The fragile fragments that are created in this way will only be partly retrieved by sieving soil samples

When used as food, the salt is removed by washing or soaking in water. The recovery of fruit fragments just outside some buildings in Berenike suggests local growth of the plant, which would have eliminated the need for preservation. Members of the white bryony family are very juicy in a fresh condition and cannot be dried successfully. Four species are evidenced in Berenike by their seeds: the colocynth , the watermelon , the cucumber , and the bottle gourd . The colocynth grows in the Eastern Desert, where it is easily gathered and remains fresh for at least a couple of weeks. Ripe water melons can only be stored for five up to eight days and must have been available from local kitchen gardens or harvested in an unripe condition if they were imported. The ripening process continues after harvesting if the fruits are exposed to warm air, preferably between 20 to 25°C. Methods of preserving gourds until the next yield are available, is described by Pliny . One way is to add brine and another is to store them in a trench floored with sand and covered with dry hay and earth. In this way, gourds will remain in a green condition. The fruits of the subfamily Maloideae are classified as pomes. The seeds are enclosed in a cartilaginous structure in the center of the soft, indehiscent fruit. Apples can be preserved in several ways: fresh, dried, or kept in a preservative liquid, such as honey, as described by Apicius , or in a mixture of vinegar and brine as mentioned by Columella . Fresh, juicy apples might have been the most desirable ones. Dealing with the preservation of fresh fruits, Varro, cited by Pliny , bud drying rack says that fruits such as apples should be separately wrapped in fig leaves. According to Galen , properly ripened apples can be stored for the winter and the following spring . Windfall specimens were not used for storage.

The storage life of fresh apples is prolonged if stored in a cool environment. Although this condition is not met while on the way to Berenike, the transport of fresh apples to this remote destination would not have been problematic from September to October, just after harvesting. Today, dried apples produced on a commercial scale are peeled, cored, and sometimes cut into rings. But small apples can be dried as whole specimens or simply cut into segments, as described by Columella . In the latter case, fruits evaporate quicker and spoilage is reduced. Such halved apples, preserved by charring, are frequently found in archaeological contexts. The presence of pips in Berenike leaves open all aforementioned kinds of preservation methods. The last group of fruits and nuts to be dealt with concerns false fruits. They are represented by the juniper and the fig . The berrylike fruit of the juniper is easily dried and can be kept for a considerable period. Fresh accessory fruits of the fig are juicier but can also be dried easily. Due to the high sugar content, they can be kept until the next harvest. Long-distance transport is no problem, as is evidenced by the recovery of figs throughout the Roman Empire. Fresh figs can only be kept for a couple of days, as they are vulnerable to mold, which results in lactic fermentation. It is possible to preserve fresh figs in honey, as is mentioned by Apicius . Fig paste, made from trampled figs mixed with sesame , anise , fennel , and cumin , was made into balls and kept in jars covered with pitch or was heated in vessels in order to reduce the moisture content .The archaeobotanical record of Berenike and Shenshef, based on samples from seven excavation seasons, comprises 68 cultivated plants that could be identified in most cases to the level of species. As much as 28 cultivated plant species are evidenced from Berenike but are absent from the archaeobotanical record from Shenshef. Juniper is, on the other hand, a plant species that has been found only in Shenshef. Most certainly, the discrepancy between these records is partly affected by the unbalanced excavations at both sites. The wild plants are represented by some 110 wild plant taxa and could only partly be identified to the level of species. Some of the botanical remains are still unidentified, and it is assumed that they concern both cultivated and wild plant species. A part of these still-unidentified remains have clear diagnostic features and may even concern whole seeds or fruits. Their identification is hampered by the incompleteness of the reference collection.

The cultivated plants were obtained from an extended geographical area, and for this reason a large number of economic plants from remote areas have to be taken into consideration. As far as the reference collection of the wild plants is concerned, the incompleteness is caused by the difficulties met in collecting the seeds and fruits of desert plants, their presence largely dependent on the fluctuations in the humidity. The first step to a successful identification of the unidentified cultivated plants would be the compilation of lists of potential plant species that were traded from specific areas. Such a valuable checklist was presented by Warmington , who selected medicinal plant products traded by the Arabs on the assumption that they were perhaps also known to the earlier Romans. Although the number of species is limited, this list includes the coconut , the emblic , and the tamarind , three species evidenced at Berenike by the current archaeobotanical research as real trade products. Although these three species were already present among the botanical remains unearthed in Berenike during the first excavation seasons, the identification of emblic was only successful several years later, when a sample of this food plant containing some of the characteristic endocarp fragments was obtained from a spice shop in Cairo. Consultation of Warmington’s list at an earlier stage would have been of great help. Recently, a more extensive list of plant species that might have been traded from tropical Asia and China has been compiled. The matter is urgent as spice shops in the Near Eastern bazaars are becoming increasingly threatened. Their broad and exotic selection is being replaced more and more by a limited and more standardized supply as a result of globalization. But it is still possible to get a glimpse of the range of goods offered for sale during the past and to collect relevant reference material. From the uneven distribution of many plants over the trenches as well as the still untouched large habitation and industrial areas of both settlements, we can expect more plant species to be unearthed in future excavations, which in turn may shed new light on the subsistence and trade economy. This is not to say that a more elaborate sampling will ultimately result in a representative picture of subsistence and trade. Despite the excellent preservation conditions, archaeobotanical research is faced with the transfer and consumption of plants that have left no visible marks and of which all traces are vanished. Wastage of expensive commodities would have been the exception rather than the rule.

Food may also have been prepared in such a way that allowed for no inedible leftovers. Other food is completely digested, and therefore no identifiable remains would have been excreted in the feces. Despite their botanical richness, a similar warning is called in for the analysis of dump areas. Most of the botanical remains that have been investigated originate from organic dumps, which appeared to be predominantly middens and fl at dump areas in the proximity of stone buildings. It should be realized that in the course of time even such concentrated and well-preserved archives become only a faint reflection of the organic waste once deposited. Several processes are responsible for the reduction of this organic archive. Although it is conceivable that at least part of the habitation area was inaccessible to straying animals, as was the case, for example, with locally constructed garden plots, it seems most likely that organic dumps in particular were frequently visited by browsing animals, as is evidenced by the many dung pellets especially of sheep and goats and, to a lesser extent, from camels. Archaeozoological research and recent observations of trash deposits have attested to the presence of the following animals: cattle, pigs, chickens, sheep, goats, donkeys, black rats, dogs, Egyptian vultures, vertical grow rack system and brown-necked ravens . On a smaller scale, small creatures, such as beetles and larvae, also eat organic material. This kind of reduction may even last for thousands of years, as could be observed when special attention was paid to this process while analyzing a Roman trash deposit in Karanis . It was found that such small animals are present at least in the upper 10 cm of the trash. The reduction of the archaeobotanical archive may also be the result of burning the trash. During the excavations campaigns, such burning is still frequently practiced to get rid of the stench and to control as far as possible the number of flies, rats, and other such vermin, which in turn attract predators such as snakes. For safety reasons, such burning would mainly have taken place outside the settlement. This explains why most of the plant remains from the trash dumps inside the settlement have been preserved by desiccation, those used for offering ceremonies being an exception. The wind would also have been a factor in the reduction of the trash deposits. When trash dries out during high temperatures, it may be easily blown away. Only those remains that were sufficiently covered would have been protected from further losses due to wind and other factors, and these deposits constitute the actual archive that has been sampled . Some prudence is called for when numbers of plant remains are compared. A quantitative approach is not only obscured by the different plant parts we are dealing with, which is even true for seeds and fruits, but also because numbers of seeds and fruits differ highly among plants. Additionally, taphonomic processes bias the number of sub-fossil remains. Some fruits may serve as examples. It appeared that the endocarp fragments of the almond easily fall apart in an arid environment. This is caused by the relatively thick veins that are present in the middle of the inner fruit layer. Differences in the fruit anatomy also cause a discrepancy in counts. Each consumed date and olive , for example, will produce only a single leftover, either a seed or a fruit stone . Other fruits contain many seeds, such as a pomegranate with has some 250 to 400 seeds and the fig , whose number of “seeds” ranges from about 700 to 1,800 . A simple conversion of the number of sub-fossil seeds that have been found to the original number of fruits that were consumed is therefore impossible. The comparison of the retrieved plant remains is further complicated by differences in the consumption of fruits. Seeds of the date and fruit stones of the olive are not consumed and have a good chance of ending up in the trash deposits. On the other hand, the small seeds of the pomegranate and the fig are consumed and excreted and are therefore less likely to be found in trash layers. The consumption of whole, destoned fruits at the sites proper would have been the main source of recovered fruit-stone fragments. The concentration of these remains in middens or dump areas will partly depend on the way the fruits are eaten. Eating olives with an Egyptian colleague in a courtyard, I observed that I just threw the fruit stones on the ground, while my colleague collected all the fruit stones carefully but simply discarded the packing material of his cigarettes. From an archaeobotanical point of view, a possible recovery of my discarded fruit stones during a hypothetical excavation is most unlikely, whereas in excavating the trash deposit, one would have a good chance of finding some of my companion’s, which most probably will have been deposited there by the kitchen help.

Two cluster II Frankia genomes contain close homologs of rhizobial nodABC genes

There has been only one report of nodulation of a compatible host from soil that was devoid of native host plants , although only after 18 months of bait plant incubation. Recently, a cluster II Frankia strain was isolated in pure culture from a nodule of Coriaria japonica collected in Japan . These two examples indicate that not all cluster II Frankia strains are obligate symbionts. That being said, it seems clear that the cluster II Frankia strains form a strongly restrictive association with their host plants. Our study adds to the complex nature of cluster II Frankia strain association with their hosts. Two degrees of ecological association between a microsymbiont and its host may be considered: persistence and enrichment. Members of cluster I and III Frankia spp. have been known to persist in soil devoid of host plants for many years and to readily nodulate their respective hosts . Moreover, the number of infective units of cluster I Frankia spp. has been known to increase under Alnus host plants up to 30-fold, which makes the presence of the host plants a major factor in ampliftying Frankia populations . interestingly, the rhizosphere of Betula sp., a non-host closely related to the genus Alnus, showed enrichment of cluster I Frankia spp. compared to the rhizosphere of Alnus spp. at a host-plant-present site and abundance comparable to Alnus spp. at a host-plant-absent site . In our study, as evidenced in the microbiome analysis, an enrichment effect of cluster II Frankia was present but was much subtler. Cluster II Frankia spp. were detected in the host-plant-present sites and were also detected in rhizosphere soil of a non-host, H. arbutifolia, in the host-plant-present site, but no cluster II Frankia spp. were detected in the host-plant-absent site. In contrast, plastic flower bucket strains of the other clusters of Frankia were detected in both host-plant-present and -absent sites.The relatively greater abundance of cluster I Frankia spp. among the typical symbiotic and nitrogen-fixing subgroups that we observed is in congruence with results of previous studies . On the other hand, we found that cluster III was the least abundant: 6 reads total and only two samples with any reads detected.

This contrasts with previous findings where cluster III Frankia spp. was the dominant or codominant subgroup in a noncompatible host rhizosphere . This difference may be due to the fact that the soil conditions among these studies are at the opposite end of the ecological spectrum. In reference 35 the soil was moist and in a university campus arboretum, whereas our MiSeq samples were collected in dry nutrient-poor serpentine soil in summer. It is possible that there was a specific effect of serpentine soil conditions on cluster III Frankia spp. For example, Oline showed that while microbiomes from serpentine soils were similar to those of nearby non-serpentine soils at the phylum level, distinct subgroups were adapted to the specialized environment of serpentine soil. Not very much is known about the population distribution of the atypical cluster IV Frankia strains in soil. However, since they comprise the most abundant subgroup inour samples, it is clear that at least some strains of cluster IV Frankia are adapted to serpentine soil. The observed trend of decreasing clusters I and IV Frankia spp. in host-plant-present sites compared to that in host-plant-absent sites suggests that the host plant may be influencing the population sizes of strains of clusters I and IV as well as of cluster II Frankia. This may be related to an inhibitory factor in the host soil ecosystem, as discussed below. Nearly 34% of the reads that mapped to the genus Frankia mapped to OThus of indeterminate identity. Thus, depending on the true identity of these OThus, our results are subject to change. However, all of the reads that mapped to indeterminate OThus were  1% different from any of the cluster II OThus; with the mapping threshold set to 99%, no reads that mapped to these OThus would map to cluster II OThus, and all indeterminate OThus are more closely related to known cluster I or III strains than to cluster II strains.While the relative abundance of cluster II Frankia spp. in the microbiome was significantly higher in the host-plant-present site, it is still clear that this group of OThus is quite rare in the microbiome, representing only 0.05‰ in any sample.

This suggests that the host factor is not a source of energy to sustain a large population; i.e., it is not promoting the prolifteration of cluster II Frankia outside the host, in contrast to some strains within the cluster I Frankia that can utilize a host-derived soil carbon source . Further, permutational MANOVA and factorial ANOVA both showed that the presence of the host plant did not significantly affect the relative abundances of all OThus in the microbiome overall nor the -diversity; in fact PCoA with an outgroup showed that the microbiomes of the samples from the host-plant-present and host-plant absent sites were very similar to each other. This minimal influence of host/non-host on the overall microbiome together with the trend observed for clusters I and IV Frankia OThus suggests that a host-derived factor has an effect specifically on cluster II Frankia OThus. Cluster II Frankia OThus were found in the rhizosphere soil of the non-host H. arbutifolia in host-plant-present sites, but not in the rhizosphere of H. arbutifolia in host-plant-absent sites. Thus, the host-plant effect was not limited to the host-plant rhizosphere but rather extended to the level of the site where the host plant was present. Extracts from host-plant roots and shoots have also been found to enhance growth of Frankia strains in culture particularly strongly if the strain is compatible . Taking this into account, we propose that possible mechanisms for the dispersal of this host-dependent factor might be diffusion of a compound or compounds originating as root exudates, leaf litter decomposition and leaching into the soil profile, or chemical signals dispersed via mycorrhizal networks . A plant factor with specific targets that might have a strong impact on Frankia strain presence in the rhizosphere might be a plant-derived secondary compound related to nodulation signaling, such as a flavonoid. Flavonoids are known signaling molecules in rhizobial symbioses . Additionally, in Myrica sp. symbioses, a suite of hydroxy-chalcones, termed myrigalones, has been shown to promote infective Frankia and inhibit noninfective Frankia strains . Alternatively, this factor might be a terpenoid with effects similar to strigolactone, known to stimulate spore germination and limited hyphal growth of fungal symbionts in the rhizosphere in arbuscular mycorrhizal endosymbioses . The host signaling pathway that initiates AM symbiosis is ancestral to signal exchange in RNS; and the microsymbiont signal molecules are also similar between these two types of endosymbiosis . These nod gene homologs have been shown to be expressed in developing root nodules of D. glomerata . In summary, our data demonstrate the existence of a host site enrichment of cluster II Frankia strains that is independent of host plant species, vegetation type, geographical location, climate, soil type, and soil pH. The dependency of Frankia spp. on theirhosts for survival and persistence has been postulated, based on in vitro experiments . Host and non-host rhizospheres were recently compared with respect to cluster I Frankia strains in one location , showing no significant host rhizosphere effect. Cluster III Frankia strains were shown to have a distribution independent of the host rhizophere, implying a broad physiological adaptation in soil . The current report is the first comprehensive study showing a significant host plant influence on cluster II Frankia strains in natural soil environments, an effect that is distinct from that of other ecological parameters. Identification and testing of potential specific host-plant compounds and transmission pathways remain to be carried out in more controlled environment experiments.All soil, leaf, flower buckets wholesale and root nodule samples were collected from three locations in northern Califtornia: in and near Anderson Lake County Park, Santa Clara County, Califtornia , McLaughlin Natural Reserve, Lower Lake, Califtornia , or Sagehen Experimental Forest, Truckee, Califtornia , as shown in Fig. 6 . Maps of these locations were generated with R with the ggmap package .

These locations varied considerably in altitude, monthly precipitation, monthly temperature, and soil type. The altitudes were approximately 200 m, 650 m, and 1,950 m above sea level in ALCP, MNR, and SEF, respectively. Multiple sampling sites were selected within each location: ALCP, n 4 sampling sites; MNR, n 5 sampling sites; and SEF, n 2 sampling sites. The main factors that were tested across these sites were host plant presence/absence , host plant species , soil type , soil pH, vegetation type, and climate type. Factors 1 to 5 are described in Table 2. The soil type in each site was determined by methods described below, and the vegetation types were determined to the macro group level according to the U.S. National Vegetation Classification using A Manual of Califtornia Vegetation . The annual patterns of temperatures and precipitation at each location were determined according to the WorldClim database . Sites within each location were paired according to host-plant presence/absence for comparison. In ALCP, AR, WA, and K were compared with CC on serpentine and non-serpentine soil . In MNR, SE02 was compared with SE05 , both on serpentine soil, and NS02 was compared with NS01 , both on non-serpentine soil. In SEF, SH06 was compared with SH01 , both on non-serpentine soil. Vegetation composition was similar for pairs in ALCP and in MNR . In the Sagehen Creek site, the pairing was between sites with dissimilar but adjacent vegetation types . In ALCP, the host plant Ceanothus ferrisiae was present in AR, WA, K but absent in CC. In MNR, the host plant Ceanothus jepsonii was present in SE02, and the host plant Cercocarpus betuloides was present in NS02. In SEF, Ceanothus velutinus was present in SH06. Site SE06 in MNR was used to collect leaf samples of Avena fatua for 15N comparison .Strawberry production in Califtornia accounts for more than 80% of total U.S. production, with an annual farm gate value of $1.10 billion , which is four times greater than all other states combined . In addition, Califtornia produces nearly one billion strawberry transplants each year in nurseries, and these transplants must meet strict phytosanitary standards for local production and export. Such a profitable industry in Califtornia has been made possible by the fumigation technology developed in the 1950s with methyl bromide and chloropicrin . Since then, preplant fumigation with methyl bromide and chloropicrin has become an integral part of the Califtornia strawberry production industry , and nearly all conventional strawberry production occurs in fumigated soils . Annual soil fumigation has contributed to the control of soilborne pathogens, nematodes, and weeds while also boosting the yields of strawberry plants. Historically, this also allowed breeding programs to focus on improving horticultural characteristics of strawberry cultivars in lieu of emphasizing disease resistance. Because of the negative effects of methyl bromide on stratospheric ozone, the fumigant was designated as a class I stratospheric ozone depleting substance by the Montreal Protocol and as a significant risk to human health . The continued availability of this efficient fumigant for agricultural soil fumigation beyond the 2005 phase-out date will be through critical-use exemptions. It has been estimated that annual losses in short term net farm income in Califtornia will be more than $162 million, with strawberry accounting for more than 60% of these losses . Over the past 10 years, research has focused on identifying alternative fumigants with efficacy comparable with methyl bromide . Alternative fumigants such as chloropicrin and Telone C35 have been identified, and improved application techniques have been developed to reduce emissions . Although chloropicrin is as efficacious as methyl bromide + chloropicrin at high rates, these are not feasible for the growers due to regulatory limits placed on application rates. Regardless, chemical alternatives to methyl bromide will be subjected to increasing review and regulation and they may not be readily available over the longer term. It has been estimated that soilborne diseases caused by Pythium, Phytophthora, Cylindrocarpon, Macrophomina, Rhizoctonia, and Verticillium spp. result in 20 to 30% strawberry yield losses in the absence of fumigation .

The green plant can be eaten as a vegetable and the seeds are used as a spice and as a medicine

In Egypt it has a tradition of being added to bread, mixed with the fl our of sorghum in particular . A popular dish in Yemen is “hulbah,” which is made from pounded fenugreek mixed with meat broth and various vegetables . Fenugreek is often eaten with white lupine , which is already mentioned by Galen . The seeds are also used for making fenugreek tea , which is still served in Egypt. Medical treatments include promoting lactation; healing inflammations; regulating digestion; relieving coughs, asthma, and emphysema; and its use as an aphrodisiac . Duke states that in Punjab , whole plants are added to stored grains as insect repellents and that seeds are used locally for a yellow dye. It is not known if this was also practiced in ancient Egypt. During a botanical field trip, a sample of fenugreek seeds was found by the author among the belongings of a dead Ababda sheikh, who was buried on a terrace of Wadi Shenshef near the Roman ruins . The author discovered these belongings high up between the branches of an Acacia tree, well safeguarded against incidental water currents. I visited the tree again the following day with some Ababda nomads who allowed me to inspect part of the find, including a tin that appeared to contain an almost-empty packet of Nefertiti cigarettes, indicating that the belongings must have been deposited in the tree at least ten years earlier. Also present were three boxes of matches, an illegible piece of paper, a small medicine bottle, plastic pot manufacturers and two tied-up pieces of cloth in the form of a purse. The fenugreek seeds were present in one of these purses; the other one contained seven topped cowrie shells.

These shells could be used by gifted people for fortune-telling and for consulting to plot out a proper route during a journey. It could not be determined whether the fenugreek seeds were used as a spice or as a medicine. The archaeobotanical records of fenugreek are scattered and include Spain, Bulgaria, Germany, Egypt, Israel, Jordan, Iraq, and India. The oldest finds originate from Iraq , Israel , and Jordan . Archaeobotanical records from Egypt almost equalize the number of all other finds, the oldest one originating from predynastic Ma’adi . In Berenike, fenugreek might have been cultivated in locally constructed garden plots. It is adapted to sandy soils and can even withstand some salinity. Seeds can be harvested about three to five months after planting. Alternatively, the seeds could have been imported from the Nile Valley.The general picture from the Egyptian archaeobotanical record is that the hulled emmer wheat predominated during the pharaonic period and was largely replaced by free-threshing wheat during the Ptolemaic period. A reexamination of the published records revealed that we are predominantly dealing with hard wheat in the GrecoRoman period, and not with bread wheat . Both free-threshing wheats can easily be distinguished by their threshing remains, whereas the identification of fruits can be problematic . Although hard wheat is well adapted to relatively warm climates, and for that reason is the dominant wheat in the Near East and the Mediterranean area, it has quite recently been replaced in Egypt by bread wheat on a large scale. The cultivation of hard wheat is now confined to small areas in the Delta and the Fayum. Nevertheless, a new variety of hard wheat has recently been brought into cultivation. The yield of durum wheat is larger than that of emmer wheat, but less than that of bread wheat. The name hard wheat refers to the very hard endosperm, which makes it unsuitable for bread making. After grinding, the gluten-rich flour is made into a stiff, unleavened dough that is used for all kinds of pastas, such as spaghetti, noodles, and macaroni. The identification of the Berenike specimens is based on the morphological features of the rachis fragments .

It appeared that almost all the rachis nodes unearthed belong to durum wheat. Only a few spikelets of emmer wheat and some rachis fragments of bread wheat were found. Both species were always found mixed up with the predominant durum wheat and are therefore considered as contaminants. Durum wheat has been recorded in reasonable quantities in both early and late Roman deposits of Berenike and would have been one of the staple grains for human consumption. Most probably, the hard wheat was obtained from the productive Nile Valley. The delivery of mainly modest quantities of wheat from Koptos to Berenike is evidenced by an ostraka archive found at Koptos, dated from 18 BC to AD 69 .The bitter vetch is a pulse crop native to the Near East and the Mediterranean area. Also from Egypt, it has been recorded as a wild plant species, where it grows on cultivated grounds in the Mediterranean coastal area . From this area it moved westward to the Balkan area and to the countries along the Mediterranean Sea, where it has been recorded from many Neolithic sites. Records from Egypt date back as far as predynastic times . The bitter vetch belongs to the first group of domesticated crops of the Fertile Crescent and central Anatolia, although it is considered of less importance than the lentil or the pea . Seeds of the bitter vetch can be eaten by ruminants such as cows and sheep and by birds such as chickens, but they are toxic for other animals such as donkeys, horses, and pigs. The seeds are only fit for human consumption if they are soaked in water for some time. According to Zohary and Hopf , bitter vetch was mainly used as forage for animals, at least from Roman times onward and was only eaten by man in times of famine. Originally a weed in lentil fields, the bitter vetch is still difficult to eliminate once it has become established in such fields.

The correlation between the bitter vetch and the lentil in samples from Berenike and Shenshef is weak and assumes that the bitter vetch had been purposely brought to the site. Considering the availability of a broad spectrum of high-quality food products in both these sites, the bitter vetch was probably imported as animal fodder from the Nile Valley. According to Becker-Dillingen , seeds of the bitter vetch have also been used to adulterate black pepper , but such a practice at Berenike is not conceivable.The wild ancestor of the fava bean has not yet been identified and the archaeological evidence from early Neolithic settlements is also still scanty. For these reasons the fava bean is not considered as one of the founder crops of early agriculture . The oldest finds are recorded from the Levant, from where the crop spread westward into the Mediterranean area and eastward into India and China. The oldest Egyptian archaeobotanical find, unequivocally identified to the level of species, is dated to the Old Kingdom and originates from Abusir . Today, the fava bean belongs to the most important food legumes produced in Egypt . The fruits have a white and velvet coating on the inside and contain some 3 to 6 seeds. Dry beans have a protein content of about 25 percent, black plastic plant pots wholesale which is comparable with that of the pea . Fava beans are grown for both green and dry consumption. If consumed as immature green beans, the plants are harvested several times. To obtain dry beans, the whole plant is harvested as soon as the first fruits turn black. Green beans are eaten after boiling, whereas dried beans are often baked first, a meal that is called foul midamis . Taamiah is a fried ball of paste made of germinated fava beans, and is a popular snack in Egypt. In the eastern Mediterranean, a part of the human population is hypersensitive to the consumption of fava beans, which can cause a condition known as favism. Pollen may also bring on this disease, and it is probably for this reason that Pythagoras advised against eating flowers of this plant . A variety of chemical compounds is responsible for the oxidation of red blood corpuscles and hemoglobin, resulting in anemia and jaundice . Unless they are stored in a cool environment, harvested beans will quickly deteriorate. This may explain why the fava bean was frequently found among crops cultivated in the small kitchen gardens present in settlements in the Eastern Desert. Obviously, this pulse is easily grown under controlled conditions, and it is most likely that it was also cultivated in local kitchen gardens in Roman Berenike and Shenshef. Such plants might concern special garden cultivars, which are distinguished from field cultivars. The garden cultivars are especially grown on a small scale throughout Africa, Southeast Asia, and parts of India for its green pods . The green pods can be harvested for human consumption, and the remaining plant parts are suitable for use as stock feed. Despite their relatively thick seed coat, only a few small fragments of the fava-bean seeds have been unearthed in Berenike and Shenshef, indicating that their presence in the archaeobotanical records is probably underrepresented. This could be explained in several ways. First, large seeds will have less chance of getting lost in the sieving process in food preparation. This is in line with the predominance of small, aborted fava-bean seeds frequently found in waterlogged samples from The Netherlands. In addition, it may be assumed that if large seeds do indeed get lost, they have a better chance of being eaten by browsing animals than smaller ones such as lentils that will easily sink down in the trash. Furthermore, large seeds of pulses easily disintegrate and, in this state, are easily susceptible to further organic decay.

As only seed fragments were found, it was not possible to measure the size of the seeds, which would enable an identification to the level of variety.The mung bean is a domesticate of the Indian subcontinent. Its wild progenitor, Vigna radiata ssp. sublobata , is found in the western Himalayan foothills and extends throughout the Western Ghats into Sri Lanka. Apart from the discovery of the mung bean in Roman Berenike, the only other archaeobotanical records so far originate from India and date at least to the start of the second millennium BC . Together with black gram , the mung bean is highly valued, the former being particularly prized by high-caste orthodox Hindus . Today, the mung bean is mainly produced in Southeast Asia, including India. According to Westphal , the mung bean has been introduced relatively late in East Africa and is still not fully accepted there as a pulse crop. Mung beans are grown primarily for their dried pulses. They can be eaten whole, split into dhal, or ground into fl our. Whole or split seeds are eaten after boiling, and the fl our of mung beans can be used for all kinds of baking products. Additionally, germinated bean sprouts are eaten, which are obtained from the green-colored seeds. Their protein content is well over 25 percent. Almost 70 mung beans have been unearthed from several loci in two trenches, located in the northwest part of the center of Berenike. They were traded as whole seeds. sub-fossil mung beans from India that coincide with the Roman trade with India have been recorded from Narhan and Taradih , both located in north India, and from Nevasa , east of Bombay . This would suggest the import of mung beans from Barygaza. On the other hand, it has to be realized that the Indian archaeobotanical record is still biased in favor of prehistorical sites found in the southern part of India. Prehistorical finds of the mung bean are recorded from Hallur and Sangankallu in Karnataka . Assuming a continuation of the mung bean cultivation, this means that import from the more-southerly ports of Muziris and Nelkynda must also come into consideration.The progenitor of the cultivated grape is Vitis vinifera ssp. sylvestris, which grows in the Mediterranean area and eastward to Turkmenistan and Tadzhikistan where isolated populations are present. As far as the North African Mediterranean coastal strip is concerned, the distribution of this wild grape is confined to Morocco and Algeria. Grape cultivation started in the Levant in the early Bronze Age, making it one of the first classical fruits of the Old World .

The groves in Abu Nechle were not used anymore when Schweinfurth visited the area

Olive oil can be kept for a long time if not exposed to air and is mainly used in cooking. The first archaeobotanical evidence of olive dates back to the Eighteenth Dynasty and includes leaves found in the tomb of Tutankhamen. Not until the fourth century BC, however, did the olive become a substantial part of the diet of the Greeks in Ptolemaic Egypt . A further increase of its use in Egypt is dated to the Roman period, as is evidenced by the numerous references in literary sources and the substantial archaeobotanical record. Both the number and size of the fruits found at Berenike and Shenshef indicate that we are dealing with the cultivated subspecies and that they were imported from the Mediterranean area or the Fayum. The fruits of the wild species are smaller than those of the cultivated ones. Also the relatively small population at Gebel Elba makes it improbable that olives were imported from this area. The presence of the stony endocarps at Berenike and Shenshef indicate that they were available as table fruits. Salted olives must have been a valuable food in the warm desert as it supplements the loss of salt via evaporation.Two different species of rice have been domesticated: the Asian rice Oryza sativa L. and the African rice O. glaberrima Steud. Cultivation of Asian rice began in many parts of south and Southeast Asia, including northeastern India. In fact, rice cultivation in India goes back to 2500 BC, raspberry grow in pots making it one of the oldest regions of rice cultivation. The cultivation of African rice, on the other hand, is probably not more than 3,500 years old .

Although the original distribution area of the annual Oryza glaberrima Steud. spp. barthii J. M. J. de Wet, the wild progenitor of Oryza glaberrima, coincides with the savanna zones south of the Sahara and even extends towards the center of Sudan, its domestication area is limited to West Africa . Therefore, it is plausible that the rice that has been found in the excavations was imported from India late in the history of Berenike. The Periplus Maris Erythraei frequently mentions grain as a trade item together with rice, suggesting that the taxonomic relationship of rice with wheat and barley was not clear to the author of the Greek text. See for example the enumeration of export commodities of Syrastrênê : “The region, very fertile, produces grain, rice, . . .” . The separate status of rice was also expressed in its use in beauty treatments . Casson notes that with the exception of textiles, not all items traded from India to Arabia and Africa are mentioned in the list of exports of the Periplus. According to Casson, this implies that merchants from Roman Egypt were only interested in the real luxury products that were available at these ports. The presence of reasonable amounts of rice in both early and late deposits at Berenike, however, makes it very likely that at least for this cereal an indirect trade with India did exist. According to Feliks , who wrote about rice cultivation in the Roman period, a highly prized large kernel rice variety was cultivated in Israel, probably in the Hula Valley. Feliks’s main source is the Mishnah , which is considered to be very authentic. This assumption is, however, not yet evidenced by archaeobotanical research. Greek and Roman writers only mention rice cultivation in Syria and Mesopotamia . Strabo , for example, mentions that rice grew in Bactria, Babylonia, Susis, and lower Syria Others, such as Dalby , are of the opinion that rice was never grown within the Roman Empire. Because fragments of amphoras were found at Berenike that originated from Syria and possibly also from the Gaza-Negev area , it is possible that the rice found at Berenike was imported from the Near East.

With Berenike’s southern and southeastern maritime orientation, however, it seems more likely it was an item of the distributive trade with Arabia. According to the Periplus, rice was exported from the country of Parsidai and the Gulf of Terabdoi and the district of Ariakê , which are identified by Casson as respectively the area around modern Karachi in Pakistan and the area between Broach and Surat in northwest India . From there it was exported to the island of Socotra , which is located northeast of Somalia, and to the so-called far-side ports on the northern coast of Somalia . The Periplus is less clear about the export of rice from Limyrikê, the present-day state of Kerala, which is located along the southwest coast of India. It is mentioned that sailors from both Barygaza and Limyrikê by chance put in at Socotra and would have exchanged rice, among other commodities . This is in line with modern rice cultivation in India, which stretches along the western coast from Gujarat southward to Sri Lanka. Rice is mentioned as one of the trade items of Sri Lanka by Ptolemy . Ships that called at ports along the Somalian coast could be loaded up with Indian rice brought there by Indians or Arabs. In this way rice was indirectly traded to Egypt. It is even possible that for some Roman ships these ports in Somalia were the final destinations . The claim that rice import from India must have been expensive, as put forward by Dalby , is questionable in the light of intermediate involvement of Indian and Arab traders. Pliny states that rice was imported from the East without further specification and does not mention a price. Rice could be obtained from harbors along the northern coast of Somalia and from the island Socotra . The import of rice to Berenike is in concordance with its earlier mentioned use within the Roman Empire, although the export of grain, probably wheat , from Berenike to Muziris and Nelkynda in southwest India seems to contradict this. According to Casson , grain exported from Egypt to Indian ports was destined for Westerners permanently established in those places, while Indian merchants living in Berenike ate rice.

Conversely, perhaps, the import of rice to Berenike indicates the presence there of Indian or other South Asian residents who consumed it. Although rice cultivation probably began in Egypt between the Arabic conquest and the Turkish invasion , the archaeobotanical evidence from Berenike indicates that it was already being consumed during the Roman period. Intact spikelets as well as loose chaff remains show that rice was transported in the husks. In addition to the rice found at Berenike, only three other archaeobotanical records of rice are known from sites within the Roman Empire so far. Small quantities of rice have been found at Quseir al-Qadim, both from the Roman and the medieval period . A still unpublished record is mentioned by A. R. Furger from Zurzach in Switzerland . Roman rice consumption has also been documented by Knörzer for the first quarter of the first century AD in Novaesium . In this fortress along the Rhine River, altogether 196 charred kernels were found. Whether this rice originated from the East or was imported from Arabia, perhaps via Berenike, cannot be determined. All other archaeobotanical records from Europe are dated to the Middle Ages and modern times. Papyrological evidence of the availability of rice in Egypt is extremely scanty. Only four out of the approximately 34,230 published documents, dating from the third century BC to the eighth century AD, mention rice. Three of them are dated to the early Roman period, the other to the sixth century AD . Two of these documents originate from the Fayum; one mentions the purchase of rice, the other the control of the rice trade. According to Konen, such a monopoly indicates that, at least temporarily, rice trade must have been quite important. Strabo states that most of the Indian food consisted of rice porridge and that Indians made a beverage from rice that is known as arak. In India, 30 planter pot rice has also a tradition in being used in offerings at all kinds of religious and auspicious ceremonies. Only new rice is used for this purpose, whereas old rice is used for cooking. It is also believed that rice gives strength and makes one more fertile, which is, for example, expressed in throwing rice on a bridal couple . Possibly, rice was only used on a limited scale within the Roman Empire and in particular for medical reasons. In some of the recipes in the Apicius cookery book, rice is used as a thickening agent for sauces, making use of the waxy consistency of rice fl our. A kind of rice cake is mentioned by Chrysippus of Tyana . Additionally, it was mixed with beans and used by women for preserving the smoothness of the skin .

The main distribution area of the date palm lies between lat 15°N and 30°S and extends from the Spanish Sahara toward Pakistan. To the north it follows the coastal area of the Mediterranean Sea as far as southern Spain, and to the south it is recorded from the coastal area of Sudan, Eritrea, and Somalia. In this southern part of its distribution area, however, most of the date palms grow in the northern provinces of Sudan . The date palm is recorded from all the phytogeographical districts of Egypt, although the most numerous groves can be found in the oases of the Western Desert and in the cultivated land along the Nile. Date palm groves along the Red Sea coast are recorded by Schweinfurth from the Wadi Gimal estuary, some 100 km north of Berenike, and from Abu Nechle south of Ras Hadarba , some 300 km south of Berenike. Also in the near vicinity of Berenike, several date palms were observed by the author recently. All these specimens were, however, still immature and not fruit bearing. Being a monocotyledon, the date palm has only adventitious roots, which do not grow deeper than 2 m. Its presence, therefore, is confined to localities where sufficient fresh water is available. Although it is not a halophyte, the date palm can withstand considerable concentrations of salt and can be found near the sea, as is the case with some specimens in the estuary of Wadi Gimal. Several freshwater springs are present in this coastal area, which enable the date palms to survive. Schweinfurth links the palm grove in the Wadi Gimal estuary with human interference, suggesting a relict vegetation from an ancient settlement or a plantation initiated by pilgrims or sailors. Mandaville , however, is of the opinion that such palm groves along the coast might also concern true relict populations of wild specimens. Propagation from seeds is indicated by equal numbers of male and female plants. The date palm is dioecious, just like the doam palm. One single male date palm can fertilize on average about 25 females date palms. Cross-pollination is achieved either by wind, and eventually insects, or by humans. The advantage of artificial pollination over wind pollination is that a higher yield is obtained. This is not only because more flowers become fertilized, but also because artificially pollinated flowers produce on average larger dates. Unfertilized flowers of the date palm are also capable of producing fruits, a phenomenon that is known as parthenogenesis. Although dates from such flowers are as sweet as the ones that develop in fertilized flowers, they are significantly smaller . artificial fertilization has been described as early as ca. 2300 BC in a cuneiform text of Ur . The knowledge of artificial fertilization was probably introduced into Egypt during the Middle Kingdom . This assumption partly rests on the rare occurrence of date fruits or seeds dated before the Middle Kingdom, whereas dates are frequently found from this period onward. We must bear in mind, however, that the total number of archaeobotanical records concerning the remains of the date palm of the pre-Middle Kingdom period is still relatively small, which is, for example, also true for the sycamore fig , another fruit tree native to Egypt. Another problem is that no comprehensive analysis of the variation in seed size is available on seeds dated from the Middle Kingdom onward.

Two genera within this family are important for the production of aromatics

Job’s tears is native to tropical Asia where it grows in forest margins and swamps and has been introduced throughout the tropics . Cultivated soft-shelled forms are grown as a cereal in northeastern India and southeastern Asia, though it has been replaced now by rice and maize on a large scale . Forest dwellers in northeastern India gather the hard fruits of wild forms and bring them to tribal and village markets where they are sold for ornamental purposes, contributing to the local economy of the forest villages . The hard-shelled form also grows along the eastern coast of India, where it is found as a weed in rice fields. It seems possible that Job’s tears is also mentioned in historical sources, but the translations of the concerning text fragments refer to wheat instead. In his enumeration of the wheat varieties, Theophrastus mentions a variety that grows not far from Bactria and produces kernels that grow as large as olive stones. Pliny includes this Greek reference in his description of wheat varieties, but this time the size is even equated to that of the ear of a cereal. Together with barley , rice , and millet , Job’s tears is native to this area, and it is the only one that comes into consideration when a large kernel is concerned. The glossy bract varies both in size and shape, and it may reach the length of a small olive,container raspberries although most specimens are smaller. So far, sub-fossil remains of Job’s tears have only been found at Berenike and in India.

Although fruits are actually found on many sites in India, they have only been published from eight sites, including early historical Ahichchtra and Balathal. Other plant remains that have been used as beads in ancient Egypt include seeds of grape , accessory fruits of the sycamore fig , seeds of rosary pea , unripe fruits and female flowers of the date palm , endocarps of the Egyptian plum , leaves of grasses , fruits from Polygonum senegalense Meis., and possibly pierced tubers from earth almond . The stone fruit from olive that has been previously described as a bead is now considered as a gnawed specimen.The balsam tree is a member of the incense-tree family . Some species of Boswellia produce frankincense, which is also traded as incense, and include the olibanum tree , whereas some members of the genus Commiphora produce myrrh. Usually, C. myrrha Engl. is considered as the tree from which common or hirabol myrrh is obtained, although this is doubted by Wood . Scented myrrh is produced by C. guidottii Chiov. and is also known under a variety of names, including bissabol, habak hadi, perfumed bdellium, scented bdellium, coarse myrrh, East Indian myrrh, false myrrh, perfumed myrrh, sweet myrrh, and opopanax . Whereas frankincense and myrrh are produced as solid resins, some of the Commiphora species produce balsams, which are aromatics that remain liquid. One of these species is C. gileadensis, whose distribution in Egypt is restricted to the Gebel Elba area . The balsam can be obtained from three different parts of the tree: the dried fruits , the dried wood , and the bark from which an unguent is collected .

Judging by the many sub-fossil fruit fragments, it is assumed that balsam was probably available as carpobalsamum in Berenike. According to Pliny , the quality of opobalsamum was better than that of carpobalsamum and xylobalsamum, its price comparable to that of the best-quality frankincense. Theophrastus , on the other hand, states that the fruit is more fragrant than the gum, the latter probably extracted from the stem and thus similar to Opobalsamum. Carpobalsamum was used in the production of the Egyptian perfume metopium, which constitutes an oil extracted from the seeds of bitter almonds , mixed with some other plant materials . The balsam extract was also used for making ointments, such as the balm of Gilead, which is also called Mecca balsam and opobalsum . In more than 60 different samples from Berenike, fruits and seeds have been found from Commiphora gileadensis. The only other member of this genus that is native to Egypt is C. quadricincta Schweinf. ex Engl. This species is characterized by four longitudinal wings along the fruit stone, as is indicated by its Latin name. C. quadricincta has only been recorded in 1929 from the Hailab area and also from the Red Sea coastal area more to the south. Because of this, identifying the sub-fossil specimens as C. quadricincta is not plausible. The aromatic fruits C. gileadensis were probably frequently imported from the Gebel Elba area, where this tree is still quite abundant together with acacia trees. Because C. gileadensis is native to both sides of the Red Sea, it is also possible that the fruits were sometimes imported from more remote trade centers on the East African coast and the Arabian Peninsula. Both the presence of the fruits and seeds and their morphology exclude the possibility that we are dealing with the Indian bdellium tree .

The Periplus Maris Erythraei mentions the export of bdellium from northwest India, so the presence of fruits and seeds is not in keeping with this kind of commodity. Moreover, the fruits of the Indian bdellium tree are two-celled, whereas the specimens from Berenike are without exception one-celled, being quite similar to those from the balsam tree. Today, the Bisharin nomads who live in the Gebel Elba area do not exploit the balsam tree for its balsam or fragrant fruits . The same is also true for Yemen: the branches are used here instead as firewood . Two other records of the balsam tree from Egypt have been published, although both undated and only one with a reference to a site, namely, Thebes. Additionally, several resins are recorded that belong to Commiphora .The Egyptian plum has fruits of about 2 cm in diameter. Only the outer, soft layer of the fruit is edible, comparable with, for example, plums and cherries. The edible pulp is so mucilaginous that it is also used as birdlime. According to Pliny , the fruits were also used in Egypt for making wine, and Theophrastus states that cakes are made from the stoned fruits. Also the seeds can be eaten, but this has not been practiced by the inhabitants of Berenike and Shenshef. The calyx is persistent, and this explains why it is sometimes found in addition to the fruit stones. The genus was formerly mentioned as Sebestena, and the old name is still in use in the name of the New World species Cordia sebestena L., the geiger tree. Dried fruits of C. myxa and C. latifolia Royle are still offered in spice markets as sapistan . They are used as a medicine for chest complaints and inflammations of the urinary tract . Despite the well-documented archaeobotanical record of C. myxa from Egypt, it is thought to be a native of the Indian subcontinent. Its exotic status in Egypt is supported by the cultivated specimens grown in gardens and the absence of naturalized specimens. Most probably, C. myxa had been introduced at an early stage into western Asia and northeast Africa, which is expressed in several of its common names. Theophrastus , for example, states that the tree grows in abundance in Thebes. Classical writers refer to it as ἡ κοκκυμηλέα [plum] or “myxis” , although there is some confusion by Pliny with respect to the plum , which is common to Europe. The English translation of Theophrastus adds “sebesten” and “Egyptian,” whereas “Indian cherry” is used in India . Stone fruits of this species are well represented in early and late deposits from Berenike and also in late deposits from Shenshef. Occasionally, whole fruits have been found. The fruit also occurs at other Egyptian sites, dated to the pharaonic period, such as Saqqara, Deir el-Medineh, and Thebes. The stone fruit from Saqqara, dated to the Third Dynasty, shows that the tree was introduced and cultivated in Egypt from at least as far back as the Old Empire . Thus, berries of C. myxa may have been imported to Berenike from the Nile Valley or the Mediterranean area, but also long-distance supply from India may not be ruled out. Because the berries are susceptible to fermentation,draining pots only fresh ones may have been available from Egypt proper. Indian berries may have been dried or pickled, methods that are still in use.In addition to Cordia myxa, C. sinensis has also been found at Berenike and Shenshef. Recently, the latter species has been split into two distinct species: C. sinensis Lam. and C. nevillii Alston . C. Sinensis, which grows in moist habitats, is widespread in Africa, though its distribution in Egypt is limited to the southern part and one location in Sinai. In Egypt, only its presence in the Gebel Elba area may be considered as indigenous, whereas in other parts of the country it only grows in gardens. C. nevillii, on the other hand, is adapted to more-arid conditions and has a Somalia-Masai distribution in Africa, the Gebel Elba area in which it is found being an isolated northern outpost. Drar collected plants of both species from Gebel Elba .

According to Drar , who uses the old name C. gharaf Ehrenb. ex Ascherson for both species, Cordia grows, but is not common, in the higher reaches of the rocky gorges of the Gebel Elba. Both species have edible fruits that are about half the size of those of C. myxa. According to Schweinfurth , the berries of C. sinensis are of a better taste than those of C. myxa. It is most likely that the fruits from Berenike and Shenshef were imported, at least partly, from the nearby Gebel Elba area. Additionally, the fruits might have been traded from Ethiopia and Eritrea, where substantial populations of both species occur. Both species are also found in the coastal areas of the Arabian Peninsula and the Indian subcontinent, including Sri Lanka. The trees are also valued for their wood. In the Gebel Elba area, the Bisharin nomads make walking sticks from the branches, which are also products of trade. For that reason, each tree is assigned to a specific family or clan . As far as generative features are concerned, both species can be distinguished by their accrescent calyx but not from their fruits. As only fruits were found in Berenike and Shenshef, the identification is made to the level of a combined taxon. Consequently, previous records of C. sinensis from Abusir and Thebes may refer to C. nevillii as well.Coriander is a garden herb with a quite unpleasant smell. Ripe and dry fruits, however, acquire a pleasant smell and are therefore more in demand than the young plants or the leaves in the use of seasonings. Because the ripe fruits easily scatter from the plant at this late stage of development, harvesting the fruits is therefore very precise work . Coriander probably originates from the eastern Mediterranean, where the oldest finds are recorded from Greece, Jordan, and India. The oldest Egyptian find originates from predynastic Adaïma . Coriander fruits are still much appreciated in Egypt, where the plant is cultivated in the Mediterranean coastal area, the Nile Valley, the oases of the Libyan Desert, and the Sinai . Also, this condiment was in great demand by the Romans, and it was imported in large quantities from Egypt. Pliny states that in his time the best coriander came from Egypt. Coriander is mentioned, for example, in many recipes of Apicius, who lived in the time of Augustus-Tiberius, who collectively reigned from 27 BC–AD 37. The predominant use of coriander’s fruits in preference to its leaves explains why this plant is often found in archaeobotanical research. Fruits and seeds of coriander were scattered in small numbers over several samples, which is especially true for those of Berenike. Coriander is one of the plants that could have been locally cultivated in well-protected and irrigated kitchen gardens. Besides its use as a condiment for flavoring foods and beverages, coriander is also used in medical treatments. It is recommended, for example, by Pliny for all kinds of complaints, such as the treatment of various skin disorders, the retardation of the menstrual period, and as an antidote to snake poison.

Pod-tanned leather is still obtained by the Ababda nomads by using ripe pods

Plant remains may become charred by accidental fire, for example, during cooking. They also may be the result of deliberate fire, for example, in the case of burning rubbish and offering food in religious ceremonies. The preservation of organic materials is also affected by salts. The proximity to the sea results in relatively high salt concentrations, and even salic horizons are present in the soil around Berenike. Salt has certain conserving properties for organic remains, as it dehydrates microorganisms responsible for decay, but it also may cause damage as a result of mechanical stress resulting from crystallization and hydration . The latter effect is especially perceptible at the level where the salic horizon is present. For the reconstruction of trade routes, it is important to know the possible areas of origin. Unfortunately, the assignment of a certain plant species to a particular area of origin is problematic. The main reason is, that during Roman times, many cultivated plants were introduced to new areas via old land and sea routes. Long before the Greek and Roman conquests of Egypt, transoceanic trade between India, the Arabian Peninsula, and East Africa had already been established. It is suggested that via such old trade routes African species were introduced into India, such as Lablab purpureus, Vigna unguiculata, Sorghum bicolor, and Pennisetum glaucum . Conversely,plant bench indoor species from the Indian subcontinent found their way to Africa. Scientific names may, in this respect, be misleading.

Those of sesame and tamarind , for example, suggest an Indian origin for both species, whereas this is probably only true for the former one. The unraveling of the migration and trade routes is especially complicated in the area under consideration as it is linked to the Near East where domestication started some 10,000 years ago. Information on the origin and expansion of a certain species can be derived from a variety of sources, in which evidence from archaeobotanical research, plant geography and ancient written sources all play an important role. If necessary, these sources will be discussed to determine the possible trade routes.The genus Abrus belongs to the Leguminosae and contains twelve tropical species. The best-known member of this genus is the rosary pea . The seeds of this species are normally red with a black eye around the hilum. This is interpreted as an imitation aril, but it is puzzling why birds would eat this hard seed because it does not provide any nutrition . Besides the scarlet seeds, entirely white seeds and black seeds have been reported from India. Although India is frequently mentioned as the geographical origin of this genus, its probable origin is Africa, where most species of Abrus are found . Seeds of the rosary pea are used in several ways. Besides its application as a stimulant and its use in medicinal preparations , the rosary pea is also valued for its ornamental properties. In India its seeds have a tradition of being used as weights by goldsmiths, as beads in rosaries and necklaces, and for the decoration of objects such as baskets . Their use in weighing precious metals is attributed to the seeds’ uniform size and weight.

Each seed has a weight of 0.1 g. Because the seed coat is extremely hard and glossy, it is not possible to tamper with the seeds. Nevertheless, some variety in seed weight exists, and it happens that heavier seeds are used for buying and lighter ones for selling . Seeds are not edible because they contain the poisonous protein abrine. The consumption of a few pounded seeds is lethal. Poisoning by rosary pea is also recorded from drinking tea in which several seeds have been immersed for a while. In Egypt the seeds of rosary pea are also used as an additive in mixtures of incense and are offered on markets all over Egypt. An incense sample bought in November 1996 at the Khan al-Khalili bazaar in Cairo contained seeds or fruits from the following plant species: Abrus precatorius, A. cf. fruticulosus, Coriandrum sativum, Phalaris paradoxa, Trigonella foenum-graecum, Nigella sativa, and Senna holosericea. The use of rosary pea seeds in Egyptian incense is also mentioned by Kamal . Additionally, unmixed seeds of the rosary pea are offered for sale . Such supplies together with coins are often used to scatter over bridal couples. In both instances, it is believed that the rosary pea seeds protect against the evil eye. One of the Arabic names, “Any-al afrit,” refers to this belief. It holds that certain people have the power to bring misfortune that could affect other people, cattle, or even houses. The evil eye can only be effective if it strikes the victim’s eye at first glance. A protection against the evil eye can either be offensive or defensive in character. An example of the first approach is the hand of Fatimah, which is used as an amulet. The use of rosary pea seeds in, for example, incense mixtures can be classified among the second approach.

It was also possible, though with some difficulty, to buy an incense from Morocco at the Khan al-Khalili bazaar whose special purpose is protection against the evil eye. Contrary to normal incense samples, this gives off a ghastly smell. That completely black seeds of Abrus sp. can also be used for religious purposes is evidenced by the presence of two such seeds in the well-hidden, covered hole at the back of a Yao wooden statue, which originates from either China, Thailand, or Laos. Together with a piece of animal skin, metal, a paper fragment with text, and bark, these seeds allowed the statue to become gifted with a soul. A similar use is applied to wild rue , a plant species indicative of disturbance. According to Horne , in Iran, the fruits are, among other things, worked into wall hangings and burned as a deterrent to the evil eye. The specimens from Berenike are completely black, just as the ones that originate from grave no. 35 in the mortuary temple of Seti 1 in the necropolis of Thebes and the four seeds stored in the Louvre, which are not further specified . The seeds from the temple of Seti 1 are threaded on a string and have been identified by Schweinfurth. The suggestion by Germer that at the time of Schweinfurth’s identification the red color was still visible, seems very unlikely as it implies that the discoloring of the seeds only started after more than 3,000 years. It is more likely that, taking its use as a bead into consideration, Schweinfurth assumed that he was dealing with the decorative rosary pea. Another possibility is that we are dealing with an African species, in which the following ones come into consideration: A. precatorius ssp. africanus Verdc. , A. somalensis Taub. , A. schimperi Bak. , A. pulchellus Thw. , A. bottae Defl ers , and A. fruticulosus Wall. ex Wight & Arn. . Seeds of these species are greenish or black and lack the decorative red color of the rosary pea and are only rarely seen at Egyptian markets. With respect to the seeds from Thebes that have been used as beads,greenhouse rolling racks it would make sense that they belong to the rosary pea indeed. Although there was no direct trade during the New Kingdom with India, it is possible that we are dealing here with indirect trade via the east coast of Africa south of the Sahara. Because the specimens of Berenike are not pierced, and thus do not indicate a decorative feature, it might well be that these seeds belong to one of the African species. That these seeds were used as gold weights in Berenike is merely speculative as it implies that precious metals were traded at Berenike, an assumption, however, of which no clear evidence is present up to now.In addition to its valued wood, Nile acacia is exploited for its reddish gum, which exudes spontaneously from the trunk and principal branches and is stimulated by artificial incisions. Additionally, it is exploited for its bark, which contains high concentrations of tannin and can be used for both tanning and dying, and for the pods that serve several purposes. Young pods can be eaten as a vegetable, and roasted seeds serve as a spice. Although green, unripe pods contain even 50 percent more tannin than the bark of the tree, its concentration is halved when the pods are ripe . At Berenike, the Nile acacia is only represented by ripe pods. Although bark and unripe fruits contain more tannin than the ripe fruits, obviously only ripe fruits were processed. The reason is a practical one. Harvesting the bark would kill the tree, whereas unripe fruits are difficult to pound. In fact, ripe pods are still offered for sale in large quantities in spice markets for tanning purposes.

The following procedure was described and partly demonstrated by an Ababda woman. First, pods are pounded with a stone on a piece of cloth or plastic . The pulverized fruits are mixed with water in a proportion of 1 k fruit to 10 L of cold water. A skin is then soaked in this mixture for two days. Next, the hairs are removed by hand. Then the skin is soaked again in a qarad mixture for about another week. Finally, the skin is sun-dried, after which it is suitable for all kinds of processing. Other Acacia species that can be used for tanning are A. oerfota Schweinf. and A. etbaica Schweinf., both well represented in the Gebel Elba area . Pod segments and seeds of the Nile acacia have been found in Egypt from predynastic times onward. Today, the Nile acacia grows in the Nile Valley; the desert west of the Nile, including the oases; and in the Sinai. According to Zahran and Willis the Nile acacia probably also once grew in the southern areas of the Egyptian Red Sea coast. Its exploitation, rather than ecological factors, would have been responsible for its absence in this area today. As only fruits have been attested from the late habitation period, the most likely option is that they were imported from the Nile Valley or at least from a remote distant place. In a reverse situation, namely that the fruits found were only from early deposits, over exploitation of a nearby population could be considered.Twisted acacia can easily be distinguished from Nile acacia by its fruits, even if only small fragments are preserved. The former has small, spirally twisted fruits, which are only slightly constricted between the seeds and bear a clear veining pattern. Nile acacia, on the other hand, is characterized by more or less glabrous pods, which are deeply constricted between the seeds. Another difference concerns the spines that are present on the branches. Twisted acacia has both long straight spines and small curved spines, whereas Nile acacia has only long straight ones. Both straight and curved spines were found in the samples of Berenike and Shenshef. It has not been tried to distinguish between the two subspecies of twisted acacia, namely, ssp. tortilis and ssp. raddiana Brenan, which were formerly treated on a species level. The twisted acacia is a common species that grows in the midstream part of almost every wadi branch around Berenike. Leaves, young shoots, and especially fruits provide a valuable, nutritious forage for all domestic animals . Trees are browsed by passing camels and sheep as well as goats that eat the pods that fall onto the ground. Drar states that Ababda nomads pull down ripening fruits with long, hooked sticks. Another way of harvesting could have been cutting off whole branches that were fed to animals. The disadvantage of this kind of exploitation is, however, that it reduces the tree population, which in this arid environment cannot be compensated by regeneration. Today, the Ababda nomads mainly exploit A. tortilis by harvesting dead branches to make charcoal. Wood of A. tortilis has a high calorifi c value and is the leading supplier of charcoal. Exceptionally, fresh branches are cut to make the skeleton of small houses that are covered by skins or, as is practiced nowadays, by mats and cloth. The presence of seeds, fruits, and spines in many loci of Berenike cannot be explained unambiguously.

The first class represents the number of species recorded from the particular region only

The first two sources refer to the major phytogeographical districts only, whereas the grass monograph uses a detailed subdivision. Because of this, the assignment of some grasses that are recorded for the Isthmic Desert remains problematic and should be added either to the Sinai district or to the Eastern Desert. The number of these grass species is low, however, and therefore has no significant influence on the calculations . In each phytogeographical district, plants have been divided into three classes. The second class shows the number of species also recorded from the Eastern Desert and Red Sea coastal strip. For the Eastern Desert this second class refers only to the number of plants also found in the Red Sea coastal plain, and for the Red Sea coastal plain the case is vice versa. The third class represents plant species found in the concerning region and other regions but excluding the Eastern Desert and the Red Sea coastal strip. Four phytogeographical districts are characterized by a reasonable number of indigenous plants, which give them their individual status: the Mediterranean coastal strip , the Nile region , the Gebel Elba , and the Sinai . In the other four districts, these percentages vary between 2 to 5 percent, indicating that on a presence/absence analysis the flora in these districts does not differ much. The flora of the Sinai comprises no less than 61 percent of the Egyptian flora. This high percentage, together with the rather high number of indigenous plant species,blueberry pot size reflects the variety of habitats in this district and may also justify a further subdivision of the Sinai phytogeographical district as has been published elsewhere .

The Mediterranean area, where plant growth is favored by the relatively large annual precipitation and mild temperatures in winter and summer, represents 49 percent of the Egyptian flora. Although the Egyptian deserts are characterized as hot and hyperarid, the desert flora is quite rich and diverse in plant numbers. Remarkably, the number of species in the Western Desert is considerably smaller than that of the Eastern Desert. With 39 percent, the number of species in the mountainous area of the Eastern Desert is more than twice as much as that of the Western Desert and almost equals the species richness of the Nile Valley and Delta. Even the number of species recorded for the small coastal plain bordering the Red Sea is considerable: 15 percent. The proximity to the sea, the variation in environmental conditions, and the effective concentration of rainwater in the wadi system seem to be favorable factors in this respect. The flora of the coastal plain is, for the most part, also present in the Red Sea mountains. At least 800 plant species occur in both these landscapes. Of these species, 32 percent is present in both areas and only 3 percent is unique for the coastal plain . The small percentage that is specific for the coastal plain can be attributed to the presence of salt marshes, an ecosystem that is characterized by its limited number of species. Certain species of the flora of the Red Sea mountains and the coastal plain are also present in the other phytogeographical districts. This is also true for the Gebel Elba area, where the Sudanian flora penetrates into the north. A group of large mountains, including the Gebel Elba, is located near the sea and causes much orographic rain. This is especially true for the north and northeast flanks, where even several moss species and ferns are quite abundant on the higher latitudes. A total of 443 different vascular plant species have been recorded for the Elba mountains, and its botanical richness is in marked contrast with that of the mountains north of this area. The barren Red Sea mountains north of lat 23° N have only a limited plant cover.

On rock surfaces where some soil has accumulated and in the many small affluents through which the runoff water drains into the main wadi beds, short-lived plants are present in particular. Most of the vegetation is, however, concentrated in the wadi branches and consists of both ephemeral and perennial vegetation. The number of flowering plants in a particular wadi is estimated by Schweinfurth at 60 to 100 species. The vegetation in these wadis also penetrates the coastal plain close to the Red Sea, where it is finally replaced by a salt-marsh vegetation. These salt marshes can be quite small, as is the case in the delta of Wadi Gimal, or may penetrate inland for several kilometers. Plant growth in wadis is determined by the catchment area as well as the thickness and composition of the wadi filling. Small, shallow wadis are dominated by drought-tolerant plant species. Broad and deep wadi beds on the other hand, which are filled with sand deposits and are fed by a large mountainous area, act as natural reservoirs for large quantities of water. In such deep wadis, only the upper layers dry up completely. Remarkably, perennials with long roots can reach the deep water table and are able to survive in these restricted environments year round. Trees, such as Acacia tortilis, are mainly found in the transitional zone between the elevated terraces and the central part of the wadis and along the convex sides as this fringe optimizes the availability of groundwater and protection against erosive flood waters. Smaller perennial species, such as Chrozophora plicata and Cornulaca monacantha, that have successfully established themselves in the central part of the wadi are often badly damaged by previous floodings. The thick, woody stems indicate that such specimens have a considerable age. The terraces of the coastal plain support almost no vegetation: only ephemeral vegetation is present, this being dependent on rainfall.To facilitate a better interpretation of the sub-fossil record of wild plant species from Berenike and Shenshef, a small-scale inventory of the recent natural vegetation has been made.

This inventory of the vegetation is based on plants collected around Berenike, Kalalat, Khesm Umm Kabu, Hitan Rayan, Shenshef, and Qariya Mustafa ‘Amr Gama and on the description of 16 relevés located at Berenike, Kalalat, and Shenshef . A relevé is a vegetation sample which is deliberately chosen as being a uniform and representative sample of the plant community being described . It is stressed that, apart from the descriptions of the relevés, these inventories are biased by differences in the duration of the visits paid to these sites. A short visit will certainly result in an incomplete inventory, as many plants are relatively small in size due to an adaptation to moisture shortage in particular and may easily be overlooked if they are present in low frequencies. Gisekia pharnaceoides may serve as an example. This plant species is the only member of the Gisekiaceae in Egypt and is recorded for the Eastern Desert and the Gebel Elba area . Several specimens of this prostrate annual were found around Fort Kalalat and measured sometimes hardly more than 1 cm , whereas in the wadis of Gebel Elba this plant species is considerably larger. Under more favorable conditions, the stems may reach a length of 10 to 40 cm . The possibility that this particular plant species has been overlooked at other sites cannot be ruled out. The Roman settlements that have been less-intensively investigated are Khesm Umm Kabu, Hitan Rayan, and Qariya Mustafa ‘Amr Gama. The first two settlements were visited twice and the last one only once. In all of these locations, most of the plants that have been recorded originate from the wadi branches. Berenike, Shenshef, and Kalalat have been visited during all excavation seasons. Berenike was visited during the excavation seasons from 1994 to 1998, making it possible to look at the vegetation from late December until early March. The inventory covers the salt-marsh vegetation and the vegetation in the branches of Wadi Mandit west of Berenike and Wadi Umm el-Mandit south of the site. The inventory of the vegetation in the surroundings of the watering station Kalalat is based on the plant species that grows inside the fort and on the sand that has accumulated along the outside of the external walls. Additionally,plant raspberry in container plants from the wadi branches around the watering station and on the fl at coastal plain were listed. A thorough study of the plants around Shenshef could be made because one of the visits lasted for two weeks and took place after heavy rainfalls. Plants were collected from the main wadi branch and its affluents, from the slopes of the mountains, and from the terraces occupied by the former settlement . The completeness of the inventory is also affected by the time of the year. Although all sites were visited during the excavation seasons, which lasted from late December to early March, and the life cycle of most annuals coincide with this relatively mild part of the year—due to lower temperatures and availability of water— plant species may still have been missed because they had not yet germinated or were already withered. Identification of very young plants, lacking characteristic features of flowers and fruits was, however, still possible in most cases by using leaf characteristics and by looking at the germinated seed or fruit still attached to the root, which was easily gathered from the loose sand.

This last method proved to be successful especially with respect to grasses with a resistant fruit coat, such as Centropodia forsskalii, Aristida spp., and Stipagrostis spp. All Egyptian members of both last genera can easily be identified to the level of species on the basis of their fruit characteristics . Some other young plants lacked such characteristic features, such as Fagonia spp., and could therefore not be identified beyond the genus level. Finally, the sampling is further distorted by differences in the vegetation between years, which in turn are related to variations in precipitation. This seasonal phenology is witnessed in nonsaline ecosystems of the deserts . Some plant species, such as Arnebia hispidissima and Zilla spinosa, were observed every season, though in varying density, whereas others, such as Senna italica and Cleome amblyocarpa, were infrequently present in the same wadi branches.The plant composition in the relevés is summarized in Table 2.1. Although the size of the relevés is relatively small for a desert environment, the number of species is surprisingly high. It varies from 5 to 19 with a mean value of 13. Also the plant cover can be reasonable high as it varies from 10 percent to 85 percent. It has to be realized, however, that these inventories were made after heavy rainfall, resulting in exceptionally rich vegetation, which found its expression in both plant cover and in the proportion of ephemeral species. Ephemeral species have a short life cycle and need water only during a part of the year. The distinction between ephemerals and perennials is not exclusive, as some perennials such as Zilla spinosa, Pulicaria undulata, Farsetia ramosissima, Citrullus colocynthis, and Trichodesma africanum can also behave as ephemerals if there is a shortage of water. Common ephemeral species are Zygophyllum simplex, Lotononis platycarpa, Triraphis pumilio, Eragrostis ciliaris, Arnebia hispidissima, and Astragalus vogelii. Zilla spinosa is a perennial species that is present in most relevés, though mostly at a young stage of development as perennial species tend to develop more slowly. Z. spinosa is absent in the vicinity of the sea. The diversity of grasses is striking. A total of 16 species has been recorded, of which 15 grasses are also present in the relevés. With the exception of Rostraria pumila, which grows as a small grass in a wadi branch near the watering station Kalalat, all these grasses are C4-species. Such species are adapted to arid conditions by a different photosynthetic pathway and a regulation mechanism that minimizes evaporation. The distribution of the C4-grass species seems to be influenced primarily by temperature . In the southern part of the Eastern Desert C4-grasses make up 81 percent of the species, in the Red Sea coastal zone 93 percent, and in the southerly located Gebel Elba area even 95 percent. The high proportion of C4-grass species in the vicinity of the Roman installations corresponds quite well with these percentages. Most grass species become reasonably tall and produce a substantial amount of biomass for grazing, such as Aristida spp, Stipagrostis spp., Dichanthium foveolatum, Cenchrus ciliarus, and Panicum turgidum. In years with limited or no autumn rain, those species are heavily grazed .

This is expected as the 28S-D2 gene region is generally more highly conserved than the COI region

Total DNA was isolated using an EDNA HiSpEx Tissue Kit , following the manufacturer’s protocol. This method is non-destructive, allowing slide mounting and morphological examination of the specimen after extraction. After DNA extraction, two separate gene regions were amplified using PCR: the conserved 28S-D2 domain of the large rRNA subunit and the cytochrome c subunit I of mitochondrial DNA . Post DNA extraction, specimens were transferred into 70% ethyl alcohol to prepare for slide mounting. Specimens were cleared in 5% NaOH for 12 h, processed through an alcohol dehydration series, placed in clove oil, and slide mounted in balsam . Specimens were identified to species using the online and interactive key of the “Thrips of California” . A representative from each haplotype generated from DNA analysis for this work was slide mounted and placed at the University of California, Riverside Entomology Research Museum as vouchers, and their collection numbers are included in Table 6-1. Representative species-specific sequences are deposited in GenBank. A BLASTN 2.2.19 search was used to compare sequences to existing Scirtothrips and Neohydatothrips sequences deposited in GenBank. Sequences were initially aligned manually in Bio-Edit version 6.0.7 . The COI sequences were all of equal length and contained no stop codons, with the first position equal to codon position one,blackberry container making alignment simple. The 28S-D2 included six ambiguous regions that were variable enough to make alignment difficult.

The six regions were coded as single multi-state characters in the analysis, or INNASE coding . This mixed alphabetical and numeric multi-state coding was treated as unordered and combined with the other data. Parsimony analyses using 1,000 random addition sequences and random starting trees were done on 1) the complete molecular data set , 2) molecular data with only 28S-D2 and 3) molecular data with only COI. All analyses were performed using PAUP 4.0* , with the complete matrix verified using TNT with a new technology search . Bootstrap values were generated in PAUP 4.0* using 1,000 BS replicates with two random heuristic searches for each replicate. Maximum Likelihood analyses and associated bootstrapping, were conducted on the complete molecular data set with RAxML v.7.2.7 via the CIPRES Science Gateway . Two gene partitions were included , and data were analyzed with 1000 rapid bootstrap replicates. None of the specimens collected from citrus in Texas were Scirtothrips; all thrips in the vials collected from multiple types of citrus were Frankliniella occidentalis and were therefore not included in any analyses for Scirtothrips citri or near citri. All specimens from California and Arizona keyed to Scirtothrips citri. The distinguishing features for S. citri include: 1) both sexes fully winged, 2) body mainly yellow without dark markings, 3) antennae 8-segmented with segments III – IV having forked sensorium and segments III-VIII grey, 4) head wider than long with ocellar triangle and postocular region having closely spaced sculpture lines, 5) three pairs of ocellar setae present with pair III close together between the anterior margins of the hind ocelli, 6) pronotum with closely spaced sculpture lines and the posterior margin with four pairs of setae, 7) metanotal posterior half with irregular longitudinal reticulations and median setae originating behind the anterior margin, 8) first vein of the forewing with three setae on the distal half , the second vein with three widely spaced setae and the posteromarginal cilia wavy, 9) abdominal tergites III – VI with median setae close together, tergites II – VIII with lateral thirds covered in closely spaced rows of fine microtrichia, these microtrichial fields had four discal setae, the posterior margin had a fine comb and the lateral discal microtrichia extended medially and lastly 10) abdominal sternites were without discal setae and the posterior margins were without a comb of microtrichia .

Morphological differences from the above on specimens from Florida , Mexico and Nicaragua could not be found and all were keyed to S. citri. The Florida citrus thrips collected from Mimosa sp. did not fit well into the Thrips of California key. The metanotal median setae arising at the anterior margin are in contrast to that of S. citri, where the metanotal median setae arise behind the anterior margin. All other characters appeared to be in congruence with S. citri. The thrips collected from Mimosa in Miami, Florida were keyed to Scirtothrips near citri. The specimens from Turkey were keyed to Scirtothrips citri using the Thrips of California key. Relationships were largely congruent among all analyses with the following exceptions: the group ‘Turkey_Hatey_citrus1_5’ in the combined tree and in the 28S-D2 alone tree as a sister group to the S. citri clade but these were placed without support along with S. aff dorsalis . This group switched positions, depending on the type of analysis. Results regarding S. citri will be discussed in terms of the combined parsimony analysis. The parsimony analysis of the complete molecular data set, including ambiguous region coding, resulted in 114 most parsimonious trees . The overall = consensus solution of all of the trees had no significant areas of conflict and collapsed with strong support for branches and monophyly of the groups. Further, analyses based upon exclusion of one gene region at a time resulted in trees with the following monophyletic groupings 1) Arizona and California, 2) Mexico-Nicaragua and 3) Florida . The group collected from Miami, Florida was sister to Scirtothrips bounites Mound & Marullo in all analyses. The RAxML analysis resulted in a single tree with a final Ln likelihood of -9,623.28. The parsimony and RAxML analyses both produced similar trees for the specimens collected in Arizona, California, Florida , Mexico and Nicaragua.

Clades Arizona and California, Mexico-Nicaragua, Florida and Florida , while morphologically indistinguishable, are highly molecularly divergent differing by 10-40 base pairs. The analyses conducted on molecular data confirmed that Scirtothrips citri in the Americas is not a single species but a complex of at least three molecularly divergent groups . The parsimony and maximum likelihood analyses both produced similar trees for the specimens collected in Arizona, California, Florida , Mexico and Nicaragua with the specimens collected from Turkey switching positions, depending on the type of analysis. Arizona and California specimens were morphologically identified using the Thrips of California key as ‘California’ citrus thrips and the Quincy , Mexico and Nicaragua specimens also keyed out as ‘California’ citrus thrips but this is not surprising as the key was developed for the known thrips found in California. The Miami citrus thrips collected from Mimosa sp. did not fit well into the Thrips of California key, which is again not surprising. Our molecular results suggested the specimens collected from Mimosa in Miami are S. bounites, although these specimens were identified as S. citri by a collaborator . To our knowledge, S. bounites has not been found in California to date but was collected from mango in Mexico . This suggests that without any consistent morphological differences, a cryptic species complex is likely present. The specimens collected from Turkey were morphologically keyed to S. citri but were grouped differently based on parsimony analyses of the two gene regions separately. The Turkey specimens in the 28S-D2 alone analysis were placed as the sister group to the overall ‘Scirtothrips citri’ clade however, in the COI alone analysis, the Turkey specimens are grouped within the Scirtothrips dorsalis Hood clade. These specimens do not share S. dorsalis specific characters such as microtrichia covering the sternites and straight wing cilia. In the parsimony and maximum likelihood combined gene region analyses,planting blueberries in a pot the Turkey specimens are placed as the sister group to the overall ‘Scirtothrips citri’ clade. The Turkey specimens did not appear to be different than S. citri morphologically and were included in the analysis because the group may be a new/related species to the ‘Scirtothrips citri’ clade. A comparison of bootstrap values between the individual gene trees suggests that 28S-D2 is driving the backbone of the combined tree. There is a need for further research into the relationships between the groups of S. citri presented here. Based on our molecular data, citrus thrips in California and Arizona are clearly different than those found in Quincy and Mexico-Nicaragua but morphological data suggests they are the same. These conflicting pieces of information illustrate the fact that morphological identifications may not be accurate enough especially when dealing with some organisms of economic importance. For example in a hypothetical situation, if specimens were detected and incorrectly morphologically identified as a species of economic concern but actually are a part of a non-economic clade, not distinguishing these groups could have serious import/export implications. Utilizing molecular markers for species identification and separation could be an immensely useful tool if/when morphological identifications are unclear. There are a few instances of cryptic species investigations with thrips and the development of molecular keys for many different types of thrips pests have aided in our understanding that thrips populations from various areas, or thrips collected from different host plants, are not necessarily all the same species, even if morphological analysis suggests this is the case.

Between 1994 and 2001 a joint expedition of the University of Delaware and Leiden University conducted eight seasons of excavations at the Ptolemaic-Roman emporium of Berenike on the Red Sea coast of Egypt. It was a great opportunity to work in an area that is usually closed for research, because it is part of Egypt’s border area with Sudan and is protected by the military. The Berenike Project was created as an interdisciplinary research effort involving a large number of archaeological specialists, who discussed their findings and interacted both on and off the site. Since many of the specialists were present on-site at the same time, preliminary results and conclusions could be shared and discussed often while excavations were in progress. This valuable collaborative effort could then be shared with those conducting the actual excavations. The annual reports on the excavations and specialists’ results have been published in six volumes, while the seventh, and last, is in preparation . These reports provide comprehensive interpretations and conclusions that have emerged over the years. At the end of this phase of fieldwork, however, it is time to publish an overview, and in the coming years several specialists will produce final reports. Although such final publications concentrate on specific artifact or material groups, they have the benefit of ongoing discussions among various Berenike team members. The result, in each case, will be a specialist’s report rooted in a broad basis of multidisciplinary insights . As the present volume amply demonstrates, the archaeobotanical research at Berenike provides essential information for our understanding of the subsistence patterns of the inhabitants, as well as the long-distance trade. The occurrence of exotic plant remains at Berenike changed through its history, with clear peaks in the first/second and fourth centuries CE. The results of the archaeobotanical research, therefore, provide extremely important information that supplements, complements, and, in some cases, changes the image that emerges from the ancient written sources. The variety of plants traded was larger than expected. Analysis of the well-preserved archaeobotanical remains found in our excavations at Berenike also indicates the regions with which the traders were in contact and when. This book will not only be interesting for archaeologists and archaeobotanists, but the comparison with written “classical” sources provides a fascinating read for historical and classical scholars as well. In some cases the Berenike finds provide evidence to propose new translations for Greek or Latin botanical terms, and correct current interpretations. Perhaps the most important result of the archaeobotanical research conducted at Berenike is the attested correlation between the harbor function, where goods were transferred from ship to caravan, and the amount and variety of goods that can be recorded. The items lost, purchased, or pilfered over the centuries at Berenike represent a micro version of the goods that were in demand throughout the Roman Empire as a whole. To research the imports used in Rome effectively, it is effective to start the investigation at the periphery of her empire: at Berenike, the harbor town through which many of these products once passed.

A laurel sumac seedling was placed into each of eight styrene cages per block

Citrus thrips are adapted to hot and dry environments and thus, they are less likely to have evolved natural tolerance to fungi, whereas, avocado thrips thrive in a very wet environment where exposure to fungi is more likely. The differences may be due to different habitat adaptations and the different origins of the two thrips species . We find it interesting that two congenerics have such widely different habitat preferences and this may explain differences in fungal tolerance. Differences were seen when citrus thrips and avocado thrips were placed on leaves of their associated host plants, then placed separately in sealed zip-lock bags where the moisture that condensed in the bags was lethal to citrus thrips but not to avocado thrips. Thus, it is possible that avocado thrips, due to their adaptation to living in cool and wet climates , have a higher tolerance to fungal pathogens, as they may encounter them more frequently than citrus thrips, which prefer a hot and drier climate . Many researchers have investigated alternatives to traditional insecticides such as biopesticides, i.e. natural or organismal methods of controlling pest populations. The utilization of entomopathogens against thrips is not a new concept; entomopathogenic fungi, such as, Metarhizium anisopliae Sorokin , Neozygites parvispora Remaudière & Keller , Verticillium lecanii Viegas , and Paecilomyces fumosoroseus Brown & Smith have also been used in laboratory and greenhouse trials with much success,large plastic pots for plants whereas field trials have shown limited successes.

However, various strains of B. bassiana have been shown to effectively control western flower thrips on greenhouse ornamentals and peppers , and several reports indicated that F. occidentalis, Thrips palmi Karny and T. tabaci Lindeman were successfully controlled under field or laboratory conditions . In conclusion, both citrus and avocado thrips can be infected by B. bassiana but high doses may be required, especially for avocado thrips. These high doses are difficult to obtain outside the laboratory and application of such doses would be costly. We believe B. bassiana is not a sufficiently effective alternative to traditional insecticides to warrant further study with avocado thrips, particularly because the commercially available strain GHA gave poor control on avocado thrips, but it may have potential against citrus thrips in an integrated pest management program. Further studies are warranted to determine if GHA could be used in field control of citrus thrips. Citrus thrips, Scirtothrips citri , has been recognized as a major pest of California citrus since the 1890s and is also known to scar mango fruits . Historically, highbush varieties of blueberries could only be grown in regions too cold for citrus production . However, breeding efforts to cross the northern highbush blueberries with several other Vaccinium species led to the development of heat-tolerant highbush blueberry varieties . This has enabled the establishment of a blueberry industry in the San Joaquin Valley, a region where both citrus and citrus thrips flourish . The known host range of citrus thrips has broadened and in recent years, they have become a significant pest of blueberries planted in the San Joaquin Valley of California .

Citrus thrips feed on blueberry foliage during the middle and late portions of the season causing distorted, discolored, and stunted flush growth and poor development of fruiting wood required to obtain the subsequent crop. Repeated pesticide applications of the few effective and registered pesticides to reduce thrips populations pose a concern regarding pesticide resistance management, and this issue is relevant not only to the blueberry industry but also for the 108,665 ha of California citrus which has experienced repeated documented cases of pesticide resistance in citrus thrips populations . Currently, there are no integrated pest management plans available for control of citrus thrips in blueberry, probably due to the recent nature of this crop-pest association. With a limited number of pesticides available for thrips control and the frequency of insecticide resistance shown by thrips, populations should be monitored carefully, treatments limited to populations of economic concern, and applications timed optimally . Appropriate cultural practices and conservation of natural enemies should be practiced in concert with the use of pesticides only on an as-needed basis. Understanding citrus thrips’ life history in the blueberry system to determine where and if susceptible stages could be exploited, is one of the first steps in the development of alternative methods to the use of traditional insecticides. In citrus, citrus thrips pupation occurs on the tree in cracks and in crevices, however, the majority of thrips drop as late second instars from trees to pupate in the upper layer of leaf litter below trees and move upward onto the plant after adult eclosion. Propupae and pupae are rarely seen, move only if disturbed, and do not feed.

Pupation in the upper layers of the soil surface may create the ideal interface for control using the entomopathogenic fungus Beauveria bassiana Vuillemin due to this vertical movement of the citrus thrips. However, blueberry plants have much different plant architecture than citrus trees and citrus thrips pupation behavior has yet to be studied on blueberries. In the U.S., pressure is increasing to move away from broad-spectrum insecticides and focus on alternative methods of control. Earlier work with B. bassiana determined that the commercially available strain, GHA , was the most effective of six strains tested in laboratory trials against citrus thrips . The goal of this study was to determine if this strain of B. bassiana could be utilized effectively against citrus thrips in California blueberry production. To achieve this objective, several factors of importance to fungal efficacy were evaluated before commencement of our field trial: 1) location of citrus thrips pupation in commercial blueberry plantings, 2) field sampling locations and methods, 3) fungal formulation and timing of application, and 4) density of product used and method of thrips infection. We then conducted a field trial evaluating the potential utility of the GHA strain of Beauveria bassiana in commercial blueberries for citrus thrips management as a possible alternative to the use of traditional insecticides. Because of the complex arrangement and number of blueberry canes arising from the rhizome of commercial blueberry plants, we first evaluated movement of second instar citrus thrips on potted single cane blueberry plants in the laboratory. Known numbers of late second instar citrus thrips were released onto the leaves of potted blueberry plants in the lab. Paper sprayed with Tangle Trap sticky coating was placed a) at the base of the plants with a ring of sticky tape around the base of the stem and partially on the stem of the plant to capture any insects crawling down, and b) extending from the base of the plant horizontally outward above the pot surface to ensure complete coverage of the area covered by the plant canopy . This experiment was replicated on a single potted plant over time on 7 dates . Data were analyzed using Fisher’s exact test using SAS 9.2 . At our planned field trial site that would later be used in the B. bassiana trial, pupation emergence cages were used to sample insects moving off foliage towards pupation sites and later emerging out of the soil after pupation. Cages were made from Schedule 40 white PVC pipe with a diameter of 10.2 cm with cages cut to a height of 5.1 cm. The cage was then topped with a double-sided sticky card cut to fit, which was fixed into place with two elastic bands. Four lines of four cages were pushed into the soil to a depth of approximately 1 cm immediately adjacent to each other at the base of a blueberry plant and oriented in a cardinal plane to determine which direction showed the most thrips activity. The four adjacent cages in a particular plane were used to assess thrips movement in the understory of the blueberry plant in each directional. The study was replicated on 5 plants on a single date and conducted just prior to the commencement of the field trial. Data were analyzed with a nested ANOVA using SAS 9.2. In a greenhouse trial, Mycotrol O ® was applied directly to the soil surface as raw spores and compared to the same product colonized onto millet seed,plant pots with drainage also using soil application. Millet seed colonization used the Stanghellini and El-Hamalawi method as described below. The colonized millet seed, when allowed to imbibe water and incubate in the laboratory, can support 1.0 x 106 conidia/seed . Based on Stanghellini et al. with modification, we held the GHA colonized millet seed in containers such that the seed mat was at a depth of no greater than 2.54 cm.

The seeds were wet with the consistency of very thin slurry and were gently stirred three times per day for four days to ensure they imbibed water properly so that mycelial growth and sporulation would occur. Sporulation was confirmed by slide mounting random sections of mycelia and checking for condia formation under the microscope. Once spores were initially observed, the seed was held an additional three days so that sporulation could continue before use of the colonized seed in the field study. Mycotrol O® was applied in the maximum recommended field rate for high thrips levels of 2.84 L of material in 378.5 L of water. The colonized millet seed was tested in the greenhouse to determine if late second instar citrus thrips would become infected if they crawled over or through the seed when it was placed at the base of a laurel sumac seedling. A single small laurel sumac seedling, about ~10 cm tall, was placed into each of ten, 9.5 x 9.5 x 18 cm styrene cages with 6 cm diam air holes on all four sides that were covered with ultra fine mesh screening . Small holes were made in the bottom of the container and covered with pebbles to allow for drainage, then soil was added to a depth of 7.62 cm and the top of the container was covered with a removable lid. The base of each plant was completely surrounded by either B. bassiana colonized millet seed or with uncolonized seed . A minimum of 20 late second instar thrips were released onto the leaves of each plant, and were left until enough time had passed for the thrips to molt to the propupal stage. The seedling was then cut at the soil line and examined for pupating thrips; the removable lid of the cage was sprayed with Tangle Trap sticky coating to collect any emerging adults after 5 days so infection could be measured. The study was replicated on 5 dates . Data were analyzed using 1-way ANOVA with time as a factor and means were separated using Tukey’s Least Significant Difference test using SAS 9.2. To determine the optimum number of colonized millet seeds needed for close to 100% infection when thrips were seeking pupal refuges off the plant, varying amounts of colonized seed were evaluated in a greenhouse trial based on the size of the seed once it had imbibed water and sporulation had occurred. After water inhibition, nine seeds completely filled one square cm of soil surface. There was a 0.5 cm buffer area around all sides of the cage, which was kept clear of seed to provide a 9 x 9 cm grid of seed on the soil surface below the plant. All but two leaves were plucked from the seedling. Small holes were made in the bottom of the container, which was covered with pebbles to allow for drainage. The 9 x 9 cm2 grid was created from wire screen and differing amounts of sporulating seed or seed alone were placed on the light imprint made from the wire screen on the soil surface. Two replicate seedlings per treatment were set up per date in a complete block design . Plants were watered every third day. A minimum of 20 late second instar thrips were placed onto the leaves of the plant, and were left until enough degree-days had passed for the thrips to molt to the propupal stage, typically about 5 days. The seedling was then cut at the soil line and examined for pupating thrips; the removable lid was sprayed with Tangle Trap sticky coating to collect any emerging adults after another 5 days.

Dormant-season cover crops in the middles minimize runoff from winter rains

Although anthocyanin content was not affected, anthocyanin composition was modified by treatments from mid-ripening to harvest . Berry skins of ST and LRST grapevines showed a lower 3’4’5’/3’4′ ratio leading to increased proportion of cyanidins and peonidins in detriment of malvidins which was the most abundant anthocyanin found in berry skins . During the monitored period, different canopy management practices modified berry flavonol content . The berries from LRST grapevines showed the greatest berry skin flavonol content, while, at harvest, the flavonol content of LR, ST, and LRST was similar and greater when compared to the UNT content. Not only canopy management practices modified flavonol content but they also affected their composition. The LRST treatment had a higher proportion of kaempferol and quercetin from midripening to harvest and lower of proportion of myricetin after veraison . As expected, berry IBMP content decreased throughout ripening with all the canopy management practices tested in this study . However, we found the significant differences among treatments after veraison and at harvest. The LRST treatment resulted in the lowest IBMP content from mid-ripening to harvest. Correlation analysis between the monitored variables at harvest revealed a strong relationship between canopy architecture variables and berry flavonol content . Moreover,drainage for plants in pots canopy porosity was strongly correlated to the kaempferol proportion in berry skins . On the other hand, a lower yield due to canopy management practices was related to decreased IBMP and increased flavonol content .

Finally, a strong relationship was found between TSS and TA with the leaf to fruit ratio . Finally, a higher solar exposure estimated as the kaempferol proportion was strongly correlated with decreased anthocyanin berry contents and yield .Analysis of labor operations cost of canopy management practices indicated that the most expensive canopy management practices was the LRST where growers received a 53% lower income per hectare. Thereby, productivity data provided evidence that the cost of producing a kg of anthocyanin and removing a µg of IBMP was 10-fold greater in LRST compared to UNT per ha .Yield components were mainly affected by shoot thinning practices, decreasing the number of clusters and yield per vine leading to unbalanced vines according to the previous studies . Yield per meter of row is increased quasilinearly with the increase in shoot density per meter of row as indicated by previous studies . The lack of effect of LR on yield was corroborated by several studies when a late leaf removal was applied. Moreover, Yu et al. and Cook et al. reported that grapevines may produce more leaves than required, especially in warm climates, therefore, the increase in canopy gaps and the diminution of external leaf layers did not elicit decreases in yield as they were not severe enough reductions to the functional leaf area. The RI between 5 and 10 is considered optimum for vine balance . Therefore, RI and leaf area to fruit ratio data reported with the grapevines subjected to shoot thinning were under cropped that led to lower yields. In our study, Cabernet Sauvignon vines were not able to modulate their vegetative biomass in response to canopy management practices applied. Previous studies showed that pruning mass values up to 1 kg/m of row were considered optimal under warm climate . In our experiment the pruning mass per meter of all treatments ranged from 0.5 to 0.7 kg/m without differences between treatments. Moreover, although the shoot counts were obviously different between treatments, we did not find differences in the pruning mass, that suggested lower lateral expansion and/or reduced shoot diameter with an increasing number of shoots as previously reported Brillante et al. . Consequently, we found that the mass of each shoot ranged from 28 and 25 g in UNT and LR, respectively, to 45 and 42 g in ST and LRST, respectively, corroborating work by Brillante et al. .

Martınez-Lüscher et al. reported negligible variation of berry mass of Cabernet Sauvignon due to higher solar exposure under irrigated viticulture. Similarly, berry masses remained unaffected by a higher solar exposure of the cluster due to canopy management practices unless they were directly exposed to sunlight where berries may suffer dehydration as previously reported by Mijowska et al. . This has been attributed to the effect of the higher temperatures with subsequent increases in berry transpiration that affected cell division and elongation . Under our experimental conditions, shoot thinning treatments hastened berry ripening by enhancing the TSS to ca. 2.5°Brix and decreasing must titratable acidity by 0.6 g•L−1 at harvest. Thus, overexposure has been related with higher pH due to the elevated temperature that berries overcome and the subsequent organic acid degradation . Nevertheless, Wang et al. recently suggested that changes on the source-to-sink ratio induced by shoot thinning might have more influence on berry maturity than the change in the microclimate they reported.Cultural practices have been related to increased anthocyanin content . However, in agreement with other studies , under our experimental conditions, berry anthocyanin content did not increase due to LR, ST or LRST. Similarly, anthocyanin content was not affected by mild exposure in berries collected from the commercial vineyardeither. Increasing exposure was detrimental for anthocyanin content as the overexposed berries were subjected to higher temperatures that may have impaired their accumulation . The anthocyanin berry content at harvest is the result between synthesis and degradation rates. It was reported anthocyanin synthesis may be up-regulated by greater exposure . Therefore, ST and LRST increased the anthocyanin content at mid-ripening because of the increasing solar exposure . Additionally, it was recently highlighted that some members of the dihydroflavonol reductase and UFGT genes required for anthocyanin biosynthesis were moderately up-regulated in LR treated berries leading to increases of anthocyanin content at mid-ripening . However, at harvest, no significant effect of canopy management practices on anthocyanin content was found, and this result is corroborated by Pastore et al. who reported no beneficial effect due to higher cluster exposure in warm climates.

Although cultural practices may induce different cluster temperatures by increasing exposure, we did not find a clear relationship between exposure and cluster temperature when kaempferol proportion are low suggesting that results of this work were mainly explained by different exposures. Nevertheless, under elevated temperatures, a down-regulation of anthocyanin biosynthesis and enhanced rates of degradation have been reported . Those authors suggested that high temperature induced anthocyanin degradation by enhancing the expression of VviPrx31 and consequently the peroxidase activity. Likewise, overexposed berries with kaempferol proportions greater than 10% were subjected to higher temperatures that dramatically decreased anthocyanin content. Matus et al. reported that flavonol content increased by two-fold in exposed berries compared to non-exposed. Our results corroborated this finding partially, depending on the level and duration of exposure, canopy position of the berries, and orientation of the vineyard. Therefore, when flavonol proportion was below 10% of kaempferol, flavonol content increased; but would decrease after this inflection point due to degradation. Matus et al. further indicated that this increase in flavonol may be driven by the up-regulation of MYB12 and flavonols synthase 4 due to the greater exposure suggesting that FLS4 could be a target of MYB12 in grapevine. Accordingly, Sun et al. found that increased accumulation of flavonols in light exposure berries, were accompanied by the up-regulation of several genes of the FLS gene family suggesting that they may be functionally redundant in response to light signal. During the experiment conducted in the 2019 growing season, the kaempferol proportion increased in LR and ST treatments,growing raspberries in pots but largest increase was measured when ST and LR were applied concurrently. Likewise, the higher the degree of exposure degree a greater kaempferol accumulation was observed during the 2017 growing season. The increase in kaempferol in total proportion of flavonols was accompanied with a concomitant decrease of quercetin and myricetin proportions. These results are corroborated with our previous work performed on Merlot and Cabernet Sauvignon. , and by others on Cabernet Sauvignon, Nero d’Avola, Raboso Piave, and Sangiovese in Italy . We previously reported the proportion of kaempferol was a feasible tool for accounting the solar radiation received by berry due to the greater canopy porosity and this corresponded to the 1930 W·m−2 of global radiation accumulated at the research site in Experiment 3. On the other hand, the higher proportion of quercetin derivatives in detriment of myricetin derivatives found in LR vines has been related to down regulation of F3’5’H family genes . Previous work on red grapevine berries, indicated that IBMP content decreased with greater solar exposure due to the canopy management practices during berry ripening . In our work, the lowest IBMP content was measured in LRST berries. Our results indicated a negative and linear relationship between leaf to fruit ratio and IBMP content. Conversely, the relationship between kaempferol proportion and IBMP was not significant. Therefore, our data suggested that the decrease of IBMP content was better explained by changes in the source-sink balance rather than differences in solar exposure. Likewise, Koch et al. provided evidence that solar exposure affected IBMP content to a greater extent when canopy porosity was enhanced before fruit set and not during berry ripening corroborating our results.

The lower berry IBMP content was explained by a diminution of the accumulation rates rather than increased rates of degradation due to canopy management practices and restriction of applied water between fruit set and veraison in a warm climate.Vineyard-fl oor management strategies, such as weed control and cover-cropping, have wide-ranging impacts both inside the vineyard, in terms of crop management and productivity, and outside the vineyard, in terms of runoff and sediment movement into streams and rivers. The increasing importance of water-quality issues statewide, including in Monterey County where the Salinas River drains into the Monterey Bay National Marine Sanctuary, highlights the need for management strategies that limit environmental impacts. Growers are interested in alternative weed-control practices and cover crops, but they need information in order to balance benefits with the economic realities of wine-grape production. We established a 5-year experiment in a commercial vineyard in Monterey County with the intent of identifying effective practices that can be integrated into the cropping system without negatively affecting winegrape production. The vineyard floor consists of two zones: the rows, a 2- to 4-foot-wide swath underneath the vines, which are managed primarily to control weeds by herbicide applications or cultural practices ; and the middles, interspersed between the rows, which are vegetated by cover crops or resident vegetation in the dormant season, and are tilled or left untilled in spring. Growers manage weeds in rows to reduce competition for water, nutrients and light , and to prevent tall-statured weeds such as horse weed  from growing or climbing into the canopy, where they interfere with harvest. Growers transitioning to more sustainable production systems need information on how management practices affect the physical properties, health, organic matter and water retention of soil. We monitored soil microbial activity for arbuscular mycorrhizal fungi and soil microbial biomass, since weed control and cover-cropping can affect populations of beneficial soil microbes in annual crops . Many California growers are also willing to plant cover crops because they protect soil from nutrient and sediment loss in winter storms , suppress weeds , harbor beneficial arthropods , enhance vine mineral nutrition and increase soil organic matter . Competition between vines and cover crops for soil moisture in spring, when both are actively growing, can lead to severe water stress and reduce grape production . However, wine-grape production is distinct from other cropping systems because water stress may be imposed to enhance wine composition ; this practice has been studied mostly in high-rainfall regions of California. The vineyard production region of Monterey County, in contrast, has low rainfall , and growers must weigh the benefits of cover crops with the possible need to replace their water use with irrigation. In addition, growers must decide on the type of vegetation to utilize in the middles. Resident vegetation is cheap and generally easy to manage. Cover crops can provide specific benefits such as nitrogen fixation or high biomass production and vigorous roots . There are many choices for cover crops in vineyard systems, ranging from perennial and annual grasses, to legumes . Each species has strengths and weaknesses, as well as associated seed and management costs.