The date and grape have a long tradition in preservation by drying

Black pepper is the most important one found in Berenike. Black peppers are either sun-dried or dried in special smoke rooms. Black pepper is very vulnerable to damage from mold and should be dried quickly and thoroughly. If peppercorns are stored in a dry environment, they may be kept for many years without losing their quality. Maybe it is for that reason that traders of Berenike chose to leave behind a large supply of black pepper safely stored in a buried dolium on the assumption that they would return within a few years.The group of fruits and nuts is well represented in the archaeobotanical record of Berenike, both qualitatively and quantitatively. The classification of raw food is mostly based on its use in food preparation, but its categorization is not unequivocal, as can be seen for food products classified as fruits and nuts. Some fruits, such as okra and tomatoes are even classified as vegetables. The matter is even more confused by the inclusion of seeds such as stone pine , pigeon pea , and chickpea within the group classified as nuts . From a biological point of view, nuts are a special group of fruits. A classification of fruits, nuts, and a single seed based on their anatomy and moisture content makes more sense with respect to their storage life and will serve here as a guideline. A group that is easily preserved concerns nut like seeds and nuts: stone pine , hazel , and sacred lotus . The first two were imported from the Mediterranean and the last one from the Nile Valley. Long-distance transport of these food products would not have been problematic as they can be stored for considerably long periods without spoilage. It is striking that very often seed coats of stone pine were found together with cone scales, drain trays for plants implying that they were transported as whole cones on a regular basis.

A reason for this might be that in this way all seeds could be transported, whereas after threshing some seeds would become damaged and unsuitable for transportation. It should be realized that import and consumption of seed contents only cannot be ruled out, but their possible share cannot be established as no traces are left. The baobab and the bentree have dry fruits that contain many seeds. Trading whole fruits of the baobab makes sense as both seeds and pulp are edible, and the indehiscent fruit does not easily shed its seeds. The fruit of the bentree is a three-valved capsule that bursts open when ripe. To prevent seed loss, it is believed that fruits were threshed to release the seeds. This kind of processing is still practiced by the Ma’aza bedouins living in the northern part of the Eastern Desert . No information on storage life is available, but it is assumed that both fruits can be stored for considerable periods. Most fruits found at Berenike are classified as drupes. This fruit type is characterized by one or more seeds enclosed by a fruit wall that consists of three different layers. A dichotomy between dry and fleshy drupes is based on the moisture content of the middle layer of the fruit: the mesocarp. The coconut , the almond , the walnut , and the fruit of the balsam tree are classified as dry fruits. Only the first three are edible, and it is the seed that is consumed. All three species are evidenced by their hard endocarps, indicating that these fruits were, at least partly, traded destoned. The presence of the hard endocarp is advantageous as it protects the seeds against decay and insects. The fibrous mesocarp of the coconut is highly valued as crude material for mats and ropes and explains the import of whole fruits to Berenike. The coconut was traded over long distances as whole fruits and maybe also as dried copra, although the presence of the latter could not be demonstrated. With or without the fibrous husks, coconuts can be kept until the liquid endosperm has completely transformed into solid “meat,” as is mentioned for example by Cosmas. For long-distance transport, fruits must therefore have been harvested in an unripe condition.

The drying process of coconuts depends on the temperature to which they are exposed, but will take at least several months. Although coconuts should be harvested at intervals of two months from a particular tree, in practice a yield of coconuts can be obtained from a coconut plantation every month. The logistics of transporting coconuts from India to Rome were, therefore, not problematic with respect to the short, fixed period during which Roman ships visited the Indian ports but may have been critical with respect to the duration of the voyages from Muziris to Rome, which would have taken some three months without any unforeseen delays. The fruits of the sugar date and doam palm are semisoft and take up a middle position in terms of preservation. The mesocarp of both species is sugary and edible. The archaeobotanical remains of both species indicate that they were transported as whole fruits. Although the seeds of both species also have an economic value, there is clear evidence for the latter species only that the seeds were extracted from the fruits. Both species are adapted to desert conditions and could be obtained from reasonable short distances. Hence their preservation might not have been problematic. It should be noticed, however, that fruits of the sugar date were partly infected by an insect that feeds on the seeds. Such infected fruits are still among the fruits offered for sale at markets today, which implies that the consumer does not discriminate in this case. Fruits having a soft mesocarp make up the largest part of those categorized as a drupe. The only fruit that could have been gathered from short distances, even from locally cultivated specimens, is nabq . Three other species originate from the Gebel Elba area or beyond: Cordia nevillii/sinensis, Grewia, and Cocculus pendulus. All others must have been brought from more-remote areas. Transported from the Mediterranean were the cherry plum , the domestic or bullace plum , the olive , the apricot , the peach , and the Egyptian plum . The last four species might also have been obtained from orchards in Egypt, for example in the Fayum. Finally, the soft fruits of the emblic had their origin in India. Although the high vitamin C content of emblic fruits has an antiscorbutic property and may have been transported in syrup, it is more likely that the dried fruits were traded.

Very perishable fruits are the apricot and the peach, ones that are only sweet and juicy if harvested in a ripe condition. Galen states that the juice and flesh of apricots rot easily and that these fruits are bad in general . He even advises against the consumption of these perishable fruits after other foods, as they will initiate rotting. A similar warning is given by Pliny , cited earlier. From his remark it can be concluded that fresh, sweet peaches were offered for sale, and people must have appreciated their sweet taste. The only way to transport such a fruit in an unspoiled condition is to add preservatives. Today, pickled peaches still consist of whole fruits that are peeled but not destoned. When apricots and peaches are dried, the stones can be removed. Dried apricots that are sold in Egypt today are available with or without stones. Those containing the stone are strongly dehydrated and have lost their characteristic color and are cheaper as a result. This kind of preservation also occurs quite easily under natural conditions after windfalls. Many well-preserved fruits can be collected under trees without any extra effort. The presence of a stone fragment in Berenike from a peach could be indicative of pickled fruit which must have been a rare item judging by the single fragment found so far. Considering the same fragment as a contamination in a load of dried peaches, it would leave open the possibility that they were available on a more regular scale. The apricots could originate from either pickled or dried specimens. The few remains indicate that the fruit probably was not commonly available. Plum stones, especially those of the cherry plum, are better represented in the archaeobotanical record of Berenike than peach and apricot and indicate that plums most probably were traded with their stones. Whole plums are easily dried but were also preserved in honey, as is mentioned by Apicius . The storage life of dried plums depends on the moisture content. Well-dried plums, having lost two-thirds of their weight, can be stored for about three years. Members of Cordia would have been preserved in a similar way. Some whole specimens were found, in addition to calyxes and huge numbers of stones, 4 x 8 grow tray and support the assumption that they were transported as dried fruits. Dried fruits were among the products offered for sale in Iran , but are probably no longer part of the assortment in Egyptian bazaars. According to Theophrastus , the inhabitants of Thebes made dried destoned fruits of the Egyptian plum into cakes. Fruits of the emblic can be preserved by drying or by pickling in a sugary syrup. Pickled fruits concern whole specimens, whereas recent samples of dried fruits that have been analyzed consist for the most part of fragments of the mesocarp. Segments of the stony endocarp were only present as a contamination. Thus, the few inedible stone fragments found in Berenike indicate a substantial import of the emblic if we assume that they were traded in a dried condition. Berries are fleshy, mostly indehiscent fruits and are represented by the date , the pomegranate , the grape , the persea , the caper bush and members of the Cucurbitaceae. The date contains a single seed; with all other berries the seed count ranges from a few to many. The less-juicy varieties of the date are easily sun-dried and have a suitable sugar-towater proportion of two to one. When they are kept for a long period, they become candied and lose some quality.

Dry varieties are known from the Sudan and are easily transported and stored. Mostly, dried dates still have their seed inside, which explains the large quantities of seeds in the trash dumps of Berenike. Seeds may also be present in a paste made from compressed, soft dates, which is called agwah . The storage life of fresh grapes is limited, but raisins and currants, being a Greek variety from Corinth, can easily be transported and stored. Whole fresh bunches stored in clay jars with chaff were also transported in classical times and they could keep for a long time. However, both jars and such fruits were vulnerable, which is why we may consider the grape more of a luxury item. Due to their sturdy leathery skin, pomegranates can be stored for several weeks despite their juicy content. It is important that carefully picked, unbruised fruits are selected for long-distance transport. Special treatments, such as dipping the fruit in tar or seawater before drying, will benefit the longevity of the fruit . Saltwater also prevents discoloration of the fruit. Fruits that are kept too long will become completely dehydrated and inedible. Transport of pomegranates by caravan trade was possible over quite long distances, as is demonstrated by the trade between Kabul and Peshawar in Afghanistan, a distance of about 250 km . No particular information is available on the preservation of the persea fruits. Judging by their less-juicy consistency and their availability within relatively short distances, it is assumed that preserving the persea would not have caused many problems. Galen mentions that fruits were transported from Persia to Egypt, and were ripe and became edible when they reached the land of destination . Although it is the unripe flower buds of caper bush that enter the commercial trade today, ripe fruits are also edible. Gathering ripe fruits should be done before they burst open. Once the pulp is exposed, the fruits are eaten by animals, most probably by birds that are attracted to the red-colored pulp. Caper fruits are juicy and easily rot when they are ripe. Galen mentions that fruits subjected to transport are sprinkled with salt as a means of preservation .

Plants engage in symbiotic associations with microorganisms that vary in their degree of association

Biological nitrogen fixation as a bacterial process in symbiosis with plants accounts for a major fraction of nitrogen input in terrestrial ecosystems . The spectrum ranges from obligate endosymbioses, such as arbuscular mycorrhizal fungi , to generally facultative nitrogen-fixing symbioses; the latter, including the rhizobia-legume nodules , Frankia-actinorhizal nodules , and cyanobacteriumGunnera associations . Nitrogen-fixing root nodule symbiosis began with one evolutionary event that occurred in the common ancestor of a clade of angiosperms known as the nitrogen- fixing clade approximately 100 million years ago . The Frankia-actinorhizal symbioses are of particular interest in the interpretation of the evolutionary underpinnings of RNS because these symbioses developed, probably independently, within several distinct lineages across the NFC . Genetically distinct subgroups within the genus Frankia form compatible associations with different lineages of plants within the NFC , while cluster IV includes atypical non-symbiotic and/or non-nitrogen-fixing strains . Of these subgroups, cluster II Frankia strains have the broadest host range among nitrogen-fixing plants: species in four different families in two orders form RNS with cluster II Frankia strains. Representatives of these families are particularly abundant in western North America, where the current study has been carried out. Comparisons of representative sequenced genomes from these subgroups suggested a general correlation of Frankia genomic features with the corresponding host bio-geographic distribution . That being said, since RNS is first established between the host-plant root and the bacterial symbiont by the two partners coming into physical contact with each other in the rhizosphere, procona valencia it is to be expected that all of the symbiotic Frankia subgroups must have at least some capability to persist in a free living state in soil, in addition to living in symbiosis.

Clusters I, III, and IV are known to have a free-living phase in soil: infective units of clusters I and III are ubiquitously distributed in soils . Moreover, infective units of cluster I Frankia can be as abundant, or more abundant, in nearby non-host rhizosphere soil than in hosts . Similarly, strains of cluster III Frankia are well known to occur in soil independent of host-plant presence . Cluster II Frankia strains, however, have been characterized as highly host dependent, with only one report of nodule formation from soil devoid of a host, 18 months after inoculation . Previous DNA-based surveys of cluster II Frankia strain distribution have been conducted using field-collected nodules . Thus, there is only limited knowledge about the soil ecology of cluster II Frankia strains in relation to their host plants, particularly concerning the degree of association with the host presence. Since the cluster II Frankia strains have been shown to be phylogenetically early divergent within the genus , a better understanding of host association in this subgroup may shed light on the process of evolutionary acquisition of RNS. The paucity of evidence for cluster II Frankia strains living independently of host plant influences compared with the other subgroups led us to hypothesize that cluster II Frankia strains may be unusually dependent on the unique environment provided by the host plant for their long-term persistence . To test this, we conducted a comparative survey to determine the soil distribution pattern of free-living cluster II Frankia strains in three geographical locations in northern Califtornia; in different soil types, including serpentine soils, a depauperate soil type that harbors several actinorhizal hosts; and under different climatic conditions. The distribution of cluster II Frankia strains was assessed through detection of DNA sequences unique to cluster II Frankia strains in host and non-host rhizosphere soils, sampled in sites with species of host and non-host plants. The microbiome of a subset of these soil samples was also analyzed via high-throughput sequencing.

Because of the seasonally arid climate and soil environment, which has been shown to limit nitrogen fixation activity in Ceanothus , nitrogen-stable isotope analysis was conducted to confirm an active state of nitrogen-fixing symbiosis in each host plant sampled. In addition, stable isotope analysis enabled us to assess the degree of site-level influence of nitrogen-fixing host plants on cluster II Frankia strain distribution.Our study has demonstrated, based both on presence/absence of glnA sequences and relative abundance of OThus in rhizosphere soil, a significant correlation between the presence of cluster II Frankia strains and the presence of host plants in a particular site, that is consistent across all three experimental locations. There was complete correlation between cluster II strains and host plant presence in two of the three locations . Even though a high proportion of host-plant-absent site samples in MNR were positive for cluster II Frankiastrains , this was nevertheless statistically significantly different from the host-plant-present site samples . This confirms and expands upon previous observations of increased abundance of cluster II Frankia strains in soil of one host-plant-present site relative to the soil of an adjacent host-plant-absent site, using a bait plant bio-assay . While the effect of host-plant presence was consistent, the following ecological factors did not correlate with cluster II Frankia strain presence/ absence across the three locations: host species present , soil type , elevation, geographic location, summer or annual precipitation, average monthly temperature, or soil pH. The consistency of the host influence is evidence that a host-plant-derived factor present in the soil of host-plant-present sites is the major determinant of cluster II Frankia distribution in these ecosystems. We found that pH was not a significant ecological factor determining presence/ absence of cluster II Frankia strains. The pH differed significantly among the three locations, but within each location, the pattern was consistent between sites, regardless of host presence or absence. Earlier studies found pH to have an influence on Frankia nodulation units in cluster I strains , but, on the other hand, both cluster I Frankia and cluster II Frankia strains have been reported to nodulate successfully in a wide range of pH between 3.5 to 8.0 and 5.0 to 9.0 , respectively. The range of soil pH in our study was between 5.0 and 7.2. Although no studies to date have examined the effect of pH directly on cluster II Frankia strain populations in soil, in culture, the optimal growing condition seems to be neutral to basic.

The cultured cluster II Frankia strain BMG5.1 was shown to grow optimally at pH between 9.0 and 10.0 , and in another study, Coriaria myrtifolia had optimal growth and nodulation at pH between 7.0 and 9.0 . It seems, therefore, that in general Frankia spp. can tolerate a range of pH while having a more specific optimum. It is possible that at a more basic pH, where cluster II Frankia spp. are known to grow optimally, there will be a difference in cluster II Frankia abundance; however, Frankia presence/absence is not affected.ANOVA of the rhizosphere microbiomes showed that while host-plant presence/absence at a given site significantly correlated with the relative abundance of cluster II Frankia spp., this correlation did not differ significantly between the rhizospheres of C. jepsonii and H. arbutifolia , when H. arbutifolia was growing in the host-plant-present site. This suggests that the effect of the host plant is not limited to its rhizosphere but can extend to the site level. A site-level influence was also observed for nitrogen fixation, determined by the ANOVA of 15N. While host plants all showed significantly more negative 15N values than the co-occurring non-host plants, flower bucket indicating active engagement in nitrogen fixation via RNS , the non-host species in host-plant-present sites also showed significantly more negative 15N values in host-plant-present sites than in the same species in host-plant-absent sites. This pattern of 14N stable isotope enrichment at the site level is similar to the facilitative effect of nitrogen-fixing plants on primary succession, reported in other nitrogen-fixing ecosystems . It was shown previously that Thissue extracts from both roots and shoots of a host plant Casuarina cunninghamiana enhanced growth of a range of Frankia spp. grown in culture but particularly enhanced the in vitro growth of the Casuarina-compatible strains . This suggests that some factor derived from the host plant, either as leaf litter or root exudate, can explain the site-level effect observed in our experiments.Our findings suggest that the abundance of cluster II Frankia spp. in a given site is associated with the strength of the influence that a host plant exerts. In the case of MNR, where the host-plant-absent site had a relatively close proximity to the nearest host plant growing in a similar vegetation type , cluster II Frankia spp. were detectable; however, no cluster II Frankia spp. were detectable in the CC site in ALCP where no host plant was proximate , despite similarities in vegetation types. At the SEF site, cluster II Frankia spp. were not detected in the wetland site despite the high proximity to the nearest host plants ; however, the wetland site had dramatically different soil conditions and vegetation type than the adjacent montane chaparral. Cluster II Frankia itself can survive in moist wetland soil: DaThisca glomerata, another host for cluster II Frankia spp. found in Califtornia, Nevada, and Baja Califtornia, grows and nodulates in seasonally flooded streambeds or washes throughout its range . Rather, it is likely that cluster II Frankia spp. were absent in the wetland site at SEF because the more xeric-adapted host plants could not grow in wetland soil, thus eliminating a host effect. The strength of the influence that the host plants may have on a given site might be affected by factors other than proximity or vegetation type. Since the cluster II Frankia enhancement factor may be able to persist in the soil over time, a legacy effect from host plants that were previously present might play a role. For example, cluster II Frankia spp. have been found in sites devoid of host plants for more than 100 years ; Elaeagnus-compatible Frankia spp. were present at a site that had not had Elaeagnus for many years , and Comptonia peregrina seed was viable after being in a pine-dominated forest seed bank for more than 70 years . Alternatively, since Ceanothus and Cercocarpus support mycorrhizal symbiosis , the mycorrhizal mycelia may be a means of conveying a chemical signal.There was low glnA sequence variability throughout this study with all samples within 0.01 substitution/site from each other; in fact, of the 74 sequences generated in this study, 56 had at least one other sequence that was identical .

Low genetic diversity of cluster II Frankia spp. based on DNA from field nodule collections, rather than from rhizosphere soil DNA, has been well documented in Oregon with repetitive sequence PCR or PCR-restriction fragment length polymorphism and in Califtornia and Nevada using single-sequence analysis . interestingly, a glnA sequence identical to one found in the current study has been detected in DNA extracted from a nodule of Dryas drummondii collected in Alaska . On the other hand, DNA from nodules collected in the southeastern United States showed a substantial divergence, based on RFLP, from that of nodules from western North America . A study of nodules collected in a circumscribed area in southern coastal Califtornia found diversity of cluster II Frankia at the species level, using rep-PCR . The sampling area in the southern coastal Califtornia study corresponds to a zone of biodiversity for the genus Ceanothus as documented previously . Taken together, these findings suggest that a suite of closely related strains of cluster II Frankia is dominant in major portions of western North America, with some localized diversity. Various factors have been suggested to account for the observed strain diversity within cluster II, however low it may be. Previously, a weak geographic pattern of strain diversity by Ceanothus collection sites was found . In reference 15, diversity was attributed to two ecological groupings of the host plants , but the sequence groupings were separated by 1%. Both of these findings reflect the generally low cluster II Frankia strain sequence diversity in western North America. Whole-genome sequencing has been carried out on strains of cluster II Frankia from Pakistan and Japan ; additional whole-genome sequencing from a greater number of sources will be valuable for assessing the degree of global geographic and genetic diversity within this clade.It has been postulated that cluster II Frankia is an obligate symbiont, based on the difficulty of isolating any strain in this group and the difficulty of detecting it in soils devoid of host plants .

Papyrological evidence of sesame in Egypt does not exist before the fourth century BC

The fruits are produced in the axils of the leaves and on the upper portion of the stem and branches where they ripen in an upward direction. Owing to the long period of fruit ripening and the fact that the capsules remain open during ripening, seed losses can be sizeable during harvesting. Strictly speaking, varieties that still have open capsules are not fully domesticated. To prevent seed losses, the plants can be uprooted before the first fruits start to scatter their seeds. The uprooted plants are shocked and left to dry, a process in which the water and minerals present in the stem allow further ripening. Drying and threshing of sesame is done at suitable spots that may be found at quite some distance from the fields before harvesting. This is still practiced in Egypt today. The color of the seeds varies from white to black. Fried seeds can be eaten in soups and, after removal of the seed coat, they are also frequently used to garnish bread and the like. From the dried ground white seeds, a paste called tehinah or halva is made. The oil, traded as oil of benne, can be kept for a long period owing to its resistance to oxidation and rancidity, subject to temperatures being rather stable. Also the high vitamin E content, high grade protein, and good taste attribute to its quality . Sesame oil is used as a salad and cooking oil. Furthermore, both seeds and oil are also used in medicine. Pliny , for example, recommends them for a variety of disorders, including the bite of a gecko and inflammation of the ears. Strabo mentions that it was a custom in Mesopotamia to anoint the body with sesame oil. The uprooted sesame stems, which are leftover after threshing, have little stock-feeding value and are used as fuel, 30 litre pots bulk as is still practiced, for example, in Egypt and Syria. In India, sesame seeds have a long tradition in religious ceremonies. Depending on the associated food products with which it is offered, it gratifies ancestral souls for one month, with for example rice and barley , and for infinity if combined with rhinoceros flesh .

Literature dealing with the origin of sesame is incomplete in its compilation of archaeobotanical records, that of Egypt being no exception. All Egyptian finds are dated from the New Kingdom onward with the exception of two records from Naqada, located some 25 km north of Luxor. One record concerns unspecified remains and is dated to the First Intermediate Period or even to predynastic times . The other record concerns pollen from either S. indicum or S. alatum Thonning and is dated to the predynastic period . S. alatum is a wild species from Sudan, but is also locally cultivated for its edible seeds, which are sometimes used as an adulterant of sesame . If both records from Naqada concern Predynastic S. indicum indeed, they predate the one from Harappa. Historical sources indicate that the Greek and the Romans also were very well acquainted with sesame. During the Greco-Roman period, sesame was especially cultivated for the consumption of the whole seeds rather than for oil production . According to the Periplus Maris Erythraei, only sesame oil could be obtained from the so-called farside ports along the Somalian coast, this oil originating from Skythia, the alluvial plain of the Indus in northwest India . Obviously, the import of sesame oil to Berenike would have been on a modest scale. The seeds found at Berenike were probably imported from the Nile Valley and used as a garnish, among other things. Of particular interest is a piece of what might be a sesame cake, found in trench 33. It consists of an indefinable matrix in which many sesame seeds are incorporated. According to Galen , such a cake was made of raw honey and sesame seeds .Sorghum is a typical domesticated grain in the African savanna belt, south of the Sahara where the wild sorghum is still widely distributed . The plant has a C4-metabolic pathway, making its photosynthesis very efficient in an environment characterized by high temperatures and intense light. In comparison with other cereals, such as barley, wheat, and rice, it has a considerable yield.

Two subspecies are distinguished: wild sorghum and domesticated sorghum . Additionally, four races of wild sorghum and five races of domesticated sorghum have been described. The start of sorghum domestication is still disputed. Some are of the opinion that sorghum is a latecomer and opt for a domestication not earlier than the late first millennium BC . Others, such as Harlan and Fuller , do not rule out the possibility of a much earlier African domestication and accept the validity of some of the records from Pakistan and northern India, dating to the first half of the second millennium BC. This would imply that soon after its domestication, sorghum reached this area through coastal trade. The oldest record of Egyptian sorghum comes from the early Neolithic site Nabta Playa and is considered as wild sorghum . Several records of sorghum are dated to the New Kingdom , without further indication on its wild or cultivated status. From several of these finds, the dates are doubted. It should be realized, however, that the tough and glossy husks of sorghum may retain a surprisingly fresh appearance, comparable with that of well-preserved rachis fragments from barley and wheat, as is the case with many specimens from Berenike. The abundant remains of cultivated sorghum that have been excavated from Berenike indicate that this cereal played a substantial role in the food supply in the city from the fifth century AD onward. There are several possibilities with regard to the origin of the sorghum found at Berenike: import from the East African coast south of the Sahara, import from the Nile Valley, and from local cultivation. Import from Ethiopia and Eritrea might have been possible via Adulis, which according to the Periplus was the nearest port south of Berenike. It was the only port of the Axsumite kingdom. Recent excavations at Axum, the capital of the kingdom and located some 150 km southwest of Adulis, have so far revealed only one single grain of Sorghum cf. bicolor . As far as the Nile Valley is concerned as an area from which sorghum was obtained, it is possible that sorghum originated from Nubia. Archaeobotanical evidence from this area is relatively rich and indicates that sorghum was cultivated here at the start of the Christian era.

Clapham and Rowley-Conwy found cultivated sorghum, race bicolor, in Qasr Ibrim from the Roman period onward. Trade contacts between Berenike and this important urban site is also evidenced from black pepper . Furthermore, most of the amphora vessels unearthed from trench BE95/96/97-5 are very similar to those recorded from Qasr Ibrim . Direct or indirect import of sorghum from India seems unlikely, as we are dealing with a cereal that predominates in late Roman Berenike and is, therefore, probably related to the presence of native populations. Moreover, the crop was available from neighboring areas and was probably also cultivated locally on a small scale. Both humans and animals may have consumed sorghum. Because the seeds do not contain gluten, it cannot be used for making bread. Mostly, the fl our is mixed with water or fat and made into a porridge. Schweinfurth mentions that camels thrived if fed on sorghum . The sorghum that is occasionally cultivated near Arab Saleh, wholesale plant containers some 10 km northwest of Berenike is also used as camel fodder. An old necklace obtained from an Ababda nomad, made of woven leather strands and decorated with glass beads and braided tassels, proved to be filled with sorghum seeds . Other specimens were stuffed with fibers or fenugreek could be the tamarind is only based on a description of the leaves and is not convincing as this description also applies to other members of the legume family . Also the archaeobotanical record is scanty. The specimens unearthed at Berenike are the first ones from the African continent. Additionally, only two archaeobotanical finds are recorded from India. They originate from Nevasa and Kolhapur, both located in the northwest coastal area of India in the vicinity of Bombay and are dated to the Satavahana period . The 66 tamarind seeds from Berenike were found in 11 different loci spread over 3 trenches. Assuming that the fruit part was consumed, the presence of its seeds indicates that the fruits of tamarind were at least partly traded without the seeds removed. Ripe fruits can be kept for a long time and are easily traded if compressed together in cakes or balls. Such tamarind balls are still offered for sale in Egyptian bazaars. According to Von Maydell , it is recommended that besides the inedible fruit wall and the fibers, the seeds also should be removed from fruits meant for storage over a long period of time, as the seeds may contain seed feeding insects that can cause heavy losses. Because the seeds themselves also have an economic value, and the production of pulp without seeds would have increased the price, it seems likely that whole fruits were brought to Berenike. The fruits might have been imported from either northeast Africa or India. Deducing from the information available, it seems most likely that they were obtained from the southern savanna of Sudan and Ethiopia, from where they might have been shipped to Berenike via Adulis.Ajowan occurs in south Europe, northeast Africa, and west and central Asia, but it is only recorded as a cultivated plant or as one that has strayed from cultivation. Archaeobotanical remains could be helpful in identifying its area of origin, however, records are scarce, probably due to the small size of the otherwise characteristic fruits. The only sub-fossil remains of ajowan that are known so far originate from Egypt. Besides the fruit from Berenike, it has been recorded from Mons Claudianus in the Eastern Desert , Amarna in the lower Nile Valley , and there is a sample of unknown origin and date stored at the Florence Museum .

The find from Mons Claudianus is contemporary with the one from Berenike, whereas the fruits from Amarna are dated to the Eighteenth Dynasty . Although the find of ajowan at Amarna suggests that the plant probably already had a long tradition in Egyptian horticulture, it remains unclear whether the plant once belonged to the native flora. The many synonyms that have been used for this plant indicate its close resemblance with several other umbelliferous plant species, such as khella , carrot , caraway , black caraway , and stone parsley . Most of these plants, including ajowan, are still used for flavoring food. Among others, Dalby assumes that ajowan is already mentioned by classical writers such as Pliny, Galen, and Dioscorides. But the descriptions of the concerning plant mainly deal with its uses, which are quite universal, and do not mention diagnostic features specific to ajowan. Nevertheless, we cannot know for sure if ajowan was meant indeed. Ajowan is mainly cultivated for its fruits, which are used both as a flavoring spice for food and as a medicine, for example, in the treatment of kidney calculi and dysentery. Its medical properties can be attributed to the presence of thymol, a powerful antiseptic that can be used both internally and externally . Several plant species are exploited as a source of thymol, but ajowan is the most favored one as it yields the highest percentage of this valued compound. Today, however, thymol can be prepared synthetically, which decreasing the demand for natural sources. Ajowan could have been imported from the Nile Valley, but it is also possible that it was grown in local kitchen gardens at Berenike. Although it takes 5 to 6 months before the fruits are ripe, they can be harvested earlier to prevent loss due to seed dispersal, without a reduction in quantity of the fragrant oil .The wild relatives of fenugreek grow in the Mediterranean area and in the Near East, and it is assumed that this pulse crop had been cultivated somewhere within this area .

It is believed that its extraction is only successful if no other person is present

Ben oil does not become rancid, and, due to its high melting point, it is suitable for use in a hot climate. From both Ababda and Bisharin nomads, it has been recorded that they extract the oil from the seeds that they use for cooking. To extract the oil, seeds are boiled using wood from the same tree. The oil is skimmed off from the surface. In ancient Egypt, ben oil was used for cosmetics and cooking. In modern times, the high-quality oil has also been in demand for lubricating precision instruments. Classical writers use the word myrobalan in connection with oil-producing plants in particular, but also with tannin-producing plants . As myrobalanum literally means “perfume-nut,” its original use makes more sense. According to Orta, the name myrobalan has been introduced for the tannin-producing plants by Serapio in translating Avicenna work from Arabic into Latin. It is difficult to determine the real identity of the bentree in classical sources. In Pliny’s Historia Naturalis , for example, its mere enumeration in the ingredients of perfumes and unguents provides us with little grip on the subject. W. H. S. Jones, who compiled an index of plants mentioned in Pliny’s Natural History, is probably right in his suggestion that balanos, normally a term for an acorn, is most probably a word used for the sugar date in sections 12.121, 13.8, 13.12, 13.13, and 13.15. Translating balani olea as “behen oil” instead of “balanos oil,” however, confuses the issue,plants in pots ideas as behen oil seems to refer to oil obtained from the bentree . In only one of the sections dealing with perfumes and unguents, namely, 13.18, Pliny uses the words constat myrobalano , and it is suggested by W. H. S. Jones that here the bentree is meant indeed.

It is in this particular section that the regal ointment for the Parthian kings is described and obviously the more-esteemed oil from the bentree was in demand. A description of the myrobalanum is given by Pliny in sections 12.100–103 of his Natural History. It is stated that the tree grows in the country of Trogodytis, the Thebaidi, and parts of Arabia that separate Judea from Egypt. Although this area roughly coincides with the present distribution of the bentree and excludes another species that is cultivated for its oily seeds , it is anything but clear that the bentree is meant indeed. The description does not include recognizable features and it is very likely that information of different species have become mixed up. The same is true for section 23.98, which is even more confusing, as the myrobalanum is connected with a palm tree, which excludes all species known by the name of myrobalan, namely, bentree , emblic , myrobalans from Terminalia spp. , beleric myrobalan and citrine myrobalan , and myrobalan plum . According to Hobbs , the Ma’aza bedouins still collect the benseeds and sell them to middlemen who, in turn, sell them to druggists in Cairo. Hobbs mentions that two Ma’aza bedouins collected 150 kg of benseeds in 17 days. Over exploitation of bentrees during the drought of the 1950s probably reduced the number of these trees substantially. The combined harvest of benseeds in 1983 was estimated at about 2,000 kg . Although dead wood from bentrees is used, among some other woody species, for making charcoal to prepare tea and meals, Zahran and Willis are of the opinion that the tree is not endangered by this practice. Drar , however, states that the tree had to be protected because it was heavily exploited for the feeding of cattle. Thus far, there are only very few archaeobotanical records of the bentree and all are confined to Egypt. The oldest seed, which is identified as Moringa cf. peregrina by de Vartavan and Amorós originates from the tomb of Tutankhamen.

The only other Roman record is from Hawara . The current distribution in Egypt suggests that the benseeds from Berenike and Shenshef might have originated either from the northern part of the Eastern Desert, which is currently the territory of the Ma’aza bedouins, or from the Gebel Elba, which is now inhabited by the Bisharin nomads.The genus Nelumbo is closely related to Nymphaea, and, although sometimes even classified within the Nymphaeaceae, it is distinctive by, for example, the form and structure of the rhizome, the arrangement of the leaves, and its large fruits and typically shaped receptacle. Flowers and leaves of this water plant are long-stalked and rise above the water surface. About 20 to 30 fruits develop in the flat apacial area of the receptacle, which becomes swollen and spongy and functions as a dispersal unit as it separates from the stalk when the fruits are ripe. The fruits are only released when the floating receptacle gradually disintegrates. The brown fruit coat encloses a large seed, which consists of two cream-colored, sweet-tasting cotyledons enclosing a green, bitter-tasting embryo in a central cavity. The genus Nymphaea is represented by two species in Egypt: N. lotus L. and N. caerulea Savigny . Most probably, both these species have to be considered as the sacred lotus of ancient Egypt and were used as the symbol of the Kingdom of Upper Egypt. The papyrus reed , on the other hand, represented the Kingdom of Lower Egypt. Papyrus is indigenous to tropical central Africa at the upper parts of the Bahr el Abiad , from where it penetrates westward into Lake Chad and the Niger region . In Egypt, papyrus reed was restricted to the Nile Delta, where it grew in huge populations, as is for example mentioned by Strabo . The genus Nelumbo is the only genus of the Nelumbonaceae and is represented by two species that have a discontinuous distribution in the tropical zone. Nelumbo lutea Willd is found in North America and Mesoamerica, whereas the other species, the sacred lotus, is confined to Southeast Asia and northeast Australia. The sacred lotus is not indigenous to Egypt . Nevertheless, historical sources as well as archaeobotanical remains indicate that during the Greek-Roman time the sacred lotus was probably cultivated in Egypt.

Herodotus, who visited Egypt as far south as Aswan, is the first one who unmistakably describes both Nymphaea sp. and Nelumbo nucifera in his section dealing with dwellers of the Egyptian marshes . Both plants are called “lilies” by Herodotus, but he also mentions that the Egyptians use the name “lotus” for Nymphaea. Theophrastus also describes both plants in his section dealing with plants of rivers, marshes, and lakes, especially in relation to Egypt . Theophrastus did not visit Egypt, and it is beyond doubt that his description is partly based on Herodotus. According to Theophrastus, the sacred lotus grows in marshes and lakes and is even planted in these suitable habitats. That Egypt is mentioned, indeed, is supported by using the name “Egyptian bean” for its fruits. Other sources, such as Strabo, who visited Egypt in 24–20 BC, Pliny, Columella, and Dioscorides are less clear in their descriptions, and it seems as if both plant genera got mixed up or were probably even confused with other species. The complete absence of depictions of the characteristic leaves and flowers of the sacred lotus from the pharaonic period indicates that the plant was most probably introduced into Egypt during the first Persian period , as is suggested by Schweinfurth . This is supported by the archaeobotanical remains from Egypt,container size for blueberries which seem to be confined to the Greco-Roman period. Some earlier finds are doubted because of their uncertain identification or possible contamination with more recent material . The only archaeobotanical records from the sacred lotus outside Egypt originate from Cyprus, from where it is recorded from the fourth century BC . Both seeds and rhizomes are edible. The roots are dried and made into fl our from which bread can be baked. The seeds are eaten raw or cooked. Because the embryo is bitter, a feature already mentioned by Theophrastus , it has to be removed before it can be consumed. Today, seeds from the sacred lotus that are exported from China are divided into halves and have their bitter embryos removed. Ripe seeds are very hard, and a strong tool is required to break them. Additionally, rhizomes and fruits are also used as medicines. Today, dried receptacles are used in bouquets of dried flowers. Two whole seeds of the sacred lotus were found in the trash dump. The presence of whole seeds in Berenike implies that its residents knew what to eat. It is quite likely that during the Roman period the sacred lotus was still cultivated in Egypt. The most plausible center of origin of the fruits found in Berenike is therefore the Nile Valley or the Delta, but import from India cannot be ruled out.Some classical sources mention the local cultivation of the olive tree in Egypt. Theophrastus states that the olive was common in Thebaid and according to Strabo , the olive only grew in the Arsinoite Nome and in gardens near Alexandria. Elsewhere, Strabo , in referring to the mangrove tree along the coast of the Red Sea, mentions its resemblance to the olive tree and the laurel tree . Strabo would have also meant the mangrove tree in his description of the entrance of the harbor of Myos Hormos and of the coastal region of Cape Deirê in Djibouti , despite the translation as “olive trees.” This interpretation is supported by Strabo’s description of the Fayum , in which it is explicitly mentioned that the presence of olives in Egypt is restricted to this fertile area and the gardens near Alexandria. Most probably, the cultivation of olives in Upper Egypt, as mentioned by Theophrastus some centuries before, had been discontinued by then. The wild ancestor of the olive, now treated at the level of subspecies or variety , grows in the coastal areas of the Mediterranean Sea, but is not recorded from Egypt . The Sudanian flora, however, includes a wild olive, which is recorded from the Sudanian territory in, for example, Yemen and the Gebel Elba area, some 180 km south of Berenike. This species has a high water requirement and in both areas it is, therefore, limited to the higher zones of the mountains.

The taxonomic status of this wild olive is still disputed. It is treated both as a separate species and as a subspecies of the cultivated olive . It is not quite clear from the Periplus Maris Erythraei whether unripe olives from Egypt were exported via Berenike. The Greek word ὄμφακος, which is mentioned in the Periplus means either unripe grape or olive. Fabricius translated the passage in the Periplus as grapes: “der Saft herber diospolitischer Weintrauben” . Casson chose unripe olives, following the argument of Giangrande, who assumes that the climate of the area rules out the transportation of grapes. This argument is, however, not convincing. Even if unripe olives and grapes were harvested with immature endocarps and seeds, the presence of a reasonable numbers of mature endocarps and seeds of the olive and grape from Berenike and Shenshef still indicates that transport of both fruits was possible. In fact, the Periplus itself supports the possibility of grapevine by mentioning Diospolis, present-day Thebes, as the place from which the unripe ὄμφακος came. Diospolis was located about 40 km south of Koptos , one of the Nile termini of the Berenike–Nile roads. This close proximity to the trade route would have facilitated the supply of grapes to Berenike. Olives have an oil content varying from 14 to 25 percent and are used both as table olives and for oil production. Olives contain a bitter substance that is extracted by soaking in lye, water, or a salty brine. Sweet-tasting olives are recorded from the Gebel Elba and parts of southwestern Saudi Arabia . Full-grown but still unripe olives yield the green variety, whereas black olives are obtained from the ripe fruits. Olives were served as a whole fruit; as crushed fruit, known as columbades; or as stoned and chopped fruits, known as epityrum . If used for oil production, ripe olives are dried before crushing. Because both the fleshy mesocarp and the seed inside the stony endocarp contain a similar kind of oil, dried fruits are crushed without being first destoned.

Coconut oil was regularly used for oiling the ropes used for sewing together the planks of ships

The correct name for the liquid endosperm is “coconut water.” It is also called “coconut milk,” but this name concerns a derived product made from a mixture of grated endosperm and milk. The solid edible layer is often referred to as “meat” or “flesh,” words that wrongly suggest animal origin. The fibrous, buoyant mesocarp takes up the largest part of the coconut fruit and is adapted to dispersal by water. It yields coir, a fiber that is used for making ropes, carpets, and mattresses. Among others, the coir is made into rigging and cordage for ships and it is also used for caulking. Coconut has the advantage of not rotting in saltwater. Coconuts can be harvested either by cutting them from the tree or by collecting fallen fruits. Both methods can be applied for the production of copra, although the former has the advantage of removing dead leaves and detecting damage by pests. Because ripe nuts will hang on the tree for another two or three months before falling, the amount of coconut water will be too low for long-distance trade. Therefore, harvesting the nuts by cutting is necessary if they are meant for export. Mostly, coconuts are husked so that the fibrous mesocarp and exocarp is separated from the inner part of the fruit, consisting of the endocarp and the seed . Further processing concerns the drying of the seed into copra. The hard “shells” are split and the “meat” is dried until the moisture content is reduced to about 6 percent. During this process, the “meat” becomes loose and can be separated from the “shell,” raspberry container size which is more effective for drying. If the copra is properly dried, it can be stored almost indefinitely. Coconut oil can be obtained from the copra. Today, the Ababda nomads occasionally use coconuts as containers .

Such containers are called garras or lozes and are mostly made from gourds . They are used for storing meat, animal fat, milk, and rancid butter used as a kind of hair gel. The rancid butter is mixed with additives that enhance the smell. Such containers may be decorated with leather, textile, basketry shells, and beads. The whole-coconut lozes are obtained by removing the “meat,” when it has rotted, through a hole bored through one of the eyes. The small hole is often closed with a stopper made by rolling up a piece of leather adjusted to the size of the hole.The origin of the coconut is still unknown. The possible area of origin is Southeast Asia and Melanesia, whereas India is considered as a possible secondary center of origin . The present distribution comprises the wet tropical lowlands, and the present-day coconut cultivation lies between lat 20° N and 20° S, although the noncommercial growth may extend as far as 27° N and 27° S . There is still scholarly disagreement concerning the introduction of the coconut to East Africa, both with respect to period and dispersal agent. Its spread from India to East Africa by oceanic currents, preceding the distribution by humans, is only based on assumptions, although crossing the Indian Ocean by floating seems to be possible. An early introduction by humans, on the other hand, is plausible, based on the existence of ancient trade contacts between both continents. A reference made by Schuiling and Harries to the Periplus Maris Erythraei is not tenable because it is based on a mistranslation by Schoff of the Greek word nauplios [ναύπλιος]. Egypt fringes the northern distribution line of the coconut. According to Täckholm and Drar , all attempts of cultivating the coconut in Egypt were futile because of unsuitable climatic conditions. The cultivation of coconut palms with the aid of irrigation on the island of Elephantine, near Aswân, is mentioned by Walter .

A striking record concerns the presence of a coconut palm growing near Berenike published by D’Abaza . An English naval officer, who had spent many years in the tropics and must have known the tree very well, made the observation, according to D’Abaza. The observation was confirmed in 1907 by the captain of one of the Khedivial mail boats. Distinguishing the coconut palm from the date palm is, however, quite difficult if no ripe fruits are present. Both trees have a similar shape with scattered, pinnate leaves. Another problem may be the identification of the location, The archaeobotanical record from the Roman Empire is extremely scanty. Excavations at Arikamedu on the southeast coast of India, which cover the period of about 300 BC–AD 200 and later, have yielded remains of the coconut . The presence of all kinds of Roman period remains that have been unearthed attest contact with the West. According to Casson , it could well be that a colony of Westerners resided here who were chiefly engaged in forwarding goods not all the way to Egypt but only to intermediate stations at Muziris and Nelkynda on the southwestern coast of India. Both these sites have not been excavated yet, so nothing can be said about the possible presence of the coconut palm or other plant species there. Anyway, the record of coconut from Arikamedu makes it very likely that the Westerners, putatively present in Arikamedu from the early years of the first century AD until ca. AD 200, would have known it. Unfortunately, no botanical research was carried out in Arikamedu so that only species are recorded that could not have been overlooked: the coconut palm and the palmyra palm . Further botanical evidence of the connection between Berenike and this site, therefore, remains unknown. With the exception of a few husk segments from trench BE99-33, all fragments of coconut unearthed in Berenike concern endocarp fragments. The unbalanced representation of different kinds of fruit fragments in the sub-fossil record suggests that most coconut fruits were traded as dehusked, whole fruits. It is also possible that part of the coconut had been traded as copra, but the chance of recovering such pieces is small, as no detectable waste is produced and recovery relies on discarded pieces.

Unopened coconuts can be kept for consumption until all the liquid endosperm has disappeared. As soon as the coconut is opened, the production of free fatty acids starts and consumption cannot be delayed too long. To exclude the possibility of contamination, a radiocarbon date has been obtained from one fragment, originating from BE96-13.002 and giving 1935 ± 35 BP . The calibrated date coincides with the first two centuries AD. No attempt has been made to convert coconut fragments to whole specimens. Taking into account the scattered distribution of these fragments in trash deposits and the estimated expected finds in the unexcavated area, we can deduce that the coconut was not a rare commodity in either early or late habitation periods. This interpretation is supported by the fact that most coconut fragments were not worked after consumption of its “meat.” So far, only five worked fragments have been unearthed. Two of them, which have holes measuring 7.5 and 13 mm, were found in the same trench . It is possible that they were used for making beads or buttons. In trench BE99-21, a fragment was found with a carved decoration showing a pattern of lines at a right angle. Trench BE99-33 yielded two worked fragments that looked like cups with a diameter of approximately 6.5 cm. One of these fragments still had some resin present along part of its rim. Actually, this low frequency of worked fragments of such an exotic fruit,raspberry plant container having an attractive size and a hard endocarp, making it suitable for fashioning into all kinds of decorative and useful articles, is striking. Even so, it is to be expected that future excavations at Berenike or any other Roman settlement will unearth worked fragments of this nut. As far as the Indian subcontinent is concerned, it is very likely that the coconuts from Berenike would have been imported from ports along the Malabar coast. The most productive area of coconuts in India is confined to its interior and the present name of this state is Kerala, which literally means “the land of the coconut.” The abundance of coconut palms is, for example, expressed in the description of the landscape of Tyndis in a Tamil poem cited by de Romanis , saying that “In its immense fields grow coconut palms laden with fruit, . . .” Due to the activity of pirates, Tyndis was abandoned, as the people moved to the more southerly Muziris and Nelkynda. The cultivation of the coconut palm at isolated spots along the strait of Bab el-Mandeb and the Gulf of Aden, as recorded from the last centuries, leaves open the possibility of a shorter trade route for the coconut. If so, coconuts might have been obtained from Muza or Kanê. Today, this area falls outside the main coconut producing areas and it is, therefore, merely speculative to assume this was an alternative trade route. Coconut fragments belonging to several specimens were also found in Quseir al-Qadim, though dated to the Mamluk period . The only other reference of coconut concerns an Egyptian specimen of unknown age and origin, kept at the Florence museum.

Initially, E. Bonnet did not preclude the possibility of a Greco-Roman date, suggesting that the coconut was merely a curiosity from India . Later, however, he expresses doubt about its ancient date. The coconut is mentioned for the first time on a list that presents all the objects that were collected by Dr. Alessandro Ricci during his travels in Egypt in 1818 and 1822. This list was compiled by M. Arcangelo Migliarini in 1932 on the occasion of the acquisition of the collection by the archduke of Toscane. The coconut is described in the last part of this catalogue as “Un frutto di Cocco. Una mezza scorza di un piccolo cocco” . Because this last part of the list deals with Ricci’s anthropological collection, it is assumed that it is of recent date, despite the fact that a contemporary is of the opinion that it is an old coconut indeed . The reliability of Italian inventories from the nineteenth century and the fact that Riccipays no special attention to the coconut argues for a recent date. But the curious thing is that an ordinary fragment of a coconut is considered as an anthropological object. A definite answer concerning its age, therefore, can only be obtained by radiocarbon dating. It has frequently been suggested that coconut oil was present in Egyptian graves. According to Lucas and Harris , however, this assumption is wrong, because the strong smell is due to a very small proportion of nonoic acid, which had been formed as a result of decomposition of fatty materials.Among the many beads that were found at Berenike, four specimens could be identified as Job’s tears. This bead is made of the false fruit of Job’s tears, a grass species that is closely related to maize . In maize, male and female flowers are arranged in separate inflorescences, in which each female inflorescence consists of a number of fertile flowers and is enclosed by a series of almost bladeless leaves. In Job’s tears, however, each inflorescence bears both female and male flowers, and only the lower fertile flower of each inflorescence is enclosed by a tough and modified sheathing bract or involucre. The slender axis of the inflorescence that bears the male flowers protrudes through the somewhat oblique opening of the callous bract. Once the fruit is detached from the stem, Job’s tears is already effectively a bead, as the basal area has an opening as well as the top. Moreover, the grain is folded lengthwise around the axis, creating a natural groove that facilitates its piercing. The specimen from Berenike has a hard shell, which is indicative of the wild species that is used for bead making. The decorative fruits are mostly tear-shaped to globular in shape, have a white to brown, shiny coating and are 6 to 12 mm long, making them applicable to all kinds of bead work. Although the glossy surface of the Roman specimen from Berenike had disappeared, the shape, the basal perforation, and the characteristic orifice at the top of the fruit justify the identification.

A further complication results from the incorporation of vegetative parts in the effective dispersal unit

Even when all the water has been absorbed by the soil, which may take a couple of weeks or even months, depending on the amount of water and the soil structure, such play as are still easily recognized because their surfaces are sealed with silt, which bursts open after drying. When all the water has been absorbed, small holes are made in the soil, which are then filled with water. The next day, several seeds of sorghum are put in each hole. Only during the first week, extra water supply is given if necessary. From then on, the farmer only does the weeding . The plants can be harvested after six months and are used as forage for camels. One would assume that the cultivation of cereals such as sorghum and barley is only meaningful if fair sized fields are sown. Evidently, present cultivation of cereals in the Eastern Desert is mostly practiced on a small scale and has an opportunistic character. Sorghumis a crop domesticated in Africa and has a C4-metabolic pathway making its photosynthesis very efficient in an environment characterized by high temperatures and light intensities. Being C3-crops, barley and wheat lack this photosynthetic adaptation to arid conditions and, as a consequence, irrigation is necessary. Long-term irrigation is a disadvantage as the soil may become salty, a condition to which only barley has a considerable tolerance, as far as cereals are concerned. The reason that barley is the preferred crop in drier environments but still suitable for agriculture is that it has a relatively short life cycle. In this way, it succeeds in avoiding water stress. Additionally,seedling grow rack its roots are well developed in the upper soil layer, enabling the plant to benefit from rain showers and morning dew.

Besides problems dealing with water and thermal stress, cereals also have to survive diseases and predation. Crops may be affected by flying animals and microbial diseases such as smut. The leaves of sorghum plants near Arab Saleh, for example, were partly damaged by caterpillars. A fungal disease that was found on sub-fossil remains of barley will be discussed in Chapter 4. Although the grains of sorghum are mostly used as animal fodder—the absence of gluten making it is unsuitable for bread making—it can only be fed in a fresh condition to animals after seed ripening. The presence of the poisonous dhurrin, a cyanogenic glucoside, in the leaves during the growing season and also after drying , makes it necessary to fence off the fields and excludes the possibility of hay making. The dried culms can be used as fuel or as building material. The competition for water in an arid environment results in relatively large distances between plants, so that the spatial distribution of the roots of each plant ensures the uptake of sufficient amounts of water. The relatively large distance between plants is apparent in sorghum fields and kitchen gardens. To reduce water competition, special attention has to be paid to weed control. Nevertheless, in most kitchen gardens, desert plants with a weedy character, such as Sonchus sp., Zygophyllum simplex, and Caylusea hexagyna, were found. In the sorghum field south of Arab Saleh, the following weed plants were recorded: Forsskålea tanacissima, Farsetia ramosissima, Aristida funiculata, A. adscensionis, Tribulus terrestris, Dichanthium foveolatum, Astragalus eremophilus, A. vogelii, Trichodesma ehrenbergii, Zilla spinosa, Acacia tortilis, and Morettia sp. All these species are desert plants and are also found outside the cultivated areas.With respect to the exploitation of plants by man, two terms need to be addressed: cultivation and domestication.

Cultivation concerns the exploitation of plants in its broadest sense, including the harvest of useful products of wild growing plants. Domestication, on the other hand, is restricted to a human-induced process in which plants become dependent on humans for their reproduction . It follows logically from these definitions that domestication implies cultivation. Domesticated plants are mostly recognizable, as such, in an archaeobotanical context, since generative parts, especially, have undergone morphological changes. It may be difficult however to pronounce upon the cultivated status of wild plant species that are considered to be part of the surrounding vegetation in former times. As far as edible plants are concerned, the presence of leftovers unequivocally indicates a cultivated status. But more often, remains of wild plant species are found that do not show clear evidence of a special use. Sometimes, a specific use is known from present-day dwellers, eventually from a more remote area, but mostly no traces of such a use are detectable. In such cases the plant might have been deliberately collected by man but may also have been transported to the site by natural dispersal agents. In the Eastern Desert, both strong winds and irregular occurring torrents are responsible for the transport of plant remains over long distances. Obstacles such as buildings and depressions are excellent seed traps. This process was continuously evidenced during the excavation seasons, where a trench sweep is a daily procedure, and old trenches are filled with sand intermixed with all kinds of plant remains, including tumbleweeds and plants that germinate in such protected safe sites. It may thus be clear that the status of some wild plant species that have been found at Berenike and Shenshef is still obscure.

Some of them have been categorized as cultivated ones, such as beet and Johnson grass , although conclusive evidence is lacking for such a labeling. On the other hand, the potential use of several wild plant species will be mentioned in the next section to compensate for the assumption that they were dispersed by natural dispersal agents. It is assumed that most desert plants with an economic value will have been exploited without intentional planting in gardens or plantations. For this reason, all cultivated plants that grow wild in the Eastern Desert, the Red Sea coastal area, and/or the Gebel Elba area will also be dealt with in the next section. This section presents all cultivated plants from Berenike and Shenshef that have been evidenced by at least non-woody remains. Plants evidenced by wood identification are described by Vermeeren . Some of them will be discussed briefly in the summarizing section. The cultivated plants are described separately in alphabetical order. Each description starts with the scientific name and, as the occasion arises, relevant synonyms are also mentioned. Synonyms that are linked to wild progenitors, such as Olea oleaster Hoffm. & Link. and Vitis sylvestris C.C. Gmel, have been omitted. Next, English, Arabic, and Indian names are presented alphabetically, followed by a description of the plant parts, the trenches in which the plant has been evidenced, the period, the state of preservation, and a reference to a picture of the plant remain. The scientific plant names are mainly based on Wiersema and León . Tentative identifications on a species level are indicated with “cf.” . For example, Monsonia cf. nivea means that only the identification on the genus level is certain, but that the identification to the level of species is only tentative. The common names were obtained from a variety of sources, in which the preferred common English plant name follows Wiersema and León . The Arabic and Indian names are taken mainly from Bedevian , Kamal , Kay , Mandaville , and Dey . These common names refer to whole plants as well as particular plant parts of economic value. All variant spellings have been listed that were found in literature. Doubtless this compilation is by no means complete. The common names are arranged alphabetically and are also indexed. It should be realized that the same name may be used for several plant species, including non-cultivated ones. For each plant, the official names of the recovered remains are mentioned. Sometimes such labels deviate from the names that are commonly used. Such established names may, however, be incorrect and confusing, which is caused by several factors. The main objective of using the official botanical names is that it facilitates an unequivocal description. This, in turn, enables an interpretation with respect to the processing of the plant remains,greenhouse growing racks including preservation techniques that might have been used and a comparison with present-day samples of these plants. To avoid an unreadable text, the botanical names are explained when necessary and eventually also linked to more-popular terms. For the same reason, specific names of fruit types, such as pods and legumes, have been omitted in the description of the botanical remains. First of all, the labeling of “seed” and “fruit” is not always correct.

Strictly speaking, an ovule develops into a seed and an ovary develops into a fruit. This means that a seed is always enclosed by a fruit. Unfortunately, seeds and fruits are not always clearly separated, as for example, the members of the mustard family and the legume family . If only one seed is produced in a fruit, very often the fruit coat is fused with the seed coat. What appears to be a seed is in fact a fruit. Such fruits are produced, for example, by members of the buttercup family and the daisy family .A second factor that obscures the issue is the great variety of fruit types. Thus, the so-called diaspore may either be a seed, a fruit, or even a false fruit. The fig may serve here as an illustrative example. Very often, sub-fossil records mention that seeds or fruits of the fig have been retrieved, but the official labels should be “fruits” and “accessory fruits,” respectively. If the description is unequivocal, for instance in the case of dried fruits, it is not problematic. Unfortunately, this is mostly not the case, as can be deduced, for example, from the codex of ancient Egyptian plant remains by de Vartavan and Amorós , which lacks a sound botanical standardization. Another complication arises from the taphonomic processes and preservation conditions, both resulting in the fragmentation of botanical remains. A correct identification of such fragments should ideally be based on the anatomy of the diaspore. But again, popular names, partly derived from the description of fruit types, are often used instead. Examples of such names are shell, flesh, meat, pip, and kernel. The word stone is used, for example, both for the seed of a date palm fruit and for the endocarp of fruits, which are classified as “drupes,” such as the discarded inner part of an olive fruit. Standardization of descriptions for the archaeobotanical remains of cereals Triticum and Hordeum, including threshing remains, has recently been recommended . Trench numbers, recorded in a consecutive series throughout the whole excavation period, indicate the origin of the botanical remains from Berenike. In Shenshef, on the other hand, the numbering of trenches starts at one in both seasons. A complicating factor in the numbering is that a particular midden has been excavated in three different trenches . Therefore, the numbers of the middens from Shenshef have been used instead to designate the origin of their botanical remains. Only in particular cases, where a more detailed description is considered as informative, are complete locus identifications given. The locations of the trenches and middens are indicated in Figure 4.3 and Figure 4.4. Plant remains can be preserved in different ways: waterlogging, mineralization, desiccation, and charring. Additionally, imprints in pottery, for example, may indirectly maintain the presence of a species in the course of time. The type of preservation depends on environmental conditions and human activity. The plant remains from Berenike and Shenshef are preserved by desiccation and charring. Owing to the extreme arid conditions, plant remains can be preserved by desiccation without special treatments. Basically, this nondestructive mode of preservation is valid for all kinds of plant remains. Differences in fragility, however, allow for bias in favor of the more-solid remains such as seeds and fruits. Since these generative plant parts are, by nature, adapted to survival in the soil, they offer resistance to both mechanical hazards and decay by microorganisms. This kind of preservation is sometimes erroneously mentioned as “mummification,” a qualification that is, however, only applicable to organic subjects that have been treated in a special way to improve preservation. Even plant remains that have been found in tombs and are stored as offerings for the dead do not deserve this special qualification. Charring, on the other hand, is mostly human induced and destructive in character .

The absence of soil and plants results in a quick surface runoff toward the wadis

In the Ma’aza territory, north of the road between Quseir and Qift, such wet years are reported, among others, 1928–1932, 1949–1951, 1956–1958, and 1977–1985. Seven cloudbursts are recorded by Springuel over the past 15 years in the southern part of the Eastern Desert. The most recent one was in November 1996 and fell over much of the Western and Eastern deserts. However, winter rains mostly result in limited amounts of rainfall. Because the main rain-producing winds come from a northeasterly direction, rainfall is concentrated on the northeastern sides of the mountains. This orographic influence of the Red Sea coastal mountains results in a comparatively well-developed flora on the slopes. The Elba area in particular, located some 260 km south of Berenike, catches much water in this way, and a unique Sudanese flora characterizes its slopes. Most mountain slopes north of the Gebel Elba are, however, almost barren. Especially during nighttime, the evaporation is minimal, and much water sinks into the coarse wadi sand. Some water is caught in rock depressions and is used by animals and humans as a source of drinking water. Even if such rocky clefts are difficult to inspect by eye, the presence of water can easily be checked by throwing a piece of stone. For the Eastern Desert it should thus be realized that the almost-barren orographic relief affects principally the redistribution of rainfall. As a result, the water supply of wadi branches is actually much more than what is calculated on the basis of the mean annual precipitation. This concentration of water is also supported by the presence of impervious surface crusts,blueberry plant pot which are found in hyperarid regions in particular and are virtually water resistant .

Water that penetrates deep into the wadi escapes evaporation and will be available for plants with deep taproots. The amount of water that can sink into the ground will largely depend on the texture of the soil, in which penetration and the coarseness of the soil texture are positively correlated. For this reason, sandy soils offer better water supplies in dry regions than clay soils . Silty depressions are an exceptional case as the concentration of the water may compensate the high field capacity, which hinders water penetration. Groundwater will be drained by an underflow toward the Red Sea or the Nile. Such an underflow was probably used by the Romans in the watering station Kalalat as a source of water for the inhabitants of Berenike. The persistence of this water fl ow is demonstrated by the permanent presence of slow running water in a narrow gorge of Wadi Shenshef, even when preceding winter rains have not been recorded. From the Greek period onward, the groundwater in Arabia, stretching from Kuwait through eastern Arabia to Bahrain, has been exploited via a network of aquifers, which could be tapped for drinking water and irrigation .Atmospheric humidity is an important source of water for plants growing in the Red Sea coastal zone. Evaporation of seawater during daytime replenishes the water content of the air. This water partly condenses again as the temperature of the air sufficiently drops during winter and early spring. Considering the fact that air temperature is lowest just before sunrise, morning dew may therefore be responsible for damp soils. Unlike precipitation, morning dew is a much more steady water source for plants. Referring to agricultural practices in Bactria, Africa, and Cyrene, Pliny even states that crops depend on the dew fall at night for their nourishment. Also Theophrastus emphasizes the contribution of dew to plant growth in regions with no rainfall, including Egypt.

According to Jabbur , who studied the bedouin life in the Syrian Desert, there is so much dew on plants in spring, that some women collect the water in their water skins. This atmospheric humidity is of value for both the fl at coastal plain adjacent to the Red Sea proper and the mountains, which intercept the clouds blown inland. Especially high mountain massifs may receive considerable amounts of rain in this way. This also explains the presence of several mosses and ferns on the higher levels of the Gebel Elba . Fertilization of these plants is only possibly under moist-to-wet conditions.The adaptation of desert plants to water stress is diverse. Several principles can be distinguished and specific combinations of such principles may find their expression in desert plants. One possible adaptation that has arisen by natural selection is the storage of water in leaves, stems, or roots. Such plants are called succulents and are adapted to arid habitats as well as to saline environments. Succulent plants that grow in the Eastern Desert, including the environment of Berenike, are Suaeda monoica Forssk. ex J. F. Gmel, Halopeplis perfoliata Bunge ex Asch., and Aizoon canariense L. A good example of a halophytic succulent is Zygophyllum coccineum, which has swollen stems and leaves and is able to abort part of its leaves and even young shoots to prevent excessive water loss. Other plants have leaves with xeromorphic features. They are called xerophytes and are, by definition, adapted to arid regions. A common feature is the reduction in size and/or number of the leaves, as can be observed by Crotalaria aegyptiaca Benth. and Moringa peregrina Fiori. An extreme example of leaf reduction is demonstrated by Tamarix aphylla H. Karst, a shrub that has only sheath-like leaves. Another way of drought survival is shortening the life cycle. Annual plants that take advantage of the availability of water independent of the season, being either a rain shower or a damp period caused by dewfall, are called ephemerals. 

They include succulents, such as Zygophyllum simplex L.; nonsucculents, including grasses such as Aristida funiculata Trin. & Rupr.; and nongrasses such as Lotononis platycarpa Pic. Serm. Such plants can survive by a quick absorption of the rainwater and dew. Some of them have reasonably long roots, such as species of Aristida, whereas other plant species have only very small and shallow root systems, such as stunted but flowering specimens of Triraphis pumilio R. Br., Eragrostis ciliaris R. Br., and Coelachyrum brevifolium Hochst. & Nees. These plants were frequently found in the environment of Berenike and Shenshef. Some plants may develop temporarily so-called rain or extension roots, which grow in the upper part of the soil and absorb the water from a rain shower or dew . Perennials may also be have as ephemerals. Conversely, some species, such as Zygophyllum simplex, that normally behave as ephemerals, can prolong their life span under favorable conditions . They are as resistant to drought as a seed. Plants that obtain their water directly or through the capillary fringe from the groundwater are called freatophytes. Because in most cases groundwater is only available at considerable depths, freatophytes are predominantly perennial species that are able to develop deeply penetrating taproots. Regeneration of freatophytes will be especially successful during years with heavy showers, resulting in a moist soil from the surface down to the water table over a considerable time span. During this period, long taproots must grow to the water table. As soon as this is reached, the plants will have become independent of the unpredictable rainfall and can survive in an apparent unfavorable environment. A good example of a desert freatophyte is the twisted acacia , which may have a taproot of more than 40 m. Owing to its deep, penetrating root,plastic gardening pots this tree is capable of growing in the fringe of wadi branches. Their presence is even indicative of the many wadi branches that dissect the fl at coastal plains along the Red Sea. The growth of the root starts soon after germination. A seedling under a full-grown tree in Wadi Shenshef of only 2 cm high could be dug out to a length of 25 cm before its fragile root broke off. Examples of herbaceous species with long roots are Aerva javanica Juss. ex Schult., and Aizoon canariense L. Some plants, such as the Nile tamarisk , are capable of initiating condensation of atmospheric moisture by means of special hygroscopic salt crystals under conditions of high humidity . For this reason, drops of water on the surface of the Nile tamarisk taste rather salty. The water condensation is triggered when the humidityof the air exceeds 76 percent . Like dew deposition on the plant surface, which depends on a lower plant surface temperature in relation to the dew point temperature of the adjacent air, this salty water drips from the plant and can be absorbed by the shallow roots. The effectiveness of this self-induced sprinkling is evidenced by the many imprints of raindrops in the damp soil beneath such excretive plants.

The availability of water is, however, no guarantee for the presence of desert plants. This is especially true of regions that do not profit from the dewfall. Here, the irregularity of rainfall is at odds with the presence and longevity of seeds in the soil. Seed banks in desert soils are characterized by a concentration of seeds in the upper 2 cm of soil, with a high degree of spatial heterogeneity and a great seasonal and annual variability. Seeds tend to be concentrated in depressions where water collects and in wind shadows, such as below established plant species . As dispersal distances are generally short, a specific type of ephemeral vegetation may exist for several years . Isolated spots that are completely dependent on unpredictable showers may lack a viable seed bank, and consequently no vegetation will develop after rainfall .The major phytogeographical districts that are distinguished in Egypt are: the Mediterranean coastal strip, the Nile region, the deserts, the mountainous region of the Sinai proper, the oases, and the Sahelian scrub region in and around Gebel Elba . A further division is possible with respect to the Mediterranean area, the Nile Valley, and the deserts . Because the desert vegetation is of prime interest, only the division of the desert vegetation will be discussed in more detail. The Egyptian desert is divided by the Nile Valley into the Western or Libyan Desert and the Eastern or Arabian Desert. Additionally, the Nubian Desert is distinguished, being present on both sides of the Nile south of Aswan. Both the Libyan and Nubian deserts are flat and have several large depressions . On the eastern side of the Nile, three different deserts are distinguished. The Isthmic Desert is located south of the Mediterranean coastal strip on both sides of the Suez Canal. South of this desert, fringed by the Nile and the Gulf of Suez, lies a small desert area consisting of limestone mountains called the Galaga Desert. The Eastern Desert is the third one in the row and extends southward from about lat 26° N to the Gebel Elba region . This large desert is characterized by rugged, igneous mountains that run parallel to the Red Sea on the east side and a rocky plateau on the west side. The Red Sea mountains are dissected by a web of branched wadis that drain off into either the Nile or the Red Sea. Although a detailed labeling of the distribution of the complete Egyptian flora is presented by Täckholm , including a dichotomy between the northern and southern part of the Eastern Desert, which coincides with the territories of the Ma’aza and Ababda nomads, respectively, it has not been used in this study because it is too out-of-date due to many taxonomical revisions and corrections of geographical distribution records during the last decades. Between the mountains and the Red Sea there is a gently sloping coastal plain. Near Berenike this plain is only 8 km wide, but elsewhere it can extend as far as 35 km. This fl at plain is dissected by many wadis draining off into the Red Sea, and its flora differs so much from the flora of the mountainous areas that it represents a separate phytogeographical district. The Egyptian flora comprises almost 2,100 plant species. The distribution of these species over the major phytogeographical regions is presented in Figure 2.8. The analysis is based on the checklist of the Flora of Egypt , the Flora of Egypt and the Key to the Egyptian Grasses .

The highest numbers of larvae present in clipcages were on the citrus, sumac and Star blueberry variety

The first and second instar larvae were grouped into a single ‘larvae’ observation and recovered males and females were counted separately as well as grouped into an ‘adults’ observation . There were clear differences among the mean number of eggs and larvae found across the different plant types . ANOVA indicated a significant interaction between the plant and the developmental stage present on the respective plant . The interaction indicates there was a differential treatment impact with respect to the number of eggs laid and the number of larvae found. The presence of a significant effect of developmental stage indicates that the number of eggs and larvae in clip cages were not similar, i.e. egg mortality was higher with some plants versus others . A correlation coefficient of 0.75 was calculated between the mean numbers of eggs and hatched larvae. Tukey’s HSD rendered a significant plant effect for the total number of eggs and larvae present in the clip cages . More eggs and larvae were found on four types of plants, i.e. citrus , sumac , Star and Jewel . The blueberry varieties Showchaser and O’Neal had significantly lower total number of eggs and larvae and numbers on Emerald and O’Neal were intermediate . The Jewel variety data were slightly skewed, however,planting blueberries in pots as the mean number of eggs and hatched larvae were negatively correlated . Roughly 25% of males and less than 50% of female citrus thrips were recovered from the plants in the bug dorms 14 days post release. The between replicate bug dorm adult survivorship was relatively low.

Of ca. 60 adults of each sex released, male survivors per bug dorm ranged from 13-21 with a mean of 15.4 ± 3.3 [SE]; female counts ranged from 20-37 with a mean of 28.2 ± 6.4 [SE]. The Star variety had the highest mean numbers of adults collected in comparison with other varieties . Data from the bottom two thirds of the plant were pooled together, as there were few thrips collected and eggs found on the lowest third. There were more eggs located on the top third of the plant than the bottom two thirds of the plant across all plant varieties. There were also more larvae found on the top third of the plant than the bottom two thirds, but no differences in larvae found on the bottom two thirds of the plant than eggs present in those areas, for all plant varieties . The Star variety had the numerically highest numbers of total eggs and larvae per plant compared with any other variety. The likelihood ratio chi-square test indicated a significant effect of egg count on the top third of the plant as well as a significant effect of larval count on the two plant locations, i.e. the top third also having the highest count . More eggs were found on the top third of the blueberry plants and there was a higher percent of larvae present on the top third of the plant , regardless of plant variety. However, in the choise test, the Star variety had the highest percent of eggs found and larvae present. The Snowchaser variety had the lowest numbers of thrips counted overall, with fewer eggs, larvae and adults collected on that variety. The Misty variety had fewer larvae present than eggs laid and the number eggs laid were comparable to levels on Emerald.

Under the conditions of these trials , citrus thrips egg hatch typically would take place within 4-9 days , therefore numbers of motile insects counted in both studies would be larvae from the first generation and surviving adults. The primary focus of this work was to determine if there was a difference in the number of eggs laid on the different types plants and our data clearly show this was the case . The no-choice oviposition test showed that the highest number of eggs and larvae were found on citrus and sumac and the blueberry varieties Star and Jewel. Intermediate levels were found on Emerald and Misty, and the lowest numbers on Snowchaser and O’Neal. The Tukey HSD separation for total counts of eggs and larvae group Jewel with the citrus, sumac, and Star, but these data may be skewed due to the negative correlation between the number of eggs laid on the Jewel variety and the number of larvae present. The choice oviposition test also showed a correlation between the number of eggs located and the number of larvae present on the plant. Evaluation of count means showed a difference in citrus thrips numbers on the Star variety over the other blueberry varieties in the test. When given the choice to move about the plants, the citrus thrips’ activities at the time of isolation and counting appear to be similar for Emerald and O’Neal with a mean of less than 10 adults recovered from those plants. In the choice oviposition tests, the negative correlation between egg and larval counts is also apparent for the Jewel variety. Some explanations for this negative correlation could be: first instars eclosing from the egg suffered high mortality, the larvae did not or cannot feed and/or develop well on this variety, and/or the larvae dispersed away from the plant onto other plants. If the latter were the case, then one would expect to see one or more other blueberry varieties with higher numbers of larvae present, but that was not the case and given the fact the canes did not remain in contact with one another, it is unlikely that high levels of larval dispersion to different pots took place.

There are several possibilities which explain the results: 1) females chose to lay eggs preferentially in some varieties over others, 2) egg survivorship influenced the numbers counted/found, 3) first instar egg eclosion varied across variety, 4) differential larval survivorship and development on the plants, and 5) larval dispersion, possibly to other plants, but possibly off the plants with mortality ensuing in unevaluated portions of the bug dorm. The Chi-square values for plant location were significant indicating that there were disparate numbers of eggs laid and larvae counted between the two locations, with the top third of the plants, regardless of variety, preferentially chosen over the bottom two-thirds. This was likely due to the amount of flush growth present on the top third of the plants. Also, given data in our studies, thrips activities appear higher on the top third of the plant; this is likely a good area to sample when looking for thrips on blueberries. Combining all the data, our studies clearly show that the Star variety had more citrus thrips than the other blueberry varieties, and numbers of oviposited eggs found on Star, Jewel, citrus and sumac were similar. This quite possibly is due to Star’s parentage. The Star variety is a variety that produces flowers the earliest and for the longest period during the season and while yield may not be as high as some of the other varieties, it is planted as a season long fruit bearer. Because the high bush blueberries are hybrids of one another and the nurseries have proprietary rights to the genetics of these varieties, we are therefore unsure how closely related the varieties we tested are. Snowchaser was the variety least preferred by the citrus thrips based on our data, however, to our knowledge it is not a variety currently grown commercially in California. There are countless factors that need to be considered to fully understand insect host choice, including but not limited to, host plant finding, host plant acceptance, host plant relatedness, resource concentration, resource availability and host use . The insect’s ability to locate the host plant, settle, feed, reproduce and successfully produce the next generation that achieves reproductive maturity involves countless steps and subtle interactions that are beyond the scope of the work presented here. We provide a platform for further work on the distribution, abundance and avenues for research related to citrus thrips as a pest of blueberries in California. We anticipate citrus thrips will remain a pest of concern to the blueberry industry of California as the industry continues to expand. Citrus thrips, Scirtothrips citri , is a significant insect pest of citrus and mango fruits and has been recognized as a major pest of California citrus since the 1890s . In the USA,blackberries in containers citrus thrips are known from Arizona, California, Texas and somewhat recently, possibly Florida , whereas in Mexico they are reported only from northern Mexico . Based on its past distribution, several authors have reported that citrus thrips is native to southwestern North America and northwestern Mexico . Citrus thrips is primarily a pest of citrus in California, particularly in the San Joaquin and Coachella valleys. They can have a broad host range, including, but not limited to, alfalfa, rose, grape, laurel, cotton, date, fir, lucerne and various grasses, pecans, and other ornamentals. Citrus thrips have been collected from over 55 different plant species .

Their native host plant is hypothesized to be Quercus or more likely Malosma laurina Abrams which was likely one of citrus thrips more common native host plants in southern California and northwestern Mexico prior to the introduction of citrus. In the SJV of California, S. citri has recently broadened its known host range and become a significant pest of high-bush blueberries . Scirtothrips citri was not known in Florida until 1986 where it was first detected in grape surveys . A collaborator was aware that in Florida, S. citri is not often collected from or abundant in several crops it is notorious for attacking in other regions of the Americas , but it is the most common thrips species he has collected from native vegetation and weeds. Species identifications from slide-mounted specimens can be unreliable or inconsistent and alternative or additional methods of identification may be necessary. Morphological identification suggests that S. citri is present in California, Arizona and Florida, but given that it is not a pest on several crops one might expect in Florida, further investigation is necessary to determine if S. citri is actually a cryptic species complex. The development of molecular genetic techniques , predominantly analysis of mitochondrial DNA , has significantly contributed to an understanding of natural genetic diversity and speciation . Genetic markers offer additional methods of species determination and delineation, especially when coupled with morphological identifications . These approaches are especially useful in groups that demonstrate a mixture of diverse ecological traits coupled with a conserved morphology. Given the distribution of S. citri in major citrus growing regions of North America and the level of its pest status in those regions, re-evaluation of morphological and molecular identifications was deemed necessary. The goals of this work were to investigate the haplotypic variation among S. citri populations based on phylogenetic analysis of the mitochondrial and ribosomal DNA, and to identify possible cryptic species complexes within the Scirtothrips attacking citrus. The collection records for all specimens used in this study are listed in Table 1. Specimens were collected from various parts of California, Arizona, Texas, Florida , Mexico, Nicaragua and Turkey. Specimens from Turkey were included in this analysis as it is an under-represented area of the world and at the time of collection from citrus, the collector believed the specimens to be citrus thrips. Specimens were collected into 95% ethanol by beating the live thrips onto a white piece of paper, touching a clean 5/0 Princeton paint brush into the ethanol filled collection vial, touching the ethanol imbibed paint brush tip to the live insect so that the insect stuck to the paint brush tip and then depositing it passively into the collection vial. After collection, all specimens were stored at -20°C until analysis. Some of the collections contained Frankliniella occidentalis and Neohydatothrips burungae but these collected groups were not included in our analysis.spinetoram and this may result in reduced E. hibisci mortality. Growers should be aware of the data presented herein when deciding upon a pesticide rotation management plan, which reduces avocado thrips resistance evolution. Each of the four recommended products have different features with respect to the efficacy of thrips control, concurrent control of avocado mite pests, and persistence of impacts on predaceous mites and other natural enemies . Thrips were removed from ethanol and allowed to air dry on filter paper for 2 min.

Mortality was determined by lack of movement after gently probing each thrips with a small brush

The upper and lower parts of the Plexiglas sandwich were solid and between the lower base and test arena a piece a piece of filter paper was placed to allow moisture exchange and to extend the life of the leaf during the bio-assay. Airflow through the test arena was provided through two holes drilled through the center cell layer directly opposite one another, with fine-mesh screening melted onto the interior of the test arena to prevent escape. The Plexiglas sandwich was held together with four binder clips positioned such that the airflow was not covered. Once dry, the leaves were placed on the filter paper in Munger cells and the respective thrips species was added. The lid was placed on the cell but leaving the cell arena exposed, so that once the thrips were added, the cells could be closed quickly. Female and late second-instar avocado thrips and citrusthrips were then placed on treated leaves of their respective host plants inside the Munger cell. Control leaves for both species were treated with a mixture of the same suspension ingredients minus the protein. Bioassays were conducted concurrently in the following manner for both species: adult female thrips were placed on leaves coated with activated or inactivated forms of both Cyt1A and Cry11A, immature thrips were also placed on leaves coated with activated or inactivated forms of both Cyt1A and Cry11A, and all combinations for adults and immature thrips were carried out along with the corresponding control cells. The Munger cells were closed and placed in an environmental chamber at 28ºC, 55% RH,raspberry container and long daylight conditions . Each cell was carefully removed daily and the filter paper doused with water to prevent leaf desiccation. The bio-assay was replicated on two separate dates .

A minimum of 10 individuals was placed into each Munger cell and thrips were checked daily for eight days to assess mortality. Post seven days, the integrity of the leaves was questionable and in all but one bio-assay, mortality was observed before seven days; thus data were analyzed using day 7 mortality. Six strains of B. bassiana were obtained from the USDA-ARS Western Integrated Cropping Systems Research Unit located in Shafter, CA. GHA is the commercially available strain found in the field formulation of B. bassiana, Mycotrol O and the greenhouse formulation BotaniGard ES, and each of the other five strains were obtained via isolation from soils in Kern County by USDA-ARS collaborators in 2000. They were stored at – 80ºC. Culture methods for the thrips experiments were similar to those described previously for Lygus hesperus Knight bio-assays and were conducted by collaborators from USDA-ARS, Shafter, CA. Briefly, isolates were grown on SDAY media, or Sabouraud’s dextrose agar plus yeast extract . The conidia were harvested from culture plates after 10–14 days incubation by scraping with a sterile rubber policeman into a 0.01% solution of Silwet L- 77 . The conidia were then enumerated with a hemocytometer. For preservation and storage, glycerol was added to the conidial suspension and stored in aliquots of 2 × 108 in a 2 ml solution at −80°C until needed for bio-assays. Conidial viability was assessed following incubation for 16 h in potato dextrose broth just prior to use in experiments. Viability was determined by adding a sample of approximately 107 conidia to 20 ml potato dextrose broth and incubating ca. 16 h in a rotary shaker at 28°C.

Conidia germination was examined under a compound microscope at 400× and scored as viable if the germ tube was at least twice the length of the conidium. Percentage viability was measured on 250 conidia of each isolate. All bio-assays were conducted on the basis of the number of viable conidia measured after thawing and the desired concentrations were formulated by serial dilution. The strain from Mycotrol was isolated and cultured exactly as above to eliminate possible effects of production methods and formulation ingredients on insecticidal activity. Glycerol was not removed prior to using the conidia in bio-assays. All six B. bassiana strains were suspended in 0.01% Silwet in a de-ionized water solution and evaluated on the same date at four concentrations for each thrips species. The control consisted of 0.01% Silwet in de-ionized water solution. Each of the 25 treatments was evaluated using five Munger cells , which contained a minimum of ten adult female thrips. These bio-assays were repeated on 10 dates with both species tested simultaneously on each date . Groups of thrips were anesthetized by exposure to CO2 for 15-30 sec, and each strain was administered to the dorsum of the abdomen of each knocked out thrips quickly and carefully in a 1µl drop with a Burkard Hand Microapplicator over filter paper. The droplet spread the length of the thrips immediately and the thrips was then deposited, still knocked out, onto the leaf tissue in the Munger cell. Once a minimum of 10 treated thrips were added, Munger cells were closed and sealed with binder clips and placed in an environmental chamber at 28ºC, 55% RH, and long daylight conditions . Each cell was checked daily for seven days to observe infection by the fungus. Each cell was carefully removed daily and the filter paper doused with water to prevent leaf desiccation. Individuals infected with B. bassiana were defined as those whose natural activity was retarded and/or showed arrestment and subsequently produced mycelia, which was confirmed post bio-assay.

Mortality caused by mycosis was confirmed on the basis of visual observation and then crushing individuals to reveal the presence of mycelial growth. When mycelial growth was not apparent, crushed individual thrips were placed on potato-dextrose agar plates for 5 days and then re-examined for the presence of mycelial growth. Data were analyzed after Abbott’s correction for control mortality using log-probit analysis with PROC PROBIT on SAS 9.2 and using the Raymond Statistics package . The purpose of the probit analysis was strictly for gross strain comparison. Probit analysis was used to estimate the LC50 and LC95 levels, confidence intervals, and χ 2 values for each strains. Lethal concentrations with overlapping 95% confidence intervals were not considered significantly different. The daily check data were analyzed as non-cumulative counts per day via the Survival Distribution Function on SAS 9.2 , where observation time represented the probability that the experimental unit from the population would have a lifetime exceeding that time with the variables strain and concentration. Assessments for each variable by species were done with Log-rank and Wilcoxon tests and multiple comparisons for the log-rank test were adjusted by using Tukey-Kramer method. Data were then plotted as estimates of the survivor function for the different strains separately for each species. Bacillus thuringiensis israelensis produces two groups of toxic proteins, the Cry and Cyt toxins that have different modes of action. In this investigation, results with Cyt1A and Cry11A were disappointing as both activated and inactivated forms of both proteins showed little effect against adult and second instar citrus thrips and avocado thrips. To our knowledge, there have been no reports of Bt endotoxins with activity against Thysanoptera, although Cyt1Aa was found to be toxic to the non-target species Chrysomela scripta Fabricius . Many hypothesize that because thrips feed with a punch and suck method, rather than direct chewing and mastication of leaf tissues, they do not receive toxic amounts of the Bt proteins . Alternatively, they may not possess the proper binding receptors for the Bt proteins tested to date and thus, no pore can be formed in the midgut lining and the Bt proteins are excreted . The literature indicates the latter hypothesis is more likely based on findings from life table parameters where development, fecundity,growing raspberries in container and adult longevity or relative abundance are not significantly different from thrips reared on Bt positive versus Bt negative corn, cotton, or potato plants. The aforementioned studies were not specifically looking at Bt effects on thrips nor were the Bti toxins tested here involved in previous studies involving thrips. The combinations of proteins used in this study were, to date, unique pairings with thrips. It is indeed possible that there are no Bt endotoxins currently available that cause mortality to Thysanoptera. The LC50 with strain GHA was 8.61 x 104 conidia/ ml and was two orders of magnitude lower than for the other five B. bassiana strains tested . GHA also gave the only statistically valid dose-response values in probit analysis, and provided the only data that fit the probit model. The other B. bassiana strains failed to provide a linear relationship based on their p-values , i.e. the probit regression lines were of poor quality, except for GHA. Therefore, data were evaluated based on line slopes as is commonly seen in the scientific literature with other biological agents where data lines are not straight and do not fit the model . Strains 1741ss, SFBb1, S44ss, and NI1ss showed a flat dose-response between concentrations, did not fit the model, and LC50’sranged from 2.7 x 106 – 9.6 x 108 . Assessment of Beauveria strain while adjusting for concentration, in both Logrank and Wilcoxon tests showed that strain and concentration had a highly significant effect on the infection rate. Multiple comparisons for the Logrank test to assess the strain effect while adjusting for the concentration differences showed that strains 1741ss, S44ss, 3769ss, and NI1ss infection rates were not distinct from one another.

Strain GHA and SFBb1 had infection rates different from each other as well, and GHA had the fastest infection rate and SFBb1 showed the slowest kill rate . The Survival Distribution Function analysis coupled with the probit analysis clearly shows that GHA would be the best strain choice for citrus thrips control. Results with avocado thrips. The LC50 for strain GHA was 2.2 x 106 conidia / ml and was similar to that obtained with the other five B. bassiana strains tested . Again, because a strong linear response was not observed, the performance between strains was rated based upon the LC50 and relative linearity of the response. Based on overlap of confidence intervals, there were no significant differences between any of the strain LC50’s or LC95’s . Assessment of Beauveria strains while adjusting for the concentration, using both Log-rank and Wilcoxon analysis showed that strain did not have an effect on the infection rate. The multiple comparisons for the Log-rank test to assess the strain effect while adjusting for the concentration differences showed infection rates for all 5 strains were not distinct from one another . The Survival Distribution Function analysis coupled with probit analysis indicated there was no one best strain to select for avocado thrips management. Citrus thrips were more susceptible to Beauveria than avocado thrips; citrus thrips LC values were much lower for the most active strain, GHA, indicating that significantly lower dosages of strain GHA were required to infect and kill citrus thrips compared with avocado thrips. The overall survival analysis results showed a similar pattern to the results of the probit analysis; GHA had the fastest infection rate and SFBb1 had the slowest rate . Infection rates for the other three strain’s fit in between the rates for GHA and SFBb1, and 1741ss, S44ss, 3769ss, and NI1ss infection rates were not separable. This low dosage association and having the fastest infection rate suggest GHA is the best candidate for field-testing among the strains examined. Except for the worst performing strain, SFBb1, the performance of all of the strains with avocado thrips were similar. The LC50 value for citrus thrips was 8.6 x 104 conidia/ml, which may suggest economical feasibility in some cases, e.g., for use on organic products. The maximum recommended field application rate is 5.0 x 1012 conidia/ha. Therefore, 8.6 x 1011 conidia/ha of GHA is needed based on the estimated LC50 of 86 conidia/µl and this amount is reasonable to obtain in a field setting. Conducting the same analysis for avocado thrips control using GHA, with an LC50 of 2.2 x 106 , 2.2 x 1013 conidia/ha would be required. This is 4.4 times greater than the standard field use rate of GHA. We hypothesize that differences in susceptibility between citrus and avocado thrips may be due to the different habitats in which they evolved.

The LR and ST decreased leaf area index and increased canopy porosity

These treatments are provided visually in Figure 1.In the second experiment, individual grapes from different cluster positions were collected from two cultivars grown in a commercial vineyard in Oakville, CA in 2017. Cabernet Sauvignon grapevines and Petit Verdot grapevines were 21 and 9-years old, respectively. The exposure of each individual grape was estimated with fish-eye lens photography from the grape perspective pointing the zenith. The images were processed in R . After applying a thresholding condition to the blue channel of all images, they were converted into binary pixels . Thus, the percent of binary pixels capturing the sky was used to calculate the percentage of canopy porosity as reported previously . Then, those berries were collected at harvest, and their flavonoid content was analyzed with reversed-phase high performance liquid chromatography.The experiment was conducted in 2019 in Oakville, CA with row orientation NW-SE. The vineyard was spaced 2 m × 2.4 m with Cabernet Sauvignon grapevines on 110R root stock. The grapevines were trained to a vertically shoot-positioned system with a cordon height 96 cm above vineyard floor, trained to a bilateral cordon, and pruned to 1-bud spurs. Plants were irrigated weekly with 2-drip emitters per vine, with the capacity to deliver 3.8 L of water per hour. The experiment was designed as a randomized complete block with three canopy management practices: removal of 5 to 6 basal leaves on the NE side ; thinned to 24 shoots per vine ; and a combination of LR and ST and an untreated control ,square plant pots with four replicates each consisting in 5 grapevines, 3 of which were sampled and the 2 on distal ends were treated as border plants.

The ST and LR treatments were applied on 11 June 2019. Harvest commenced when the berry TSS reached to ca. 24°Brix on 23 September . The sampling time points were as follows: 2 weeks before veraison , veraison , 2 weeks after veraison , 3 weeks after veraison , 5 weeks after veraison , and harvest , were chosen to cover the response of the berry metabolism to cultural practices and the concomitant increase in exposure.Leaf area index was measured on 21 June to characterize grapevine canopy growth and converted into leaf area on by a smartphone based program, VitiCanopy, coupled with an iOS system . The gap fraction threshold was set to 0.75, extinction coefficient was set to 0.7, and sub-divisions were 25. A “selfie-stick” was used for an easy access to place the device about 75 cm underneath the canopy. The device was positioned with the maximum length of the screen being perpendicular to the cordon, and the cordon being in the middle of the screen according to previous work . In each experimental unit, three images were taken to capture half canopy of each vine, and analyzed by the software. The relationship between leaf dry mass and area was determined on a sub-sample of leaves of different sizes using a leaf area meter . Total leaf area was calculated by defoliating one grapevine per treatment replicate after harvest and using the regressive relationship between leaf dry mass and leaf area. At harvest, clusters were manually removed, counted, and weighed on a top-loading balance. Leaf area to fruit ratio was calculated by dividing leaf area with crop weight. Dormant pruning weight was collected during the dormant season ; and crop load was calculated as the ratio between yield per vine and the pruning mass of each vine. Labor operations costs and gross income per hectare were calculated based on yield and net returns per hectare and methods presented elsewhere . Anthocyanin productivity was calculated as reported by Cook et al. .

At each sampling point and experiment, 55 berries were randomly collected from the middle of each treatment-replicate and kept on ice until they were measured. Berries were weighed, and mean berry mass was determined as the average mass of the counted berries. These berries were used to determine the total soluble solids , the pH, and the titratable acidity . TSS was measured as °Brix, with a digital refractometer . The juice pH and TA was determined with an autotitrator using sodium hydroxide to titrate to an end point of pH 8.3, and it was expressed as g•L−1 of tartaric acid.For each sampling point in each experiment, 20 berries were collected, gently peeled, and berry skins were freeze-dried . Dried tissues were ground with a tissue lyser . Fifty mg of the resultant powder was extracted in methanol: water: 7 M hydrochloric acid to simultaneously determine flavonol and anthocyanin concentration and profile as previously described Martınez-Lüscher et al. . Briefly, extracts were filtered and analyzed using an Agilent 1260 series reversedphase high performance liquid chromatography system coupled to a diode array detector. Separation was performed on a reversed-phase C18 column LiChrospher® 100, 250 mm × 4 mm with a 5-µm particle size and a 4-mm guard column of the same material at 25°C with elution at 0.5 ml per minute. The mobile phase was designed to avoid co-elution of anthocyanins and flavonols consisted in a constant 5% of acetic acid and the following gradient of acetonitrile in water: 0 min 8%, at 25 min 12.2%, at 35 min 16.9, at 70 min 35.7%, 65% between 70 and 75 min, and 8% between 80 and 90 min. The identification of flavonoid compounds was conducted by determining the peak area of the absorbance at 280, 365, and 520 nm for flavan-3-ols, flavonols and anthocyanins, respectively. Identification of individual flavan-3-ols, anthocyanins, and flavonols were made by comparison of the commercial standard retention times found in the literature. Commercial standards of epicatechin, malvidin-3-O-glucoside, and quercetin-3-Oglucoside were used for the quantification of flavan-3-ols, anthocyanins, and flavonols, respectively.

The determination of proanthocyanidins was performed using an Agilent HPLC-DAD after an acid catalysis in the presence of excess phloroglucinol , with minor modifications described in Martınez-Lüscher et al. .The 3-isobutyl-2-methoxypyrazine was quantified by a stable isotope dilution assay using headspace solid phase microextraction coupled to a gas chromatograph and a mass spectrometer as described Chapman et al. and Koch et al. with some modifications. Briefly, 20 berries per treatment-replicate from Experiment 3 were randomly collected from the clusters of three vines in the middle of each treatment-replicate on both side of the canopy, by cutting the pedicel with a pair of scissors and frozen at −80°C until analysis. Pedicels were removed by hand and berries were placed in 50 ml conical tubes. 10 ml of pure water and 100 ml of deuterated IBMP isotope were added into the tube. Then, samples were ground with a tissue homogenizer Power Gen 1800D and centrifuged at 3000 rpm for 10 min. 10 ml of the supernatant was pipetted into 20 ml SPME vials containing 3 g of sodium chloride.Statistical analyses were carried out using the R-Studio version 3.6.1 for Windows. All data were subjected to Shapiro-Wilk’snormality test . Correlations between variables were calculated with the Pearson’s test by using the same software. Segmented regression analysis was used to determine the point of inflection the in the relationship between increasing exposure and the berry skin anthocyanin and flavonol content with “segmented” 0.5-0.3 R package . Data were normally distributed and, subsequently, were submitted to an analysis of variance to assess the statistical differences between the treatments applied in each experiment performed. Means ± standard errors were calculated, and when the F value was significant , a Duncan’s new multiple range post hoc test was executed using “agricolae” 1.2-8 R package . When data were not normally distributed, a Kruskal-Wallis test was conducted. Percentage data were transformed according to the suggestion of the most likelihood test,plastic pots for planting into arcsine root square before ANOVA or Kruskal-Wallis tests.The growing season of 2017 was warmer and drier compared to the reference data for the same period within the last 20 years . Thereby, average daily temperature was 4°C higher and rainfall was 18 mm less. Grape berry mass differed significantly depending on the degree of exposure . Overexposed berries were the smallest due to overexposure resulting in dehydration thereby reducing berry mass. Neither total soluble solids nor titratable acidity changed regardless of the degree of exposure to which berries were subjected. However, the juice pH of the Exp+ Deg+ and Exp+ Deg++ berry must was greater compared to Exp− and Exp+ Deg− berries. Berry skin flavonoid content and composition were also affected by the degree of exposure . The berry anthocyanin content of Exp− was similar to Exp+ Deg−. However, overexposed berries resulted in berry anthocyanin content that was 70% and 90% lower when compared to the Exp− berries. Grape berry exposure to solar radiation not only affected the anthocyanin content but also modified the ratio between the tri- and di-substituted anthocyanins leading to a less stable profile in all treatments with exposed berries. Likewise, berry skin flavonol content and composition were strongly affected by the degree of exposure to solar radiation. Therefore, in Exp+ Deg− flavonol content was two-fold greater than Exp−, albeit they abruptly decreased in overexposed grapes where flavonol content was 25% and 50% lower when compared to Exp− berries. Furthermore, in overexposed berries the proportion of kaempferol and quercetin significantly increased while the proportion of myricetin decreased. Regarding proanthocyanidins in berries, mild exposure did not affect their content in Exp+ Deg− compared to Exp− berries.

However, greater solar exposure decreased proanthocyanidin content in berries but to a lesser extent compared to Exp−. Finally, the content of flavan-3-ols was severely reduced in Exp+ Deg++ berries .The analyses performed on single berries from two varieties confirmed the obtained response in anthocyanins and flavonols in Cabernet Sauvignon . Thus, exposure affected the accumulation/degradation of these flavonoids. Exposed berries from the East side of the canopy decreased 8%and 36% of the anthocyanin content in Cabernet Sauvignon and Petit Verdot, respectively. Thus, Petit Verdot seemed to be more sensitive to higher level of solar exposure and degraded anthocyanins. Overexposed berries of Cabernet Sauvignon resulted in an 87% decrease of the berry skin anthocyanins when compared to the interior berries . Berry skin anthocyanins and increasing exposure showed a significant trend below the 22% of kaempferol . Conversely, analysis of the segmented regression on Petit Verdot berries did not show a clear trend below the 3.2% of Kaempferol and after the point of inflection, anthocyanins started to degrade . Regarding flavonol content, no differences were observed between cultivars . Conversely, when exposure increased to ca. 60% the content of flavonols in exposed berries of both canopy sides and in both cultivars; the overexposed berries had the lowest flavonol content . Thus, our data revealed a strong positive relationship between the berry skin flavonols and the percentage of kaempferol until 8.6% of kaempferol proportion for Cabernet Sauvignon and 7.2% Petit Verdot . However, beyond these thresholds, flavonols started to degrade, and there was an indirect relationship between the flavonol content and the percentage of kaempferol for both cultivars, this relationship being significant only for Cabernet Sauvignon .The weather conditions during the execution of this experiment were highlighted by greater maximum daily temperatures when compared to the reference period . This was more prominent during the driest months . Moreover, global solar radiation received at the experimental site was to ca. 200 W m−2 greater than the total solar radiation recorded within the reference period . The combinatory effect of LR and LT treatments caused a 58% reduction of LAI and a 45% increase of canopy porosity . However, neither leaf area nor pruning mass showed significant differences between treatments. On the other hand, yield components were mostly affected by the shoot thinning treatments . Thus, shoot thinned vines showed lower number of clusters, yield, and Ravaz Index , and increased leaf area to fruit ratio per vine as expected. The extent of yield reductions was 55% and 47% for ST and LRST vines, respectively . Berry mass was not significantly affected by canopy management practices during the berry ripening although vines subjected to LRST tended to result in smaller berries . The most influential effects observed on berry chemistry were due to shoot thinning treatments . Therefore, shoot thinned vines had greater total soluble solids and lower titratable acidity from mid-ripening to harvest. However, no significant effect was observed on the must pH . Shoot thinned grapevines had higher anthocyanin content at veraison . However, we did not measure any changes to anthocyanin content at harvest as affected by the canopy management practices applied.