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Prestigious fruit trees in ancient Israel: firstpalynological evidence for growing Juglans regia andCitrus medicaDafna Langguta

a The Laboratory of Archaeobotany and Ancient Environments, The Sonia and Marco NadlerInstitute of Archaeology, Tel Aviv University, P.O. Box 39040, Tel Aviv 69978, IsraelPublished online: 01 Oct 2014.

To cite this article: Dafna Langgut (2014): Prestigious fruit trees in ancient Israel: first palynological evidence for growingJuglans regia and Citrus medica, Israel Journal of Plant Sciences, DOI: 10.1080/07929978.2014.950067

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Prestigious fruit trees in ancient Israel: first palynological evidence for growing Juglans regia

and Citrus medica

Dafna Langgut*

The Laboratory of Archaeobotany and Ancient Environments, The Sonia and Marco Nadler Institute of Archaeology,Tel Aviv University, P.O. Box 39040, Tel Aviv 69978, Israel

(Received 18 March 2014; accepted 28 July 2014)

This article describes the earliest evidence for the growing of two prestigious fruit trees: Juglans regia (Persian walnut) andCitrus medica (citron) in ancient Israel. The study also tries to identify the origin of these trees as well as their influence onJewish tradition and culture. The palynological information from the Southern Levant supports the hypothesis of the survivalof J. regia during the Last Glacial period in some areas of Eurasia. Accumulating palynological information as well asarcheobotanical evidence of J. regia plant remains from northern Israel from »1800 years BCE suggests the beginning ofhorticulture of walnut in the Southern Levant. The growing of walnut within Israel probably started in the north, and nearlyone millennium later, palynological evidence indicates that J. regia cultivation had spread also to the Judean Mountains.Walnut is mentioned only once in the Bible, in Song of Solomon (6:11). From the interpretation of this text as well otherJewish texts and the available palynological diagrams, it is clear that since the Persian period (fifth to fourth centuries BCE),J. regia was well established in ancient Israel. Citron, although being one of the four species of the Jewish feast of Tabernacles(Sukkot), is not native to the flora of the Near East. The earliest archeobotanical evidence of the growing of C. medica in Israelwas recently discovered in a Royal Persian garden in Ramat Rahel near Jerusalem, dated to the fifth to fourth centuries BCE.C. medica seems to have made its way to Ramat Rahel from India via Persia. From that point on, citron gradually penetratedthe Jewish culture and tradition. The citron is not mentioned in the Bible, and the association between the citron and theP€urı `€ec h€ad€ar (Leviticus 23:40), translated “fruit of the goodly tree,” was only made during the first century AD.

Keywords: Juglans regia; Persian walnut; Citrus medica; citron; pollen; Last Glacial refuge; horticulture; SouthernLevant; ancient Israel; Four Species

Introduction

The introduction of a specific species of tree can be traced

by its first appearance in fossil pollen records and/or by its

presence in botanical assemblages extracted from archeo-

logical sites. While the latter may only point to the impor-

tation of a specific tree (for its timber, fruits, etc.), the

occurrence of pollen grains of a particular plant in a paly-

nological spectrum is direct evidence that it actually grew

in a given region. The Southern Levant natural flora is

reflected in Pleistocene and Early Holocene fossil pollen

diagrams (Horowitz 1968, 1979; Niklewski & van-Zeist

1970; Weinstein-Evron 1983; Kadosh et al. 2004; van-

Zeist & Bottema 2009; van-Zeist et al. 2009; Langgut

et al. 2011; Litt et al. 2012; Aharonovich et al. 2014).

Within the palynological records which cover the second

half of the Holocene, the rise of pollen percentages of

local tree cultivation (e.g., Olea europaea) and the intro-

duction of non-indigenous trees are clearly evident. The

aim of this study is to trace the first palynological evi-

dence of the growing of two prestigious fruit trees in

Ancient Israel: Juglans regia (Persian walnut) and Citrus

medica (citron). The study also tries to shed light on the

origin of these trees, as well as their links to Jewish tradi-

tion and culture in later periods.

Juglans regia

Origin

Juglans regia (Persian walnut) is considered one of the

most important traditional nuts of Old World agriculture

(Zohary et al. 2012). This species is a mesophytic tree

which grows on rich, slightly moist and deep soil on

mountain slopes, in ravines and in the valleys of rivers

and streams (Browicz 1996). As with other old and wide-

spread cultivated plants, it is not easy to reconstruct its

original distribution and determine the borders of its past

natural geographical ranges (Hegi 1981). J. regia is cur-

rently native to the mountain ranges of Central Asia � the

Balkans, eastern Turkey, Armenia, Azerbaijan, north Iran,

south Caspian region, the Caucasus, Pakistan, northern

India, Nepal and Tibet up to the Xinjiang province of

western China (Browicz 1996; Zohary et al. 2012). Today,

*Corresponding author. Email: langgut@post.tau.ac.ilThis paper has been contributed in honor of Professor Daniel Zohary

� 2014 Taylor & Francis

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the species is also found throughout southern and western

Europe, although it is more frequent in south-central and

southeastern Europe (Huntley & Birks 1983).

It is common wisdom that J. regia disappeared from

central Europe, the Balkans and southwest Turkey during

the Last Glacial period (the W€urm, approximately between

75,000 and 15,000 years BP), and reappeared in the Bal-

kans and southwest Turkey not before the middle of the

fourth millennium BP (Zohary et al. 2012). This very late

reappearance suggests that J. regia did not return to these

areas as a post-glacial wild element, but was reintroduced

by humans. Its wild occurrence today in the Balkans and

central Europe therefore represents, according to Zohary

et al. (2012), feral derivatives of cultivated walnut intro-

duced by humans. Browicz (1996) also suggests that the

occurrence of J. regia in the Balkans, Caucasus, Lebanon

and Iraq can be considered to be of anthropogenic origin.

In general, refuge (D a relict � a species that exists as

a remnant of a formerly widely distributed group in an

environment different from that in which it originated) is

by definition always difficult to demonstrate. There is an

ongoing debate whether J. regia populations are from nat-

ural or anthropogenic origin (e.g., Carri�on & S�anchez-Gomez 1992; Fornari et al. 1999; Carri�on et al. 2003;

Beer et al. 2008). This study gathered all the palynological

evidence available for the Southern Levant, which covers

the late Pleistocene and the early Holocene, in order to

cheek the presence of J. regia prior to the beginning of

horticulture.

The palynological evidence

The usefulness of J. regia pollen to indicate its actual

presence in the Southern Levant

The pollen of the walnut tree is relatively large (equatorial

diameter 43�53 mm) and is very easy to identify. The

grain is disc-shaped with a psilate wall (D a smooth sur-

face) and it is considered to be periporate, but it is very

characteristic in that the 10�15 pores are all found on the

equator and in only one hemisphere (heteropolar) (Bot-

tema 2000; Beug 2004). Unlike the pollen of other air-

borne fruit trees (e.g., Olea europaea; Langgut et al.

2014), J. regia pollen does not spread very far despite its

recognizable anemophily (Huntley & Birks, 1983; Bot-

tema & Woldring 1990; Bottema 2000). Its low pollen

dispersal efficiency is ascertained by the investigations of

its “recent pollen rain” in relation to the vegetation (Bot-

tema 2000): several meters from J. regia orchard, pollen

values of this species drop to 0.1�0.2% of the total arbo-

real pollen or even to zero. When values of J. regia

exceeded 2%, a fairly large number of these trees must

have been present close by and/or other tree species may

have been reduced to a minimum. That means that in the

fossil pollen records J. regia pollen is usually under-repre-

sented and its presence, even in very small numbers, indi-

cates the occurrence of this species somewhere in the

vicinity of the sampling location. Wild and cultivated pol-

len grains of J. regia are palynologically indistinguishable

(Bottema & Woldring 1990).

Late Pleistocene�Early Holocene Southern Levant

pollen diagrams

Fossil pollen of J. regia was found in several Southern

Levant palynological studies from the Ghab Valley (Last

Glacial and Early Holocene: Niklewski & van-Zeist

1970), Birkat Ram (Early Holocene, ca. 7000 years BP:

Neumann et al. 2007a), Lake Hula (Last Glacial and

Deglaciation period: Horowitz 1968; Last Glacial: Wein-

stein-Evron 1983; Mid Holocene: van-Zeist et al. 2009),

Sea of Galilee (Early Holocene: Horowitz 1968) and the

Carmel coast (ca. 10,000 years BP: Kadosh et al. 2004)

(Table 1). In all these pollen records, the occurrence of

Table 1. Palynological evidence of the occurrence of J. regia during the late Pleistocene�Early Holocene in the Southern Levant.

Location Chronology of pollen appearance Reference Period being covered in the study

Ghab Valley (Syria) Appearance in several spectraduring the last glacial and earlyHolocene

Niklewski & van Zeist1970

Last Glacial and Early Holocene

Birkat Ram Early Holocene (»7000 years BP) Neumann et al. 2007a » the last 7000 years BP

Lake Hula (Borehole K Jam) Appearance in two spectra: the firstis dated to the Early W€urm andthe second is dated to the endof the Pleistocene, slightly beforethe transition to the Holocene

Horowitz 1968 The Pleistocene and theHolocene

Lake Hula (LO7 borehole) Last Glacial Weinstein-Evron 1983 Last Glacial

Lake Hula (The 1987 Hula core) Early Bronze (»5000 years BP) van Zeist et al. 2009 » the last 12,000 years BP

Sea of Galilee (Borehole D-1016/2) Early Holocene Horowitz 1968 Holocene

Dor, Carmel coast (D-Dor drilling) Beginning of the Holocene(»10,000 years BP)

Kadosh et al. 2004 Early Holocene

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J. regia is usually of a single pollen grain in one pollen

spectrum. However, in the relatively longer diagrams,

sometimes it does appear in several spectra (Table 1).

Because J. regia is poorly dispersed and tends to be

under-represented in the pollen assemblages (Huntley &

Birks 1983; Bottema & Woldring 1990; Bottema 2000), it

is not likely that the pollen grains presented in Table 1

arrived from areas outside the Levant. During the Last

Glacial and the Deglaciation period, J. regia also survived

in several locations in Turkey (e.g., van-Zeist et al. 1975;

Litt et al. 2009). A similar picture is emerging from paly-

nological information from other regions in Europe that

reveals that J. regia was present there in earlier intergla-

cials but that the last glacial severely reduced its numbers.

The earliest pollen evidence for European Juglans comes

from Lower Eocene, France (Gruas-Cavagnetto 1978).

Paleobotanical information indicates that from the Ter-

tiary onwards through the Quaternary, this species became

more frequent in Western Europe (Renault-Miskovski

et al. 1984). In the Deglaciation period and during the

Early Holocene, it is also present in Europe, but not in

impressive numbers, growing as indigenous before the

appearance of horticulture (e.g., Renault-Miskovski et al.

1984). Although the conclusions that various authors

draw are somewhat contradictory (e.g., Carri�on et al.

2003 versus Zohary et al. 2012), it seems that there are

several areas where J. regia possibly survived the last gla-

ciations as refugia: southern Europe, the Near East, China,

and the Himalayas (Beer et al. 2008 and references

therein). For example, palynological data from the Cari-

huela Cave (Granada, southeastern Spain) suggest the

presence of J. regia in that region throughout the early

and middle stages of the last glaciation (Carri�on &

S�anchez-Gomez 1992). An analysis of genetic diversity

and differentiation in European and Asiatic J. regia popu-

lations supports the theory of a native origin of European

walnut in post-glacial times (Fornari et al. 1999). It there-

fore seems that in limited areas, most probably where cli-

matic conditions were milder, J. regia survived the cold,

dry Last Glacial period. The palynological information

from the Southern Levant which is presented in Table 1

lends support to the hypothesis of survival in this region.

Mid-Holocene Levantine palynological diagrams

Since the Middle Bronze Age I (1950�1750 years BCE)

there is increasing palynological evidence for the occur-

rence of J. regia in the Southern Levant. In the Sea of

Galilee pollen diagrams, there are rising percentages from

»1800 years BCE (Figure 1). This finding is corroborated

by archeobotanical data � the earliest J. regia remains in

the land of Israel were found in a Middle Bronze Age

stratum at Tel Megiddo (Liphschitz 2000). According

to stratigraphy (Ilan et al. 2000: 78�79) and the

general chronology of the Tel (Finkelstein, personal

communication) the stratum should be dated to the late

Middle Bronze Age I to early Middle Bronze Age II,

meaning that this archeobotanical finding is most proba-

bly from the »1800�1700 years BCE time interval. Other

archeobotanical remains come from the upper Galilee at

Hazor in a Late Bronze Age stratum (1550�1150 years

BCE; Liphschitz 2000). Walnut remains were also found

at the Caves of Nahal Mishmar (Zaitschek 1961); how-

ever, their context is not clear, as they have not yet been

directly dated (e.g., by AMS radiocarbon) to confirm their

stratigraphy (Chalcolithic or Roman level). In the pollen

records of northern Israel, the appearance of J. regia from

the Middle Bronze Age and into the Iron Age is inconsis-

tent (Neumann et al. 2007a: Birkat-Ram; van-Zeist et al.

2009: Hula Valley; Sea of Galilee: Figure 1). Pollen of

J. regia was also found in an Iron Age stratum at Beth-

saida (Geyer et al. 2009). Since the fifth to fourth centu-

ries BCE, during the Persian period, the Juglans pollen

curves are continuous, showing that the walnut tree was

well established in northern Israel by that time (Neumann

et al. 2007a; van-Zeist et al. 2009). Nearly a millennium

later, there is the first palynological evidence of the occur-

rence of J. regia in pollen diagrams from southern Israel

(the Dead Sea area), showing that it probably grew on the

eastern slopes of the Judean Mountains; The earliest pol-

len evidence comes from an Iron Age II layer, dated to

approximately the ninth century BCE, from a palynologi-

cal study conducted in the Ze’elim Gully (Langgut et al.

2014). Other evidence is dated slightly later: in Ramat

Rahel J. regia pollen is present in assemblages from the

Royal Persian garden, dated to the fifth to fourth centuries

BCE (Langgut et al. 2013a) and in the Ein Feshkha pollen

record, it appears in the Hellenistic period (late fourth

century to late first century BCE; Neumann et al. 2007b).

The appearance of a continuous pollen curve of J. regia in

the Ghab Valley in Syria precedes the evidence from

Northern Israel. According to Bottema and Woldring

(1990), the start of growing of walnut trees for economic

reasons is dated to the early second millennia BCE.

Until now, it is unknown where and when the domesti-

cation of J. regia took place (Zohary et al. 2012). The pal-

ynological evidence does not give a clear picture of the

origin of domestication of the walnut or the direction of

its spread (Bottema 2000). Zohary et al. (2012) claim that

walnut cultivation most likely started in northern Iran,

northeastern Turkey and the Caucasus. Increasing percen-

tages of pollen and more continuous appearances of J.

regia pollen curves since the beginning of the second mil-

lennium BCE are indicated in diagrams from Turkey and

Greece, suggesting that the walnut tree was most likely

planted by farmers (van Zeist et al. 1975; Bottema 1980,

2000; Bottema & Woldring 1984, 1990; Eastwood et al.

1999). In the Zeribar region in Iran, J. regia appears only

at about 500 years BCE (van-Zeist & Bottema 1977). It is

therefore more likely that the idea and knowledge of

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growing J. regia for economic reasons spread to ancient

Israel from the northern Levant.

The walnut in the Bible and Jewish tradition

Its Hebrew name ( זוגא - �egoz) appears only once in the

Bible (Song of Solomon 6:11): “I went down to the nut

orchard, to look at the blossoms of the valley, to see

whether the vines had budded, whether the pomegranates

were in bloom”.

According to Jewish tradition, the book Song of Solo-

mon was written during the time of King Solomon (tenth

century BCE). However, based on its linguistic interpreta-

tions, it is accepted today, among many scholars, that

most of the songs within the book were actually written

several centuries later, during the Hellenistic period (most

probably during the third century BCE; e.g., Zakovitch

1992: 19). Indeed, according to the verse from Song of

Solomon, the walnut grew in a garden with some other

fruit trees such as grape and pomegranate. This means

that by the time the text was written, the walnut tree was

well established as a cultivated fruit tree in the region.

The idea of a garden which includes walnut trees among

other fruit trees is reflected in the archebotanical

reconstruction of the Royal Persian garden in Ramat

Rahel, where in addition to walnut the following fruit

trees were identified palynologically: fig (Ficus carica),

grape (Vitis vinifera), citron (Citrus medica), and olive

(Olea europaea). Although some scholars interpreted the

garden in the Song of Solomon as a metaphor, most

researchers believe that the meaning is of a garden in

which the walnut tree is its main element (Stoop-van-Pari-

don 2005 and references therein). In the Mishnah (dated to

the third century BCE) the walnut is mentioned as a tree

which requires peaa ( האפ D a form of charity � the corner

of a field or orchard left unharvested for the poor to come

and take what they need). Josephus Flavius (first century

AD) praised the fruitful valley near the Sea of Galilee for

its abundance of walnut trees, among other plants (Zohary

1982: 64).

Citrus medica

C. medica (citron, Etrog) is not mentioned in the Bible

among the four species (“arba’at ha-minim”) of the Jew-

ish feast of Tabernacles, Sukkot), and the association

between the citron and the P€urı `€ec h€ad€ar (Leviticus

23:40), translated “fruit of the goodly tree,” was made

hundreds of years later. Additionally, and as opposed to

Figure 1. Palynological diagram of the Sea of Galilee presenting a selection of pollen curves during the Bronze and Iron Ages. A 10-fold exaggeration is used to show changes in taxa percentages (shaded gray). The chronological framework and the division into histori-cal periods are presented in details in Langgut et al. 2013b.

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the other three species (willow, myrtle, and frond from a

date palm), the citron is not native to the Israeli flora

(Zohary et al. 2012). Scholars argue over the question

when the citron first appeared in this region and was iden-

tified as the biblical “fruit of goodly tree.” Biger and

Liphschitz (1987) claimed that citrus fruit generally

arrived in the region during the tenth century AD at the

earliest, and the identification of the citron as the “fruit of

goodly tree” only came about at a much later period. They

believed that the citrus fruit mentioned in the ancient

scripts is not the citron, but rather the Lisbon Lemon,

which also has an elongated shape and a pedicel. Felix

(1987), however, following Gur (1974) and Moldenka

and Moldenka (1952), claimed that the citron existed in

Israel since biblical times. Tolkowsky (1966) proposed a

compromise, arguing the citron made its way to the region

during the conquest of Alexander the Great (late fourth

century BCE).

New recently published pollen findings from Ramat

Rahel near Jerusalem shed new light on the earliest possi-

ble dating of the cultivation of C. medica in ancient Israel

(Langgut et al. 2013a; Figure 2a�c). A Persian Royal gar-

den was excavated in Ramat Rahel, on the western side of

an extravagant palace (Lipschits et al. 2012). While exam-

ining one of the plastered pools in the garden, dating to the

fifth or fourth century BCE, fossilized C. medica pollen

was identified that had been trapped in one of the plaster

layers (various facilities within the garden were plastered

in several layers, most probably due to ongoing mainte-

nance). The unique palynological spectra extracted from

Figure 2. Citrus medica pollen grains: a�c D fossil, e�kD recent. Fossil pollen grains were recovered from the ancient Persian gardenof Ramat Rahel (fifth to fourth centuries BCE). Three varieties of recent pollen grains of C. medica were collected from citron groveslocated in the Sharon area: Halamish variety, Braverman variety and Organg variety. The recent pollen grains were submitted to a simi-lar chemical procedure as the fossil ones. While all C. Medica fossil pollen grains identified in the Ramat Rahel’s pollen assemblageswere colporate with a colpus number of four, within the recent pollen some appeared with four colpus (the majority) while otherappeared with five colpus (2i). Each bar D 10 mm.

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this plaster layer included, in addition to C. medica, other

palynological evidence of trees introduced from remote

parts of the Persian empire of special and highly valued

trees by the ruling Persian authorities [e.g., the cedar of

Lebanon (Cedrus libani)]. The pollen evidence of these

exotic trees in the Ramat Rahel palatial garden suggests

that they were probably brought to flaunt the power of the

imperial Persian administration. Native fruit trees and

ornamentals that were also grown there include the fig

(Ficus carica), grape (Vitis vinifera), olive (Olea euro-

paea), willow (Salix), poplar (Populus), myrtle (Myrtus

communis), and water lily (Nymphaea) (Langgut et al.

2013a). It should be noted that although Citrus pollen is

hardly ever airborne, it appears in high frequencies within

the Ramat Rahel’s palynological assemblages (up to

32%). Therefore, it was argued that at least one citron tree

was located near the sampling location (Langgut et al.

2013a).

Origin

C. medica (etrog) originated in eastern India and south

China, and is the first Citrus to spread west, apparently

through Persia (Zohary et al. 2012). Gur (1974) suggested

that C. medica made its way from India to Afghanistan,

Persia, Syria, Israel and Egypt. In the Southern Levant, C.

medica requires watering (mainly during summer) and

much care in order to thrive (Oppenheimer 1955).

The name etrog ( גורתא ) indicated its origin. In Hindi, the

citron is called torange. In Persian it is called toronge and

later etronge; in Aramaic it’s called etronga ( אגנורתא ) or

etroga ( אגורתא ), in Arabic turug or eturug ( جرتألا ). In Coptic

the citron is ghitri, in Greek k�ıtrion and in Latin citreum or

citrium. The Latin word medicamay also suggest its Persian

(Median) origin or its use for medical purposes. Tolkowsky

(1966) mentions that during the 1950s and 1960s, C. medica

trees which match Theophrastus’s description were grown

in the Gilan province in Persia, which was part of ancient

Medes [Theophrastus of Eresos (287�372 BC) � the great

Greek botanist who wrote the Enquiry into Plants

(Theophrastus 1916) which contains elaborate and accurate

illustrations of East Asian flora; in the fourth chapter of the

book, descriptions of the citron tree are given; the book is

dated to approximately 310 BCE].

The citron is one of the first three ancestral species of

the genus Citrus, along with the pomelo and the mandarin.

This argument was first made by Bart and Rhodes (1976)

and is the popular approach in today’s research, supported

by the latest genetic studies proving that other species of

Citrus are in fact crossbred. Nicolosi et al. (2000) exam-

ined over 40 “species” of Citrus and found that genes

originating in C. medica are a key part in all kinds of

domesticated Citrus, concluding that the citron gave rise

to the various kinds of Citrus known to us today, that are

in fact natural crosses between C. medica and other Citrus

species. One must note that this study examined not only

the nuclear DNA, but also the chloroplast DNA. Based on

the results it was concluded that C. medica was the male

participant in all these citruses, meaning the citron pro-

vided the pollen, as opposed to the female participant (see

also Nicolosi et al. 2005).

As far as we know, C. medica is the first species of

Citrus to spread west (e.g., Zohary et al. 2012), while other

species arrived later. The lemon, the lime and the pomelo

reached the Mediterranean only during the Muslim con-

quest (starting during the seventh century AD), whereas

the orange and mandarin reached the area over the past

few hundred years following developments in maritime

trade (Ram�on-Laca 2003). However, Jashemski et al.

(2002: 101) suggested that the depiction of yellow fruits in

two Roman paintings dated to the first century AD from

the Vesuvius area should be identified as lemon trees.

Palynological and other archeobotanical evidence

Archaeological excavations in Nippur in the south of Bab-

ylonia revealed Citrus seeds dating to the Sumerian period

(4000 years BCE; Bonavia 1894: 68). Because the seeds

found in the excavation were charred, they can only be

identified as Citrus, and specific species cannot be deter-

mined. Tolkowsky (1966) claimed that these seeds were

probably citron seeds brought to Nippur either as an offer-

ing to a divinity or as a gift to a king. He argues that find-

ing seeds in an ancient site is not evidence of the tree

being cultivated in the region, but simply indicates the

fruit was imported. Moreover, the seeds were not dated,

their age being assessed only by their archeological

context.

The French archeologist Loret (1891) claimed that

citrons are evident in the mural at the Karnak Temple,

Egypt, which was built in the time of Thutmosis III

(1490�1450 years BCE). In my opinion it is not possible

to clearly define what was depicted in this mural. Other

scholars reached the same conclusions when they failed to

find a clear connection between the Egyptian murals and

the citron (e.g., Amar 2009).

In the archeological site Hala Sultan Tekke (Cyprus),

seeds that resemble Citrus were discovered in a layer

dated to the twelfth century BCE; however, the exact spe-

cies could not be identified (Hjelmqvist 1979). Based on

Lortet’s observations from the Karnak Temple, Hjelmqv-

ist suggested that these seeds were in fact citron seeds, as

this is the first Citrus species to be grown in Egypt. How-

ever, these remains have not yet been directly dated to

confirm their antiquity (e.g., by AMS radiocarbon; Zohary

et al. 2012). Furthermore, those seeds were found in an

unsealed stratum. In addition and as mentioned above, the

presence of seeds or fruit remains can point to fruit impor-

tation rather than tree cultivation. Better identified and

dated finds have come from three different Roman sites in

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Egypt (van-der Veen 2001, 2003; van-der Veen & Tabi-

nor 2007). Moreover, recent studies indicate that Citrus

seeds can be difficult to recognize due to their morpholog-

ical variability and the changes caused by preservation

processes; traditional methods of seed identification,

therefore, tend to be problematic (Coubray et al. 2010;

Pagnoux et al. 2013). Hence, more advanced identification

methods are recommended when identifying Citrus seeds

such as microtomography technology which enables 2D

and 3D simulation to a scale of microns. The problem

also lies in the similarity of the general morphology of

Citrus seeds and other seeds (e.g., Maloideae types), and

especially when mineralized (Pagnoux et al. 2013).

The archeological evidence mentioned above (Karnak,

Nippur and Cyprus) seems to indicate that the citron fruit

was imported but was not necessarily grown locally. This

is the case even if we accept their indefinite identification

and inconclusive dating and context. However, identifying

citron pollen is a clear sign of a tree grown nearby, espe-

cially in the case of an insect-pollinated species that has

low pollen dispersal efficiency. Furthermore, identifying

seeds, grain or pulp may point to the fruit having existed,

but does not prove that a tree was growing in the area.

It appears that the citron was considered a valuable

commodity since antiquity due to its healing qualities,

symbolic use and pleasant smell on the one hand and its

rarity on the other (e.g., Zohary et al. 2012), possibly mak-

ing the citron known to the people in the region by reputa-

tion throughout antiquity. Liran (2013) reached a similar

conclusion, suggesting that the citron was grown by the

wealthy and represented the rich as it was a rare commod-

ity that only they could afford.

From this overview, one can conclude that the remains

of citron pollen found in the Ramat Rahel Royal Persian

garden are so far the oldest archeobotanical findings indi-

cating that citron was grown in the Near East. Perhaps in

the future, with the development of research and further

excavations, more evidence will be discovered predating

the fifth to fourth centuries BCE findings from Ramat

Rahel. Since the Roman period, the citron became well-

known throughout the Mediterranean (Zohary et al.

2012). As of the first century AD, palynological evidence

from the Vesuvius area indicates that Citrus trees were

probably cultivated in gardens owned by the affluent

(Mariotti-Lippi 2000).

Textual evidence

A precise description of the citron was first written by The-

ophrastus, in his book Enquiry into Plants (1916 English

translation by Hort):

[. . .] And in general the lands of the East and Southappear to have peculiar plants, as they have peculiar ani-mals; for instance, Media and Persia have, among many

others, that which is called the “Median” or “Persianapple”. This tree has a leaf like to and almost identicalwith that of the Arbutus, but it has thorns like those of thepear or white-thorn, which however are smooth and verysharp and strong. The “apple” is not eaten, but it is veryfragrant, as also is the leaf of the tree. And if the “apple”is placed among clothes, it keeps them from being moth-eaten. It is also useful when one has drunk deadly poison;for being given in wine it upsets the stomach and bringsup the poison . . .

The text goes on describing exact instructions on how

to grow the tree along with two key observations: the first

being the tree’s unique quality in bearing fruit during sev-

eral seasons (this means, new fruit may grow on the same

tree alongside fruit that first grew during the previous

year). This character made the citron tree a symbol of an

eternal spring, fertility and inspiring many poets and

artists. The second observation concerns the tree’s flow-

ers, having a prominent pistil, making them more fertile

as opposed to other flowers found to be sterile. From a dif-

ferent piece written by Theophrastus, it appears that the

discovery of sterile flowers with no pedicle was first made

by Persian gardeners, who informed the Greek botanists.

However, they and perhaps Theophrastus himself first rec-

ognized this trait’s significance in identification (Tolkow-

sky 1966). The pedicle of the citron develops from the

style and the stigma. In other citrus species this part

degenerates, whereas in the citron it develops into a pedi-

cle (Felix 1987). Nowadays some citrons no longer pro-

duce pedicles due to cross-breeding with other Citrus

types. In isolated, faraway places, where other species of

Citrus are not grown, all citrons grow a pedicle (for fur-

ther discussions on the topic of the pedicle, see: Felix

1994). The Arbutus mentioned in Theophrastus’ text is

related to the eastern strawberry tree, Arbutus andrachne.

Tolkowsky (1966) holds that the description of the citron

leaf as having a round base and a point end much like the

arbutus eliminates any intent to perhaps refer to a different

Citrus as they all have either winged petioles or very nar-

row chisel-shaped leafs. The pear within this text is

related to the wild Syrian pear, Pyrus syriaca.

Scholars argue that Theophrastus emphasizing that the

tree grew in Media and Persia is further evidence that

prior to »300 years BCE the citron was not widely culti-

vated outside Media and Persia (e.g., Tolkowsky 1966).

Theophrastus’ descriptions in Enquiry into Plants are

based on observations by a number of Greek scholars who

accompanied Alexander the Great’s camp and were given

the task by him to examine the geography, the people and

the natural resources of the countries that he was yet to

conquer. These scholars accompanied the king and his

army in all their campaigns and conquests through Asia

Minor, Syria, Israel, Egypt, and Persia and even reached

the region that is today’s Pakistan. Around 310 BCE, The-

ophrastus published his book Enquiry into Plants,

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including the observations of those accompanying the

king in his journeys. If so, they passed twice through

the area west of Persia and did not mention observing the

growing of citron trees. This leads to the conclusion that

the citron tree was limited to the Iranian plateau and was

not cultivated west of there yet. On the other hand, Theo-

phrastus not describing the fruit itself but rather its charac-

teristics may suggest the citron fruit was known to the

Greeks. In approximately 35 BCE the citron is still

described as an exotic fruit: the Roman author and poet

Virgil mentions the citron as antitoxic and having scented

oil. He names the citron the “Median Apple”.

The citron in the Bible and Jewish tradition

The use of the noun “h€ad€ar” � רדה and the meaning of

“glory” or “grandeur” are typical of the poetry and proph-

ecy in the Bible, and is mentioned six times in the phrase

“hod ve’ h€ad€ar” � רדהודוה (“Splendor and majesty”:

Psalms 21:6, 96:6, 104:1, 111:3, Job 41:10, Chronicles 1

16:27). Other phrases such as “glory and majesty”

(Psalms 8:5) or “strength and dignity” (Proverbs 31:25)

appear in the Bible only once. Usually “h€ad€ar” will referto God’s strength and grandeur (may be compared to Isa-

iah 35:2, for example). Note the parallel phrases

“majesty” and “glory” (“The glory of Lebanon will be

given to it, the majesty of Carmel and Sharon. They will

see the glory of the LORD, The majesty of our God”).

The verse in Leviticus 23:40 instructing the holding of

the four species over the feast of Tabernacles (“And ye

shall take you on the first day the fruit of goodly tree,

branches of palm trees, and the boughs of thick trees, and

willows of the brook; and ye shall rejoice before the Lord

your God seven days”) clearly refers to two known spe-

cies (willow and palm); however, researchers are finding

it difficult to determine whether “the fruit of goodly tree”

and “boughs of thick trees” refer to specific species or can

be summarized as general instructions.

As for the “fruit of goodly tree,” the Septuagint (third

century BCE) determined the phrase consists of a noun

referring to a grand and delightful fruit. This appears to be

the intention in Leviticus 27:30 and in Nehemiah 10:36,

where the verses do not refer to any specific kind of tree.

The phrase “fruit of goodly tree” is not mentioned in the

description on the feast of Tabernacles in Nehemiah

8:13�15 where five different species are mentioned:

On the second day of the month, the heads of all the fami-lies, along with the priests and the Levites, gatheredaround Ezra the teacher to give attention to the words ofthe Law. (14) They found written in the Law, which theLord had commanded through Moses, that the Israeliteswere to live in temporary shelters during the festival ofthe seventh month (15) and that they should proclaim thisword and spread it throughout their towns and in Jerusa-lem: “Go out into the hill country and bring back branches

from olive and wild olive trees, and from myrtles, palmsand shade trees, to make temporary shelters” � as it iswritten”.

Within the description in the Books of the Maccabees

2 10:6�7 there is no mention of “the fruit of goodly tree”,

but rather “ivy-wreathed wands and beautiful branches

and also fronds of palm” (Schwartz 2005). Following this,

a number of scholars have interpreted the phrase in Leviti-

cus 23:40 as a request to bring forth a variety of goodly

fruit in addition to the three other species mentioned

above, or as a general instruction demanding the three

species that follow to be of goodly quality (see also Breen

1972). The Septuagint (third century BCE) refers to a

goodly fruit of a tree and later interpretations (following

the first century AD) mention the citron by its name, usu-

ally following a noun to replace the phrase goodly. For

example, the Targum Onkelos clearly mention “And you

shall take to you on the first day the fruit of the citron

tree.” Despite this, the Vulgata (fourth�fifth centuries

AD), much like the Septuagint, translates the verse liter-

ally, with no clear identification.

Since the first century AD there is a significant change

in texts in which the four species mentioned in Leviticus

23:40 are defined: palm, willow, myrtle and citron. In

Antiquities of the Jews 13 13:5 372); late first century

AD), Josephus describes how the Jews threw citrons at

Alexander Jannaeus for disrespecting of the libation ritual

(compare to Mishna tractate Sukkah 4:9). The citron

became a fixed element in the feast of Tabernacles since

the first century AD in visual sources as well as textual

ones. The citron appears on coins alongside the palm

branch during the fourth year of the great revolt (Hebrew

months of Nissan 69 AD�Adar 70 AD) and is coined

about 60 years later again in the days of Simon bar

Kokhba. The documents from the days of Chazal indicate

that the citron was fully accepted as part of the holiday

tradition with no mention of any kind of objection (Amar

2012: 108�109). Therefore, it seems that before the days

of Chazal, other traditions were not practiced.

Fossil Citrus medica pollen grains versus recent one

Based on morphologic characteristics it was possible to

distinguish the Ramat Rahel Citrus pollen into its species

level, C. medica. In the genus Citrus, C. medica grains are

larger than all other Citrus species and characterized also

by slightly coarser surface ornamentation (Xianghong

1982). The Ramat Rahel C. medica pollen grains are sub-

prolate-spheroidal in shape with average circularity of

1.2. All grains are colporate with a colpus number of four

and their average polar axes was 33.9 mm while the equa-

torial axis average was 28.2 mm (Figure 2a�c and

Table 2).

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During the spring of 2012, pollen grains of three varie-

ties of C. medica were collected from citron groves

located in the Sharon area: Halamish variety, Braverman

variety and Organg variety. The recent pollen grains were

submitted to a similar chemical procedure as the fossil

ones (that includes acetolysis process and mounting in

glycerin) and then were measured using the software

Image J (Table 3). The measurements of recent pollen

grains yielded some differences in comparison to the

Ramat Rahel fossil pollen grains: the recent ones are more

rounded; therefore, their calculated average circularity

was 1.1 (Table 3). All grains are colporate with a colpus

number of four (Figure 2d�k). However, more than 15%

of the grains included colpus number of five (Figure 2i).

In all the three recent varieties of C. medica both averages

of polar axis and equatorial axis were smaller than the cal-

culated average of the fossil grains (Tables 2 and 3).

Table 2. Measurements (in microns) of C. medica fossil pollengrains from the Ramat Rahel’s assemblages.

Order�Polaraxis (P)

(mm)Equatorial diameter (E)

(mm)Circularity(P/E ratio)

1 35.6 28.5 1.2

2 35.7 28.8 1.2

3 29.8 26.1 1.1

4 34.2 29.5 1.2

5 34.3 28.5 1.2

6 34.1 30.4 1.1

7 34.6 27.7 1.2

8 35.4 27.8 1.3

9 33.3 30.9 1.1

10 35.1 26.3 1.3

11 32.1 28.2 1.1

12 32.6 25.8 1.3

Average 33.9 28.2 1.2

�Each measurement represents one pollen grain.

Table 3. Measurements in microns of C. medica recent pollen grains.

C. medica Halamish variety C. medica Braverman variety C. medica Organg variety

Order�

Polaraxis (P)(mm)

Equatorialdiameter (E)

(mm)Circularity(P/E ratio)

Polaraxis (P)(mm)

Equatorialdiameter (E)

(mm)Circularity(P/E ratio)

Polaraxis (P)(mm)

Equatorialdiameter (E)

(mm)Circularity(P/E ratio)

1 31.2 28.8 1.1 29.2 27.1 1.1 29.7 28.2 1.1

2 30.9 26.7 1.2 28.3 27.2 1.0 28.5 26.9 1.1

3 25.6 18.1 1.4 28.4 27.1 1.0 26.6 25.3 1.1

4 30.0 25.8 1.2 36.4 31.1 1.2 29.1 26.5 1.1

5 27.4 23.9 1.1 28.9 27.4 1.1 28.1 25.3 1.1

6 30.7 27.4 1.1 26.7 25.9 1.0 28.3 27.1 1.0

7 28.5 25.5 1.1 30.4 24.1 1.3 28.0 27.0 1.0

8 30.5 27.4 1.1 24.0 22.2 1.1 29.9 29.5 1.0

9 25.2 24.4 1.0 29.7 28.3 1.0 27.5 27.0 1.0

10 27.8 24.2 1.1 27.3 26.6 1.0 29.2 27.6 1.1

11 29.4 27.3 1.1 29.2 27.6 1.1 28.7 28.5 1.0

12 23.2 19.2 1.2 29.1 26.7 1.1 26.1 24.7 1.1

13 28.4 27.6 1.0 29.0 26.1 1.1 28.1 26.8 1.0

14 30.6 29.1 1.1 26.9 23.6 1.1 28.9 28.2 1.0

15 28.2 23.9 1.2 32.2 27.5 1.2 26.3 24.4 1.1

16 34.7 32.7 1.1 26.9 26.4 1.0 28.4 27.1 1.0

17 28.0 24.2 1.2 26.9 25.6 1.1 29.1 26.5 1.1

18 31.6 30.4 1.0 27.8 26.4 1.1 26.8 25.7 1.0

19 22.0 19.2 1.1 31.1 28.7 1.1 28.4 27.5 1.0

20 34.4 31.9 1.1 27.3 26.6 1.0 29.5 27.2 1.1

21 36.6 32.4 1.1 26.6 24.8 1.1 28.7 28.1 1.0

22 26.9 25.2 1.1 30.3 23.2 1.3 32.7 29.4 1.1

23 28.5 24.0 1.2 30.9 24.9 1.2 28.8 27.8 1.0

24 27.4 24.0 1.1 30.0 25.2 1.2 25.4 25.0 1.0

25 30.0 29.7 1.0 28.5 27.4 1.0 28.1 26.0 1.1

26 29.5 24.3 1.2 29.7 28.4 1.0 26.0 25.5 1.0

27 28.8 27.7 1.0 28.3 22.0 1.3 30.3 28.9 1.0

(continued)

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Conclusions

This study evaluated the origin and the beginning of horti-

culture of two prestigious and culturally important fruit

trees into the land of Israel: Juglans regia (Persian wal-

nut) and Citrus medica (citron), mainly based on palyno-

logical information from our region. The presence of

pollen grains of a certain insect-pollinated plant in a paly-

nological spectrum is direct evidence of its actual growing

in the region, unlike other archeobotanical remains (seeds,

grains, pulp charcoal and wood remains) which can come

from importation of fruit or timber rather than the actual

growing of the tree. Because the two investigated trees

are characterized by low pollen dispersal efficiency, their

occurrence within the fossil Israel pollen assemblages

most probably indicates that they grew in the region.

Based on the Southern Levant palynological evidence,

it seems unlikely through common wisdom that J. regia is

native to our region. Its presence in Late Pleistocene�Early Holocene regional pollen records, although in small

percentages, shows that J. regia survived as a refuge in

the Levant during the Last Glacial period, indicating the

native character of the species in the area. Since the Mid-

dle Bronze Age I, around 1800 years BCE, there is

increasing evidence of pollen embedded both in lake sedi-

ments in northern Israel and in remains in archeological

sites, suggesting the beginning of the growing of J. regia

for economic reasons. Within Israel, the growing walnut

trees probably started in the north, and only nearly a mil-

lennium later, around the ninth century BCE spread to the

Judean Mountains. The nut tree is mentioned only once in

the Bible in Song of Solomon (6:11). From the interpreta-

tion of this text as well other Jewish texts and the avail-

able palynological diagrams, it can be concluded that

since the Persian period (fifth�fourth centuries BCE) J.

regia was well established in ancient Israel. The palyno-

logical information from Eurasia does not give a clear pic-

ture of the origin of domestication of the walnut or the

direction of its distribution. Yet, it seems that it spread to

our area from the northern Levant.

Identifying grains of C. medica pollen from the Royal

Persian garden in Ramat Rahel dating to the fifth to fourth

centuries BCE suggests the citron first appeared in the

Table 3. (Continued )

C. medica Halamish variety C. medica Braverman variety C. medica Organg variety

Order�

Polaraxis (P)(mm)

Equatorialdiameter (E)

(mm)Circularity(P/E ratio)

Polaraxis (P)(mm)

Equatorialdiameter (E)

(mm)Circularity(P/E ratio)

Polaraxis (P)(mm)

Equatorialdiameter (E)

(mm)Circularity(P/E ratio)

28 24.4 23.9 1.0 29.6 21.6 1.4 26.3 25.7 1.0

29 26.4 23.4 1.1 30.6 29.4 1.0 29.0 24.1 1.2

30 22.8 21.6 1.1 26.0 25.2 1.0 25.5 25.3 1.0

31 23.1 19.4 1.2 26.3 24.1 1.1 27.4 25.8 1.1

32 25.6 18.2 1.4 26.9 25.0 1.1 29.2 28.0 1.0

33 27.4 24.1 1.1 28.4 25.1 1.1 29.2 24.2 1.2

34 28.5 24.0 1.2 28.9 27.6 1.0 25.3 24.4 1.0

35 26.9 25.2 1.1 25.9 20.6 1.3 28.8 27.8 1.0

36 27.8 23.9 1.2 26.7 21.2 1.3 26.1 25.9 1.0

37 29.5 24.3 1.2 24.6 21.3 1.2 28.7 27.1 1.1

38 28.8 27.7 1.0 32.0 23.3 1.4 27.0 26.7 1.0

39 24.4 23.9 1.0 34.5 27.4 1.3 28.6 27.6 1.0

40 26.4 23.4 1.1 26.8 26.5 1.0 24.9 23.1 1.1

41 22.8 21.6 1.1 31.2 24.4 1.3 26.8 24.7 1.1

42 23.1 19.4 1.2 27.1 25.7 1.1 26.0 25.5 1.0

43 27.8 23.9 1.2 28.1 22.8 1.2 26.9 25.6 1.1

44 23.2 18.9 1.2 30.1 22.3 1.3 24.6 24.0 1.0

45 27.8 24.2 1.1 26.7 24.6 1.1 28.2 27.1 1.0

46 26.1 26.0 1.0 24.8 23.5 1.1 28.0 27.5 1.0

47 22.3 18.9 1.2 26.4 24.5 1.1 27.1 26.6 1.0

48 24.3 22.7 1.1 27.7 22.5 1.2 26.3 25.9 1.0

49 23.5 22.5 1.0 22.3 20.7 1.1 29.8 28.9 1.0

50 26.0 21.7 1.2 26.3 24.4 1.1 28.0 25.1 1.1

Average 27.5 24.6 1.1 28.4 25.2 1.1 27.9 26.5 1.1

�Each measurement represents one pollen grain.

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region probably due to the local Persian ruler’s desire for

an extravagant garden. The purpose of the garden was to

portray the inhabitant as a powerful ruler as in addition to

ruling his subjects, he has also the ability to control over

nature. As a product of society which represents status, the

citron perhaps gradually penetrated the Jewish culture and

tradition. Regarding this, keep in mind that among the pol-

len grains found in Ramat Rahel’s garden, the other two

species out of the “four species” of the Jewish feast of Tab-

ernacles (the myrtle and the willow) were also identified in

the palynological spectrum. However, this is beyond the

scope of this study and will be discussed elsewhere.

C. medica was the first citrus crop that was grown in

southwest Asia and the Mediterranean basin. Several

other citrus crops (e.g., lemon, bitter orange, lime and

pomelo) apparently arrived in the Mediterranean basin

only much later, in early Islamic times (seventh century

AD). C. medica seems to have made its way to Southern

Levant from Persia. The citron (named etrog in Hebrew, a

word of Persian origin) is not mentioned in the Bible, and

the association between the citron and the P€urı `€ec h€ad€ar(Leviticus 23:40), translated as “fruit of the goodly tree,”

was made hundreds of years later. The citron became a

fixed element in the feast of Tabernacles since the first

century AD, as is evident from both textual and visual

sources. Based on the comparison between fossil C.

medica pollen and recent cultivated varieties, it seems

that some evolutionary changes occurred during the last

two millennia: the fossil pollen grain is slightly larger in

size and has more elliptic shape than the recent varieties.

Acknowledgments

I am grateful to Israel Finkelstein for his help in setting the exactdate of the Juglans regia remains from Tel Megiddo and toOded Lipschits for his assitance with the interpretations of bibli-cal and Jewish texts. I wish to thank Yirmy Freud for his help inlocating citron orchards and to Yosef Ludmir for his permissionto conduct my research in his orchards. Michael Kitin isacknowledged for pollen samples preparation. Mordechai Stein,Frank Neumann and Thomas Litt and his team from the BonnPalynological Laboratory are thanked for their help in the extrac-tion of the Sea of Galilee and the Dead Sea (Ze’elim) cores. Iwish also to thank Ehud Weiss and Zohar Amar for their assis-tance in gathering the materials for this study and to NaomiLiran for the helpful exchange of thoughts and ideas.

Funding

This study was funded by the European Research Council underthe European Community’s Seventh Framework Program (FP7/2007�2013)/ERC grant agreement no. 229418.

References

Aharonovich S, Sharon G, Weinstein-Evron M. 2014. Palyno-logical investigations at the middle palaeolithic site of NahalMahanayeem Outlet, Israel. Quat Int. 30:149�166.

Amar Z. 2009. The four species anthology. Tel Aviv.Amar Z. 2012. Flora of the Bible. Rubin Mass. Jerusalem.Bart HC, Rhodes AM. 1976. A numerical taxonomic study of

affinity relationships in cultivated citrus and its close rela-tives. Syst Bot. 1:105�136.

Beer R, Kaiser F, Schmidt K, Ammann B, Carraro G, Willy EG.2008. Vegetation history of the walnut forests in Kyrgyzstan(Central Asia): natural or anthropogenic origin? Quat. Sci.Rev. 27:621�632.

Beug HJ. 2004. Leitfaden der Pollenbestimmung f€ur Mitteleuropaund angrenzende Gebiete. M€unchen: Friedrich Pfeil. German.

Biger G, Liphschitz N. 1987. Is the citron the biblical “hadarfruit”? Beit Mikra. 148:28�33. Hebrew.

Bonavia E. 1894. The flora of Assyrian monuments and its out-comes. London: Westminster.

Bottema S. 1980. On the history of the walnut (Juglans regia L.)in southeastern Europe. Acta Bot Neer. 29:343�349.

Bottema S. 2000. The Holocene history of walnut, sweet-ches-nut, manna-ash and plane tree in the eastern mediteranean.PALLAS. 52:35�59.

Bottema S, Woldring H. 1984. Late quaternary vegetation andclimate of southwestern Turkey, part II. Palaeohistoria.26:123�149.

Bottema S, Woldring H. 1990. Anthropogenic indicators in thepollen record of the eastern Mediterranean. In: Bottema S,Entjes-Nieborg G, van-Zeist W, editors. Man’s role in theshaping of the eastern Mediterranean landscape. Rotterdam:Balkema; p. 231�264.

Browicz K. 1996. Chronology of trees and shrubs in southwestAsia and adjacent regions. Supplement. Poznan: PolishAcademy of Sciences; p. 9�10.

Breen G. 1972. Citrus fruit. Biblical encyclopedia. Jerusalem:Bialik Institute; p. 582�583. Hebrew.

Carri�on JS, S�anchez-G�omez P. 1992. Palynological data in sup-port of the survival of walnut (Juglans regia L.) in the west-ern Mediterranean area during last glacial times. J Biogeogr.19:623�630.

Carri�on JS, Yll EI, Walker MJ, Legaz AJ, Cha�ın C, L�opez A.2003. Glacial refugia of temperate, Mediterranean andIbero-North African flora in south-eastern Spain: new evi-dence from cave pollen at two Neanderthal man sites. GlobEcol Biogeogr. 12:119�129.

Coubray S, Zech-matterne V, Mazurier A. 2010. The earliestremains of a citrus fruit from a western Mediterraneanarchaeological context? A microtomographic based re-assessment. Palevol. 9:277�282.

Eastwood WJ, Roberts N, Lamb HF, Tibby JC. 1999. Holoceneenvironmental change in southwest Turkey: a palaeoecologi-cal record of lake and catchment-related changes. Quat SciRev. 18:671�695.

Felix Y. 1987. Citrus fruit � the citron. Beit Mikra. 148:288�292.Hebrew.

Felix Y. 1994. Fruit trees � in the biblical and talmudic litera-ture. Jerusalem: Rubin Mass Press.

Fornari B, Cannata F, Spada M, Malvolti ME. 1999. Allozymeanalysis of genetic diversity and differentiation in Europeanand Asiatic walnut (Juglans regia L.) populations. ForGenet. 6:115�127.

Geyer PS, Larson T, Cyr C, Mansour A. 2009. Pollen analysis atet-Tell, 1996�2006: laboratory report on the Iron Age Gate.In: Arav R, Freund R, editors. Bethsaida: a city by the north-ern shore of the Sea of Galilee (4). Kirksville: Truman StateUniversity Press; p. 180�192.

Gruas-Cavagnetto C. 1978. Etude palynologique de l’Eoc�ene dubasin Anglo-Parisien. M�em Soc G�eol Fr. 131:1�64.

Israel Journal of Plant Sciences 11

Dow

nloa

ded

by [

Tel

Avi

v U

nive

rsity

] at

21:

13 0

3 O

ctob

er 2

014

Gur A. 1974. Fruits of the land of Israel. Jerusalem: HakibbutzHameuchad.

Hegi G. 1981. Illustrierte Flora von Mitteleuropa. Pteridophyta,Spermatophyta, Band 3: Angiospermae, Dicotyledones 1,Teil 1. Dritte €uberarbeitete und erweiterte Auflage. PaulParey Verlag. Berlin: Hamburg; p. 3�15.

Hjelmqvist H. 1979. Some economic plants and weeds from theBronze Age of Cyprus. Stud Mediterr Archeol. 45:110�117.

Horowitz A. 1968. Upper Pleictocene-Holocene climate and vege-tation of the Northern Jordan valley, Israel [PhD dissertation].Jerusalem: Hebrew University. Hebrew, with English abstract.

Horowitz A. 1979. The quaternary of Israel. New York. Aca-demic Press.

Huntley B, Birks HJ. 1983. Atlas of past and present pollen mapsfor Europe, 0�13,000 years ago. Cambridge: CambridgeUniversity Press.

Ilan D, Franklin N, Hallote S. 2000. Area F. In: Finkelstein I,Ussishkin D, Halpern B. Megiddo III, editors. The 1992-1996 seasons. Monograph series 18. Tel Aviv: Tel Aviv Uni-versity; p. 75�103.

Jashemski WF, Meyer FG, Ricciardi M. 2002. Plants � evidencefrom wall paintings, mosaics, sculpture, plant remains, graffiti,inscription and ancient authors. In: Jashemski WF, Meyer FG,editors. The natural history of Pompeii. Cambridge: Cam-bridge University Press.

Kadosh D, Sivan D, Kutiel H, Weinstein-Evron M. 2004. A latequaternary paleoenvironmental sequence from Dor, CarmelCoastal Plain, Israel. Palynol. 28:143�157.

Langgut D, Almogi-Labin A, Bar-Matthews M, Weinstein-Evron M. 2011. Vegetation and climate changes in the southeastern Mediterranean during the last glacial�interglacialcycle (86 ka): new marine pollen core. Quat Sci Rev.30:3960�3972.

Langgut D, Gadot Y, Porat N, Lipschits O. 2013a. Fossil pollenreveals the secrets of royal Persian garden in Ramat Rahel(Jerusalem). Palynol. 37:115�129.

Langgut D, Finkelstein I, Litt T. 2013b. Climate and the LateBronze collapse: new evidence from the Southern Levant.Tel Aviv. 40:149�175.

Langgut D, Neumann FH, Stein M, Wagner A, Kagan EJ, Boar-etto E, Finkelstein I. 2014. Dead Sea pollen record and his-tory of human activity in the Judean Highlands (Israel) fromthe Intermediate Bronze into the Iron Ages (»2500�500BCE). Palynol. 38(2):1�23.

Langgut D, Lev-Yadun S, Finkelstein I. 2014. The impact ofolive orchard abandonment and rehabilitation on pollen sig-nature: an experimental approach to evaluating fossil pollendata. Ethnoarchaeol. 6:121�135.

Liphschitz N. 2000. Archaeobotanical finds. In: Finkelstein I,Ussishkin D, Halpern B, editors. Megiddo III: the1992�1996 seasons. Monograph series 18. Tel Aviv: TelAviv University; p. 487�495.

Lipschits O, Gadot Y, Langgut D. 2012. The riddle of RamatRahel: the archaeology of a royal edifice from the Persianperiods. Transeuphraten. 41:57�79.

Liran N. 2013. The etrog in the Jewish culture: interdisciplinarystudy of the ritual object throughout the ages. [PhD disserta-tion]. University of Haifa. Hebrew, with English abstract.

Litt T, Krastel S, Sturm M, Kipfer R, Orcen S, Heumann G, FranzSO, Ulgen UB, Niessen F. 2009. ‘PALEOVAN, internationalcontinental scientific drilling program (ICDP): site surveyresults and perspectives. Quat Sci Rev. 28:1555�1567.

Litt T, Ohlwein C, Neumann FH, Hense A, Stein M. 2012. Holo-cene climate variability in the Levant from the Dead Seapollen record. Quat Sci Rev. 49:95�105.

Loret V. 1891. Le c�edratier dans l’antiquit�e. Annales de laSoci�et�e Botanique de Lyon 17:225�271.

Mariotti-Lippi M. 2000. The garden of the “Casa delle Nozze diErcole ed Ebe” in Pompeii (Italy): palynological investiga-tions. Plant Biosyst. 134:205�211.

Moldenke HN, Moldenke AL. 1952. Plants of the Bible. Tor-onto: Waltham, Mass; p. 290�292.

Neumann FH, Sch€olzel C, Litt T, Hense A, Stein M. 2007a. Holo-cene vegetation and climate history of the northern GolanHeights (Near East). Veg Hist Archaeobot. 16:329�346.

Neumann FH, Kagan EJ, Schwab MJ, Stein M. 2007b. Palynol-ogy, sedimentology and palaeoecology of the late HoloceneDead Sea. Quat Sci Rev. 26:1476�1498.

Nicolosi E, Deng ZN, Gentile A, La Malfa S, Continella G, Trib-ulato E. 2000. Citrus phylogeny and genetic origin of impor-tant species as investigated by molecular markers. TheorAppl Genet. 100:1155�1166.

Nicolosi E, La Malfa S, El-Otmani M, Negbi M, GoldschmidtEE. 2005. The search for the authentic citron (Citrus medicaL.): historic and genetic analysis. HortSci. 40:1963�1968.

Niklewski J, van Zeist W. 1970. A late quaternary pollen diagramfrom Northwestern Syria. Acta Bot Neerl. 19:737�754.

Oppenheimer H. 1955. Citrus cultivation. Tel Aviv. The SadeLibrary. Hebrew.

Pagnoux C, Celant A, Coubray S, Fiorentino G, Zech-MatterneV. 2013. The introduction of citrus to Italy, with reference tothe identification problems of seed remains. Veg HistArchaeobot. 22:421�438.

Ram�on-Laca L. 2003. The introduction of cultivated citrus toEurope via Northern Africa and the Iberian Peninsula. EconBot. 57:502�514.

Renault-Miskovsky J. 1984. A propos de l’indigenat ou del’introduction de Juglans et Platanus dans l’ouest del’Europe au Quaternaire. Rev Pal�eobiologie. Special issue;155�178.

Schwartz D. 2005. The second book of Maccabees: introduction,Hebrew translation and commentary. Jerusalem: Yad Ben-Zvi.

Stoop-van-Paridon PWT. 2005. The Song of Songs� a philolog-ical analysis of the Hebrew book םירישהריש . Ancient NearEastern Studies Supplements 17, Louvain: Peeters;p. 338�339.

Theophrastus. 1916. Enquiry into plants. A.F. Hort, trans. Cam-bridge, MA: Loeb Classical Library.

Tolkowsky S. 1966. Citrus fruits � their origin and historythroughout the world. Jerusalem: The Bialik Institute.

van-der Veen M. 2001. The botanical evidence. In: MaxfieldVA, Peacock DPS, editors. Survey and excavations at MonsClaudianus 1987�1993, volume 2. Cairo: Institut Francaisd’Arch�eologie Orientale du Caire; p. 174�247.

van-der Veen M. 2003. When is food a luxury? World Archaeol.34:405�427.

van-der Veen M, Tabinor H. 2007. Food, fodder and fuel atMons Porphyrites: the botanical evidence. In: Maxfield VA,Peacock DPS, editors. The Roman imperial quarries, surveyand excavation at Mons Porphyrites 1994�1998, volume 2.London: Egypt Exploration Society; p. 83�142.

van-Zeist W, Bottema S. 1977. Palynological investigations inWestern Iran. Palaeohistoria. 19:19�85.

van-Zeist W, Bottema S. 2009. A palynological study of theAcheulian site of Gesher Benot Ya’aqov, Israel. Veg HistArchaeobot. 18:105�121.

van-Zeist W, Woldring H, Stapert D. 1975. Late quaternary veg-etation and climate of Southwestern Turkey. Palaeohistoria17:53�143.

12 D. Langgut

Dow

nloa

ded

by [

Tel

Avi

v U

nive

rsity

] at

21:

13 0

3 O

ctob

er 2

014

van-Zeist W, Baruch U, Bottema S. 2009. Holocene palaeoecol-ogy of the Hula area, Northeastern Israel. In: Kaptijn K, PetitLP, editors. A timeless vale, archaeological and relatedessays on the Jordan Valley. Leiden: Leiden UniversityPress; p. 29�64.

Weinstein-Evron M. 1983. The paleoecology of the early Wurmin the Hula Basin, Israel. Pal�eorient. 9:5�19.

Xianghong YYKYZ. 1982. Studies on the pollen morphology ofcitrus plants. Scientia Agric Sinica. 112:88�93.

Zaitschek DV. 1961. Remains of cultivated plants from the cavesof Nahal Mishmar. IEJ. 11:70�72.

Zakovitch Y. 1992. The Song of Songs � introduction and com-mentary (Mikra Leyisra’el � a biblical commentary forIsrael). Tel Aviv: Am Oved. Hebrew.

Zohary D, Hopf M, Weiss E. 2012. Domestication of plants inthe old world. 4th ed. Oxford: Oxford University Press.

Zohary M. 1982. Plants of the Bible. Cambridge: CambridgeUniversity Press: Cambridge.

Israel Journal of Plant Sciences 13

Dow

nloa

ded

by [

Tel

Avi

v U

nive

rsity

] at

21:

13 0

3 O

ctob

er 2

014