Holocene distribution and extinction of the moose (Alces alces, Cervidae) in Central Europe

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Review

Holocene distribution and extinction of the moose (Alces alces, Cervidae) in Central Europe By U. Schmo¨lcke and F.E. Zachos Institut fu¨r Haustierkunde, Christian-Albrechts-University, Kiel, Germany Receipt of Ms. 8.4.2005 Acceptance of Ms. 26.8.2005

Abstract Moose (Alces alces L.) were among the first large mammals to recolonize Central Europe after the last glaciation. Already during the Allerød they established themselves in most parts of the area. In the early Holocene their distribution range extended from the Pyrenees to Denmark and from Austria to Great Britain and also covered eastern Central Europe where they still occur today. In the Preboreal, the moose slowly vanished from the southwestern parts of its distribution range, leading to its extinction in France and, later, in England. During the Atlantic period, the moose died out in large parts of Denmark and population densities apparently decreased in the rest of Central Europe as well. Around the birth of Christ only relict populations were left in western Central Europe, which finally became extinct in early medieval times. In Thuringia and in the region northeast of the river Elbe as well as in central Poland, some stocks persisted until the high and late Middle Ages. The causes of the gradual extinction in Central Europe during the Holocene are complex. Changes in vegetation, climate and sea-level, the increasing fragmentation of habitat through human activities and hunting were, at different times, important factors. In the recent past, however, moose have repeatedly migrated from the east towards the west. The development of its distribution range since the end of the Second World War as well as experiences with Scandinavian populations show that moose are able to thrive in close proximity to humans and that a future expansion of its distribution range towards the west seems possible. r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. Key words: Alces alces, Holocene, Europe, archaeozoology, palaeoecology

Introduction The moose (Alces alces) is the largest living cervid and one of the biggest terrestrial mammals in Europe, second only to the European bison. Its present distribution range extends from northern, central and eastern Europe across northern Asia to North America. Moose occur almost continuously between 50 and 701N latitude and south to about 401 (Nygre´n 1986; Geist 1998).

Systematically, moose (Alcinae) are related to the New World deer and branched off in the late Tertiary, being present in the Palaearctic mainly as A. (Libralces) gallicus and A. (Libralces) latifrons (Nygre´n 1986; Geist 1998). According to the orthodox view of Alcine phylogeny, A. latifrons gave rise to the modern moose A. alces (cf. Kurte´n 1968; Heintz and Poplin 1981; Lister 1987, 1993;

1616-5047/$ - see front matter r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2005.08.001 Mamm. biol. 70 (2005) 6
329–344

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Kahlke 1990, 1994; but see von Koenigswald 2002 and Sher 1987 for criticism). The origin of the Alcine lineage is Central Asia (Kahlke 1990) or Europe (Heintz and Poplin 1981), while modern moose originated in Asia and then subsequently colonized Europe and America (Hundertmark et al. 2002). Alces alces, as compared to the fossil forms, is characterized by shorter antler beams and the loss of any contact between the premaxillary and nasal bones. It first occurred in the Riss/Saale glacial. Modern moose were absent from North America until after the last glacial maximum when they moved in from eastern Siberia via Beringia, being part of the Siberian fauna that filled the vacuum left by the late Pleistocene ‘‘megafaunal’’ extinctions some 10,000 years ago (Geist 1987; Guthrie 1990). Moose have traditionally been divided into two types: European-West Siberian (up to the Yenisei River) and East SiberianAmerican (Flerov 1952). These two groups differ in antler plan (three-pronged in European-type, four-pronged in American-type moose, Geist 1987, 1998) and also in karyotype (2n=68 for European, 2n=70 for eastern Asian and American moose, Gustavsson and Sundt 1968; Hsu and Benirschke 1973; Boeskorov 1996, 1997). The European moose, which this review is dealing with, is referred to as Alces alces alces. Based on genetic data, Hundertmark et al. (2002, following Avise 2000) suggested a recent bottleneck in European moose although the sample size for Europe in their study is too small to draw definitive conclusions. Nevertheless, this scenario is in line with the history of moose in the European part of the former Soviet Union, an area covered by Hundertmark et al. (2002), as described detailedly by Heptner et al. (1966) and Heptner and Nasimowitsch (1967). According to these authors, moose populations drastically declined in the first half of the 19th century following the rapidly increasing demand of moose leather since Peter the Great, some 100 years earlier, had decided to clothe the army almost exclusively with this material. When the use of moose leather was stopped, populations slowly recovered from this bottleneck, but after the Revolution poaching exploded and moose

Table 1. Subdivision and timescale (BP=years before present) of the Late Pleistocene and the Holocene (after von Koenigswald 2002). Holocene

Subatlantic Subboreal Atlantic Boreal Preboreal

3000 BP–present 5600–3000 BP 9100–5600 BP 10,000–9100 BP 11,500–10,000 BP

Late Pleistocene Younger Dryas 12,700–11,500 BP Allerød 13,500–12,700 BP

stocks again declined catastrophically. After that, due to strict protection measurements, moose numbers increased again. Similar bottlenecks have been reported from Scandinavia, where the species was nearly extinct at the beginning of the 19th century (Ru¨lcker and Sta˚lfelt 1986), and Poland (see below). Being present since the Allerød period (Table 1), the moose was one of the first large mammals to immigrate into Central Europe at the end of the Pleistocene (Willms 1987; von Koenigswald 2002). Studies from Denmark show that A. alces survived in a woodland or open park tundra 350 km south of the ice front (Aaris-Sørensen 1992). According to Willms (1987; see also Andersen et al. 1990; Do¨hle 1996; Szymczyk 1973), the Holocene history of the moose was as follows. During the late Pleistocene, moose were very rare north of the Alps but became more abundant (and in the Baltic countries even the most frequent ungulate species) in the Preboreal. In the Boreal, the English moose population, now separated from the mainland, declined, and Scandinavia, with the ice sheets retreating, was colonized, on the one hand, via Denmark and Scania (Sweden), and, on the other hand, from Russia via Karelia to Finland and north Sweden. At the same time, the southern edge of the species’ distribution shifted northwards with the moose-dominated zone (i.e. the area in which moose were the most abundant ungulates) already being north of Scania. From the following Atlantic period on, northern Europe was free from glaciers. At most Central European excavation sites from this period, the moose was rare while in

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Denmark and Scania, it was found in several places. In England, the moose seems to have become extinct during the Atlantic, and at the Danube’s Iron Gate between Romania and Serbia, it was also found for the last time while in the east it probably occurred up to the Black Sea region. In Central Europe, the ungulate fauna was dominated by bovids, wild boar, red or roe deer. In the Subboreal, the period with the best coverage of moose distribution, moose were found in the Netherlands, Belgium, France, Switzerland, Austria and even Hungary. Thus, moose occurred in parts of western and in most of Central Europe as well as in eastern Europe and Scandinavia. In the first stages of the Subatlantic, from the Bronze and Iron Age through the Roman period, the moose seems to have expanded southwards, but later, during the Middle Ages, its distribution area was greatly reduced, and the species vanished from

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large parts of western and southern Central Europe. In this review, we present and discuss the Holocene distribution and extinction of A. alces in Central Europe, which, in our analysis, comprises France, the Netherlands, Belgium, Luxembourg, Switzerland, Austria, the Czech Republic, Germany, Denmark and Poland (Fig. 1). The focus is on the development from the Pleistocene–Holocene border until the Middle Ages. The more recent history of the moose, if there is any, and the chances of a future re-immigration into other parts of Central Europe are outlined. For the history and status of moose in Fennoscandia see, e.g., Ekman (1922), Ru¨lcker and Sta˚lfelt (1986), Cederlund and Markgren (1987), Nygre´n (1987), Østga˚rd (1987) and Pulliainen (1987). Heptner et al. (1966) and Heptner and Nasimowitsch (1967) give a very detailed report on moose history in the European part of the former Soviet

Fig. 1. Geographic map of the Central European regions mentioned in the text. 1: western Central Europe, 2: the Alps, 3: southern Germany/Czech Republic, 4: western Germany, 5: eastern Germany, 6: northern Germany/ Denmark, 7: Poland.

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Table 2. Numbers of moose records in different central European areas (numbers in left column refer to geographic areas in Fig. 1) and Holocene periods (after the data base Holocene History of European Vertebrate Fauna; cf. Benecke 1999). Period culture area

Late Glacial palaeolithic

Preboreal mesolithic

Boreal mesolithic

Atlantic meso-/ neolithic

Subboreal metallic

Subatlantic Roman / medieval

Total

1 2 3 4 5 6 7

6 3 1 3 1 9 1

4 2 1 1 1 4 1

5 30 3 1 0 27 9

9 19 7 5 0 28 7

6 7 6 3 3 0 11

11 13 15 7 12 14 72

41 76 33 20 17 82 101 370

Union, Tomek (1977) gives an overview of the present occurrence of moose in Poland, and Peterson (1955) presents a readable outline of the American history of A. alces. This review is based on historical sources (Prell 1941) and a large data base of published and non-published subfossil European vertebrate findings (cf. Benecke 1999), which contains 370 dated Central European moose records (Table 2). The number of undated moose records, which were not considered in this review, is much larger. Most of the dated moose findings are hunting remains from former human settlements and graves while findings belonging to a natural taphocoenosis are rare. This point causes methodological problems when the former distribution of this species is reconstructed (see also Heinrich 1999): the distribution of archaeological excavations with moose findings is not homogeneous throughout Central Europe because archaeological research has often concentrated on specific regions. The distribution of former human settlements is not homogeneous either. They were concentrated along coasts, shores and banks and in areas suitable for agriculture. In addition, there are considerable differences among soil types in the preservation capacity of bones and antlers. In particular, sandy and acid soils are not conducive to the preservation of organic material. Consequently, there may be an artificial element in the results of studies on the temporal and spatial distribution history of Holocene mammals.

Holocene development of moose distribution in Central Europe Western Central Europe (France without the Alps, the Netherlands, Belgium, Luxembourg) In the Late Pleistocene and early postglacial period, A. alces must have been part of the fauna of western Central Europe as shown by various findings from the Pyrenees, England and Belgium. During the Preboreal, it disappeared from the Pyrenees, and in central France, there are no Holocene moose records at all. Accordingly, Caesar, in his ‘‘De Bello Gallico’’ (VI, 26), reports that the animal ‘‘quae appellantur alces’’ lives in the Germanic Hercynian Forest, but not in France (Gallia). At the same time, a stable population of A. alces established itself in the area of the Netherlands, Belgium and Luxembourg with the oldest findings dating from the late glacial (Gautier et al. 1985). Nine moose records from mesolithic and neolithic sites and six records from Bronze and Iron Age settlements show that A. alces lived here during the Atlantic and Subboreal periods. Even in the Roman period findings of moose bones are not rare in the Netherlands (n=8) and Belgium (Ervynck et al. 1999). According to Lauwerier (1988) it was the rising sea-level that formed the northern lowlands near the North Sea in a favourable way for moose: it might have caused a rising groundwater-table

ARTICLE IN PRESS Holocene distribution and extinction of the moose in Central Europe

and frequent floodings, and this could have resulted in larger areas of suitable habitat. The youngest bone and antler records in the Netherlands – a few findings from Valkenburg, Dorestad and Rijnsburg (Clason 1967; Prummel 1983) – date from early medieval times (700–850 AD), and moose are last mentioned for the Drenthe Forest during the 10th century (Gandert 1941/42). There is no unequivocal evidence of its presence in later times (Clason 1999; Ervynck et al. 1999). Alpine region (south-eastern France, southernmost Germany, Switzerland and Austria) From the Alps, 76 localities with moose bones or antlers are known, ranging from the late glacial to the Middle Ages. The moose was part of the immigration wave of diverse woodland species into the alpine region after the Allerød period (Chaix 1993). Particularly in the mesolithic and neolithic, moose were obviously not rare (Chaix 1975; Chaix and Desse 1981; Hu¨ster-Plogmann and Schibler 1997), and during the Boreal and Atlantic, they were of some economic importance as shown by hundreds of bones found in Twann (Switzerland; Becker and Johansson 1981) and Auvernier-La Saunerie (Switzerland; Stampfli 1976). However, the ecological conditions, especially the dense forestation, probably caused a lower population density (Schibler and Steppan 1999). In younger periods, the findings are less frequent and the population in southeastern France probably died out already in late neolithic times while 20 records from the Metallic Ages and the Roman period in Switzerland, southernmost Germany and Austria show that other populations in the alpine region persisted for another 2000 years. However, the formerly homogeneous alpine population seems to have split up into disjunct units during this time. The youngest known records in this area date from medieval times (northern Austria: Raabs an der Thaya and Thunau, 10th century; Pucher and Schmitzberger 1999; Kanelutti 1990; Switzerland: Frohburg/Trimbach, 1000–1500; Markert 1989).

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Southern Germany (south of the river Main) and the Czech Republic From southern Germany, 23 records of moose have been published so far. From the western Czech Republic, 10 records are known. The number of sites and the generally small number of identified bones suggest a widespread distribution of A. alces in this area during the early and middle Holocene but probably small population densities. In mesolithic and neolithic times and during the Iron Age, the records concentrate south of the Danube, whereas in the following Roman period, when most parts of this area belonged to the Roman Empire, all moose records come from sites in or near the highland region. At this time, A. alces had possibly already been pushed back into the Rhine Valley (site Sponeck-Jechtingen; von den Driesch 1986), the Black Forest, the Swabian mountains (sites Rottweil, Bondorf, RainauBuch; Gulde 1985; Kokabi 1982; Kokabi et al. 1994) and the Frankish mountains (sites Ellingen, Eggolsheim; Breu 1986; von den Driesch and Liesau 1992). There are only a few early medieval moose records from southern Germany, and the species apparently became extinct there around 600 AD; the youngest dated remains were found at a burial-place in Schretzheim. There are additional written sources claiming the existence of moose in early medieval southern Germany but their validity is debatable (Prell 1941). In the western part of the Czech Republic, only some populations survived longer as archaeozoological records and (doubtful) written sources show (Prell 1941); here, A. alces died out in the Middle Ages also. Western Germany up to the river Elbe Since its re-immigration after the ice age A. alces had been part of the fauna in this area. However, the small number of only 14 records between the late glacial and its extinction in the Middle Ages could be an indication of low population density. Four of these records, located near Koblenz and Halle (sites Miesenheim Andernach-Martinsberg, Go¨nnersdorf, Allendorf-Fuchskirche; Feusel and

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Musil 1977; Street 1995a, b; Street and Baales 1999), document the presence of the moose in late glacial and early Holocene times. Six more records come from mesolithic and neolithic settlements. A well-preserved cranium with big antlers was found in the WeserDattel-Canal in North Rhine-Westphalia; it has been dated to 5270750 BP and was part of a natural taphocoenosis (Ueckermann et al. 1970). The economic importance of the moose was low in western Germany during the mesolithic and neolithic as shown by the small number of moose remains at all sites. In the neolithic settlement Hu¨de, however, 1.5% of the determined bones belong to this cervid (Hu¨bner et al., 1988). During the time between the Bronze Age and the Roman period, A. alces possibly persisted in parts of western Germany, but the moose from the important Roman castellum Colonia Ulpia Traiana (Peters 1994; Waldmann 1967), today’s Xanten on the Lower Rhine, possibly belonged to the above-mentioned population in the Netherlands. In the main part of western Germany, A. alces became extinct before the birth of Christ. During the Metallic Ages, the Roman period and towards the Middle Ages, moose occurred in the triangle of the Harz mountains in the north, the Thuringian Forest in the southwest and the Ore mountains in the southeast as evidenced by six records from different sites (Einhornho¨hle, Eisleben, Mu¨hlberg, Naumburg, Eschwege, Groitsch; Go¨tze et al. and Sippel in Willms 1987; Mu¨ller 1977, 1980b, 1996; Teichert 1990). The latest moose records from this area are the findings from Groitsch dating from high medieval times. A second population survived for a long time in the easternmost part of this area, near the river Elbe. It seems that this stock was (at least at times) quite large and also of interest to hunters since moose are regularly found in medieval castles near the river (sites Weinberg-Hitzacker, Magdeburg-Cracau, Klein Bu¨ddenstedt, Zehren, MeiXen, Fichtenberg; Boessneck and Stork 1973; Mu¨ller 1980a, 1982; Prilloff 1993; Teichert 1979; Walcher 1978). This population, which may not have been separated from the one east of the Elbe, probably became extinct between 1000 and 1200 AD.

Eastern Germany (Berlin and Brandenburg, region east of the Elbe) The moose colonized eastern Germany in the Allerød (site Berlin-Tiergarten; Benecke and Heinrich 2003), but from the long time between this oldest record and the Metallic Ages only one more record is known. These 16 bones from the mesolithic settlement near Friesack (Brandenburg; Gramsch 2000) document that A. alces had persisted in this area. Its status in younger periods is less enigmatic (see also Mu¨ller 1966): 15 records from the end of the Bronze Age to the Middle Ages suggest a wide distribution and a more or less high population density in this comparatively small area and also, especially in Roman times and in the Middle Ages, a certain economic interest in this species. During the high Middle Ages, the moose disappeared: the youngest records from Berlin (Pohle 1961) date from approximately 1200 AD. According to Heptner et al. (1966), the moose survived considerably longer in Saxony (until 1746), but these specimens may well have been migrating individuals from the east.

Northern Germany and Denmark This area comprises the Danish islands, the Cimbric peninsula and the German state Mecklenburg-Western Pomerania. A. alces first occurred here during the Allerød as findings from the locus classicus in northern Jutland show (Aaris-Sørensen 1988). At the time, it was the dominant ungulate species in this region (Benecke and Heinrich 2003). Moose may have found suitable conditions there in the colder Younger Dryas, too, because records from Zealand date from this period (Aaris-Sørensen 1992). From the early Holocene to the Atlantic, moose were quite common, at least in Mecklenburg-Western Pomerania, the eastern part of the Cimbric peninsula and on what is nowadays the Danish islands. There are no records from the western part of the Cimbric peninsula, but this absence may be an artefact because soil conditions there are often unfavourable for the conservation of organic matter. As to the Baltic coast, it is important to

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bear in mind that at that time, the distribution of land and sea was very different from today: the whole western Baltic area was terrestric from 8800–7000 BC (Jensen et al. 1999; Lemke et al. 2002). This land bridge was an important faunal gate between southern Scandinavia and Central Europe and decisive for the recent distribution of many species (Aaris-Sørensen 1992; Lepiksaar 1986). From mesolithic and neolithic sites, altogether 59 moose records are known. The bones from Skottemarke on Lolland and Favbro on Zealand, dating from the 8th millennium BC, are of particular archaeological importance: far away from their settlements, mesolithic hunters deposited the remains of altogether eight moose skeletons. These findings are not easy to interpret, but there may have been a ritual context (Møhl 1978). The number of 59 moose remains from the Boreal and Atlantic periods in Denmark and northern Germany suggests a widespread distribution of A. alces, but in most cases, the number of individual specimens found was small, indicating low population densities. Other species like red deer (Cervus elaphus), roe deer (Capreolus capreolus) and wild boar (Sus scrofa) were obviously economically more important to the human population. Starting around 6700 BC, a significant sealevel rise occurred in the Baltic area, the socalled Litorina transgression (Lemke 2005). As a result, the formerly continental eastern Denmark was transformed into an archipelago. For terrestrial animal species, this caused a splitting of populations and led to a decrease in faunal diversity during the following centuries (Aaris-Sørensen 1980, 1985). Particularly island populations of big species with low birth rates became extinct, as was the case with the big bovid Bos primigenius (urus, Schulz and Kaiser pers. comm.). The same obviously also holds for the moose: in spite of their being capable of long-distance swimming there are only few records from the newly formed islands after the transgression. On the Cimbric peninsula, A. alces persisted as shown by findings from 16 late mesolithic sites. In the wake of the neolithization, with its changes in human

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land use after 4000 BC, the records become scarce: only one finding from the northern Danish part of the peninsula is younger than 4000 BC (site Kainsbakke, about 2200 BC, Richter 1991), and from the southern German part, only three locations of partly questionable neolithic age are known (sites Bistoft, Wolkenwehe, Heidmoor; Ewersen 2001; Johansson 1979; see also Schmo¨lcke 2001). Hunting became less important for humans after the introduction of agriculture and husbandry and therefore, the number of records and bones of hunted animal species decreased but nonetheless, there is no unequivocal evidence of moose occurring here after the neolithic, i.e., after the Atlantic or early Subboreal. In Mecklenburg-Western Pomerania, the moose survived much longer. There are records from neolithic sites (Parchim-Lo¨ddigsee, Stinthorst and Basedow; Benecke 2002; Gehl 1974, 1976) and in the central part of the state also from the Bronze and Iron Ages, the Roman period and the Middle Ages which document the presence of A. alces. The moose was last mentioned in Mecklenburg-Western Pomerania in the 13th century AD (sites Hanfwerder, Zirzow; Prilloff 1994; for details see Benecke 2000, 2001). There are some medieval records from Denmark, Schleswig-Holstein and north-western Mecklenburg-Western Pomerania, too (sites Haithabu, Schleswig, Alt-Lu¨beck, GroX Stro¨mkendorf, Næsholm; Heinrich 1991; Møhl 1961; Reichstein 1991; Schro¨der 1984; W. Mazurek pers. comm.), but these findings mainly come from important trading centres. Therefore, it is probable that these bones and antlers were commercial products. Given the lack of respective findings in the previous periods, it is also possible that they belonged to single migrating individuals. In any case, these records cannot be viewed as certain evidence of an autochthonous population on the Cimbric peninsula at that time. Poland It is almost certain that A. alces was been part of the Polish fauna from the end of the ice age (Gumin´ski 2003) until high medieval times. The scarcity of moose records from the

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late glacial to the early Holocene is probably artificial given the distribution history in the surrounding parts of Europe. Also, the number of medieval sites with animal remains in Poland is much higher than for any other time period (see Wyrost 1993). While stone age records of the moose from the middle Holocene are concentrated in the south, the situation during the Middle Ages was different: the majority of the 83 records from Bronze Age to the high Middle Ages comes from Great Poland, Kujawy and from the northwestern (Pomeranian) coast line. Certainly, moose lived in the main part of Poland until early medieval times (see Wyrost 1993; Makowiecki 2003), but then a rapid decline led to its extinction at least in the southern mountains from where no younger findings are known. The claim that moose survived in Galicia until 1760 and in Silesia until 1776 (Heptner et al. 1966) seems, against this background, untenable. According to Willms (1987), this claim is a generalization of records of single migrating individuals. However, six records from Great Poland and Kujawy dating from late medieval and early modern times indicate the presence of A. alces for another 500 years in this area. Some even younger records from northern Poland suggest that the moose

survived in the Polish Baltic area until 1830 (Heptner et al. 1966).

Possible reasons for the extinction of the moose in western and Central Europe Given the spatial and temporal distribution of extinction events (see Fig. 2), the disappearance of the moose in Central Europe seems to have come about in two different phases, one during the Middle Ages, the other one in middle Holocene times. While in medieval times, the final extinction of already small and isolated populations may well have been caused by hunting, the gradual decline during the middle Holocene was certainly a more complex phenomenon. As mentioned above, according to AarisSørensen (1980, 1985, 1992), the extinction pattern in the north is closely related to two prominent events: the transformation of the region into a group of peninsulas and islands in the wake of the Litorina transgression and the neolithic revolution. The sea-level rise caused a fragmentation and isolation of mammalian populations which became much more vulnerable to demographic random effects and extinction. This was obviously

Polish Baltic area Great Poland Southern Poland Eastern Germany Mecklenburg-Western Pomerania Thuringean Uplands Eastern Alps Southern German mountains Switzerland The Netherlands Southern German lowland Western Alps Cimbric peninsula Western Germany Danish Islands Central France Pyrenees 10000

8000

6000

4000

2000

BC

0

AD

2000

Fig. 2. Spatial and temporal pattern of moose occurrence in different Central European regions during the Holocene.

ARTICLE IN PRESS Holocene distribution and extinction of the moose in Central Europe

only relevant in the directly affected areas and thus cannot serve as a general explanation. However, the moose populations were also threatened by habitat fragmentation, husbandry and a growing human population. To what extent and in what way these human influences led to the species’ disappearance from most of Central Europe is not at all clear. Humans might have exterminated the moose either directly through hunting or indirectly through habitat fragmentation or destruction or introduction of parasites and pathogens with their livestock. Alternatively, climatic changes may have rendered the environment unfavourable for moose (see Post and Stenseth 1999; Post and Forchhammer 2001). This may particularly be true for the early postglacial of central France where the rapid increase in temperature and aridity caused a change in vegetation and landscape. For an area in northeastern Poland, the data from Gumin´ski (2003) suggest a dependence of moose as a hunting prey of humans on the occurrence of pine forests. Similarly, the decline of pine forests during the Boreal period is correlated with a decrease in the frequency of moose remains. It is improbable, however, that direct climatic influences such as changes in temperature and humidity, were strong enough to account for the species’ extinction. As to hunting pressure as a possible explanation (‘‘overkill hypothesis’’), Stuart (1982), while favouring climatic and vegetational changes as the primary cause, states that ‘‘the demise of [the moose] in the British Isles was at least partly due to hunting by mesolithic man’’ (p. 181). Ru¨lcker and Sta˚lfelt (1986) emphasize the almost complete lack of moose remains in archaeozoological findings from the palaeolithic whereas in the neolithic, the moose seems to have been an important prey species. They conclude that moose populations must have been much larger in the neolithic than in the palaeolithic. However, this only holds for Scandinavia as in Central Europe (where an even contrary development may be assumed, cf. Benecke 2001) the moose was a rather frequent although never numerous part of human food remains. Still, after the strong decline (Britain, central France, western Germany)

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and disjunction (Denmark, Alps, southern Germany) of moose populations during the early or middle Holocene in most areas in the western part of Central Europe, hunting could have accelerated the process of extinction. Parasites and diseases may also be of relevance for the question of moose extinction. In North America, the so-called ‘‘moose sickness’’, caused by nematodes, has gained notoriety. Similar diseases may have occurred in Central Europe in former times. Also, in Central Europe moose are among the most sensitive species in zoological gardens (P. Dru¨wa, pers. comm.). They are especially susceptible to livestock parasites and pathogens (Geist 1998; D. Christensson, pers. comm.). According to Geist (1998), American moose brought few parasites with them which they had shed during evolving in the cold, dry and long winters in eastern Siberia but, just like caribou and elk (wapiti), can be severely limited by the presence of whitetailed deer parasites. Geist (1998) also thinks that the loss of the ability to resist the parasites and pathogens of primitive relatives may be the universal barrier to northerners dispersing south. Heptner et al. (1966) mention 38 parasitic worms found in moose (trematodes, cestodes and nematodes) of which some may be fatal, especially Trichocephalus (in zoos) and Parafasciolopsis (in the wild). Elaphostrongylus alces is often fatal in 1–3-year old moose while older animals develop resistance (D. Christensson, pers. comm.). Experiences from captive moose may, however, not be representative of conditions in the wild since the animals seem to be especially susceptible to taking parasites with food from the ground whereas in the wild they have less contact with parasites during browsing (D. Christensson and B. O. Ro¨ken, pers. comm.). Another possible reason for the disappearance of the moose in Central Europe is loss or fragmentation of habitat (e.g., Prell 1941). The optimal moose habitat is commonly regarded as deciduous and mixed forests with swamps and lakes (e.g., Nygre´n 1986; Ru¨lcker and Sta˚lfelt 1986). Moose sometimes colonize new areas after crossing long stretches of unfavourable environment: after

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its extinction in the Caucasus some 200 years ago, moose recolonised the area in the middle of the 20th century. To do this, they had to cross 300 km of open steppe (Heptner and Nasimowitsch 1967). Deforestation, sometimes viewed as a cause of moose decline, is in itself not necessarily a limiting factor. On the contrary, in Scandinavia and Russia, deforestation resulted in the creation of rich food supplies and thus new habitat (Heptner and Nasimowitsch 1967; Ru¨lcker and Sta˚lfelt 1986) since moose are concentrate feeders seeking out highly nutritious foliage and browse (Geist 1998) of which there is plenty after deforestation measures. Ru¨lcker and Sta˚lfelt (1986) hold that fire cultivation during the neolithic was favourable to the moose until pine forests became predominant (but see Gumin´ski (2003) for a different view). Proximity to humans is also readily tolerated by moose which show no natural fear of humans (Heptner and Nasimowitsch 1967). An interesting point in the question of moose extinction is competition with other species. Heptner et al. (1966) report that in the Bialowieza primeval forest at the PolishBelarussian border, the decrease of the moose population in the 20th century was possibly caused by competition with growing populations of red deer and other ungulate species. In Scandinavia, on the other hand, the resource use overlap between moose, red deer and roe deer is generally low (Mysterud 2000). The factors mentioned in this chapter may have influenced moose density in Central Europe to different extents at different times but so far, the data obtainable is certainly not sufficient to fully explain the disappearance of the species from such a large part of its former distribution area. Therefore, attempts such as the one in the present review necessarily remain largely hypothetical.

Range expansion to the west in the near future? At present, the only countries with stable moose populations in the area covered by our analysis are Poland and the Czech Republic.

In Poland, the moose population was drastically reduced during World War I and then, having recovered to more than 1000 head, again during World War II, leaving no more than 10 animals (Bobek and Morow 1987). As a reaction to the danger of its complete extinction in the country, reintroduction projects were started, the most commonly known being the translocation of specimens from the east of the country to Kampinos near Warsaw in 1951 (Niethammer 1963). Subsequently, the moose expanded its range, and today, numbers are growing exponentially (Jezierski and Lewandowski 1987). In the Czech Republic (see Homolka 1998 for details), the first moose after the species’ final eradication in 1570 was seen in 1957, and since then, moose have repeatedly immigrated into the country as a consequence of the growing population in Poland. Since 1973, moose have been known to reproduce regularly and two local populations in southern Bohemia, some 200 km south of the Polish populations, have established themselves. Within the last decades, migrating individuals have further been recorded from eastern and western Germany, Austria, Slovakia, Hungary, Romania and Denmark (reviewed by Briedermann 1968, 1971, 1989; Homolka 1998; Nygre´n 1986 and Schro¨tter 1998; for a recent case in northern Germany, cf. Labes and Ko¨hler 2001). According to Briedermann (1989), the proportion of females among the migrating moose has increased since 1967 while out of the 12 specimens that crossed the Polish borders to the west or south between 1957 and 1966, 11 were males. This may cautiously be interpreted as a sign of a more general tendency of the moose to expand its distribution range. There have also been a number of (re-) introduction projects. Apart from the successful reintroduction in Kampinos near Warsaw, Poland, Niethammer (1963) lists several reintroduction attempts in western and central Germany since the 16th century and even one in New Zealand that eventually failed. Since moose easily get used to living close to human settlements and even cities if hunting pressure is not too high, it is tempting to

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imagine this fascinating species as a regular part of the fauna of Central Europe again. In fact, the lack of predators, especially wolves, may actually favour its re-immigration. Human population density is certainly a limiting factor but there are still very good habitats with forests, lakes and swamps in Central Europe, e.g., in eastern Germany, where moose could thrive. In contrast to the reimmigration or reintroduction of the large carnivores (brown bear, wolf and lynx), moose would not have to face public opposition but would probably be welcomed with enthusiasm as the coverage by the media and the public sympathy and interest in dispersing moose specimens in the past have shown.

339

Acknowledgements The authors wish to thank N. Benecke, Berlin, for access to the data base mentioned in the text, R. Lu¨cht, Kiel, for invaluable help in accumulating the moose data from it, B.H. Rickert, Kiel, for technical assistance in creating the map and P. Dru¨wa (zoological garden Neumu¨nster, Germany), B. O. Ro¨ken (Kolma˚rden Zoo, Sweden), H. Mu¨ller (Wildpark Schwarze Berge), and D. Christensson (National Veterinary Institute, Uppsala, Sweden) for parasitological information on captive and free-living moose.

Zusammenfassung Holoza¨ne Verbreitung und Aussterben des Elches (Alces alces, Cervidae) in Mitteleuropa

Elche (Alces alces L.) geho¨rten am Ende der letzten Eiszeit zu den ersten GroXsa¨ugern, die wieder nach Mitteleuropa einwanderten. Bereits im Allerød etablierten sie sich in weiten Teilen dieses Gebietes. Im Fru¨hholoza¨n reichte ihre Verbreitung, abgesehen von ihrem heutigen Areal in OstMitteleuropa, von den Pyrena¨en bis nach Da¨nemark und von O¨sterreich bis nach GroXbritannien. Im Pra¨boreal setzte von Su¨dwesten her eine Ru¨ckzugsbewegung ein, die zuna¨chst Frankreich und dann England erfaXte. Wa¨hrend des Atlantikums verschwanden Elche aus weiten Teilen Da¨nemarks, und auch im u¨brigen Mitteleuropa scheint ihre Populationsdichte in dieser Zeit abgenommen zu haben. Um Christi Geburt existierten im Westen Mitteleuropas nur noch Reliktpopulationen, die im fru¨hen Mittelalter ebenfalls erloschen. Im thu¨ringischen Raum und nordo¨stlich der Elbe wie in Zentralpolen u¨berdauerten einige Besta¨nde bis ins hohe und spa¨te Mittelalter. Die Gru¨nde fu¨r das allma¨hliche Verschwinden des Elches aus Mitteleuropa wa¨hrend des Holoza¨ns sind vielschichtig. Vegetationsund Klimavera¨nderungen, die Entwicklung des Meeresspiegels, die zunehmende Zersiedelung der Landschaft durch den Menschen sowie dessen Jagdgewohnheiten waren zu verschiedenen Zeiten entscheidende Faktoren. In der ju¨ngsten Zeit sind jedoch wieder Ausbreitungstendenzen von Osten her festzustellen. Die Arealentwicklung des Elches in Ost- und Ostmitteleuropa seit dem Ende des Zweiten Weltkrieges wie auch Erfahrungen mit den autochthonen Besta¨nden in Skandinavien zeigen, daX Elche menschliches Kulturland nicht meiden und deshalb ku¨nftig eine Ausweitung des Verbreitungsgebietes nach Westen mo¨glich erscheint. r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved.

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Authors’ address: Ulrich Schmo¨lcke, Frank E. Zachos, Zoologisches Institut—Haustierkunde, Christian-Albrechts-Universita¨t zu Kiel, Olshausenstr. 40, D-24118 Kiel, Germany (e-mail: [email protected].)