Research Projects

Integrative Taxonomy, Molecular Phylogeny and Plant Biodiversity

My research focuses on plant systematics, integrative taxonomy, molecular phylogeny, biogeography, plant morphology and biodiversity, with a particular emphasis on the flora of Iran and the Mediterranean region.

My work combines classical botany and modern systematic approaches: field observations, herbarium studies, morphological and micromorphological analyses, molecular data, phylogenetic reconstruction and biogeographical interpretation. I am particularly interested in understanding how plant species are delimited, how they are related to each other, and how their diversity has evolved through time and space.

A central part of my research has been devoted to the genus Verbascum L. (Scrophulariaceae), but I have also worked on several other plant groups, including Onosma, Cousinia, Prunus subg. Amygdalus, Centaurea, Teucrium, Astragalus, and different members of Boraginaceae and Asteraceae.

My research is strongly connected with herbarium collections, botanical fieldwork, molecular systematics and student supervision. It has been developed through collaborations with universities, herbaria and botanists in France, Iran and other countries.

Main Research Themes

1. Systematics and Evolution of Verbascum L.

The genus Verbascum L. has been the central model of my research since my PhD. My work on this genus focuses on species delimitation, taxonomy, molecular phylogeny, morphology, micromorphology, biogeography and the evolutionary history of the genus, especially in Iran and the Irano-Turanian region.

Verbascum is a taxonomically complex genus. Many species show high morphological variability, and some groups are difficult to delimit using morphology alone. For this reason, my work combines herbarium taxonomy, field observations, molecular data and comparative morphology.

My research on Verbascum includes:

  • taxonomic revision of selected species and species complexes;
  • description of new species and new taxa;
  • discovery of new records for the flora of Iran;
  • typification and synonymic clarification;
  • molecular phylogenetic analyses using nuclear and chloroplast markers;
  • study of herbarium specimens, including historical and type material;
  • comparison between morphological groups and molecular results;
  • study of seed morphology and micromorphological variation;
  • contribution to a broader understanding of the diversification of the genus.

Taxonomic Contributions in Verbascum

Several new taxa and taxonomic contributions have resulted from this work, including:

  • Verbascum shahsavarensis Sotoodeh, Attar & Civeyrel
  • Verbascum parsana Sotoodeh, Attar & Civeyrel
  • Verbascum birjandense Sotoodeh
  • Verbascum urumiense Sotoodeh
  • Verbascum carmanicum subsp. kermanense (Hub.-Mor.) Sotoodeh, Attar, Andalo & Civeyrel

I have also contributed to the recognition of new records for the flora of Iran, such as:

  • Verbascum songaricum subsp. subdecurrens
  • Verbascum oreophilum var. oreophilum
  • Verbascum cheiranthifolium var. asperulum

Other taxonomic contributions include typification and synonymic clarification, especially concerning Verbascum aspinum and Verbascum stachydiforme.

One important aspect of my research on Verbascum is the use of herbarium specimens to clarify taxonomy, verify identifications and compare historical material with newly collected specimens. This is particularly important in a genus where morphological variation, geographical isolation, possible hybridization and incomplete phylogenetic resolution can make species delimitation difficult.

2. Molecular Phylogeny and Biogeography of Iranian Angiosperms

A major part of my research concerns the evolutionary history and biogeography of angiosperms in Iran and neighbouring regions. Iran is located at the intersection of several major phytogeographical zones and represents an important region for studying plant diversification, endemism and ecological adaptation.

My PhD research focused on the biogeographical and evolutionary history of the genera Verbascum and Artemisia in Iran using molecular phylogeny. The aim was to understand how evolutionary lineages are structured across different geographical and ecological regions, and how historical and environmental factors may have influenced their diversification.

The main approaches used in this research include:

  • DNA sequence analysis;
  • nuclear and chloroplast molecular markers;
  • phylogenetic reconstruction;
  • comparison between molecular clades and morphological groups;
  • interpretation of distribution patterns;
  • biogeographical discussion in relation to the Irano-Turanian and Mediterranean regions.

This research contributes to a better understanding of plant evolution in biodiversity-rich regions and transitional zones.

3. Integrative Taxonomy of Onosma L.

Another important part of my research concerns the genus Onosma L. (Boraginaceae), especially Iranian species belonging to complex groups within the genus.

Species delimitation in Onosma can be difficult because several taxa show subtle morphological differences. Reliable identification often requires the combination of several lines of evidence, including morphology, micromorphology, indumentum characters, herbarium comparisons and molecular data.

My work on Onosma includes:

  • detailed morphological observations;
  • study of indumentum and trichome characters;
  • micromorphological analyses;
  • comparison with type specimens and herbarium material;
  • molecular data analysis;
  • species delimitation;
  • description of new taxa;
  • revision of species complexes.

Research on Onosma has led to the description and clarification of several taxa, especially in groups such as Asterotricha, Heterotricha, Haplotricha and Rostellatae. These studies are important for improving our understanding of Boraginaceae diversity in Iran and neighbouring regions.

Examples of research topics include:

  • new species of Onosma from Iran;
  • revision of species complexes;
  • comparison between Iranian, Turkish and regional taxa;
  • clarification of diagnostic morphological characters;
  • integration of molecular and morphological evidence.

This research is also closely connected with student supervision and collaborative work with the University of Tehran.

4. Taxonomy and Micromorphology of Asteraceae

I have also contributed to taxonomic and micromorphological studies in Asteraceae, especially on the genera Cousinia and Centaurea.

Cousinia Cass.

Research on Cousinia has focused on the description of new species from Iran and the use of morphological characters for species delimitation. Important characters include phyllaries, flower number, stem morphology, winged stems and inflorescence structure.

Two new species of Cousinia were described from western and northeastern Iran:

  • Cousinia bornmulleriana
  • Cousinia semnanensis

These studies contribute to the understanding of diversity and endemism within one of the most complex and species-rich genera of Asteraceae in the Irano-Turanian region.

Centaurea L.

Research on Centaurea focused on the taxonomic significance of achene morphology. Several achene characters were examined, including shape, size, surface ornamentation, hilum, pappus colour and pappus length.

This work showed that achene morphology can provide useful diagnostic characters for distinguishing sections and species within the genus. It also highlighted the importance of fruit micromorphology in Asteraceae systematics.

5. Boraginaceae: Floral and Nutlet Micromorphology

I have contributed to studies on the tribe Cynoglosseae within Boraginaceae, with a focus on the taxonomic value of floral and nutlet characters.

This work included several genera, such as:

  • Paracaryum
  • Mattiastrum
  • Microparacaryum
  • Rindera
  • Cynoglossum
  • Solenanthus
  • Trachelanthus
  • Lindelofia

The main objective was to evaluate the usefulness of macro- and micromorphological characters for identification and species delimitation. Nutlet morphology, surface ornamentation, wings, glochids and floral structures were particularly important in this work.

This research highlights the importance of microscopy and detailed morphological observation in plant systematics.

6. Prunus subg. Amygdalus and Spiny Almonds

I have also been involved in research on Prunus subg. Amygdalus, especially shrub-thorn almonds in Iran. This work focuses on species complexes, pollen morphology, leaf micromorphology and taxonomic delimitation.

The main research questions include:

  • how to distinguish closely related taxa;
  • which pollen and leaf characters are taxonomically informative;
  • how micromorphological characters can support species delimitation;
  • how morphological variation is structured within and between species.

This research is connected to collaborative work and student supervision at the University of Tehran.

7. Other Taxonomic Contributions

In addition to my main work on Verbascum, Onosma, Cousinia, Centaurea and Prunus, I have contributed to taxonomic and systematic studies on other plant groups.

Teucrium L.

I contributed to the description of Teucrium elymaiticum, a new endemic species from western Iran. This work focused on morphological comparison, diagnostic characters and the taxonomic position of the new species within the genus.

Astragalus L.

I also contributed to a phylogenetic and systematic study of Astragalus, one of the largest genera of flowering plants. This work used nuclear ribosomal ITS data and proposed a new infrageneric grouping based on molecular results and morphological characters.

Herbarium-Based Research and Natural History Collections

Herbarium collections are central to my research. They provide essential material for taxonomy, nomenclature, species delimitation, historical biogeography and the comparison of past and present plant diversity.

I have worked with specimens from several important herbaria and botanical institutions, including:

  • Central Herbarium of the University of Tehran;
  • Conservatoire et Jardin botaniques de Genève;
  • Royal Botanic Gardens, Kew;
  • Natural History Museum, London;
  • University of Montpellier collections.

My work with herbarium material includes:

  • taxonomic revision;
  • study of type specimens;
  • morphological comparison;
  • digital photography and documentation;
  • metadata management;
  • comparison between historical and recent collections;
  • use of collections for molecular and morphometric studies;
  • valorization of herbarium collections for research and teaching.

I consider herbarium specimens not only as reference material for taxonomy, but also as a powerful source of information for biodiversity research, historical biogeography, ecological interpretation and long-term changes in plant diversity.

Methods and Approaches

My research combines several complementary methods.

Field Botany

Fieldwork remains an essential part of my research. It allows direct observation of plants in their natural habitats, collection of ecological and geographical information, documentation of morphological variation and better understanding of species in their environmental context.

Herbarium Taxonomy

Herbarium work is used to compare specimens, verify identifications, examine type material and clarify taxonomic problems. This is especially important in plant groups with complex nomenclature, high morphological variability or insufficient molecular resolution.

Morphology and Micromorphology

Morphological characters are studied at different scales, from general plant habit to detailed structures such as seeds, pollen grains, achenes, nutlets, trichomes, calyx, corolla, leaves and epidermal characters.

Microscopy and imaging techniques are used to document fine-scale variation and identify diagnostic characters.

Molecular Systematics

Molecular data are used to infer phylogenetic relationships, test taxonomic hypotheses and compare molecular lineages with morphological groups. My work has used nuclear and chloroplast markers, especially in Verbascum, Onosma, Prunus and related plant groups.

Phylogenetic and Biogeographical Analysis

Phylogenetic results are interpreted in relation to morphology, geography, ecology and evolutionary history. This approach helps to understand how plant lineages have diversified and how species distributions may have been shaped through time.

Collaborative Research, Academic Network and Student Supervision

A significant part of my research has been developed through long-term academic collaborations with botanists, taxonomists, phylogeneticists and herbarium specialists in France, Iran and other countries. These collaborations have played an important role in the development of my work on plant systematics, molecular phylogeny, taxonomy, herbarium studies, micromorphology and biodiversity.

My doctoral research was developed between the Université Paul Sabatier – Toulouse III / EcoLab and the University of Tehran, under the supervision of Laure Civeyrel, my PhD supervisor, and Farideh Attar, my PhD co-supervisor. This work focused on the biogeography and molecular phylogeny of Verbascum and Artemisia, with a particular emphasis on Iranian and Mediterranean plant diversity.

Another major part of my research has been carried out through collaborations with the University of Tehran and the Central Herbarium of the University of Tehran, especially for field-based taxonomy, herbarium studies, student supervision and the study of complex plant groups from Iran.

These collaborations have resulted in several taxonomic and phylogenetic publications, including descriptions of new species, new records for the flora of Iran, typifications, synonymic clarifications and micromorphological studies. They also illustrate the importance of combining field botany, herbarium collections, molecular data and student training in plant systematics.

Student Supervision and Research Training

Student supervision is an important part of my academic activity. I have contributed as an advisor, co-supervisor or scientific collaborator to several research projects involving PhD students, Master students and undergraduate interns. These projects are mainly focused on plant taxonomy, species delimitation, micromorphology, molecular systematics, seed morphology and herbarium-based research.

Examples of supervised or co-supervised projects include:

  • F. Ghamari – PhD project, University of Tehran: species delimitation in the Haplotricha group of the genus Onosma in Iran using morphological and molecular characters.
  • M. Advay – PhD project, University of Tehran: species delimitation in the Asterotricha and Heterotricha groups of Onosma using morphological characters, nrDNA ITS and chloroplast DNA sequences.
  • N. Bayat – PhD project, University of Tehran: investigation of species complexes in shrub-thorn almonds, Prunus subg. Amygdalus, using micromorphological and taxonomic approaches.
  • P. Fouladi – Master project, University of Tehran: micromorphological study of pollen grains, leaf epidermis and indumentum in selected species of Verbascum from Iran.
  • F. Ghamari – Master project, University of Tehran: taxonomic studies on the Podonosma orientalis species complex in Iran.
  • A. Ramezani – Master project, University of Tehran: taxonomy and phylogenetic relationships in the Onosma bulbotricha complex using morphological and molecular evidence.
  • A. Hadidchi – PhD project, University of Tehran: systematic studies on the genus Delphinium sensu lato in Iran.
  • S. Molina – L2 internship, Université de Bourgogne: morphometric analysis of plant seeds, with a comparative study of shape and size.

The following internships at the Université de Bourgogne were jointly supervised in collaboration with Beryl Laitung:

  • O. Dehaye – L3 internship, Université de Bourgogne: germination and emergence of two species of Delphinium.
  • M. Govin – L3 internship, Université de Bourgogne: seed morphological diversity in two species of Delphinium.
  • K. Goret-Iihashi – L1 internship, Université de Bourgogne: phenological monitoring of plum and cherry trees in the conservation orchard of the Montmuzard campus.

These projects show the continuity between my research and teaching activities. They also reflect my interest in training students in practical botanical methods: observation, specimen comparison, microscopy, morphometry, molecular data interpretation, scientific writing and the use of herbarium collections.

Main Academic Collaborations

My research has been developed through collaborations with several researchers, supervisors, colleagues and institutions in France, Iran and other countries. These collaborations have contributed to different aspects of my work, including plant taxonomy, molecular systematics, herbarium studies, micromorphology, phylogeny and student supervision.

  • Laure Civeyrel – Université Paul Sabatier / Toulouse, my PhD supervisor and long-term collaborator, especially on the molecular phylogeny, biogeography and taxonomy of Verbascum.
  • Farideh Attar – University of Tehran, my PhD co-supervisor and long-term collaborator in the taxonomy and systematics of Iranian angiosperms, especially on Verbascum, Onosma, Cousinia, Centaurea, Teucrium and other plant groups.
  • Mansour Mirtadzadini – University of Kerman, collaborator on the taxonomy of the Iranian flora, especially in Boraginaceae and related taxonomic studies.
  • Christophe Andalo – EcoLab / Université Paul Sabatier – Toulouse, collaborator in evolutionary biology, molecular systematics and plant phylogeny during my research period in Toulouse.
  • Masoud Sheidai – Shahid Beheshti University, Iran, collaborator in plant systematics.
  • The late Ali Asghar Maassoumi – Iranian botanist and specialist of Astragalus. I had the honour of collaborating with him on the phylogeny and infrageneric classification of Astragalus.
  • Shahrokh Kazempour Osaloo – Iran, collaborator on phylogenetic studies and the classification of Astragalus.
  • Hossein Maroofi – Kurdistan / Sanandaj, Iran, collaborator in floristic and taxonomic studies of the Iranian flora.
  • Asghar Zamani – University of Guilan, Iran, collaborator in taxonomic and floristic studies.
  • Riza Binzet – Turkey, collaborator in taxonomic studies, especially on Onosma and Boraginaceae.

Through these collaborations, I have contributed to research linking classical taxonomy, molecular phylogeny, herbarium studies, micromorphology, student training and the study of plant diversity in Iran, France and the Mediterranean region.

Link Between Research and Teaching

My research activities are closely connected with teaching and training. In my teaching, I often use examples from plant diversity, systematics, ecology, field botany, herbarium work and data analysis.

This connection between research and teaching allows students to understand botany not only as a descriptive discipline, but also as a scientific approach based on observation, comparison, evidence, hypothesis testing and interpretation.

My teaching and supervision activities include:

  • plant diversity and systematics;
  • field botany and floristic observation;
  • laboratory and herbarium methods;
  • morphology and micromorphology;
  • ecology and plant-environment relationships;
  • data organization and statistical analysis;
  • scientific writing and oral presentation.

This integration of research and teaching is an important part of my academic identity.

Future Research Directions

My future research aims to further develop integrative approaches in plant systematics, taxonomy and ecology. I am particularly interested in connecting classical taxonomy and phylogeny with broader ecological questions.

Future research directions include:

  • integrative taxonomy of complex plant groups;
  • DNA barcoding and molecular identification of herbarium specimens;
  • phylogeny and biogeography of Mediterranean and Irano-Turanian taxa;
  • seed, pollen and micromorphological traits as tools for species delimitation;
  • valorization of herbarium collections for research and teaching;
  • plant responses to environmental change;
  • phenotypic plasticity and ecological adaptation;
  • relationships between plants, climate, soil and biotic interactions;
  • conservation of plant diversity in biodiversity-rich regions.

My long-term objective is to contribute to a better understanding of plant diversity, evolution and conservation, while maintaining a strong link between field botany, herbarium research, molecular systematics and education.

Selected Publications Related to These Projects

Attar F., Mirtadzadini M., Binzet R. & Sotoodeh A. 2025. New taxa of Onosma L. (Boraginaceae, Lithospermeae) from Iran. Adansonia 47: 299–310.

Bayat N., Attar F. & Sotoodeh A. 2024. Pollen and leaf micromorphological characteristics of spiny almonds, Prunus subgenus Amygdalus, in Iran. Microscopy Research and Technique.

Advay M., Attar F., Hama S., Ahmad A. & Sotoodeh A. 2024. Onosma safaie-fari (Boraginaceae), a new species from western Iran. Iranian Journal of Botany 30: 12–27.

Attar F., Mirtadzadini M. & Sotoodeh A. 2023. Onosma deltiphylla and O. humilis (Boraginaceae), two new species from Iran. Annales Botanici Fennici 60: 161–166.

Attar F., Sotoodeh A., Mirtadzadini M., Daemi M. & Civeyrel L. 2023. New findings in Onosma section Protonosma (Boraginaceae) using morphological and molecular evidence. Botany Letters 170: 285–302.

Sotoodeh A., Attar F., Laitung B. & Civeyrel L. 2022. Verbascum birjandense and V. urumiense (Scrophulariaceae), two new endemic species for the flora of Iran. Phytotaxa 538: 35–44.

Attar F., Maroofi H., Mirtadzadini M. & Sotoodeh A. 2022. Two new species of Cousinia from western and northeastern Iran. Nova Biologica Reperta 9: 144–152.

Attar F., Sotoodeh A., Sheidai M., Esfandani-Bozchloyi S. & Mirtadzadini M. 2021. Taxonomic importance of nutlets and flowers in tribe Cynoglosseae. Iranian Journal of Botany 27: 146–163.

Attar F., Sotoodeh A. & Mirtadzadini M. 2020. Four new species in subsection Asterotricha of the genus Onosma from Iran. Feddes Repertorium 131: 146–158.

Rakizadeh S., Attar F. & Sotoodeh A. 2019. Taxonomic significance of achene morphology in Centaurea. Nova Biologica Reperta 6: 352–366.

Attar F., Sotoodeh A. & Mirtadzadini M. 2019. Teucrium elymaiticum, a new species for the flora of Iran. Mediterranean Botany 40: 203–207.

Sotoodeh A., Attar F., Andalo C., Mirtajaldini M. & Civeyrel L. 2018. Focusing on three Verbascum taxa of the flora of Iran. Adansonia 40: 171–181.

Sotoodeh A., Attar F. & Civeyrel L. 2017. Verbascum songaricum subsp. subdecurrens: a new record; typification and true identity of V. aspinum as a new synonym of V. stachydiforme in the flora of Iran. Kew Bulletin 72: 24.

Maassoumi A. A., Kazempour Osaloo Sh. & Sotoodeh A. 2016. A new infrageneric grouping of Astragalus. Iranian Journal of Botany 22: 88–100.

Sotoodeh A., Civeyrel L. & Attar F. 2016. A new species of Verbascum from Gilan province, Iran, based on morphological and molecular evidence. Adansonia 38: 127–132.

Sotoodeh A., Civeyrel L. & Attar F. 2015. Verbascum shahsavarensis, a new species for the flora of Iran. Phytotaxa 203: 76–80.

Sotoodeh A., Civeyrel L., Zamani A. & Attar F. 2014. Verbascum oreophilum var. oreophilum and Verbascum cheiranthifolium var. asperulum, two new records for the flora of Iran. Phytotaxa 173: 205–210.

Keywords

Plant systematics; Integrative taxonomy; Molecular phylogeny; Biogeography; Herbarium collections; Plant biodiversity; Flora of Iran; Mediterranean flora; Irano-Turanian region; Verbascum; Onosma; Cousinia; Prunus; Centaurea; Boraginaceae; Asteraceae.