Genet Resour Crop Evol DOI 10.1007/s10722-017-0530-4

OBITUARY

Daniel Zohary (1926–2016) Aaron Rottenberg

Received: 28 February 2017 / Accepted: 31 May 2017 Ó Springer Science+Business Media Dordrecht 2017

Daniel Zohary

A. Rottenberg (&) Department of Biotechnology, Ort Braude College, P.O. Box 78, 21982 Karmiel, Israel e-mail: [email protected]

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Daniel Zohary examining a dry Cynara. Israel, 2003

Professor Daniel Zohary was an Israeli plant geneticist. He is best known to the scientific community for the seminal book Domestication of Plants in the Old World (Zohary et al. 2012). Knowing him for almost half of my life was a great and fascinating experience. Since our first meeting some two and a half decades ago, I was captured by his outstanding wide knowledge and the will to share it, his love to his homeland and its plants and his devotion to his colleagues and students, all wrapped with good nature and friendly approach.

Early life Daniel (Dani) Zohary was born to Michael and Leah Zohary in Jerusalem on the 24th of April 1926. His father Michael, a famous professor of botany, was an expert on the local flora, a prolific writer and a pioneer in the field of geobotany of the Near East

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(Zohary 1966, 1973). Young Daniel accompanied his father on numerous botanical expeditions, helping him with collecting and drying plants, notably in the Huleh Valley at northern Israel. It was then, that Daniel has acquired his great love and understanding of the wild flora, fauna and geology of Israel and the Near East. As a youngster he participated with other young friends in many hiking expeditions, some of them to less known areas of that time such as the Hermon and the Negev. In these early journeys he extended his knowledge of the local geology and phytogeography while establishing strong friendships and common biological views with researchers like Tuviah Kushnir, Daniel Raz and Eviatar (Eibi) Nevo (Zohary 2009). At the age of 17 Zohary joined the Palmach, a fighting force, and after three years he went with dozens of his Palmach associates to study at the Hebrew University in Jerusalem. Alas, his studies were interrupted by the 1948 war and he was stationed in the Jerusalem Corridor. During the heavy fighting his platoon was almost entirely wiped out and many bright students lost their lives, among them, his close friend Tuviah Kushnir (1952). Zohary wrote a short and painful testimony on that period (Levin 1998, p. 58) and in fact carried all his life the memory of his fallen friends and the heavy feelings of the missed opportunities of these gifted would-be scientists. When the war was over, he managed to finish his studies in botany and geology (Zohary 1953, 1954), and married his girlfriend Devora. In 1952 Zohary went to the University of California in Berkeley where he specialized in cytogenetics. He wrote a Ph.D. dissertation ‘‘Cytogenetic studies in the polyploid complex of Dactylis glomerata’’ under the supervision of G. Ledyard Stebbins. There, from his studies in polyploidy and meiosis he made his first steps as an evolutionary biologist (Brown and Zohary 1955; Stebbins and Zohary 1959). At Berkeley, he received the John Belling prize in genetics. Apart from evolutionary insight and deep understanding of reproduction systems, he learned and perfected the skill of preparing microscope slides. Many years later, we were spending hours at the laboratory gazing at various Cynara cells trying to detect their chromosomes. Daniel described them as ‘‘notoriously smaller than fly droppings’’, but then, through his artful skill we finally managed to count 34 tiny chromosomes.

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Professor at the Hebrew University In 1956, the Zohary family returned to Jerusalem, now with their daughter Tamar who was followed later by Ruth and Ehud. Daniel has started his long academic career at the Hebrew University in the department of botany and as one of the founders of the department of genetics. At that time, he was one of the very first geneticists in Israel, and was quickly recognized as a bright researcher and an inspiring teacher. His mastery in cytogenetics, along with his good character and his endless dedication to his students attracted to his lab a line of students with whom he published several studies in different aspects of cytogenetics in various plant genera (e.g. Zohary and Nur 1959; Zohary and Feldman 1962; Vardi and Zohary 1967; Ladizinsky and Zohary 1968; Puteyevsky and Zohary 1970; Ben Ze’ev and Zohary 1973; Zohary and Plitmann 1979; Avishai and Zohary 1980; Rottenberg et al. 1999). Many of his students became later his colleagues, and indeed some of Israel’s finest plant breeders and geneticists have graduated under his supervision (see Simchen 2014). During his career as a scientist and a teacher Zohary has developed interest in many fields including reproduction and sexual mechanisms, ecology and conservation, breeding, polyploidy, and genetic systems; but his focus for many decades to come remained the genetic affinities between related plant species and the domestication of plants and animals. Zohary has first concentrated on the genetic affinities among grass species in the genera Dactylis (Zohary and Ashkenazi 1958), Triticum and Aegilops (Ankori and Zohary 1962; Zohary and Feldman 1962; Pazy and Zohary 1965; Zohary et al. 1969), Hordeum (Tovia and Zohary 1962) and Avena (Ladizinsky and Zohary 1968). In this chain of studies, he understood the significant findings related to the cultivation of wheats, barley, oats and rye. Now, he was actually ready to write some of his pivotal articles regarding these important crops (Zohary 1965, 1969, 1971). Later, beginning in the 1970s, Zohary collaborated with his longtime friend Eviatar Nevo. They have found the genetic variation in natural populations of wild barley and wild wheats as well as other relevant implications such as correlations to ecological factors and resistance to pathogens (Brown et al. 1978; Nevo et al. 1982, 1986).

At the same time, in order to better understand the history of plant domestication, Zohary started to collaborate with archaeologists and published important studies on the origin of domesticated pulses: pea, lentil, bitter vetch, chickpea and faba bean (Zohary and Hopf 1973; Garfinkel et al. 1988). Apart from cereals and pulses, Zohary also dealt in the following years with the mode of domestication of virtually all fruit trees of the area: olive, grape, fig, date palm, pomegranate, sycamore, almond and carob (Zohary and Spiegel-Roy 1975; Zohary 1994, 1995, 2002; Browicz and Zohary 1996; Lavee and Zohary 2011), as well as plums, apples and pears (Zohary 1992, 1997a) and the genus Pistacia (Zohary 1996a). He was pleased when he was invited to write the latter, and regarded it as a tribute to his father who had written a monography on Pistacia decades before (Zohary 1952). Vegetables were also part of his main concern, among them were garlic, lettuce and artichoke (Zohary 1991a; Rottenberg and Zohary 1996, 2005). Apart from dealing with the crops themselves, Zohary concluded that an assemblage of wild species, namely the eight founder crops of the Near East (cereals, pulses and flax) were brought together into cultivation at the very beginning of agriculture (Zohary 1989, 1996b). He emphasized the importance of some biological traits of these species (e.g. self-pollination and annualism, Zohary 1996b, 1997b) and the possible mode of domestication of plants and animals (Zohary 1996b, 1999, 2004; Zohary et al. 1998).

Additional research Daniel Zohary was a gifted field botanist with a vast knowledge of the flora of Israel, the Near East and the Mediterranean basin. Actually, he was one of a handful of local botanists who could identify in nature nearly all wild plants of Israel, that is, more than 2500 species. Moreover, he knew the life traits, ecological features, wild or cultivated relatives or any other relevant aspects of most of these species. He had a sharp eye both for plants and animals and his funny skill was ‘‘AutoBotanica’’ (short for automobile botanizing), that is spotting interesting or rare plants while driving, and immediately pulling over for inspecting or collecting them. This field competence along with deep knowledge in cytogenetics,

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morphology and plant systematics led him and Joseph Katzenelson to describe a new clover species: Trifolium israeliticum (Zohary and Katznelson 1958). Also, together with Fania Kollmann he described the new species Allium truncatum (Kollmann and Zohary 1986), which he suspected that might stand as the elusive wild progenitor of domesticated garlic. In one of our journeys to the Negev he suddenly stopped the car, loped to the field and came back straightaway with this A. truncatum and cracked the bulbs, just to excite ourselves with the familiar garlic smell of this species. One of the main concerns of Zohary was nature conservation. As a boy, he accompanied his father to the wildlife-rich Huleh Valley and was impressed by its biological diversity, just to see its destruction later in the 1950s, and then to realize the importance of nature protection. Thus, he was always aware in his excursions to the maintenance of wildlife and their habitat. He was gifted with a memory of an elephant and excellent spatial orientation, so whenever discussing rare species or endangered habitats he remembered most locations of wild plants that he ever saw, including those that were long extinct. One day in the mid-1990s we were looking for an extremely rare sorrel in its last habitat near the coast. Daniel had a clear recollection of three other extinct localities where he observed this endemic sorrel many decades before. Later, upon rechecking the literature, we found that one of these extinct locations, near Binyamina, is in fact the very same spot where this rare sorrel was first collected in 1906 (Rottenberg 2001). Zohary developed a solid approach to environmental and biological conservation and particularly to the conservation of the wild progenitors of cultivated plants (Zohary 1983, 1991b, 1997a). He collaborated with other experts to create a basis for the recognition and protection of these wild progenitors and their environment (Heywood and Zohary 1995; Gabrielian and Zohary 2004). Another line of research, an applied one, involved plant breeding. Horovitz and Zohary (1966) deciphered the genetics that control Anemone coronaria various flower colours while she developed robust, colourful commercial cultivars. Later on, Zohary started a long collaboration with agronomist and friend Jehuda Basnizki on globe artichoke breeding. In painstaking crossing programmes and careful selections they developed a unique seed-planted cultivar (Basnizki and Zohary 1987, 1994). This cultivar

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transformed artichoke from a manually vegetatively propagated crop into a seed-planted vegetable, adjusted to modern mechanized cultivation. This was a revolutionizing innovation and the seeds of this novel cultivar became highly successful.

The book In the core of his academic career, Daniel Zohary accumulated vast knowledge regarding crops and their wild relatives: their geographical distribution, genetic and reproduction systems, and the modifications under domestication. To these, he added a keen interest in the rich Near Eastern archaeological record. Here, he found a great partner in Maria Hopf, an archaeobotanist from Mainz, with whom he already published an important paper (Zohary and Hopf 1973) and discussed the interesting findings from Jericho and Arad excavations during her long visit in Israel. Eventually, their partnership culminated in the coauthored book Domestication of Plants in the Old World (Zohary and Hopf 1988). In this seminal volume they reviewed the ecology, biology, wild ancestry and the archaeological evidence of the agricultural plants of the Old World. Among the plants surveyed are cereals, pulses, vegetables, fruit trees, nuts, and other crops producing oil, fibre, dye and condiments. A concluding chapter reviews the main archaeological sites with their most important archaeobotanical findings. Domestication of Plants in the Old World became immediately the basic textbook and the main reference to students and scholars in the discipline of the origin and spread of agriculture as well as other fields associated to crops and archaeology. Jared Diamond, while visiting Zohary in Israel acquired both from Zohary himself and the book a deep perception regarding the origin of agriculture. Diamond incorporated it into core chapters in his Pulitzer winner Guns, Germs and Steel, and wrote later that one can take Zohary and Hopf’s book as a gold standard (Diamond 2002). Second and third revised editions followed, but unfortunately, in 2008, during the work on the fourth edition Maria Hopf passed away. Ehud Weiss was invited to contribute to this edition (Zohary et al. 2012; and see Weiss 2015) and later, a festive seminar in honor of Daniel Zohary was held in Jerusalem to

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celebrate the publication of the fourth edition in March 2012. At that time, Zohary was weak, but with his usual good spirit he enjoyed very much his family, friends and colleagues, as well as the lectures and the warm blessings. Daniel Zohary was awarded Distinguished Economic Botanist for 2003. Also, he was a Fellow of the Linnean Society. Several journal volumes were dedicated to him (Pickersgill 2004; Parker and Kolska Horwitz 2012; Lev-Yadun and Langgut 2015), and a warm biography with an extended publication list was published by his colleague Giora Simchen (2014). A comprehensive, chronological publication list is found in www.danielzohary.com. In 1976 Daniel’s wife, Devora died, and a few years later he married Lilly. They lived together for almost four decades. In his last years Daniel was not active in research, but he was always happy to see his colleagues and friends, especially if plants were involved. Daniel Zohary passed away peacefully at home, surrounded by his family on the 16th of December 2016. He will be remembered as an outstanding scientist, curious and enthusiastic, and always happy to share his knowledge. His kind, friendly personality gained him many close students and colleagues, some of them will never forget the warm welcoming hospitality of the Zoharys at their Jerusalem home. Indeed, it was a cherished pleasure to chat with Daniel at the kitchen table and to write papers at his small round table, as it was always exciting to run with him on the hills after interesting plants, to work at the lab, and to learn from him, always with his smile and generosity. Compliance with ethical standards Conflict of interest

The author declares no conflict of interest.

References Ankori H, Zohary D (1962) Natural hybridization between Aegilops sharonensis and Ae. longissima: a morphological and cytological study. Cytologia 27:314–324 Avishai M, Zohary D (1980) Genetic affinities among the Oncocyclus irises. Bot Gaz 141:107–115 Basnizki J, Zohary D (1987) A seed planted cultivar of globe artichoke. HortScience 22:678–679 Basnizki J, Zohary D (1994) Breeding of seed-planted artichoke. Plant Breed Rev 12:253–269 Ben-Ze’ev N, Zohary D (1973) Species relationships in the genus Pisum L. Isr J Bot 22:73–91

Browicz K, Zohary D (1996) The genus Amygdalus L. (Rosaceae): species relationships, distribution and evolution under domestication. Genet Res Crop Evol 43:229–247 Brown SW, Zohary D (1955) The relationship between chiasmata and crossing over in Lilium formosanum. Genetics 40:850–873 Brown AHD, Nevo E, Zohary D, Dagan O (1978) Genetic variation in natural populations of wild barley (Hordeum spontaneum). Genetica 49:97–108 Diamond J (2002) Evolution, consequences and future of plant and animal domestication. Nature 418:700–707 Gabrielian E, Zohary D (2004) Wild relatives of food crops native to Armenia and Nakhichevan. Flora Mediterr 14:5–80 Garfinkel Y, Kislev ME, Zohary D (1988) Lentil in Pre-Pottery Neolithic B Yiftah’el: additional evidence of its early domestication. Isr J Bot 37:49–51 Heywood VH, Zohary D (1995) A catalogue of the wild relatives of cultivated plants native to Europe. Flora Mediterr 5:361–401 Horovitz A, Zohary D (1966) Spontaneous variation for perianth colour in wild Anemone coronaria. Heredity 21:513–515 Kollmann F, Zohary D (1986) Allium truncatum. In: FeinbrunDothan N (ed) Flora palaestina, vol 4. Israel Academy of Sciences and Humanities, Jerusalem Kushnir T (1952) Heterochromatic polysomy in Gryllotalpa gryllotalpa L. J Genet 50:361–383 Ladizinsky G, Zohary D (1968) Genetic relationships between diploids and tetraploids in series Eubarbatae of Avena. Canad J Genet Cytol 10:68–81 Lavee S, Zohary D (2011) The potential of genetic diversity and the effect of geographically isolated resources in olive breeding. Isr J Plant Sci 59:3–13 Levin A (1998) Testament. Artisan, New York Lev-Yadun S, Langgut D (2015) Studies in botanical archeology, ethno-botany and plant domestication: honoring Professor Daniel Zohary. Isr J Plant Sci 62:1–4 Nevo E, Golenberg E, Beiles A, Brown AHD, Zohary D (1982) Genetic diversity and environmental associations of wild wheat Triticum dicoccoides, in Israel. Theor Appl Genet 62:241–254 Nevo E, Beiles A, Zohary D (1986) Genetic resources of wild barley in the Near-East: structure, evolution and application in breeding. Biol J Linn Soc 27:335–380 Parker AG, Kolska Horwitz L (2012) Introduction to special issue on ‘‘Ancient Agriculture in the Middle East’’ dedicated to Daniel Zohary. J Arid Environ 86:1–4 Pazy B, Zohary D (1965) The process of introgression between Aegilops polyploids: natural hybridization between Ae. variabilis, Ae. ovata and Ae. biuncialis. Evolution 19:385–394 Pickersgill B (2004) The Society for Economic Botany Professor Daniel Zohary 2003 distinguished economic botanist. Econ Bot 58:3–4 Puteyevsky E, Zohary D (1970) Behaviour and transmission of supernumerary chromosomes in diploid Dactylis glomerata. Chromosoma 32:135–141 Rottenberg A (2001) Rumex rothschildianus, an endemic-toIsrael species, on the verge of global extinction. Isr J Plant Sci 49:57–60 Rottenberg A, Zohary D (1996) The wild ancestry of the cultivated artichoke. Genet Res Crop Evol 43:53–58

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Genet Resour Crop Evol Rottenberg A, Zohary D (2005) Wild genetic resources of the cultivated artichoke. Acta Hortic 681:307–312 Rottenberg A, Zohary D, Nevo E (1999) Patterns of isozyme diversity and vegetative reproduction of willows in Israel. Inter J Plant Sci 160:561–566 Simchen G (2014) Daniel Zohary: geneticist and explorer of plant domestication. Plant Breed Rev 38:1–16 Stebbins GL, Zohary D (1959) Cytogenetic and evolutionary studies in the genus Dactylis. I. The morphology distribution and inter-relationships of the diploid sub-species. Univ Calif Publ Bot 31:1–40 Tovia T, Zohary D (1962) Natural hybridization between Hordeum vulgare and H. spontaneum: corroborative evidence from progeny tests and artificial hybrids. Bull Res Counc Isr 11D:43–45 Vardi A, Zohary D (1967) Introgression in wheat via triploid hybrids. Heredity 22:541–560 Weiss E (2015) ‘‘Beginnings of Fruit Growing in the Old World’’—two generations later. Isr J Plant Sci 62:75–85 Zohary M (1952) A monographic study of the genus Pistacia. Pal J Bot Jerusalem series 5:187–228 Zohary D (1953) Ecological studies in the vegetation of the Near Eastern Deserts. III. Vegetation map of the Central and Southern Negev. Palest J Bot 6:27–36 Zohary D (1954) Notes on ancient agriculture in the Central Negev. Isr Explor J A4:17–25 Zohary D (1965) Colonizer species in the wheat group. In: Baker GH, Stebbins GL (eds) The genetics of colonizing species. Academic Press, New York Zohary M (1966) Flora Palaestina. Israel Academy of Sciences and Humanities, Jerusalem Zohary D (1969) The progenitors of wheat and barley in relation to domestication and agricultural dispersal in the Old World. In: Ucko PJ, Dimbleby GW (eds) The domestication and exploitation of plants and animals. Duckworth, London Zohary D (1971) Origin of southwest Asiatic cereals: wheats, barley, oats and rye. In: Davis PH, Harper PC, Hedge I (eds) Plant life in South-west Asia. Botanical Sosciety of Edinburgh, University Press, Aberdeen Zohary M (1973) Geobotanical foundations of the Middle East. Fischer, Stuttgart Zohary D (1983) Wild genetic resources of crops in Israel. Isr J Bot 32:97–127 Zohary D (1989) Domestication of the Southwest Asian Neolithic crop assemblage of cereals, pulses and flax: the evidence from the living plants. In: Harris DR, Hillman GC (eds) Foraging and farming: the evolution of plant exploitation. Unwin & Hyman, London Zohary D (1991a) The wild genetic resources of cultivated lettuce (Lactuca sativa L.). Euphytica 53:31–35 Zohary D (1991b) Conservation of wild progenitors of cultivated plants in the Mediterranean Basin. Bot Chron 10:467–474 Zohary D (1992) Is the European plum, Prunus domestica L., a P. cerasifera Ehrh. 9 P. spinosa L. allo-polyploid? Euphytica 60:75–77

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Zohary D (1994) The wild genetic resources of the cultivated olive. Acta Hortic 356:62–65 Zohary D (1995) Fig: Ficus carica (Moraceae). In: Smartt J, Simmonds NW (eds) Evolution of crop plants. Longman, London Zohary D (1996a) The genus Pistacia L. In: Padulosi S, Caruso T, Barone E (eds) Taxonomy, distribution, conservation and uses of Pistacia genetic resources. Report of a workshop, 29–30 June 1995, Palermo, Italy. International Plant Genetic Resources Institute, Rome, pp 1–11 Zohary D (1996b) The mode of domestication of the founder crops of Southwest Asian agriculture. In: Harris DR (ed) The origins and spread of agriculture and pastoralism in Eurasia. University College London Press, London Zohary D (1997a) Wild apples and wild pears. In: Valdes B, Heywood VH, Raimondo PM, Zohary D (eds) Proceedings of the workshops on conservation of the wild relatives of European cultivated plants. Bocconea 7: 409–416 Zohary D (1997b) Speciation patterns in predominantly selfpollinated Mediterranean annuals. Lagascalia 19:281–288 Zohary D (1999) Monophyletic vs. polyphyletic origin of the crops on which agriculture was founded in the Near East. Genet Res Crop Evol 46:133–142 Zohary D (2002) Domestication of the carob (Ceratonia siliqua L.). Israel. J Plant Sci 50(Suppl.):141–145 Zohary D (2004) Unconscious selection and the evolution of domesticated plants. Econ Bot 58:5–10 Zohary D (2009) How I first met Eibi Nevo. In: Korol A, Wasser S (eds) The evolution of Eibi Nevo (In honor of his 80th Birthday). University of Haifa, Haifa Zohary D, Ashkenazi I (1958) Different frequencies of supernumerary chromosomes in diploid populations of Dactylis glomerata in Israel. Nature 182:477–478 Zohary D, Feldman M (1962) Hybridization between amphidiploids and the evolution of polyploids in the wheat (Aegilops–Triticum) group. Evolution 16:44–61 Zohary D, Hopf M (1973) Domestication of pulses in the Old World. Science 182:887–894 Zohary D, Hopf M (1988) Domestication of plants in the Old World. Oxford University Press, Oxford Zohary D, Katznelson J (1958) Two species of subterranean clovers in Israel. Aust J Bot 6:177–182 Zohary D, Nur U (1959) Natural triploids in Dactylis glomerata L. polyploid complex and their significance for gene flow from diploid to tetraploid levels. Evolution 13:311–317 Zohary D, Plitmann U (1979) Chromosome polymorphism, hybridization and colonization in the Vicia sativa group (Fabaceae). Pl. Syst. Evol. 131:143–156 Zohary D, Spiegel-Roy P (1975) Beginning of fruit growing in the Old World. Science 187:319–327 Zohary D, Harlan JR, Vardi A (1969) The diploid progenitors of wheat and their breeding value. Euphytica 18:58–65 Zohary D, Tchernov E, Kolska Horwitz L (1998) The role of unconscious selection in the domestication of sheep and goats. J Zool Lond 245:129–135 Zohary D, Hopf M, Weiss E (2012) Domestication of plants in the Old World, 4th edn. Oxford University Press, Oxford