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Studies of the origins of agriculture in the Near East have revealed that the eight plant species known as “Founder Crops”, i.e. emmer, einkorn, barley, lentils, peas, chickpeas, bitter vetch and flax, derived from annual self-pollinating wild predecessors, were all domesticated in roughly the same period. Recent research however has prompted new d...
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Zohary and Hopf coined the term ‘founder crops’ to refer to a specific group of eight plants, namely three cereals (einkorn, emmer and barley), four legumes (lentil, pea, bitter vetch and chickpea), and a fibre/oil crop (flax), that founded early Neolithic agriculture in southwest Asia. Zohary considered these taxa as the first cultivated and domes...
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Background: Chickpea (Cicer arietinum L.) is the third most important pulse crop produced after dry bean and peas in the world. Amongst pulses, chickpea is the major crop in India but for international market the quality of this crop has to be improved which may be possible by evaluation of different planting techniques. No systematic research and...
Citations
... A geographical translocation was the case for a "new" glume wheat that was coevally cultivated at Cafer Höyük, Aşıklı Höyük, and Boncuklu. This variety was not endemic to Central Anatolia but transferred long-distance from the Upper Euphrates valley (7,55,56). Since in this part of Anatolia, early cereal cultivation went along with sheep and goat management (2,19,20), a transfer of small stock on the hoof must be taken into consideration. ...
Occupied between ~10,300 and 9300 years ago, the Pre-Pottery Neolithic site of Aşıklı Höyük in Central Anatolia went through early phases of sheep domestication. Analysis of 629 mitochondrial genomes from this and numerous sites in Anatolia, southwest Asia, Europe, and Africa produced a phylogenetic tree with excessive coalescences (nodes) around the Neolithic, a potential signature of a domestication bottleneck. This is consistent with archeological evidence of sheep management at Aşıklı Höyük which transitioned from residential stabling to open pasturing over a millennium of site occupation. However, unexpectedly, we detected high genetic diversity throughout Aşıklı Höyük’s occupation rather than a bottleneck. Instead, we detected a tenfold demographic bottleneck later in the Neolithic, which caused the fixation of mitochondrial haplogroup B in southwestern Anatolia. The mitochondrial genetic makeup that emerged was carried from the core region of early Neolithic sheep management into Europe and dominates the matrilineal diversity of both its ancient and the billion-strong modern sheep populations.
... and WesternAsia (e.g., Filipovi c et al., 2023;Kohler-Schneider, 2003;Toulemonde et al., 2015;Ulaş & Fiorentino, 2021)-as Timopheev's wheat. Today, Timopheev's wheat survives only in the Caucasus. ...
Societal Impact Statement
Agrobiodiversity is central to sustainable farming worldwide. Cultivation, conservation and reintroduction of diverse plant species, including ‘forgotten’ and ‘underutilized’ crops, contribute to global agrobiodiversity, living ecosystems and sustainable food production. Such efforts benefit from traditional and historical knowledge of crop plants' evolutionary and cultural trajectories. This review is a first attempt at systematically gauging species representativeness in studies of archaeological plant remains. Results indicate that, in addition to discipline‐specific methodological sources of bias, modern agricultural biases may replicate themselves in crop history research and influence understandings of ‘forgotten crops’. Recognizing these biases is an initial stride towards rectifying them and promoting agrobiodiversity in both research and practical applications.
Summary
So‐called ‘forgotten’ or ‘orphan’ crops are an important component of strategies aimed at preserving and promoting biodiversity. Knowledge of historical cultivation, usage, and geographic and evolutionary trajectories of plants, that is, crop history research, is important for the long‐term success of such efforts. However, research biases in the crops chosen for study may present hurdles. This review attempts to systematically identify patterns in crop species representativeness within archaeology‐based crop history research. A meta‐analysis and synthesis of archaeobotanical evidence (and lack thereof) is presented for 268 species known to have been cultivated for food prior to 1492 CE from the Mediterranean region to South Asia. We identified 39 genera with known crop plants in this geographical and historical context that are currently absent from its archaeobotanical record, constituting ‘orphan’ crops of archaeobotany. In addition, a worldwide synthesis of crop species studied using geometric morphometric, archaeogenetic and stable isotope analyses of archaeological plant remains is presented, and biases in the species represented in these disciplines are discussed. Both disciplinary methodological biases and economic agenda‐based biases affecting species representativeness in crop history research are apparent. This study also highlights the limited geographic diffusion of most crops and the potential for deeper historical perspectives on how crops become marginalized and ‘forgotten’.
... Decades have passed since the first classification of an archaeological specimen to "New Glume Wheat". It was not until numerous remains of this type of wheat were found in several Neolithic and Bronze Age archaeological sites in northern Greece and compared with other locations (Jones et al. 2000) that archaeologists were able to describe the distinctive features of this wheat (Ulaş and Fiorentino 2021). Nevertheless, identification based on grain morphology is still problematic. ...
Ancient DNA (aDNA) promises to revolutionise our understanding of crop evolution. Wheat has been a major crop for millennia and has a particularly interesting history of domestication, dispersal, and hybridisation, summarised briefly here. We review how the fledgling field of wheat archaeogenomics has already contributed to our understanding of this complex history, revealing the diversity of wheat in ancient sites, both in terms of species and genetic composition. Congruently, ancient genomics has identified introgression events from wild relatives during wheat domestication and dispersal. We discuss the analysis of degraded aDNA in the context of large, polyploid wheat genomes and how environmental effects on preservation may limit aDNA availability in wheat. Despite these challenges, wheat archaeogenomics holds great potential for answering open questions regarding the evolution of this crop, namely its domestication, the different dispersal routes of the early domestic forms and the diversity of ancient agricultural practices. Not only will this research enhance our understanding of human history, but it will also contribute valuable knowledge about ancient selective pressures and agriculture, thus aiding in addressing present and future agricultural challenges.
... We can therefore conclude from the archaeobotanical and modern wild evidence that cultivation and domestication was underway in Anatolia between the upper Euphrates Valley and the Konya plain, from which this species spread both westward to Europe and eastwards into the Zagros region. Finds in northern Iran and at Djeitun in southern Turkmenistan (Charles and Bogaard 2010;Fuller 2014;Roustaei et al. 2015); also indicate a Neolithic dispersal of timopheevii alongside emmer and einkorn in the Neolithic, while this wheat's presence in Neolithic Europe is now well established (Jones et al. 2000;Toulemonde et al 2015;Ulaş and Fiorentino (2021)) (see Fig. 13). ...
The site of Jarmo in Iraqi Kurdistan has yielded key archaeological evidence which supports its interpretation as a large PPNB village. As such, it is the perfect candidate for the study of early agriculture, plant uses, food preparation and cooking practices. In order to explore these, new excavations and intensive sampling and flotation for the recovery of archaeobotanical remains were carried out in 2012 and 2014. This study presents the results from the analysis of the newly recovered archaeobotanical assemblage from Jarmo which has provided invaluable information about early crop agriculture and plant use. Furthermore, the in-depth study of recovered remains of archaeological food by high-resolution microscopy has shed light on culinary traditions and dietary choices during the Neolithic in the Central Zagros Area.
... Remains of Triticum timopheevii found in Grotta di Pertosa could constitute a clue in this sense 89 . Indeed this is the only case for this species to be identified in southern Italy, whereas, as far as we know, its distribution range goes from Anatolia to the Balkan peninsula up to the Alpine area and Central and Eastern Europe [90][91][92][93][94][95] . Chronologically, the Mycenaean contacts start to become evident in about 1700 BCE. ...
We present direct evidence of early grape domestication in southern Italy via a multidisciplinary study of pip assemblage from one site, shedding new light on the spread of viticulture in the western Mediterranean during the Bronze Age. This consist of 55 waterlogged pips from Grotta di Pertosa, a Middle Bronze Age settlement in the south of the Italian peninsula. Direct radiocarbon dating of pips was carried out, confirming the chronological consistency of the samples with their archaeological contexts (ca. 1450–1200 BCE). The extraordinary state of conservation of the sample allowed to perform geometric morphometric (GMM) and paleogenetic analyses (aDNA) at the same time. The combination of the two methods has irrefutably shown the presence of domestic grapevines, together with wild ones, in Southern Italy during the Middle/Late Bronze Age. The results converge towards an oriental origin of the domestic grapes, most likely arriving from the Aegean area through the Mycenaeans. A parent/offspring kinship was also recognised between a domestic/wild hybrid individual and a domestic clonal group. This data point out a little known aspect of the diffusion of the first viticulture in Italy, and therefore in the western Mediterranean, which involved the hybridization between imported domestic varieties with, likely local, wild vines.
... Subsequently, through new discoveries or revision of previously ambiguous identifications of wheat remains, its presence has been confirmed at a series to this region (Jones et al. 2000;Bedoshvili et al. 2021), the ancient T. timopheevii sensu lato (Timopheev's wheat) had a much wider distribution. This is indicated by recent evidence from western Asia (Bogaard et al. 2017Ergun et al. 2018;Ulaş and Fiorentino 2021) and reviews for Europe (Kenéz et al. 2014;Toulemonde et al. 2015). ...
... It was shown that during its long cultivation history at Çatalhöyük, central Anatolia, Timopheev's wheat was undergoing domestication Roushannafas et al. 2022). Ulaş and Fiorentino (2021) suggest that, in parts of western Asia and Thrace, it was becoming adapted to local growing conditions and that, thanks to its ability to withstand cold conditions, it successfully spread across prehistoric Europe. ...
Triticum timopheevii sensu lato (‘new glume wheat’, NGW) was first recognised as a distinct prehistoric cereal crop through work on archaeobotanical finds from Neolithic and Bronze Age sites in northern Greece. This was later followed by its identification in archaeobotanical assemblages from other parts of Europe. This paper provides an overview of the currently known archaeobotanical finds of Timopheev’s wheat in southeastern and eastern Europe and observes their temporal span and spatial distribution. To date, there are 89 prehistoric sites with these finds, located in different parts of the study region and dated from the Neolithic to the very late Iron Age. Their latest recorded presence in the region is in the last centuries bce . For assemblages from the site as a whole containing at least 30 grain and/or chaff remains of Timopheev’s wheat, we take a brief look at the overall relative proportions of Triticum monococcum (einkorn), T. dicoccum (emmer) and T. timopheevii s.l. (Timopheev’s wheat), the three most common glume wheats in our study region in prehistory. We highlight several sites where the overall proportions of Timopheev’s wheat might be taken to suggest it was a minor component of a mixed crop (maslin), or an unmonitored inclusion in einkorn or emmer fields. At the same sites, however, there are also discrete contexts where this wheat is strongly predominant, pointing to its cultivation as a pure crop. We therefore emphasise the need to evaluate the relative representation of Timopheev’s wheat at the level of individual samples or contexts before making inferences on its cultivation status. We also encourage re-examination of prehistoric and historic cereal assemblages for its remains.
... One of the major questions that remains unanswered is why some crops that spread widely in the past retreated into oblivion. For example, Triticum timopheevi was a major crop in parts of Turkey before 6000 BC, spread from central Europe to Turkmenistan in the Neolithic, yet retreated in Bronze Age times, such that its historical distribution in the 20 th century was restricted to Georgia 121,122 . The expanding research on domestication through archaeology and genetics is bringing forward a new period of insights into not just domestication processes, but the long-term sustainability and resilience of agriculture and different combinations of crops. ...
Plant life defines the environments to which animals adapt and provides the basis of food webs. This was equally true for hunter-gatherer economies of ancestral humans, yet through the domestication of plants and the creation of agricultural ecologies based around them, human societies transformed vegetation and transported plant taxa into new geographical regions. These human-plant interactions ultimately co-evolved, increasing human population densities, technologies of farming, and the diversification of landraces and crop complexes. Research in archaeology on preserved plant remains (archaeobotany) and on the genomes of crops, including ancient genomes, has transformed our scientific understanding of the complex relationships between humans and plants that are entailed by domestication. Key realizations of recent research include the recognition that: the co-evolution of domesticates and cultures was protracted, the adaptations of plant populations were unintended results of human economies rather than intentional breeding, domestication took place in dozens of world regions involving different crops and cultures, and convergent evolution can be recognized among cropping types - such as among seed crops, tuber crops, and fruit trees. Seven general domestication pathways can be defined for plants. Lessons for the present-day include: the importance of diversity in the past; genetic diversity within species has the potential to erode over time, but also to be rescued through processes of integration; similarly, diversification within agricultural ecosystems has undergone processes of decline, including marginalised, lost and 'forgotten' crops, as well as processes of renewal resulting from trade and human mobility that brought varied crops and varieties together.
... The NGW of early Eurasian agriculture, grown by prehistoric farmers throughout western Asia and eastern Europe, is a cultivated member of the T. timopheevii group, as proven by ancient DNA typing (Czajkowska et al. 2020). The remains of NGW spikelet bases have been documented in two settlements in Turkey: Yenikapı (eastern Thrace) and Yumuktepe (Cilicia) (Ulaş and Fiorentino 2021). Currently, this species is held within seed banks and special collections ex situ, and is no longer cultivated as an agricultural crop (Dorofeev et al. 1979;Badaeva et al. 1994a;Czajkowska et al. 2020). ...
... Analysis of nucleotide sequence divergence in different cereal species revealed that the progenitor of T. timopheevii wheat appeared less than 0.5 million years (Huang et al. 2002;Gornicki et al. 2014). To date, four possible domestication sites have been reported about the origin of T. timopheevii in Georgia, including the Transcaucasian mountains (Jakubziner 1933(Jakubziner , 1950Nesbitt andSamuel 1996), northern Iraq (Wagenaar 1966;Nave et al. 2021), northern Syria or southern Turkey (Mori et al. 2009), and the southeastern part of the Taurus region of Turkey (Ulaş and Fiorentino 2021). ...
... Our results indicated that high genetic differentiation existed within T. araraticum and T. timopheevii populations collected from seven countries. T. timopheevii was cultivated more extensively in regions with a humid climate (average rainfall per year 900-1200 mm) at altitudes of 300-1200 m (Badaeva et al. 1994b), and adapted to the colder and wetter regions of European countries (Ulaş and Fiorentino 2021). T. araraticum was distributed in regions with a drought climate, including southeastern Turkey, western Iran, northern Iraq, and some regions of Transcaucasia: the Nakhichevan Autonomous Republic, Armenia, and northeastern Azerbaijan (Badaeva et al. 1990). ...
Genetic diversity is an important resource to improve new wheat cultivars in the breeding program. In this study, fluorescence in situ hybridization (FISH) and genotyping by sequencing (GBS) were used to evaluate the genetic diversity and population structure of the Timopheevii group (AtAtGG, 2n = 4x = 28) including 15 Triticum timopheevii Zhuk. accessions and 35 Triticum araraticum Jakubz. accessions. FISH analysis showed that there were 14 and 19 FISH signal variations in the At and G genome of T. araraticum, respectively. But there were only two signal variations in the G genome of T. timopheevii. In this study, 190,402 SNP markers were obtained from GBS, in which the lowest and highest frequencies of SNPs were found in the G and At genomes, respectively. Genetic diversity analysis of the 50 accessions indicated that the mean GD and PIC were 0.30 and 0.26, with the ranges of 0.1–0.5 and 0.1–0.4, respectively. The lowest and highest numbers of SNPs identified on the chromosomes 4G and 2At were 7748 and 17,527, respectively. Structure and cluster analyses divided 50 accessions into two subpopulations (POP1 and POP2). POP1 consisted of most T. araraticum accessions, and POP2 comprised all accessions of T. timopheevii and one T. araraticum accession AS273. In addition, the genetic variance showed that genetic variation was greater within populations (97%) than between populations (3%). The present results provided important information for the improvement of cultivated durum and hexaploid wheat production in future breeding programs of China and other countries.
... Consequently, questions about species identification, origin, spread and state of domestication of cultivated plants that previously could be addressed only through morphological and archaeobotanical studies, can today be explored directly (Schlumbaum et al. 2008). To understand plant domestication and recent plant evolution, previous researchers examined whether: (i) cereal crops were originally cultivated as common 'metapopulations' or as a core area for the origins of agriculture; (ii) whether they were associated with an early or recent domestication era; (iii) whether they originated from unknown domestication or subsequent genetic changes in an existing crop; (iv) whether they came from single or multiple domestication events or (v) whether they were still in the process of domestication (Willcox et al. 2008;Bogaard et al. 2021;Weide et al. 2021;Smith 2001;Fuller et al. 2011;Abbo et al. 2012;Blatter et al. 2002;Bilgiç et al. 2016;Castillo et al. 2016;Ciftci et al. 2019;Czajkowska et al. 2020;Ulaş and Fiorentino 2020). ...
... Recently, the aDNA study on new glume wheat (NGW), whose first use dates back to the Neolithic PPNB settlements Caferhöyük (de Moulins 1993) and Aşıklıhöyük (Ergün et al. 2018) in Anatolia and has been the subject of debate among archaeobotanists (Jones et al. 2000;Ulaş and Fiorentino 2020), has revealed that NGW wheat may belong to the Triticum timopheevii Zhuk. wheat group (Czajkowska et al. 2020). ...
... Recent archaeobotanical and aDNA reports on Neolithic wheat have revealed that several species of wheat were cultivated, in addition to the three common wheat (emmer, einkorn and free-threshing wheat). Importantly, the discovery of new glume wheat (Jones et al. 2000;Schneider and Caneppele 2009;Kenéz et al. 2014;Toulemonde et al. 2014;Czajkowska et al. 2020;Ulaş and Fiorentino 2020) has reopened discussions on the types of wheat cultivated in the Neolithic period (Ulaş and Fiorentino, 2020). In addition to new glume wheat, Hillmann suggested that T. compactum wheat was also cultivated in the PPNB Can Hasan III settlement (Hillman 1972). ...
Archaeobotanical materials subject to aDNA analysis were recovered from Yumuktepe and Yenikapı, two important archaeological sites in Anatolia and date back to the Pottery Neolithic Period i.e., 7th millennium BC. Many charred ancient seeds representing various cereal species including a great number of wheat grains were documented in mentioned sites. Among the cereal seeds, charred wheat samples were tentatively identified as Triticum aestivum subsp. spelta L. or Triticum new glume wheat (NGW) or atypical emmer or naked wheat in Yumuktepe and Yenikapı showed similarities with the morphological characteristics of T. aestivum subsp. spelta wheat, but it was difficult to reach a firm conclusion. This study aimed to provide genetic data to enable more precise identification of charred wheat seeds using an ancient DNA (aDNA) approach. aDNAs were successfully extracted from the representative charred seeds of T. aestivum subsp. spelta or NGW or atypical emmer or naked wheat. The PCR amplification of 26SrDNA and IGS gene regions with aDNA was carried out and sequenced. The expected product sizes of IGS 158 bp for the D genome and 87 bp for the A or B genomes and DNA sequence comparisons with other wheat species revealed that T. aestivum subsp. spelta or NGW or atypical emmer or naked wheat samples included the D genome from Aegilops tauschii and is more likely to be T. aestivum subsp. spelta. The discovery of T. aestivum subsp. spelta grains in the Yenikapı and Yumuktepe suggest that the cultivation of hexaploid wheat was widespread. Further, spelta hulled wheat, which is the progenitor of the hexaploid wheat, might have been cultivated in these settlements.
... Consequently, questions about species identification, origin, spread and state of domestication of cultivated plants that previously could be addressed only through morphological and archaeobotanical studies, can today be explored directly (Schlumbaum et al. 2008). To understand plant domestication and recent plant evolution, previous researchers examined whether: (i) cereal crops were originally cultivated as common 'metapopulations' or as a core area for the origins of agriculture; (ii) whether they were associated with an early or recent domestication era; (iii) whether they originated from unknown domestication or subsequent genetic changes in an existing crop; (iv) whether they came from single or multiple domestication events or (v) whether they were still in the process of domestication (Willcox et al. 2008;Bogaard et al. 2021;Weide et al. 2021;Smith 2001;Fuller et al. 2011;Abbo et al. 2012;Blatter et al. 2002;Bilgiç et al. 2016;Castillo et al. 2016;Ciftci et al. 2019;Czajkowska et al. 2020;Ulaş and Fiorentino 2020). ...
... Recently, the aDNA study on new glume wheat (NGW), whose first use dates back to the Neolithic PPNB settlements Caferhöyük (de Moulins 1993) and Aşıklıhöyük (Ergün et al. 2018) in Anatolia and has been the subject of debate among archaeobotanists (Jones et al. 2000;Ulaş and Fiorentino 2020), has revealed that NGW wheat may belong to the Triticum timopheevii Zhuk. wheat group (Czajkowska et al. 2020). ...
... Recent archaeobotanical and aDNA reports on Neolithic wheat have revealed that several species of wheat were cultivated, in addition to the three common wheat (emmer, einkorn and free-threshing wheat). Importantly, the discovery of new glume wheat (Jones et al. 2000;Schneider and Caneppele 2009;Kenéz et al. 2014;Toulemonde et al. 2014;Czajkowska et al. 2020;Ulaş and Fiorentino 2020) has reopened discussions on the types of wheat cultivated in the Neolithic period (Ulaş and Fiorentino, 2020). In addition to new glume wheat, Hillmann suggested that T. compactum wheat was also cultivated in the PPNB Can Hasan III settlement (Hillman 1972). ...
