Journal of Arid Environments 73 (2009) 754–761

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Journal of Arid Environments

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Spiny in the archaeological record of Israel

M. Ronel 1, S. Lev-Yadun*

Department of Science Education – Biology, Faculty of Science and Science Education, University of Haifa – Oranim, Tivon 36006, Israel article info abstract

Article history: The flora of Israel, like in other arid regions, has many spiny plants. We describe their existence in the Received 25 April 2008 archaeobotanical record of the last 780,000 years. Of 246 spiny/thorny species in the current flora, 19 are Received in revised form trees, 37 are shrubs, 50 are half-shrubs, 83 are perennial herbs and 57 are annuals. Forty-six (18.7%) 13 December 2008 spiny/thorny species were identified in the archaeobotanical record: 15 tree species (78.9%), ten shrub Accepted 13 February 2009 species (27.0%), six half-shrub species (12.0%), three perennial herb species (3.6%) and 12 annuals (21.1%). Available online 1 May 2009 Because humans needed , trees were taken into their habitats. Since trees are better preserved than Dedicated to Professor Aharon Horowitz on herbaceous plants and shrubs their remains are more common. Only a small proportion of the spiny/ the occasion of his retirement. thorny half-shrubs and perennial herbs was found, probably because they are not a good source of firewood, construction materials, or human food. Spiny weeds, and segetal and ruderal plants entered Keywords: the archaeobotanical remains either by chance, as seeds contaminating grain and fodder, growing at the Archaeobotany sites, or through their deliberate use by ancient peoples as supplementary foods and medicine. The Arid zone identified spiny/thorny species in the archaeobotanical record of Israel indicates that the flora was always Desert spiny even before the significant human impact in the last several millennia. Flora Ó 2009 Elsevier Ltd. All rights reserved. Herbivory Human activity Israel Spines Thorns

1. Introduction herbivores’ mouths, digestive systems (Janzen and Martin, 1982; Janzen, 1986), and other body parts. Recently, following the cloning Three terms in English are used to describe sharp defensive of a large array of pathogenic bacteria from spines and thorns, it has organs: spines when they are made of ; thorns, when been proposed that plants use spines to infect large herbivores with they are made of branches; and prickles, when they are made of such bacteria in a form of biological warfare (Halpern et al., cortical tissues (e.g., in roses) (Raven et al., 1999). Here, for conve- 2007a,b; Lev-Yadun and Halpern, 2008). Spines have additional nience, we define spiny plants as those producing any type of sharp functions, such as reducing sun irradiation (Gibson and Nobel, appendages. Because the aboveground parts of plants are 1986; Mauseth, 2006) and taking part in seed dispersal (Zohary, commonly utilized as food by various types of herbivores (Crawley, 1962), vegetative dispersal (Gibson and Nobel, 1986; Allen et al., 1983), there was a permanent evolutionary arms race between 1991; Bobich and Nobel, 2001) and camouflage (Benson, 1982; plants and their herbivores. On an evolutionary time scale, plants Mauseth, 2006). Spiny plants are more common in arid regions acquire better defenses and herbivores partly or fully overcome than in more humid environments (Grubb, 1992). Not surprisingly, them (Cornell and Hawkins, 2003). Among the many types of anti- the spiniest plant family – the Cactaceae – is mostly found in arid herbivore defenses that plants use, spines provide mechanical regions of North America (Benson, 1982). protection against herbivory (Janzen and Martin, 1982; Cooper and From the evolutionary and ecological viewpoints alike, spininess Owen-Smith, 1986; Janzen, 1986; Myers and Bazely, 1991; Grubb, increases in various ecosystems following long exposure to 1992; Gowda, 1996; Rebollo et al., 2002) because they can wound herbivory by large animals (Zohary, 1983; Janzen, 1986; Givnish et al., 1994; Sternberg et al., 2000). Even during an individual plant’s lifetime, spininess may increase following browsing as a genetically fixed induced defense (Milewski et al., 1991; Per- * Corresponding author. E-mail address: [email protected] (S. Lev-Yadun). evolotsky and Haimov, 1991; Young et al., 2003). The evolution of 1 In partial fulfillment of requirements for the degree of Ph.D. plants in an arid region like Israel, with constant pressure from

0140-1963/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved. doi:10.1016/j.jaridenv.2009.02.009 M. Ronel, S. Lev-Yadun / Journal of Arid Environments 73 (2009) 754–761 755 natural herbivory on the limited amount of vegetation, and the become a major component of the economy (Levy, 1983; Gophna millennia-old history of intensive grazing by goats, sheep, cattle, et al., 1986/1987). This must have considerably intensified the horses and donkeys as common agricultural practice, selected for impact of grazing by farm livestock above the natural level of plants that are better protected from grazing and induced an herbivory by large wild herbivores, a trend that has continued, with increase in their proportion in the plant populations (Zohary, 1983; fluctuations, ever since. Southern Samaria (The Land of Efraim, Noy-Meir et al., 1989; Perevolotsky, 1994; Perevolotsky and Selig- stretching from Shechem in the north to Beth Horon in the south, man, 1998). Zohary (1982) stated that there are more than 70 and from the foothills in the west to the desert fringe in the east, an species of spiny plants in the flora of Israel. Shmida (1981) identi- area of 1050 km2, now in the territory of the Palestinian Authority), fied 86 spiny and thorny species just in the Mediterranean element which has been studied thoroughly (Finkelstein et al., 1997), can alone of the flora of Israel, which composes about 30% of the thus be used as a test case for population increase for the whole number of species in the flora. In another study we identified 249 country and for the level of impact of human activity on the spiny, thorny and prickly plant species in the flora of Israel (Ronel vegetation. For instance, the proportion of the land used for agri- and Lev-Yadun, in preparation). culture, pasture and wood supply ranged between 0.1% and 15.4% Israel is located at the edge of the desert belt of the Old World. during the Chalcolithic and Late Bronze Age (6500–3200 years About half the territory (generally south of 31150 N, with B.P.); it was ca. 23% during Iron Age I (ca. 3000 years B.P.) and 81% a northern extension in the Dead Sea region) receives only during Iron Age II, then dropped to 22–40% during the Persian and 30–200 mm annual rainfall on average. The warmer areas in this Hellenistic periods (ca. 2400–3100 years B.P.), rose to 63% during belt are dominated by Saharo-Arabian vegetation with enclaves or the Roman period (ca. 2000 years B.P.) and to 93% during the Sudano-Decanian vegetation in oases. Cooler areas are dominated Byzantine period, and dropped to ca. 10% during the Umayyad/ by Irano-Turanian steppe vegetation (Zohary, 1962). Under such Abbasid period; it has ranged between 80% and 98% in the last 800 arid conditions agriculture is practiced only in niches where runoff years (Lev-Yadun, 1997). Palynological data from both the Sea of water accumulates and not all over the region (e.g., Evenari et al., Galilee from the last five millennia (Baruch, 1986) and the Birket 1982). Another section, north of the first (generally south of 31380 Ram crater lake in the northern Golan Heights (Neumann et al., N, with a northern extension in the Jordan Valley up to the Sea of 2007a) indicate a similar process of the degradation of the natural Galilee), receives 200–400 mm of annual rainfall (a region where vegetation, namely that the most drastic human impact started agriculture is practiced everywhere, but risks of crop failure are some 2500 years ago. high). This belt is covered by Irano-Turanian steppe vegetation in Plants were used by humans for many purposes and here we the more arid parts and by Mediterranean vegetation in the less mention only the major ones. Wood from trees (wild and domes- arid parts (Zohary, 1962). The rest is considered less arid, usually ticated) was regularly used for fire, construction and tool making with 400–800 mm annual rainfall during winter but with no (Meiggs,1982) and were collected from many of the local wild summer rains, is dominated by Mediterranean vegetation (Zohary, tree species. Certain shrubs (e.g., Retama raetam (Forssk.) Webb and 1962, 1973; Precipitation Map, 1987). These rainfall belts practically Sarcopoterium spinosum (L.) Spach) were commonly used for fire- correspond to the dominance of phytogeographical elements in the wood (Bailey and Danin, 1981; Zohary, 1982). Hundreds of plant flora. The driest belt corresponds to the Saharo-Arabian and Irano- species in the Land of Israel and adjacent countries were tradi- Turanian territory, the most humid belt corresponds to the territory tionally used for food, medicine, production of various tools, where Mediterranean maquis and forest may grow if not degraded fencing and other purposes (e.g., Moldenke and Moldenke, 1952; by human activity, and in between there is a transition zone with Thalen, 1979; Bailey and Danin, 1981; Palevitch et al., 1982; Zohary, rich, mixed vegetation types (Zohary, 1959, 1962). From early 1982; Dafni, 1984; Weinstein-Evron, 1998), and such uses intro- summer (May) to the early winter (November) the non-forested duced many of these plants into the archaeobotanical record. Plant and non-irrigated landscape, even of the less arid region of Israel, is material could also have entered the archaeological record via generally yellow without much edible green plant material. Under animal dung (Miller, 1984; Charles, 1998; Albert et al., 2008) and by the local climate, without modern methods of water pumping, growing wild at the sites. grazing was a very important practice, using natural vegetation Here, we discuss the finds of spiny and thorny plants in the resources spread at low density (Perevolotsky and Seligman, 1998). archaeobotanical record of Israel in relation to the current richness The impact of herding in the last several millennia (Sasson, 1998), of spiny plants in the flora of Israel and the Palestinian Authority. especially in the long and dry summer, followed millions of years of We consider the significance of the relative abundance of specific grazing by large mammals that thrived in the country during the groups. Pleistocene (e.g., Tchernov, 1979; Davis, 1987; Bar-Oz, 2004; Steiner, 2005). 2. Materials and methods Because of the millennia-old history of grazing, we outline the basic size of human populations and present some published As part of a broader study on the biology of plant protection by reconstructions of the level of their impact on the environment. The spines (e.g., Lev-Yadun, 2001, 2003a,b, 2006; Lev-Yadun and size of the human population of Israel down the ages has changed Ne’eman, 2004, 2006; Halpern et al., 2007a,b; Ronel et al., 2007, dramatically. A slow rise from fewer than several thousand people 2009; Lev-Yadun and Halpern, 2008) we studied the presence of in pre-agricultural periods (ending ca. 10,500 years ago), to about spiny and thorny species in the archaeological record of Israel by 100,000–200,000 in the more populated phases of the Bronze Age reviewing all the published data of the archaeobotanical remains (ca. 5500–3200 years B.P.) was followed by an increase to about from Israel and the Palestinian Authority. When a certain taxon was 1,000,000 people in the Byzantine period (ca. 1500 years B.P.) (e.g., found in more than one archaeological excavation we cite only the Broshi, 1980, 1993; Shiloh, 1980; Broshi and Gophna, 1984, 1986; earliest find. When only the genus was identified at sites of earlier Gophna and Portugali, 1988; Finkelstein, 1990; Broshi and Finkel- periods we cite finds from later periods as well if identification was stein, 1992; Finkelstein and Gophna, 1993; Zorn, 1994). The envi- on the species level. The thorny or spiny organs for each taxon are ronmental impact of these populations varied accordingly (e.g., specified in brackets when the taxon is described for the first time. Gophna et al., 1986/1987; Liphschitz et al., 1989; Lev-Yadun,1997, in We use very little palynological data because pollen identification press; Lev-Yadun and Weinstein-Evron, 2005). In the Chalcolithic is usually possible only to the family or genus level. However, of period, beginning ca. 6500 years ago, herding seems to have the spiny taxa, (some of which are spiny) Olea europaea L., 756 M. Ronel, S. Lev-Yadun / Journal of Arid Environments 73 (2009) 754–761

S. spinosum, Echinops sp., Acacia sp. and others are found in the acutus L. (pungent -sheaths). Cartwright (2005) lists wood pollen record of the Pleistocene and Holocene (Horowitz, 1971, remains in Pre-Pottery Neolithic Jericho including the half-shrub 1979; Weinstein, 1976; Weinstein-Evron, 1983; Baruch, 1986, 1990; Alhagi sp. (only the species Alhagi graecorum Boiss. grows in Israel) Baruch and Bottema, 1999; Neumann et al., 2007a,b). (thorny branches), the tree Balanites aegyptiaca (L.) Delile (thorny branches), the shrub Capparis sp., probably one of the several 3. Results species that grow in the desert and not C. spinosa, the half-shrub Noaea sp.; only the species Noaea mucronata (Forssk.) Ascherson et 3.1. Pre-agricultural period Schweinf. grows in Israel (thorny branches). In the ca. 8000-year- old Pottery Neolithic site Nahal Zehora II in the Menashe Hills, The earliest recorded assembly of botanical remains that include charred wood of the thorny shrub Genista fasselata Decaisne thorny plants identified to the species level is the 780,000-year-old (thorny branches) was found (Liphschitz, in press). assemblage of wood finds from the Acheulian site of Gesher Benot Ya’aqov (Goren-Inbar et al., 2002). Eight thorny/spiny trees and 3.3. Proto-historic period shrubs are represented in these finds: the tree Quercus ithaburensis Decaisne (spiny leaves), the tree Quercus calliprinos Webb (spiny At the ca. 6000-year-old Chalcolithic site Shiqmim in the leaves), Pyrus sp. (probably the only member of this genus in the northern Negev desert, Kislev (1987) identified among the and region, Pyrus syriaca, a tree with thorny branches), the tree seed remains the spiny annual plant C. pallescens. At another O. europaea L. (thorny branches in young or grazed trees), Amygdalus Chalcolithic site, Tel Tsaf in the Jordan Valley, Liphschitz (1988/ (probably Amygdalus korschinskii (Hand.-Mazzetti) Bornm., a small 1989) identified among the wood charcoals two spiny woody tree with thorny branches), Lycium sp. (we assume that it was Lycium plants; the tree Acacia albida Delile (stipular spines) and the shrub europaeum because of the locality) (all the shrub species in the genus Ziziphus lotus (L.) Lam. (stipular spines). Liphschitz (2005) identi- in our region have thorny branches), Rhus pentaphylla (Jacq.) Desf. or fied wood of the shrub Nitraria retusa (Forssk.) Ascherson (thorny Rhus tripartita (Ucria) Grande (both shrub species have thorny branches) in ca. 6000-year-old sediments from Mt. Sedom Caves. branches), Ziziphus sp. or Paliurus sp. (both local members of these Cartwright (2005) recorded the wood of Phillyrea sp. (spiny leaves) genera have stipular spines), Crataegus sp. (all local tree species of this (only the species Phillyrea latifolia L. grows in Israel) and Phoenix genus have thorny branches). Lev et al. (2005) who studied fruit and dactylifera L. (spiny leaflets) in Early Bronze Age Jericho. In ancient seed found in 60,000–48,000-year-old Mousterian vegetal food Lachish, the major city of the Judean foothills of the Middle and Late remains in Kebara Cave on Mt. Carmel, identified two spiny species, Bronze Ages (ca. 4000–3150 years B.P.) Liphschitz (2004) identified the half-shrub Astragalus echinus DC. (spiny petioles) and the annual among the wood charcoals of the trees P. dactylifera, Q. calliprinos Carthamus tenuis (Boiss. et Blanche) Bornm. (spiny leaves and invo- and Q. ithaburensis, the tree Acacia raddiana Savi (stipular spines lucral bracts). In the 23,000-year-old brush hut from Ohalo II on the and prickles on branches) and the tree Ziziphus spina-christi (L.) shores of the Sea of Galilee wood of the shrub sp. (we assume Desf. (stipular spines). Weeds from a cereal storage jar from Tel that it was Prosopis farcta because of the locality) (prickles on Batash, in the inner coastal plain (Shephelah) from the Late Bronze branches and spiny leaves) was found (Nadel and Werker, 1999). Age IIA, dated ca. 3300 years ago, included the spiny annuals Liphschitz and Nadel (1997) reported wood of the thorny shrub Centaurea iberica Trevis et Sprengel or Centaurea hyalolepis Boiss. Rhamnus lycioides L. (¼Rhamnus palaestinus Boiss.) (thorny branches) (spiny involucral bracts) and Centaurea verutum L. (spiny involucral fromthesamesiteandperiod.Baruch and Goring-Morris (1997) bracts) (Kislev et al., 2006). Werker (1988) identified wood of the studied wood remains in the Central Negev Highlands dated to the tree Acacia gerrardii Bentham var. negevensis Zohary in the Late end of the Pleistocene (18,000–10,000 years B.P.) and found two Bronze Age Egyptian mining temple at Timna in the southern Arava spiny taxa: the shrub Rhamnus disperma Boiss. (thorny twigs) and the Valley. small tree Paliurus spina-christi Miller. 3.4. Historic periods 3.2. Early agricultural period Among the fruits and seeds of the weed flora of Iron Age Ash- Kislev (1997), who studied fruit and seed remains from the ca. kelon on the southern Coastal Plain dating to 604 B.C. and including 10,000-year-old Pre-Pottery Neolithic A site of Netiv Hagdud in the 15 species, only a single seed of the spiny J. acutus was found (Weiss lower Jordan Valley, identified several thorny and spiny plants: the and Kislev, 2004). In the weed flora of a barley grain store from the annual Onobrychis sp. (prickles on pod) and the shrub Rubus san- Roman period at Yatir in the southern Judean mountains (135 A.D.) guineus Friv. (prickles on the branch, and spines on petiole and two of the 34 species were spiny annuals: E. spinosa and Onobrychis pedicle), Q. ithaburensis and the annual Centaurea ammocyanus crista-galli (L.) Lam./O. squarrosa Viv. (prickles on pod) (Kislev, Boiss./Centaurea pallescens Delile (spiny involucral bracts). Kislev 1986). Plant remains from a refuge cave of the Bar Kochba rebellion (1988) studied ca. 9000-year-old desiccated plant remains (fruits (135 A.D.) in the Judean desert (Simchoni and Kislev, 2004) and seeds) from the Pre-Pottery Neolithic B period layers of Nahal included six spiny taxa: the tree Z. spina-christi, the shrubs Capparis Hemar Cave in the Judean desert, and found five spiny plant sp. and R. tripartita (Ucria) Grande (thorny branches), the perennial species; the shrub Capparis spinosa L. (stipular spines), the half- herb Onopordum sp. (spiny wings of the stem, leaves and involucral shrub Anvillea garcinii (Burm. fil.) DC. (spiny leaves), the annuals bracts) and the annuals N. syriaca and Carthamus nitidus Boiss. Carduus australis L. fil. (spiny leaves and involucral bracts), Noto- (spiny leaves and involucral bracts). Baruch (1986) in his palyno- basis syriaca (L.) Cass. (spiny leaves and inflorescence) and C. tenuis. logical study of the Sea of Galilee found that the thorny shrub Hartmann (1997) identified in the fruit and seed remains from the S. spinosum (thorny branches) became more common between ca. 9000-year-old Pre-Pottery Neolithic C site Atlit-Yam at the coast 350 B.C. and 550 A.D. The tree B. aegyptiaca was found as wood in below Mount Carmel seven spiny species: the annual Emex spinosa the Roman siege rampart at Masada (Liphschitz et al., 1981) and its (L.) Campd. (spiny perianth in pistillate flowers and fruit), C. spi- (edible) fruit was found in caves in the northern Judean desert with nosa, the tree Crataegus aronia (L.) DC. (thorny branches), the shrub plant remains from the Roman period (Kislev and Hartmann, 1998; R. sanguineus, the annual Silybum marianum (L.) Gaertner (spiny Melamed, 2002). Acacia tortilis (Forssk.) Hayne, A. raddiana, leaves and involucral bracts), C. tenuis and the perennial herb Juncus A. gerrardii spp. negevensis, A. korschinskii (Hand.-Mazzetti) Bornm., M. Ronel, S. Lev-Yadun / Journal of Arid Environments 73 (2009) 754–761 757

Q. calliprinos, Q. ithaburensis and Z. spina-christi were found among maps is still premature, especially for herbaceous plants and the more than 1000 desiccated and charred wood remains from shrubs. Masada (Liphschitz and Lev-Yadun, 1989). Baruch (1986) also We address this question: Are the majority of the current spiny reported on pollen of the perennial herb Echinops sp. (spiny leaves flora indigenous to Israel, or did most of the 246 spiny species and flower heads), without giving a date. become established in the flora following damage to the primary vegetation with the advance of agriculture and herding during the last several millennia? 3.5. Total finds arranged according to plant types The high proportion of the current thorny and spiny trees (78.9%) and the ca. 30% of the thorny shrubs found in the archae- Of the 19 thorny tree species in the flora of Israel, 15 species ological data as presented here, combined with the many large (78.9%) were found in the archaeological record: A. albida, A. ger- mammalian herbivores that thrived in Israel during the Pleistocene rardii, A. raddiana, A. tortilis, A. korschinskii, B. aegyptiaca, C. aronia, (Tchernov, 1979; Davis, 1987; Bar-Oz, 2004; Steiner, 2005) and ate O. europaea, P. spina-christi, P. latifolia, P. dactylifera, P. syriaca, trees and shrubs as part of their diet, indicates that a considerable Q. calliprinos, Q. ithaburensis and Z. spina-christi. Of the 37 thorny thorny tree and shrub flora existed and characterized the landscape shrub species, 10–11 (27.0-29.7%) were found in the archae- in this land long before the massive impact of human activity obotanical record: C. spinosa, G. fasselata, Lycium sp., N. retusa, following the spread of agriculture and herding. The very short Prosopis sp., R. disperma, R. lycioides, R. tripartita (and possibly also time (in evolutionary terms) since the human impact became R. pentaphylla), R. sanguineus and Z. lotus. Of the 50 half-shrub critical, several millennia after the introduction of agriculture to the species, six (12.0%) were found in the archaeobotanical record: A. land about 10,500 years ago (Lev-Yadun et al., 2000; Zohary and graecorum, A. garcinii, A. echinus, N. mucronata, Onobrychis sp. and S. Hopf, 2000; Gopher et al., 2001), or more specifically since the spinosum. Of the 83 spiny perennial herb species in the flora of advance of pastoralism some 6500 years ago (Gophna, 1982; Levy, Israel only three species (3.6%) were found in the archaeological 1983), does not seem to enable the evolution of so many thorny and record: Echinops sp., J. acutus and Onopordum sp. Of the 57 annual spiny species from non-defended ones. Our general knowledge of spiny herb species in the flora of Israel 12 species (21.1%) were the importance of human activity in degradation of the natural found in the archaeological record: C. australis, C. nitidus, C. tenuis, vegetation of Israel and changes in its composition (Zohary, 1962, C. ammocyanus/pallescens, C. iberica or hyalolepis, C. pallescens, 1983; Naveh and Dan, 1973; Baruch, 1986, 1990; Gophna et al., C. verutum, E. spinosa, N. syriaca, O. crista-galli/Onobrychis squarrosa, 1986/1987; Liphschitz et al., 1989; Lev-Yadun, 1997; Baruch and S. marianum. Altogether 46 spiny/thorny/prickly species (18.7%) out Bottema, 1999; Neumann et al., 2007a,b) is not detailed enough of 246 such species in the current flora were found in the archae- regarding herbs and shrubs: most information, especially at the ological record. species level, is about trees. The probable changes in herb composition or distribution at the species level in recent millennia 4. Discussion and conclusions following the destruction of the forest (e.g., Zohary, 1962, 1983)are mostly cryptic in the palynological record because it is usually Some four decades of systematic identification of plant remains impossible to identify the pollen to the species level. Moreover, from hundreds of archaeological excavations in Israel allow a basic unlike pollen, the macrobotanical remains give us more qualitative understanding of the existence of spiny taxa in ancient periods and than quantitative data. Trees are better preserved and represented of some modes of human use of plants in those early times. Here we in the archaeobotanical record because (1) they were used in much analyze and discuss the wealth of archaeobotanical knowledge larger quantities as firewood and other purposes than other plants regarding the specific issue of spiny and thorny plants in the flora of were used for food, medicine, matting or tool making; (2) charred Israel from the floristic perspective and in relation to human wood fragments are commonly larger than charred tissues of activity. It is true that in certain cases the deliberate introduction of smaller plants, affording a greater chance of their retrieval in plants by human activity into the archaeological record through archaeobotanical excavations and better preserved tissues for their use by the inhabitants cannot at present be distinguished identification; (3) Israel has only a small number of tree species, from their chance entry via the contamination of grain by weeds, and some grow in specific regions. The wood anatomy of this small wind action, growth of plants in abandoned settlements before number of trees is much better known than the anatomy and a new phase of occupation, or the action of wild and domesticated morphology of other taxa (e.g., Fahn et al., 1986), permitting better animals. Spiny or non-spiny species may or may not be represented identification. All these factors result in a higher proportion of in the archaeobotanical record according to conditions of preser- identified trees than of other taxa. At any habitat in Israel the vation (charring, waterlogging, desiccation), retrieval methods number of tree species is much lower than the number of herba- during the excavation and quality of the botanical analysis. Pres- ceous species. Many of the herbaceous species may be of no use to ervation in sediments greatly depends on time passing so there is humans and have a tendency to decompose entirely under regular little chance of botanical remains being present in most non- terrestrial soil conditions. The great importance of wood for human waterlogged sites from pre-Neolithic periods. We therefore assume existence and the superior preservation of wood remains compared that the species identified are only a subset of the past flora, and we to that of herbaceous taxa has resulted in the repeated appearance will try to estimate the overall number of spiny species in the past. of wood remains in many archaeological excavations. The archae- Although tempting, calculating the diversity of the spiny plants ological plant remains offer us a much better record of thorny trees in ancient times from the archaeobotanical data with the current than of spiny herbaceous plants. knowledge may be misleading. While some wood remains from Of 19 thorny and spiny tree species currently growing in Israel, archaeological excavations in Israel started to be identified about 40 15 (78.9%) were found in archaeological contexts. By contrast, of 83 years ago, their regular identification in many (but not all) of the spiny and thorny perennial herb species currently growing in Israel excavations has been done for only about the last 25 years. For less only three (3.6%) were found in archaeological contexts, and of the woody remnants, which are preserved to a significantly lower 50 half-shrub species only six (12.0%) were found. The proportion extent, the situation is even less promising. Adding the lower of both the thorny shrub species (27.0% of the current thorny number of excavated earlier sites and the lower rate of preservation species found) and the spiny annual species (21.1% found) is in an in more ancient sites, we think that drawing regional or temporal intermediate position. Of the four thorny tree species not found in 758 M. Ronel, S. Lev-Yadun / Journal of Arid Environments 73 (2009) 754–761 the archaeobotanical record the two rare Crataegus species (Cra- Many spiny plant species were and are still used by local taegus monogyna Jacq., Crataegus azarolus L.) may actually be traditional societies. Some of them are used before the develop- present but not recognized when only their genus is identified ment of spines, or non-spiny parts are used. We list the spiny plants since the anatomical differences between the local Crataegus in use and mark* the species found in excavations in Table 1. species are slight (e.g., Goren-Inbar et al., 2002); these species are While many plants including spiny ones are grazed by the flocks very uncommon today (Feinbrun-Dothan and Danin, 1991) and in the wild and will therefore not be found in archaeological were probably so always. The small tree Amygdalus ramonensis macrofossils, ethno-botany demonstrates various traditional uses Danin is a very rare plant that grows only in the mountains of the of many plant species, including spiny ones. Bailey and Danin Central Negev Desert (Feinbrun-Dothan and Danin, 1991), where human population and thus archaeological remains, were always Table 1 sparse. The fourth tree species Prunus ursina Kotschy is also a rare Spiny species used by traditional societies in Israel (*found in archaeological plant in the mountains of the humid part of northern Israel (Fein- excavations). brun-Dothan and Danin, 1991) and this too probably resulted in its Species Bailey and Palevitch Dafni Said et al. not being found in the archaeobotanical record. Danin (1981) et al. (1982) (1984) (2002) As for species which are not trees, several hundreds were Acacia gerrardii* þ identified in the archaeobotanical record, although no review of the A. raddiana* þþ taxa found is available and many identifications are found in un- A. tortilis* þ published theses. From examining the archaeobotanical finds from Alhagi graecorum* þþ Israel published to date, we estimate the total number of wild Asparagus aphyllus þ Asparagus stipularis þ species found in archaeological remains at ca. 20–25% of the Atraphaxis spinosa þ current flora of ca. 2600 species (e.g., Feinbrun-Dothan and Danin, Balanites aegyptiaca* þþ 1991). According to this proportion (20–25%), the 46 spiny species Blepharis ciliaris þ found in the archaeobotanical record represent 184–230 spiny taxa Calicotome villosa þ in the past, a number quite close to the actual number that exists Capparis decidua þþþ Capparis sinaica þþþ today. By a different approach, if because of better preservation the C. spinosa þþþ tree finds give a better picture of past flora than herbaceous species, Centaurea aegyptiaca þ then the 15 thorny tree species found (78.9%) represent 196 thorny Centaurea behen þ species of the 246 in the current flora. Taking into account that Centaurea eryngioides þþ C. iberica* þ spiny taxa were avoided in many cases because of their spines, they C. pallescens* þþ are most probably under represented in the archaeological record. Cnicus benedictus þ Altogether, the actual archaeobotanical data and the extrapolation Crataegus aronia* þ presented suggest that the spiny nature of the flora of Israel was an Crataegus azorolus L. þ old, original character, and that from the floristic standpoint this is Cynara syriaca Boiss. þ Emex spinosa* þþ not a new phenomenon. The role of the spiny species in the land- Eryngium creticum Lam. þþ scape is a different issue from the floristic role and we discuss it Eryngium glomeratum Lam. þ below. Eryngium maritimum L. þ There are good reasons to propose that in addition to spiny Fagonia mollis Delile þþ Gundelia tournefortii L. þþ species spreading through the impact of continuous grazing, some Iphiona mucronata (Forssk.) þ spiny plants, e.g., C. tenuis, C. iberica, Centaurea procurrens, Ononis Ascherson et Schweinf. spinosa, P. farcta, Scolymus maculatus, S. marianum are weeds, Lactuca orientalis Boiss. þ ruderal and segetal plants (Zohary, 1962, 1983), and these Launaea spinosa (Forssk.) O. þ encroached on various areas they had not previously occupied or Kuntze Lycium europaeum* þ dominated because of the various types of human activity and Lycium shawii Roemer et þþ disturbances. The impact of human activity on the spread of thorny Schultes plants is demonstrated by the spread of S. spinosum about 1500 Nitraria retusa* þþþ * years ago in the vicinity of Mount Hermon (Neumann et al., 2007a), Noaea mucronata þ Notobasis syriaca* þ a time almost exactly coinciding with the rise in the pollen of this Olea europaea* þ thorny species near the Sea of Galilee (Baruch, 1986). Onobrychis crista-galli* þþ Altogether, the finds compiled here represent 18.7% of the spiny O. squarrosa* þ species in the current flora, indicating again that the spiny nature of Pallenis spinosa (L.) Cass. þ * the current flora is an old phenomenon. In the pre-agricultural age Phoenix dactylifera þ Prosopis farcta* þþ ending in Israel about 10,500 years ago (calibrated dates), 15 (6.0%) Pyrus syriaca* þ of the current spiny species were found. In the early stages of Quercus calliprinos* þþ agriculture, during the Neolithic period, 15 additional spiny species Quercus ithaburensis* þþ * were found, amounting to 12.0% of the current spiny flora. In the Rhus tripartita þþþ Rosa canina L. þ Chalcolithic period, when pastoralism intensified, three more spiny Rubus sanguineus* þ species (1.2%) were found and in the Early Bronze Age another two. Ruscus aculeatus L. þþ In the last 3200 years, since the beginning of the Iron Age, only 12 Salsola kali L. þ additional spiny species have been found, bringing the total of spiny Sarcopoterium spinosum* þ plants to 18.7% of the current spiny flora. Because the additional Scolymus hispanicus L. þ S. maculatus L. þ spiny species can be attributed both to better preservation of less Silybum marianum* þþ ancient sites and to the much higher number of excavated later Smilax aspera L. þ sites, we see no clear trend of changes in the appearance of spiny Zilla spinosa (L.) Prantl þ * species with time. Again, we interpret it as an indication for the Ziziphus lotus þ Z. spina-christi* þþþ very old spiny nature of the vegetation. M. Ronel, S. Lev-Yadun / Journal of Arid Environments 73 (2009) 754–761 759

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