An account of the genus Cistanche (Orobanchaceae) in Iraq and taxonomic considerations in the Middle East.

Species limits in the genus Cistanche are poorly understood, despite the plants' long history of use in traditional herbal medicine and food across their range. Here we present a taxonomic account for the genus Cistanche in Iraq, where several taxa have been reported, most of them doubtfully. Using herbarium specimens, images of living material, and taxonomic literature, we found evidence of only one species occurring with certainty in Iraq: Cistanchetubulosa. We found no evidence for the occurrence of other Cistanche species in Iraq, including a putative new entity reported for the region. Our work highlights inconsistencies in the literature, and underscores the importance of examining multiple stable characters for delimiting species in the genus Cistanche.

Repeated upslope biome shifts in Saxifraga during late-Cenozoic climate cooling.

Mountains are among the most biodiverse places on Earth, and plant lineages that inhabit them have some of the highest speciation rates ever recorded. Plant diversity within the alpine zone - the elevation above which trees cannot grow-contributes significantly to overall diversity within mountain systems, but the origins of alpine plant diversity are poorly understood. Here, we quantify the processes that generate alpine plant diversity and their changing dynamics through time in Saxifraga (Saxifragaceae), an angiosperm genus that occurs predominantly in mountain systems. We present a time-calibrated molecular phylogenetic tree for the genus that is inferred from 329 low-copy nuclear loci and incorporates 73% (407) of known species. We show that upslope biome shifts into the alpine zone are considerably more prevalent than dispersal of alpine specialists between regions, and that the rate of upslope biome shifts increased markedly in the last 5 Myr, a timeframe concordant with a cooling and fluctuating climate that is likely to have increased the extent of the alpine zone. Furthermore, alpine zone specialists have lower speciation rates than generalists that occur inside and outside the alpine zone, and major speciation rate increases within Saxifraga significantly pre-date increased rates of upslope biome shifts. Specialisation to the alpine zone is not therefore associated with speciation rate increases. Taken together, this study presents a quantified and broad scale perspective of processes underpinning alpine plant diversity.

Dataset of pollination traits in Fabales.

The data presented in this paper is supporting the research article "Reconstructing an historical pollination syndrome: keel flowers" (Aygören Uluer et al., 2022). We present a dataset containing information on number of species, geographic distribution, floral type (keeled or not), presence or absence of fused petals, floral symmetry, presence or absence of a pentamerous corolla (petals+petaloid sepals in Polygalaceae), androecium type, presence or absence of enclosed reproductive organs, presence or absence of three distinct petal types (petals+ petaloid sepals in Polygalaceae), flower size, corolla size (i.e., in open flower) and/or filament size (i.e., entire filament size particularly in subfamily Caesalpinioideae), flower colour, UV reflectance, habit, height, inflorescence type and inflorescence size for 758 Fabales genera. The information was obtained from hundreds of appropriate, previously published sources. This the largest morphological dataset constructed for Fabales to date, and the data presented in this article can be used for morphology, biogeography, ancestral state, ancestral area analyses of any Fabales clades.

Reconstructing an historical pollination syndrome: keel flowers.

BACKGROUND: Keel flowers are bilaterally symmetrical, pentamerous flowers with three different petal types and reproductive organs enclosed by keel petals; generally there is also connation of floral parts such as stamens and keel petals. In this study, the evolution of keel flowers within the order Fabales is explored to investigate whether the establishment of this flower type within one of the species-rich families, the Fabaceae (Leguminosae), preceded and could have influenced the evolution of keel flowers in the Polygalaceae. We conducted molecular dating, and ancestral area and ancestral state analyses for a phylogeny constructed for 678 taxa using published matK, rbcL and trnL plastid gene regions. RESULTS: We reveal the temporal and spatial origins of keel flowers and traits associated with pollinators, specifically floral symmetry, the presence or absence of a pentamerous corolla and three distinct petal types, the presence or absence of enclosed reproductive organs, androecium types, inflorescence types, inflorescence size, flower size, plant height and habit. Ancestral area reconstructions show that at the time keel flowers appeared in the Polygaleae, subfamily Papilionoideae of the Fabaceae was already distributed almost globally; at least eight clades of the Papilionoideae had keel flowers with a functional morphology broadly similar to the morphology of the first evolving Polygaleae flowers. CONCLUSIONS: The multiple origins of keel flowers within angiosperms likely represent convergence due to bee specialization, and therefore pollinator pressure. In the case of the Fabales, the first evolving keel flowers of Polygaleae have a functional morphology that corresponds with keel flowers of species of the Papilionoideae already present in the environment. These findings are consistent with the keel-flowered Polygaleae exploiting pollinators of keel-flowered Papilionoideae. The current study is the first to use ancestral reconstructions of traits associated with pollination to demonstrate that the multiple evolutionary origins of the keel flower pollinator syndrome in Fabales are consistent with, though do not prove, mimicry.

Ewé: a web-based ethnobotanical database for storing and analysing data.

Ethnobotanical databases serve as repositories of traditional knowledge (TK), either at international or local scales. By documenting plant species with traditional use, and most importantly, the applications and modes of use of such species, ethnobotanical databases play a role in the conservation of TK and also provide access to information that could improve hypothesis generation and testing in ethnobotanical studies. Brazil has a rich medicinal flora and a rich cultural landscape. Nevertheless, cultural change and ecological degradation can lead to loss of TK. Here, we present an online database developed with open-source tools with a capacity to include all medicinal flora of Brazil. We present test data for the Leguminosae comprising a total of 2078 records, referred to here as use reports, including data compiled from literature and herbarium sources. Unlike existing databases, Ewé provides tools for the visualization of large datasets, facilitating hypothesis generation and meta-analyses. The Ewé database is currently available at www.ewedb.com.

Phylogenomic Approaches to DNA Barcoding of Herbal Medicines: Developing Clade-Specific Diagnostic Characters for Berberis.

DNA barcoding of herbal medicines has been mainly concerned with authentication of products in trade and has raised awareness of species substitution and adulteration. More recently DNA barcodes have been included in pharmacopoeias, providing tools for regulatory purposes. The commonly used DNA barcoding regions in plants often fail to resolve identification to species level. This can be especially challenging in evolutionarily complex groups where incipient or reticulate speciation is ongoing. In this study, we take a phylogenomic approach, analyzing whole plastid sequences from the evolutionarily complex genus Berberis in order to develop DNA barcodes for the medicinally important species Berberis aristata. The phylogeny reconstructed from an alignment of ∼160 kbp of chloroplast DNA for 57 species reveals that the pharmacopoeial species in question is polyphyletic, complicating development of a species-specific DNA barcode. Instead we propose a DNA barcode that is clade specific, using our phylogeny to define Operational Phylogenetic Units (OPUs). The plastid alignment is then reduced to small, informative DNA regions including nucleotides diagnostic for these OPUs. These DNA barcodes were tested on commercial samples, and shown to discriminate plants in trade and therefore to meet the requirement of a pharmacopoeial standard. The proposed method provides an innovative approach for inferring DNA barcodes for evolutionarily complex groups for regulatory purposes and quality control.

Comparative phylogenetic methods and the cultural evolution of medicinal plant use.

Human life depends on plant biodiversity and the ways in which plants are used are culturally determined. Whilst anthropologists have used phylogenetic comparative methods (PCMs) to gain an increasingly sophisticated understanding of the evolution of political, religious, social and material culture, plant use has been almost entirely neglected. Medicinal plants are of special interest because of their role in maintaining people's health across the world. PCMs in particular, and cultural evolutionary theory in general, provide a framework in which to study the diversity of medicinal plant applications cross-culturally, and to infer changes in plant use over time. These methods can be applied to single medicinal plants as well as the entire set of plants used by a culture for medicine, and they account for the non-independence of data when testing for floristic, cultural or other drivers of plant use. With cultural, biological and linguistic diversity under threat, gaining a deeper and broader understanding of the variation of medicinal plant use through time and space is pressing.

Which Plants Used in Ethnomedicine Are Characterized? Phylogenetic Patterns in Traditional Use Related to Research Effort.

Plants are important resources in healthcare and for producing pharmaceutical drugs. Pharmacological and phytochemical characterization contributes to both the safe use of herbal medicines and the identification of leads for drug development. However, there is no recent assessment of the proportion of plants used in ethnomedicine that are characterized in this way. Further, although it is increasingly apparent that plants used in ethnomedicine belong to preferred phylogenetic lineages, it is not known how this relates to the focusing of research effort. Here we identify species and lineages rich in ethnomedicinal use and develop methods to describe how well they are known pharmacologically and/or phytochemically. We find 50% of plant species of the family Leguminosae used in ethnomedicine in Brazil, a geographical area where plants are an important part of healthcare, have been the focus of either phytochemical screening or testing for biological activity. Plant species which have more use reports are studied significantly more often (p < 0.05). Considering the taxonomic distribution of use, 70% of genera that include species with ethnomedicinal use have been studied, compared to 19% of genera with no reported use. Using a novel phylogenetic framework, we show that lineages with significantly greater numbers of ethnomedicinal species are phylogenetically over-dispersed within the family, highlighting the diversity of species used. "Hotnode clades" contain 16% of species but 46% of ethnomedicinally-used species. The ethnomedicinal species in hotnode clades have more use reports per species (p < 0.05), suggesting they are more frequently used. They are also more likely to be characterized pharmacologically and/or phytochemically. Research focus has followed traditional use by these measures, at least for these Brazilian plants, yet ethnomedicinal species yielding candidate drugs, raising public health concerns and more intensively studied lie outside of the hotnode clades.

Correction to: Medicinal plants used by women in Mecca: urban, Muslim and gendered knowledge.

In the original publication [1] were the Arabic letters in Table 2 incorrect. The corrected version of Table 2 can be found as Additional file 1 in this Erratum.

Medicinal plants used by women in Mecca: urban, Muslim and gendered knowledge.

BACKGROUND: This study explores medicinal plant knowledge and use among Muslim women in the city of Mecca, Saudi Arabia. Ethnobotanical research in the region has focused on rural populations and male herbal healers in cities, and based on these few studies, it is suggested that medicinal plant knowledge may be eroding. Here, we document lay, female knowledge of medicinal plants in an urban centre, interpreting findings in the light of the growing field of urban ethnobotany and gendered knowledge and in an Islamic context. METHODS: Free-listing, structured and semi-structured interviews were used to document the extent of medicinal plant knowledge among 32 Meccan women. Vernacular names, modes of preparation and application, intended therapeutic use and emic toxicological remarks were recorded. Women were asked where they learnt about medicinal plants and if and when they preferred using medicinal plants over biomedical resources. Prior informed consent was always obtained. We compared the list of medicinal plants used by these Meccan women with medicinal plants previously documented in published literature. RESULTS: One hundred eighteen vernacular names were collected, corresponding to approximately 110 plants, including one algae. Of these, 95 were identified at the species level and 39 (41%) had not been previously cited in Saudi Arabian medicinal plant literature. Almost one half of the plants cited are food and flavouring plants. Meccan women interviewed learn about medicinal plants from their social network, mass media and written sources, and combine biomedical and medicinal plant health care. However, younger women more often prefer biomedical resources and learn from written sources and mass media. CONCLUSIONS: The fairly small number of interviews conducted in this study was sufficient to reveal the singular body of medicinal plant knowledge held by women in Mecca and applied to treat common ailments. Plant availability in local shops and markets and inclusion in religious texts seem to shape the botanical diversity used by the Meccan women interviewed, and the use of foods and spices medicinally could be a global feature of urban ethnobotany. Ethnobotanical knowledge among women in Islamic communities may be changing due to access to mass media and biomedicine. We recognise the lack of documentation of the diversity of medicinal plant knowledge in the Arabian Peninsula and an opportunity to better understand gendered urban and rural knowledge.

Treating infants with frigg: linking disease aetiologies, medicinal plant use and care-seeking behaviour in southern Morocco.

BACKGROUND: Although most Moroccans rely to some extent on traditional medicine, the practice of frigg to treat paediatric ailments by elderly women traditional healers known as ferraggat, has not yet been documented. We describe the role of these specialist healers, document the medicinal plants they use, and evaluate how and why their practice is changing. METHODS: Ethnomedicinal and ethnobotanical data were collected using semi-structured interviews and observations of medical encounters. Information was collected from traditional healers, namely ferraggat, patients, herbalists and public health professionals. Patients' and healers' narratives about traditional medicine were analysed and medicinal plant lists were compiled from healers and herbalists. Plants used were collected, vouchered and deposited in herbaria. RESULTS: Ferragat remain a key health resource to treat infant ailments in the rural High Atlas, because mothers believe only they can treat what are perceived to be illnesses with a supernatural cause. Ferragat possess baraka, or the gift of healing, and treat mainly three folk ailments, taqait, taumist and iqdi, which present symptoms similar to those of ear infections, tonsillitis and gastroenteritis. Seventy plant species were used to treat these ailments, but the emphasis on plants may be a recent substitute for treatments that used primarily wool and blood. This change in materia medica is a shift in the objects of cultural meaningfulness in response to the increasing influence of orthodox Islam and state-sponsored modernisation, including public healthcare and schooling. CONCLUSIONS: Religious and other sociocultural changes are impacting the ways in which ferraggat practice. Treatments based on no-longer accepted symbolic elements have been readily abandoned and substituted by licit remedies, namely medicinal plants, which play a legitimisation role for the practice of frigg. However, beliefs in supernatural ailment aetiologies, as well as lack or difficult access to biomedical alternatives, still underlie the need for specialist traditional healers.

Comprehensive dataset of the medicinal plants used by a Tashelhit speaking community in the High Atlas, Morocco.

This dataset describes medicinal plants used in a poorly studied area of Morocco: the High Atlas mountains, inhabited by Ishelhin people, the southern Moroccan Amazigh (Berber) ethnic group, "An ethnomedicinal survey of a Tashelhit-speaking community in the High Atlas, Morocco" (Teixidor-Toneu et al., 2016) [1]. It includes a comprehensive list of the plants used in the commune, as well as details on the plant voucher specimens collected and a glossary of Tashelhit terminology relevant to the study. To collect the data, semi-structured and structured interviews were carried out, as well as focus group discussions. Free prior informed consent was obtained for all interactions. A hundred and six adults were interviewed and 2084 use reports were collected; a hundred fifty-one vernacular names corresponding to 159 botanical species were found.

An ethnomedicinal survey of a Tashelhit-speaking community in the High Atlas, Morocco.

ETHNOPHARMACOLOGICAL RELEVANCE: Traditional knowledge about medicinal plants from a poorly studied region, the High Atlas in Morocco, is reported here for the first time; this permits consideration of efficacy and safety of current practises whilst highlighting species previously not known to have traditional medicinal use. AIM OF THE STUDY: Our study aims to document local medicinal plant knowledge among Tashelhit speaking communities through ethnobotanical survey, identifying preferred species and new medicinal plant citations and illuminating the relationship between emic and etic ailment classifications. MATERIALS AND METHODS: Ethnobotanical data were collected using standard methods and with prior informed consent obtained before all interactions, data were characterized using descriptive indices and medicinal plants and healing strategies relevant to local livelihoods were identified. RESULTS: 151 vernacular names corresponding to 159 botanical species were found to be used to treat 36 folk ailments grouped in 14 biomedical use categories. Thirty-five (22%) are new medicinal plant records in Morocco, and 26 described as used for the first time anywhere. Fidelity levels (FL) revealed low specificity in plant use, particularly for the most commonly reported plants. Most plants are used in mixtures. Plant use is driven by local concepts of disease, including "hot" and "cold" classification and beliefs in supernatural forces. CONCLUSION: Local medicinal plant knowledge is rich in the High Atlas, where local populations still rely on medicinal plants for healthcare. We found experimental evidence of safe and effective use of medicinal plants in the High Atlas; but we highlight the use of eight poisonous species.

Molecular phylogeny and evolutionary history of the Eurasiatic orchid genus Himantoglossum s.l. (Orchidaceae).

BACKGROUND AND AIMS: Lizard orchids of the genus Himantoglossum include many of Eurasia's most spectacular orchids, producing substantial spikes of showy flowers. However, until recently the genus had received only limited, and entirely traditional, systematic study. The aim of the current work was to provide a more robust molecular phylogeny in order to better understand the evolutionary relationships among species of particular conservation concern. METHODS: All putative species of Himantoglossum s.l. were sampled across its geographical range. A large subsample of the 153 populations studied contributed to an initial survey of nuclear ribosomal internal transcribed spacer (nrITS) ribotypes. Smaller subsets were then sequenced for four plastid regions and the first intron of the low-copy-number nuclear gene LEAFY. Rooted using Steveniella as outgroup, phylogenetic trees were generated using parsimony and Bayesian methods from each of the three datasets, supplemented with a ribotype network. KEY RESULTS: The resulting trees collectively determined the order of branching of the early divergent taxa as Himantoglossum comperianum > H. robertianum group > H. formosum, events that also involved significant morphological divergence. Relaxed molecular clock dating suggested that these divergences preceded the Pleistocene glaciations (the origin of the H. robertianum group may have coincided with the Messinian salinity crisis) and occurred in Asia Minor and/or the Caucasus. Among more controversial taxa of the H. hircinum-jankae clade, which are only subtly morphologically divergent, topological resolution was poorer and topological incongruence between datasets was consequently greater. CONCLUSIONS: Plastid sequence divergence is broadly consistent with prior, morphologically circumscribed taxa and indicates a division between H. hircinum-adriaticum to the west of the Carpathians and H. jankae-caprinum (plus local endemics) to the east, a distinction also suggested by nrITS ribotypes. LEAFY phylogenies are less congruent with prior taxonomic arrangements and include one likely example of paralogy. Himantoglossum metlesicsianum fully merits its IUCN Endangered status. Potentially significant genetic variation was detected within Steveniella satyrioides, H. robertianum and H. hircinum. However, confident circumscription of the more derived species of Himantoglossum s.s., including local endemics of hybrid origin, must await future morphometric and population-genetic analyses.

The evolution of traditional knowledge: environment shapes medicinal plant use in Nepal.

Traditional knowledge is influenced by ancestry, inter-cultural diffusion and interaction with the natural environment. It is problematic to assess the contributions of these influences independently because closely related ethnic groups may also be geographically close, exposed to similar environments and able to exchange knowledge readily. Medicinal plant use is one of the most important components of traditional knowledge, since plants provide healthcare for up to 80% of the world's population. Here, we assess the significance of ancestry, geographical proximity of cultures and the environment in determining medicinal plant use for 12 ethnic groups in Nepal. Incorporating phylogenetic information to account for plant evolutionary relatedness, we calculate pairwise distances that describe differences in the ethnic groups' medicinal floras and floristic environments. We also determine linguistic relatedness and geographical separation for all pairs of ethnic groups. We show that medicinal uses are most similar when cultures are found in similar floristic environments. The correlation between medicinal flora and floristic environment was positive and strongly significant, in contrast to the effects of shared ancestry and geographical proximity. These findings demonstrate the importance of adaptation to local environments, even at small spatial scale, in shaping traditional knowledge during human cultural evolution.

Convergent evolution of floral signals underlies the success of Neotropical orchids.

The great majority of plant species in the tropics require animals to achieve pollination, but the exact role of floral signals in attraction of animal pollinators is often debated. Many plants provide a floral reward to attract a guild of pollinators, and it has been proposed that floral signals of non-rewarding species may converge on those of rewarding species to exploit the relationship of the latter with their pollinators. In the orchid family (Orchidaceae), pollination is almost universally animal-mediated, but a third of species provide no floral reward, which suggests that deceptive pollination mechanisms are prevalent. Here, we examine floral colour and shape convergence in Neotropical plant communities, focusing on certain food-deceptive Oncidiinae orchids (e.g. Trichocentrum ascendens and Oncidium nebulosum) and rewarding species of Malpighiaceae. We show that the species from these two distantly related families are often more similar in floral colour and shape than expected by chance and propose that a system of multifarious floral mimicry--a form of Batesian mimicry that involves multiple models and is more complex than a simple one model-one mimic system--operates in these orchids. The same mimetic pollination system has evolved at least 14 times within the species-rich Oncidiinae throughout the Neotropics. These results help explain the extraordinary diversification of Neotropical orchids and highlight the complexity of plant-animal interactions.

Phylogenies reveal predictive power of traditional medicine in bioprospecting.

There is controversy about whether traditional medicine can guide drug discovery, and investment in bioprospecting informed by ethnobotanical data has fluctuated. One view is that traditionally used medicinal plants are not necessarily efficacious and there are no robust methods for distinguishing those which are most likely to be bioactive when selecting species for further testing. Here, we reconstruct a genus-level molecular phylogenetic tree representing the 20,000 species found in the floras of three disparate biodiversity hotspots: Nepal, New Zealand, and the Cape of South Africa. Borrowing phylogenetic methods from community ecology, we reveal significant clustering of the 1,500 traditionally used species, and provide a direct measure of the relatedness of the three medicinal floras. We demonstrate shared phylogenetic patterns across the floras: related plants from these regions are used to treat medical conditions in the same therapeutic areas. This finding strongly indicates independent discovery of plant efficacy, an interpretation corroborated by the presence of a significantly greater proportion of known bioactive species in these plant groups than in random samples. We conclude that phylogenetic cross-cultural comparisons can focus screening efforts on a subset of traditionally used plants that are richer in bioactive compounds, and could revitalize the use of traditional knowledge in bioprospecting.

Combined phylogenetic analyses reveal interfamilial relationships and patterns of floral evolution in the eudicot order Fabales.

Relationships between the four families placed in the angiosperm order Fabales (Leguminosae, Polygalaceae, Quillajaceae, Surianaceae) were hitherto poorly resolved. We combine published molecular data for the chloroplast regions matK and rbcL with 66 morphological characters surveyed for 73 ingroup and two outgroup species, and use Parsimony and Bayesian approaches to explore matrices with different missing data. All combined analyses using Parsimony recovered the topology Polygalaceae (Leguminosae (Quillajaceae + Surianaceae)). Bayesian analyses with matched morphological and molecular sampling recover the same topology, but analyses based on other data recover a different Bayesian topology: ((Polygalaceae + Leguminosae) (Quillajaceae + Surianaceae)). We explore the evolution of floral characters in the context of the more consistent topology: Polygalaceae (Leguminosae (Quillajaceae + Surianaceae)). This reveals synapomorphies for (Leguminosae (Quillajaceae + Surianaceae)) as the presence of free filaments and marginal/ventral placentation, for (Quillajaceae + Surianaceae) as pentamery and apocarpy, and for Leguminosae the presence of an abaxial median sepal and unicarpellate gynoecium. An octamerous androecium is synapomorphic for Polygalaceae. The development of papilionate flowers, and the evolutionary context in which these phenotypes appeared in Leguminosae and Polygalaceae, shows that the morphologies are convergent rather than synapomorphic within Fabales.

The use of phylogeny to interpret cross-cultural patterns in plant use and guide medicinal plant discovery: an example from Pterocarpus (Leguminosae).

BACKGROUND: The study of traditional knowledge of medicinal plants has led to discoveries that have helped combat diseases and improve healthcare. However, the development of quantitative measures that can assist our quest for new medicinal plants has not greatly advanced in recent years. Phylogenetic tools have entered many scientific fields in the last two decades to provide explanatory power, but have been overlooked in ethnomedicinal studies. Several studies show that medicinal properties are not randomly distributed in plant phylogenies, suggesting that phylogeny shapes ethnobotanical use. Nevertheless, empirical studies that explicitly combine ethnobotanical and phylogenetic information are scarce. METHODOLOGY/PRINCIPAL FINDINGS: In this study, we borrowed tools from community ecology phylogenetics to quantify significance of phylogenetic signal in medicinal properties in plants and identify nodes on phylogenies with high bioscreening potential. To do this, we produced an ethnomedicinal review from extensive literature research and a multi-locus phylogenetic hypothesis for the pantropical genus Pterocarpus (Leguminosae: Papilionoideae). We demonstrate that species used to treat a certain conditions, such as malaria, are significantly phylogenetically clumped and we highlight nodes in the phylogeny that are significantly overabundant in species used to treat certain conditions. These cross-cultural patterns in ethnomedicinal usage in Pterocarpus are interpreted in the light of phylogenetic relationships. CONCLUSIONS/SIGNIFICANCE: This study provides techniques that enable the application of phylogenies in bioscreening, but also sheds light on the processes that shape cross-cultural ethnomedicinal patterns. This community phylogenetic approach demonstrates that similar ethnobotanical uses can arise in parallel in different areas where related plants are available. With a vast amount of ethnomedicinal and phylogenetic information available, we predict that this field, after further refinement of the techniques, will expand into similar research areas, such as pest management or the search for bioactive plant-based compounds.