifying reaction is inimical to bacteria. Decades ago, the senior author was involved in one prehistoric discovery of Sphagnum palustre from a Scottish Bronze Age site (Dickson and Dickson 2000). Perhaps this nd does more than hint at the very ancient use of bogmosses as wound dressings. Unique, certainly in British archaeology if not elsewhere, the fragments were found inside the kist burial of a young man whose skeleton was well preserved but, as always so far with such burials, no esh remained and so there is no way of knowing if he had died from wounds or not. It is now known that the Iceman had two wounds, one in the back (from an arrow, Gostner and Egarter Vigl 2002) and the other a deep cut in his right palm (precisely how inicted is not known with certainty, Nerlich et al. 2003). If he knew of the useful properties of bogmosses, as seems entirely plausible, then he may have gathered some to staunch the wound or wounds and so tiny pieces could well have stuck to the blood drying on his ngers and then he accidentally ingested some of them when next eating meat or bread as we know he did during his last few days (Oeggl

2000; Rollo et al. 2002). If this is true then the implication is that either or both of the wounds happened at low to only moderate altitude but not in lower Schnalstal where no species of bogmoss is known. Such an argument would have no force, however, if the Iceman had been carrying the Sphagnum. That he may have done so cannot be discounted but will never be known with any certainty. One bud of Sphagnum, tentatively ngstr. was the only other identied as teres (Schimp.) A discovery of a bogmoss with the Iceman; it was recovered from his clothes and the question again is how did it get there. It is a species well known in the Tyrol and indeed grows today a mere 4 km southwest of the site (at about 2,000 m near Kurzras) and so it may have become attached during his last journey if the Iceman had crossed boggy terrain somewhere. However, again perhaps, it had been deliberately carried. `nch (BC, day Da n Ts A rectal sample taken from Kwa Canada) contained a very well preserved branch fragment about 1.5 mm long of Sphagnum Section Acutifolia. At least four species of that large section of the genus occur in the relevant region in JHDs personal experience: S. capillifolium (Ehrh.) Hedw., S. fuscum (Schimp.) Klinggr., S. girgensohnii Russ. and S. warnstori Russ. The rst and last named can be found a little way into the alpine zone which extends from about 1,000 m a.s.l. upwards in that region. However, no specic identication can be made. No matter what the species, the same question as for the Icemans bogmosses arises. Why was the bogmoss there? Was it mere accident or did it result from an external treatment of a wound? No wounds were seen on the corpse `nch at the autopsies. However, some day Da n Ts of Kwa parts of the body such as the head and parts of the limbs were only bones retaining little or no esh and so there could have been esh wounds.

Conclusions The mosses found in the alimentary tracts of the Iceman `nch are clues to both their nal day Da n Ts and Kwa routes and the events of their last days. In the case of the `nch , as more of the tiny scraps are day Da n Ts Kwa identied so more may be revealed. If more ancient glacier mummies are found and their innards contain bogmosses it will become more and more likely that the wound dressing hypothesis will prevail.

Amendment In mid August 2006, JHD and WH, accompanied by Mlle ` ve Le crivain, recorded mosses in uppermost Genevie

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21 cereals, pollen, moss and whipworm) and stable isotope analyses. Phil Trans Roy Soc Lond 355:18431851 Dickson JH, Richards MP, Hebda RJ, Mudie PJ, Beattie O, Ramsay S, Turner NJ, Leighton BJ, Webster JM, Hobischak NR, Anderson `nch , the rst day Da n Ts GS, Troffe PM, Wigen RJ (2004) Kwa ancient body of a man from a North American glacier: reconstructing his last days by intestinal and biomolecular analyses. Holocene 14:481486 Dickson JH, Hofbauer W, Koer W, Oeggl K, Platzgummer J (2005a) How to nd the bogmoss, Sphagnum imbricatum s.l., in the South Tyrol, Italy: Microscopically examine the Icemans colon contents. Veget Hist Archaeobot 14:207210 Dickson JH, Oeggl K, Handley LL (2005b) The Iceman reconsidered. Sci Am 15:413 Dierssen K (2001) Distribution, ecological and phytosociological characterization of European bryophytes. Bryophyt Biblio 56:1 289 ll R (1991) Die Moos Tirols unter Beru cksichtigung des Pitztales/ Du ztaler Alpen. IDH-Verlag, Bad Mu nstereifel O ll R, Meinunger L (1989) Deutschlands Moose. Bad Mu nstereifel, Du IDH-Verlag Egg M, Goedecker-Ciolek R, Groeman van Waateringe W, Spindler K (1993) Die Gletschermumie vom Ende der tztaler Alpen. Jahrb Ro m-Germ Zentralmus Steinzeit aus den O Mainz 39:1128 Gostner P, Egarter Vigl E (2002) INSIGHT: report of radiologicalforensic ndings on the Iceman. J Archaeol Sci 29:323326 Grims F (1999) The mosses of Austria. Catalogus Florae Austriae, Part II. Bryophyta No 1. Musci. Austrian Academy of Sciences, Vienna Hill MO, Preston CD Smith AJE (1992) Atlas of the Bryophytes of Britain and Ireland. Harley Books, Colchester Holmen K (1960) The Mosses of Peary Land, North Greenland. Medd n 163:196 Gr Johnson LM, Vitt DH (1996) The selection of sphagnum for diapers by indigenous North Americans. Evansia 13:103108 Magny M, Haas J-N (2004) A major widespread climatic change around 5300 ca. yr BP at the time of the Alpine Iceman. J Quat Sci 19:423430 Magny M, Leuzinger U, Bortenschlager S, Hass JN (2005) Tripartitie climatic reversal in Central Europe 56005300 years ago at Arbon-Bleiche, Switzerland. Quat Res 65:319 rtensson O (1956) Bryophyes of the Tornetra sk Area, Northern Ma Swedish Lappland II Musci. Kungl Svenska Avh i Naturv 14:1 321 Mudie PJ, Greer S, Brakel J, Dickson JH, Schinkel C, Peterson-Welsh R, Stevens M, Turner NJ, Shadow M, Washington R (2005) Forensic palynology and ethnobotany of Salicornia species (Chenopodiaceae) in northwest Canada and Alaska. Can J Bot 83:111123 Nerlich AG, Bachmeier B, Zink A, Thalhammer S, Egarter-Vigl E tzi had a wound on his right hand. The Lancet 362:334 (2003) O Oeggl K (2000) The Diet of the Iceman. In: Bortenschlager S, Oeggl K (eds) The Iceman and his natural environment. The man in the Ice, vol 4. Springer, Wien, pp 89115 Oeggl K, Koer W, Schmidl A, Dickson JH, Egarter Vigl E, Gaber O tzi, the (2007) The reconstruction of the last itinerary of O Neolithic Iceman by pollen analyses from sequentially sampled gut contents. Quat Sci Rev 26:853861 Oeggl K, Koer W, Schmidl A (2008) New aspects on the diet of the tzi. Veget Hist Archaeobot (in Neolithic Tyrolean Iceman O press) Painter TL (2003) Concerning the wound-healing properties of Sphagnum holocellulose. J Ethnopharmacol 88:145148 tztal Pitschmann H, Reisigl H (1954) Zur nivalen Mossora der O Alpen (Tirol), Rev Bryol Lichenol 23:123131

Pfelderertal with the deliberate intention of nding Hymenostylium recurvirostre there. As stated in our paper, the point is that, in that area, there is extensive exposure of marble and, not only that, there are springs and streams and, consequently, there is wet limy rock. The search was successful. The precise details of the fruiting specimen, gathered by WH, are as follows: Wet rock at about 2,150 m, west of Lazins, westernmost Pfelderertal. The locality is just east of the easternmost 1 km2 shown in Fig. 3. This altitude lies in the subalpine zone and so it is no longer true that we have not seen the species in that zone in the not too distant vicinity of the Iceman site. Nobody has ever considered that the Iceman could have reached Hauslabjoch by rst travelling westwards up Pfelderertal.
Acknowledgments JHDs work on the Tyrolean Iceman is supported by the Royal Society of London, the Austrian Academy of Sciences and the Carnegie Trust for the Universities of Scotland. Dr M. O. Hill kindly examined the fragment of Sphagnum Section Acutifolia. Useful comments were made by Dr. Heinjo During of the University of Utrecht.

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