This article was downloaded by: [Aston University] On: 29 January 2014, At: 04:00 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Israel Journal of Plant Sciences Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tips20 780,000-year-old wood from Gesher Benot Ya'aqov, Israel Ella Werker a a Department of Botany, The Hebrew University of Jerusalem, Jerusalem 91904, Israel Published online: 14 Mar 2013. To cite this article: Ella Werker (2006) 780,000-year-old wood from Gesher Benot Ya'aqov, Israel, Israel Journal of Plant Sciences, 54:4, 291-300 To link to this article: http://dx.doi.org/10.1560/IJPS_54_4_291 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions
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This article was downloaded by: [Aston University]On: 29 January 2014, At: 04:00Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: MortimerHouse, 37-41 Mortimer Street, London W1T 3JH, UK
Israel Journal of Plant SciencesPublication details, including instructions for authors and subscription information:http://www.tandfonline.com/loi/tips20
780,000-year-old wood from Gesher Benot Ya'aqov,IsraelElla Werker aa Department of Botany, The Hebrew University of Jerusalem, Jerusalem 91904, IsraelPublished online: 14 Mar 2013.
To cite this article: Ella Werker (2006) 780,000-year-old wood from Gesher Benot Ya'aqov, Israel, Israel Journal of PlantSciences, 54:4, 291-300
To link to this article: http://dx.doi.org/10.1560/IJPS_54_4_291
PLEASE SCROLL DOWN FOR ARTICLE
Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) containedin the publications on our platform. However, Taylor & Francis, our agents, and our licensors make norepresentations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose ofthe Content. Any opinions and views expressed in this publication are the opinions and views of the authors,and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be reliedupon and should be independently verified with primary sources of information. Taylor and Francis shallnot be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and otherliabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to orarising out of the use of the Content.
This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in anyform to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions
780,000-year-old wood of the Early/Middle Pleistocene, from excavations at Gesher Benot Ya’aqov in the Dead Sea Rift, was examined anatomically. Twenty-eight genera of trees, shrubs, and climbers have been botanically identified. The most common among them were Fraxinus, Quercus, Olea, Salix, Pistacia, and Ulmus. Most of the identified plants represent typical Mediterranean vegetation. Remnants of almost all the identified species are still found in the region today, in spite of the dramatic changes that took place in the Holocene through the intervention of man. One species, however, could not be identified with wood anatomical data of European, Middle Eastern, or North African species. It appears to be a species that is extinct in this region.
Between the years 1989–1997, intensive, multidisci-plinary research has been carried out by Prof. Naama Goren-Inbar (The Hebrew University of Jerusalem), at a Palaeolithic site at Gesher Benot Ya’aqov (GBY) in the Dead Sea Rift. The site, assigned to the Acheulian Industrial Complex, is bedded in a Pleistocene forma-tion that bears evidence of early hominid activity at the shores of the palaeo-Lake Hula, and is dated to 0.78 Mya (Goren-Inbar et al., 2002 and literature cited there-in). The excavations yielded, in addition to thousands of stone artifacts, a great wealth of fossil faunal remains and a large amount of both woody and herbaceous plants. Over 1,400 wood specimens were recovered but only those longer than 2 cm were kept, some of which were conjoinable fragments. The origin of the wood was suggested to be mainly from two sources: from the shores of the palaeo-Lake Hula, and from driftwood transported into the lake (Goren-Inbar et al., 2002).
Trees, shrubs, and woody climbers can be identified by the anatomy of their secondary xylem, i.e., the wood. The anatomical features of wood vary between taxa in the type of the elements constituting it, their shape, size, and amount, the pattern of their distribution, their cell wall structure, etc. These characteristics may be specific to different levels of taxa—species, genus, and some-times a subfamily or a family—but most often identifi-cation up to the species level can be reached.
Preliminary results of the identification of wood from the GBY excavations were published in Werker and Goren-Inbar (2001). An extensive report on the excavations themselves, the geology of the site, wood taphonomy, wood artifacts, and wood identification were reported in Goren-Inbar et al. (2002).
Here the results of identification of the wood remains are given and the implications of these results are dis-cussed.
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MATERIAL AND METHODS
The wood samples from the GBY excavations were wa-terlogged and soft, in various degrees of preservation. They were kept in tap water to prevent desiccation. The wet segments were sectioned by hand with a razor blade in cross, tangential longitudinal and radial longitudinal directions. The sections were kept in a water–glycerin solution. Whenever needed, they were cleared with a solution of 4% NaOH. They were examined under a light microscope. The anatomical features of the wood were compared with slides of contemporary wood and with wood catalogues (Greguss, 1955, 1959; Fahn et al., 1986; Schweingruber, 1990, and a collection of current wood samples in the laboratory).
RESULTS
A total of 916 wood segments were anatomically exam-ined. Of these, 638 were botanically identified, while 181 could not be identified for the following reasons:
1. Desiccation of the wood segment prior, during, or af-ter excavation caused shrinkage and often twisting of the wood, and that usually brought about distortion of the anatomical structure beyond recognition. The extent of such changes depends on external condi-tions and on the specific anatomical characteristics of the wood of each species. Pressure by geological processes could cause similar distortions.
2. Erosion caused by biological agents such as bacteria or fungi, and by chemical agents.
3. In some samples, roots of contemporary plants pen-etrated the soft, waterlogged wood, partially obliter-ating its structure, making identification impossible or uncertain.
4. Some specimens were of young branches containing only a few growth rings. The anatomical structure of the first growth rings is often different from the typi-cal structure of more mature wood and is therefore uncharacteristic of the taxon.
5. In years of drought, trees with typically ring-porous wood may produce only pores of their early wood, making the wood appear diffuse-porous. This may lead to an erroneous identification.
Results of the identification of the wood segments are presented in Table 1.
Botanical identification of the wood specimens was reached up to various levels of taxa. A total of 25 genera of trees, shrubs, and woody climbers were identified.
Following are listed those anatomical characteristics that could be distinguished in the sections and by which the different plant taxa have been identified. Added is the present-day habitat for each taxon, including only
the regions closest to the site (mainly according to Zo-hary, 1966, 1972, and Feinbrun-Dothan, 1978, unless otherwise stated).
Anacardiaceae
Pistacia: Vessels mostly in multiples and clusters; walls of the vessels, except for the widest, and of the vascular tracheids with prominent spiral thickenings. Rays uni- and multiseriate, heterocellular, with short margins of square, upright, sometimes slightly procumbent cells, and procumbent central cells. Gum ducts present in some of the multiseriate rays. In the three species de-scribed below solitary prismatic crystals are found in marginal ray cells.
Pistaciaatlantica: Growth rings distinct. Wood ring- to semi-ring-porous, usually with one row of wide pores at the beginning of growth rings. Rays 1–5(6) cells wide, predominantly 4–5.
P. atlantica var. latifolia is an Irano–Turanian ele-ment. It grows as solitary trees or in pure stands, or as an associate in maquis or forest of the Quercus ithabu-rensis association, in the Hula Plain, Upper and Lower Galilee, the Dan Valley, and the upper Jordan Valley.
Pistacia palaestina: Growth rings distinct. Wood ring-porous with gradual decrease in pore diameter. Rays 1–4 predominantly 3 cells wide.
Only one wood segment belongs, apparently, to this species.
Pistacia palaestina is an East Mediterranean ele-ment. It grows as a tree or shrub in maquis and garigue, mainly on hills and mountains, in the Hula Plain, Upper and Lower Galilee, the Dan Valley, and the Golan.
Pistaciavera: Growth rings distinct. Wood ring- to semi-ring-porous, usually with several rows of wide pores at the beginning of growth rings. Rays 1–5 cells wide, predominantly 3–4 (Grundwag and Werker, 1976).
P. vera is a steppe tree that does not grow wild in Israel. Its orign is the Irano–Turanian region, in semiarid terrain in Iran and adjacent countries.
Rhus: Three species of Rhus grow nowadays in Is-rael, Rhus pentaphylla, R. tripartita, and R. coriaria. R. pentaphylla and R. tripartita cannot be distinguished by their wood anatomy. The examined wood specimens were identified as one of these two species.
Rhus pentaphylla/tripartita: Growth rings faint and vessels diffuse, mainly in 2–7 radial multiples, some solitary, rounded in cross section, with thin-walled ty-loses, spiral thickenings absent. Fibers mostly septate. Rays 1–3(4)-seriate, heterocellular, usually with 1–2 rows of square to upright marginal cells and procumbent to upright central cells.
Neither of these species grows in the vicinity of the
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Table 1Identification and frequency of the wood and bark*
excavation today. Rhus tripartita is an Irano–Turanian element, a shrub, growing on rocks in deserts and among shrubs in the lower Jordan Valley. Rhus pentaphylla is a South Mediterranean element, a shrub or shrublet grow-ing in Coastal Galilee and the Acco Plain.
Rhus coriaria, a Mediterranean tree or shrub, is found today in neglected places near villages, rarely in maquis, in Upper Galilee, and the Golan Heights. However, the wood of the specimens identified as Rhus do not fit the wood structure of this species, which has distinct ring-porous growth rings, with prominent spiral thickenings except for the largest vessels, and the fibers are very infrequently septate.
Apocynaceae
Nerium: Growth rings faint. Vessels diffuse, solitary and in radial multiples, and of a small diameter. Fibers thin-walled. In the examined specimens it is difficult to distinguish the vessels from the fibers. Rays 1–2(3) cells wide, heterocellular, the 2- and 3-seriates with low portions of strongly procumbent cells and tall margins of square to upright cells.
Nerium oleander is a Mediterranean element, a shrub growing on banks of lakes and streams and on stony wadi beds in the Upper and Lower Hula Plain, the Jor-dan Valley, and the Golan.
Araliaceae
Hedera: Vessels diffuse, mainly in clusters, sometimes in tangential or radial multiples of 2–4, occasionally soli-tary, often forming together with vasicentric tracheids a pattern of tangential bands. Vessel walls occasionally with spiral thickenings. Rays 1–7(14)-seriate, up to ca. 1.5 mm high, nearly homocellular to heterocellular, com-posed of procumbent central cells and slightly procum-bent, square, and sometimes upright marginal cells.
Hedera is a Euro–Siberian and Mediterranean ele-ment, growing as a climbing shrub on rocks and in maquis in Upper Galilee.
Asclepiadaceae
Periploca: Vessels diffuse, mostly solitary, sometimes in pairs, sometimes in a radial or oblique pattern, and of a small diameter. Fibers medium thick-walled. Rays mostly uniseriate, sometimes 2–3(4)-seriate up to 20 cells high; heterocellular composed of square, upright, sometimes weakly procumbent cells.
Periploca aphylla is a Sudanian element extending into the East Saharo–Arabian region. It grows as a shrub in the rocky deserts in the Upper and Lower Jordan Val-ley and the Dead Sea area.
A single segment of this species was found.
Caprifoliaceae
Lonicera: Growth rings distinct. Wood ring-porous [it is diffuse-porous in Lonicera etrusca, which grows today in the region, but ring-porous in other West Mediter-ranean species (Greguss, 1959; Schweingruber, 1990)]. Vessels generally solitary, rarely in small groups, an-gular in cross section; walls with fine spiral to annular thickenings. Rays 1–3(4)-seriate, of various heights up to more than one mm; heterocellular.
Lonicera etrusca is a Mediterranean element grow-ing as a climbing shrub in rocky places in maquis and garigue in Upper and Lower Galilee and the Golan. Lonicera nummulariifolia is a shrub that grows on Mt. Hermon; its wood anatomy was not examined.
The anatomical features of the samples are more similar to the European Lonicera species, having ring-porous wood, than to the species that grows nowadays in the region. The five segments of Lonicera were found at different levels of the excavations and they all were of thick branches. This eliminates the possibility of being misled by early secondary thickening, which may have different anatomical characteristics than that of later secondary thickenings.
Cupressaceae
Juniperus: A gymnosperm, vascular elements consist-ing of tracheids only, with uniseriate bordered pits. Axial parenchyma diffuse or in short tangential bands. Rays mostly uniseriate, 1–8 cells high, pits 2–4 per cross-field. Resin ducts absent.
Juniperus drupacea is a shrub and J. excelsa a tree; both grow on Mt. Hermon. Juniperus oxicedrus is a Mediterranean shrub or tree that grows on calcareous soil also in Upper Galilee, but is rare today. The wood anatomy of these Juniperus species is very similar and distinction between them based on wood anatomy is unreliable.
Fagaceae
Quercus: Vessels almost exclusively solitary, in a radial, flame-like, or dendritic pattern. Vasicentric tracheids present. Parenchyma scanty paratracheal, and apotra-cheal diffuse, diffuse-in-aggregates and in short narrow tangential bands. Rays of two distinct sizes, uniseriates and very wide and high multiseriates, the latter partly compound. Quercus calliprinos has diffuse-porous wood, while Q. ithaburensis, Q. boissieri, and Q. libani have ring- sometimes semi-ring-porous wood.
Quercus calliprinos is an East Mediterranean ele-ment, a tree or shrub that grows in maquis and forest from sea level to 1,500 m in Upper and Lower Galilee and on Mt. Hermon. It is the most common tree in Up-
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per Galilee, where the annual precipitation is 500 mm and more.
Quercus ithaburensis is an East Mediterranean ele-ment, a tree growing up to 500–1,000 m asl, in the Dan Valley, Upper and Lower Galilee, and in the Hula Plain, in small and scattered forest remnants.
Moraceae
Ficus carica: Fibers very thick-walled, in wide bands alternating with parenchymatous bands. Inter-vessel pits with slit-like, very often coalescent apertures. Rays 1–4 cells wide, up to 45 cells (0.8 mm) high, heterocel-lular, composed of procumbent central cells and square and upright marginal cells. The specimens are severely shrunken, especially their parenchyma bands.
Ficus carica is a Mediterranean and Irano–Turanian tree. The uncultivated male variety F. carica var. capri-ficus is native to Upper Galilee (Zohary, 1966).
Myrtaceae
Myrtus: Vessels diffuse, predominantly solitary, occa-sionally in pairs, of a small diameter, walls with fine, not always distinguishable spiral thickenings; perforation plates simple. Fibers with distinctly bordered pits. Rays 1–2(3) cells wide, up to 20 cells high, heterocellular.
Myrtus communis is a Mediterranean element, a shrub or sometimes a small tree growing in maquis and riverine thickets in Upper Galilee, the Dan Valley, and the Upper Jordan Valley.
Oleaceae
Fraxinus syriaca: Wood ring-porous. Vessels solitary, in radial multiples of 2–3(4) or rarely in small clusters. Many vessels with tyloses. Parenchyma vasicentric in the early wood, aliform and confluent towards the end of growth rings, and marginal. Rays 1–4(5) cells wide, up to 20 cells high, homocellular, composed of procum-bent cells.
Fraxinus syriaca is an East Mediterranean and West Irano–Turanian element. It grows as a tree along rivers and streams in the Hula Valley, Upper Galilee, the Bet She’an Valley, and the Golan.
Jasminum: Wood ring-porous. Vessels of late wood solitary and of small diameter. Part of the vessels and fibers with fine spiral thickenings. Fibers with bordered pits. Rays mainly uniseriate, heterocellular with upright and square marginal cells and weakly procumbent cen-tral cells. Jasminum fruticans is a Mediterranean and West Irano–Turanian element, growing as a shrub in Quercetum calliprine maquis, at an elevation of about 400–1,000 m asl in Upper Galilee and the Golan.
Olea europaea: Growth rings distinct to faint. Ves-sels diffuse, solitary, in radial multiples of 2–4(6) occa-
sionally in clusters; angular to rounded in cross section; diameter up to 70 µm. Some vessels with gummy sub-stances. Rays 1–2(3)-seriate, heterocellular; some cells crystalliferous. Among those identified as olive wood, some had typical rays up to 12(20) cells high, and others much higher rays and a larger number of uniseriate ones (cf. Werker, 1998).
Olea europaea is an East Mediterranean element, growing as a tree or shrub. Var. sylvestris, the wild ol-ive, grows in maquis in Upper Galilee and the Golan, mainly in the Quercus calliprinos—Pistacia palaestina association.PinaceaeCedrus: A gymnosperm, vascular elements consisting only of tracheids, with mostly uniseriate bordered pits with fringed torus margins. Axial parenchyma confined to the growth ring boundary. Rays mostly uniseriate, rarely biseriate (1)3–20(35) cells high, composed of marginal smooth-walled ray tracheids in single rows and fairly thick-walled ray parenchyma cells with nodu-lar end walls. Each cross-field with 2–4 taxodioid pits in early wood. Vertical resin ducts close to each other are of traumatic type, with mostly thick-walled lignified epithelial cells. Vertical resin ducts are more common than horizontal ones within rays.
Cedrus libani is an East Mediterranean element. It grows as a tree in a few relict stands (Mikesell, 1969) on mountains of Lebanon and farther away in Cyprus and Turkey.RhamnaceaePaliurus: see Ziziphus.
Ziziphus/Paliurus: Vessels diffuse; solitary, in radial multiples of 2–3, occasionally in small clusters. Fibers medium-thick to very thick-walled. Rays 1(2) up to 20(24) cells high, weakly heterocellular composed of square, upright, and infrequent weakly procumbent cells.
Ziziphus lotus is a Mediterranean and Sudanian element, a shrub growing in stony slopes and alluvial plains in Upper Galilee, the Dan Valley, and the Golan. Ziziphus spina-christi is a Mediterranean, Saharo–Ara-bian, and Irano–Turanian element, a tree growing in wadi beds and alluvial soils in Upper and Lower Galilee, the Dan Valley, and the Golan. Paliurus spina-christi is a North and East Irano–Turanian element. It grows as a tree or shrub, mostly in alluvial soils, in Upper and Lower Galilee and the Dan Valley.
The two genera, as well as the two species of Ziziphus that grow in Israel, cannot be distinguished unambiguously by their wood anatomy. Both species of Ziziphus are common today in the Hula Valley and on the Korazim basalts. Only two specimens of these spe-cies were found in the excavations.
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RosaceaePrunoideae: Semi-ring-porous. Vessels with fine spiral thickenings. Vasicentric tracheids present. Rays wide and quite high. Fibers with bordered pits.
Amygdalus korschinskii: Wood ring-porous. Vessels mostly solitary, widely spaced in late wood, with fine spiral thickenings. Fibers with distinctly bordered pits in radial and tangential walls. Rays 1(2)-seriate and 3–8-seriate, the latter up to 44 cells high, rarely compound, heterocellular, composed mainly of procumbent cells and weakly procumbent to upright marginal cells.
Amygdalus korschinskii is a West Irano–Turanian element; it grows as a tree or high shrub in impoverished maquis and maquis-steppes in Upper Galilee and the Upper Jordan Valley. It appears in association withPis-tacia atlantica on slopes facing the Hula Valley (Rabi-novitch-Vin, 1986). It has been suggested that a similar kind of steppe-forest may have covered these slopes in the past (Rabinovitch-Vin, 1977).
Cerasus: Wood ring-porous. Vessels mostly solitary, occasionally in variously directed multiples of 2–3; with prominent spiral thickenings. Inter-vessel pits round, diffuse. Fibers very thick-walled, with distinctly bor-dered pits in radial and tangential walls, occasionally with spiral thickenings. Rays 1–3-seriate, up to 0.8 mm or more in height; heterocellular, composed of square, weakly procumbent, and upright cells.
Cerasus prostrata is a Mediterranean element. It grows as a shrub on rocky ground on Mt. Hermon.
Crataegus: Growth rings distinct. Vessels diffuse, densely arranged, mostly solitary, and mostly angular in cross section. Fibers thick-walled with distinctly bordered pits in the radial and tangential walls. Apotracheal paren-chyma diffuse, sometimes diffuse-in-aggregates. Rays 1–3(4) cells wide and up to 25(35) cells high, predominantly homocellular, consisting of procumbent cells. Crystals soli-tary, prismatic in enlarged chambered parenchyma cells.
The three Crataegus species that grow in Israel are very similar in their wood anatomy and cannot be unambiguously separated. Crataegus aronia is an East Mediterranean and West Irano–Turanian element. It grows as a tree or shrub in maquis and maquis-steppes in Upper and Lower Galilee. It accompanies the Quer-cus ithaburensis—Pistacia atlantica association. Cra-taegus monogyna is a Mediterranean and Euro–Siberian element. It grows as a shrub or small tree in Quercus maquis in shady valleys at about 1,000 m asl in Upper Galilee, and is very rare today. Crataegus azarolus is an East Mediterranean element growing as a tree in more humid maquis in Upper Galilee.
Pyrus syriaca: Growth rings distinct. Vessels dif-fuse, almost exclusively solitary, rarely in pairs or small clusters, angular in cross section, many vessels
with gummy contents. Fibers with distinctly bordered pits. Apotracheal parenchyma diffuse to diffuse-in-ag-gregates. Rays1–2(3)-seriate up to 43(55) cells high, predominantly homocellular, composed of procumbent cells, occasionally with square or weakly upright mar-ginal cells. Crystals very infrequent in chambered, not enlarged parenchyma cells.
Pyrus syriaca is an East Mediterranean and West Irano–Turanian element. It grows as a tree in maquis and forests in Upper Galilee and on Mt. Hermon.
Salicaceae
Populus euphratica/Salix spp.: Growth rings distinct. Ves-sels diffuse, solitary, in radial multiples of 2–3(4), occasion-ally in small clusters. Fibers thin- to medium-thick-walled. Rays uniseriate, very rarely biseriate; ray-vessel pits in cross fields sometimes giving a reticulate appearance.
The difference between Populus and Salix is that in Populus the rays are typically homocellular, sometimes with marginal cells somewhat less procumbent than the central cells or square, while in Salix many rays are heterocellular, composed of 1–3 rows of upright, some-times irregularly shaped marginal cells and procumbent, sometimes square, or even upright central cells.
Finding an anatomical feature in a section that is absent in one genus and frequent in another is positive proof for identification of the latter (Salix), but not find-ing it is not always a reliable criterion for identification of the former (Populus). Therefore, distinction between the two genera is sometimes uncertain.
Populus euphratica is an Irano–Turanian and Saha-ro–Arabian element, a tree growing on river banks and springs; in the Upper and Lower Jordan Valley it forms riverine forests, especially on the banks of the lower course of the Jordan River.
Salix species cannot be distinguished by their wood anatomy. Salix acmophylla is an East Mediterranean and Irano–Turanian element and is more abundant than S. alba, which is a Mediterranean, Euro–Siberian, and Irano–Turanian element. Both are trees that grow near water in the Hula Valley and Upper Galilee.
Solanaceae
Lycium: Vessels form a dendritic pattern; wider vessels are clustered with very narrow ones intergrading with vascular tracheids. Rays mostly uniseriate.
It is difficult to separate the four Lycium species that grow in Israel according to their wood anatomy. Lycium europaeum is a Mediterranean element, a shrub growing along fields and wadis in Upper and Lower Galilee, the Upper Jordan Valley, and the Golan. The other species grow in more arid areas further south.
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Ulmaceae
Ulmus: Wood ring-porous. Early wood vessels large, solitary or in pairs, with vasicentric parenchyma; late-wood vessels small, in many-celled clusters forming tangential or oblique bands together with vascular tra-cheids and paratracheal parenchyma. Walls of smaller vessels and fiber tracheids with fine spiral thickenings. Tyloses may be present. Parenchyma vasicentric around large vessels and around the bands of the small ones. Rays 1–6-seriate, up to 100 cells high, homocellular, with strongly procumbent cells, sometimes weakly het-erocellular with weakly procumbent or square marginal cells. Solitary prismatic crystals are present in longitu-dinal chains of parenchyma cells.
It is difficult to distinguish between species.Ulmus canescens is a North and East Mediterranean
element. It grows as a tree in shady places near water, in Lower Galilee.
Vitaceae
Vitis: Wood ring- to semi-ring-porous. Vessels mostly in radial multiples of 2–15 and in clusters together with vascular tracheids. Perforations simple but scalariform in some of the narrow vessels. Inter-vessel pits scalari-form, in larger vessels in several rows. Vascular tra-cheids and occasionally narrow vessels with irregular, very fine spiral thickenings. Fibers septate. Rays (3)7–13 cells wide and very high, heterocellular, with mostly procumbent cells and some square or upright cells.
The wild type of Vitis vinifera, V. sylvestris, is a climbing shrub growing in humid habitats such as ripar-ian forests. It is a Euro–Siberian, Mediterranean, and Irano–Turanian element. In Israel it occurs in gorges and in the vicinity of springs and streams along the Upper Jordan River and its tributaries as well as in the Kinneret Valley (Kislev and Melamed, pers. comm.).
Unidentified specimen (Plates 1, 2)
Vessels form a dendritic, sometimes radial or tangential pattern, together with paratracheal parenchyma and vas-cular tracheids; vessels mainly rounded in cross section, with maximal radial diameter of ca. 100 µm. Walls of some vascular tracheids and vessels with spiral thicken-ings. Apotracheal parenchyma in more or less tangential bands, up to 5 cells wide, and marginal parenchyma 1–2(4) cells wide. Fibers thick- to very thick-walled, al-ternating with parenchyma bands. Rays 1–3(4) -seriate up to ca. 20 cells high, heterocellular, composed of pro-cumbent and square central cells and square to upright marginal cells. Many vessels, as well as parenchyma and ray cells, with dark materials.
Most specimens of the “unknown tree” have quite well-preserved wood. Nevertheless, it could not be identified by comparison with available wood samples, consulting wood atlases of species growing in Europe, the Middle East, and North Africa, or descriptions of ar-chaeological material from Northwest Europe (Greguss, 1959; Schweingruber, 1990; Gale and Cutler, 2000), as well as the GUESS program of computerized database (Wheeler et al., 1986) (except for a genus of the New World, Bumelia of family Sapotaceae, a family that has no representatives among the wild flora of Israel).
Of the trees and shrubs growing nowadays in the region, this type of wood resembles that of Retama raetam (Leguminosae), but it differs from it in the fol-lowing features: Its vessels and parenchyma cells are non-storied; the apotracheal parenchyma bands of the unidentified species are 1–5 cells wide while those of Retamaare 1–2(4). Retama raetam var. raetam, a Sa-haro–Arabian element, grows as a shrub. Its nearest site of growth today is the Upper and Lower Jordan Valley.
Plate 1. Unidentified tree. Upper, Cross section showing vessels in a dendritic pattern and apotracheal parenchyma in tangential bands. Scale bar = 100 µm. Lower, Longitudinal section showing spiral thickenings in tracheary elements. Scale bar = 25 µm.
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The wood of the unknown tree also resembles that of Phillyrea (Oleaceae) in its dendritic vessel arrangement and initial parenchyma, but differs from it in vessel diameter and the absence in Phillyrea of parenchyma bands.
Phillyrea latifolia is a Mediterranean element grow-ing as a shrub or tree in maquis and forest in Upper and Lower Galilee.
It must be borne in mind, however, that in spite of a certain similarity in anatomical features with these two wild species that grow in the region, a difference in even one feature between specimens may sometimes lead to remote families.
DISCUSSION
Present-day vegetation appears very different from the primary vegetation that prevailed during the Lower/Middle Pleistocene transition 780,000 years ago in the Hula Valley. During the years 1951–1959 a large part of Lake Hula and the marshes surrounding it was arti-ficially drained. Today it comprises mostly cultivated and fallow land. The wild vegetation consists mainly of segetal and ruderal plants, with Prosopis farcta dwarf shrubs scattered among the segetal vegetation. Along the water margins grow trees of Salix and scattered trees of Fraxinus syriaca. On dry land there are scattered trees of Quercus ithaburensis and Pistacia atlantica. On the border of the plain, Ziziphus spina-christi and Z. lotus are quite common.
In the wood assemblage of the GBY excavations, 28 genera of trees, shrubs, and climbers have been identi-
fied. They consist of bank and dryland woody vegeta-tion, which is mainly Mediteranean. In spite of the dra-matic changes that took place in the region because of human activity, the majority of the identified species are still found in the region today, though in very different proportions. The most common among the species was found to be Fraxinus syriaca (245). Next, in descending order, are Quercus (69) including a definite identifica-tion of Quercus caliprinos (27) and Quercus ithaburen-sis (26), Olea (45), Salix (26), Pistacia atlantica (25), and Ulmus (14). Other species occur in much smaller numbers. Only two specimens of Ziziphus (or Paliurus, see Results) were identified and none of Prosopis. The two species of Ziziphus are considered late newcomers that penetrated following human activity (Zohary, 1959; Weisel, 1984), not earlier than the devastation of the climax flora of the valley, which was most likely, ac-cording to Zohary and Orshansky (1947), the Quercus ithaburensis–Pistacia atlantica association. The results presented here indicate that Ziziphus was part of the natural flora before the degradation of the vegetation during the Holocene.
Fraxinus syriaca is known in the Dead Sea Rift, from the Pliocene (Braun, 1992), Upper Pleistocene (Liphschitz and Nadel, 1997), and many other archaeo-logical sites (Liphschitz, 1986). This species apparently constituted in the past an important component of a bank forest.
Ulmus canescens grows today only in humid and temperate regions of Galilee and Samaria. It has been suggested by Zohary (1982) thatUlmus is probably a relic of a rainier season.
The presence of a relatively large number of Quercus ithaburensis and Pistacia atlantica in the GBY exca-vations and their scarcity today in the Hula and Dan valleys confirm the view of Zohary that these species represent a continuation of the Golan forest, which once extended down to the valley (Zohary and Orshansky, 1947; Zohary, 1959).
The vegetation of the region during Lower and Middle Pleistocene must have been, therefore, very rich on the lake and swamp shores and constituting a park or closed forest on the dryland. It could sustain large herbivorous wild animals like elephants, rhinos, and hippos, the remains of which were found in the site (Goren-Inbar et al., 1992, 1994).
There are a few exceptions to the identity found between the plant species that grew in the region in the past and today. For two genera, Lonicera and Rhus, dif-ferences in some anatomical characteristics were found between the species that grow today in the area and those identified in the excavation. The 780,000-year-old identified species of Lonicera is more similar to species growing in Europe than to L. etrusca, which grows in the region today. The Rhus species identified here do not grow in the vicinity of the excavation, but in other re-gions of the country. At this stage it cannot be concluded whether the identified species have been replaced by other species of the same genus that grow elsewhere in the country or abroad, or whether these species have undergone some evolutionary anatomical changes dur-ing the past 780,000 years, or whether there is a higher intraspecific variability in certain anatomical features of the wood of these species than observed up to now.
1. It should be mentioned that the number of segments of these two species found in the excavations is rather small.
2. Thirteen samples of one species, found at various lev-els of the excavations, could not be identified. This unidentified species appears to be an extinct species, at least in this region. It could not be identified using the anatomical data of wood from European, Middle Eastern, or North African species. A search for iden-tification should perhaps be continued among more southern African arboreal plants or in the Far East.
3. Cedrus and Cerasus grow nowadays in relic stands in the mountains of Lebanon. They have never been known to grow in Israel. The ecological setting for the growth of Cedrus, as noted by Lev-Yadun et al. (1996), is not found in Israel today. A somewhat colder climate might have prevailed in the region in Early/Middle Pleistocene times (for more details on driftwood see Goren-Inbar et al., 2002). Juniperus oxycedrus, which grows in the Galilee today but is rare, may also indicate
a colder climate in the past. These three species were also found in very small numbers.
4. A few wood specimens have been identified as Pistacia vera, a species that currently does not grow wild in Israel. Within the genus Pistacia interbreed-ing between species occurs, resulting in variability in the wood anatomy of the hybrids (Grundwag and Werker, 1976). The differences in wood character-istics between P. atlantica and P. vera are mainly quantitative. Therefore, one should be cautious in drawing conclusions on the basis of wood anatomy alone as to whether P. vera grew in the past in the GBY region.
5. Remarkable is the absence in the excavations of the wood ofStyrax officinalis, which grows today in the Upper and Lower Galilee, in spite of the fact that many fruits of this species were found (Melamed, 1997). From the results reported here no explanation for this phenomenon can be reached.
A better understanding of the environment and veg-etation in the Hula region during the Lower/Middle Pleistocene times might be attained when the results recorded here on arboreal flora will be integrated with results on fruits and seeds and of pollen grains from the GBY excavations.
ACKNOWLEDGMENT
Many thanks to Prof. Naama Goren-Inbar for her valu-able comments regarding the manuscript.
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