Figure 2 - uploaded by Moslem Doostmohammadi
Content may be subject to copyright.
Morphological diversity of Veronica subgenus Pentasepalae in the Iranian plateau. (A) V. chionantha, (B) V. farinosa, (C) V. orientalis, (D) V. rechingeri, (E) V. aucheri, (F) V. czerniakowskiana, (G) V. kurdica subsp. filicaulis, (H) V. paederotae, (I) V. gaubae, (J) V. fragilis, (K) V. mazanderanae, (L) V. schizostegia, (M) V. kurdica subsp. kurdica, (N) V. kopetdaghensis, (O) V. daranica, (P) V. khorassanica and (Q) V. mirabilis. Photos (A-K and Q) by M. Doostmohammadi, (L,M,O,P) by M. Mirtadzadini and (N) by H. Moazzeni.
Source publication
Veronica subgenus Pentasepalae is the largest subgenus of Veronica in the Northern Hemisphere with approximately 80 species mainly from Southwest Asia. In order to reconstruct the phylogenetic relationships among the members of V. subgenus Pentasepalae and to test the “out of the Iranian plateau” hypothesis, we applied thorough taxonomic sampling,...
Context in source publication
Context 1
... subgenus Pentasepalae is a well-defined monophyletic subgenus, according to the phylogenetic analysis of nuclear and plastid DNA sequence data [21,25], and is the largest subgenus of Veronica in the Northern Hemisphere, comprising about 80 species [27] with several recent additions from Europe [28,29] and Turkey [30]. The species of this subgenus are distributed from Morocco and the Iberian Peninsula to the Altai Mountains in Central Asia (Figure 1), with a center of diversity in Turkey and northern Iran and comprising most of the perennial species of Veronica in SW Asia (representative taxa are shown in Figure 2). Based on molecular, morphological and karyological data, species of this subgenus are categorized into four subsections (V. ...
Citations
... The Irano-Turanian (IT) floristic region has since long been regarded as a 'donor' region of many plant taxa to 'recipient' neighboring regions (Zohary, 1973), especially as a source area for the Mediterranean flora (e.g., Manafzadeh et al., 2017;Djamali et al., 2012;Stojilković et el., 2022;Doostmohammadi et al., 2022;and references therein). Several studies provide also evidence for plant migration from Asian mountains into neighboring steppe or desert regions, as for instance from the Tien Shan into the Turan lowlands (Atraphaxis L., Zhang et al., 2014; Lagochilus Bunge ex Benth., Zhang et al., 2017) or from the Helan Mountains and northeastern Qinghai-Tibet Plateau into the neighboring steppe deserts (Stipa breviflora, Ren et al., 2022). ...
... This diversity mostly belongs to some super diverse Irano-Turanian genera such as Astragalus, Acantholimon, Allium, Cousinia, Centaurea, and Dionysia, which diversified not only in the Zagros but also throughout the highlands of the Iranian plateau, with a higher endemicity in alpine elevations. The roles of the geological events of Zagros in the diversification of some of these genera are highlighted in recent studies (Bagheri et al., 2017;Moharrek et al., 2019;Doostmohammadi et al., 2022). Along with this recently diversified species (neo-endemics), there are also some relict, paleoendemic species in the Zagros Mountains. ...
The Zagros Mountain range in western Iran is an area of species endemism within the Irano-Anatolian biodiversity hotspot. A couple of relict and morphologically isolated species have been reported for the Zagros Mountains, yet their distribution patterns in relation to the geological and climatic history of the region are not fully understood. Clinopodium kallaricum (Jamzad) Bordbar, 2019 (Lamiaceae), Hypericum dogonbadanicum Assadi, 1984 (Hypericaceae), and Iranoaster bachtiaricus (Mozaff.) Kaz. Osaloo, Farhani & Mozaff. 2018 (Asteraceae) are local endemic species restricted to Zagros with no clear affinities to the Irano-Turanian or other Northern Hemisphere temperate species. Previous studies suggested some afromontane relations for these species, beyond the Saharo-Sindian lowlands. Here, we provide dated phylogenies for these three species, to assess the most probable drivers behind this pattern of distribution. Our results represent that the split between these taxa and their relatives is not contemporaneous, implying different biogeographical histories. I. bachtiaricus originated in the Middle Miocene, while C. kallaricum and H. dogonbadanicum are relatively younger (late Miocene early Pliocene). The divergence of these taxa coincided with the major geological and climatic events of the Miocene, mainly the collision of the Afro-Arabia and Eurasia plates in 18-16 Mya and the aridification of the Sahara (started in 11-7 Mya), followed by a shift in the vegetation of the Sahara from subtropical/steppe to arid desert. The possible relict nature of these species is discussed, and the role of the Saharo-Sindian Region as a vicariant agent is highlighted, which could have subdivided formerly uniform populations and subsequently accelerated the allopatric speciation.
... However, several new taxa were added afterwards (Saeidi-Mehrvarz et al. 2001;Saeidi-Mehrvarz and Assadi 2003;Saeidi-Mehrvarz 2005, 2011 and the genus includes 61 species with 18 species endemic to the country according to the latest "Flora of Iran" (Saeidi-Mehrvarz 2011). At least two species have since been added to the flora of Iran (Doostmohammadi et al. 2021(Doostmohammadi et al. , 2022. Among these, most species occur in the Zagros and Alborz (also Elburz) Mountains of Iran, which are important centers of diversity of Veronica. ...
... 12808 HKS) was generated using ITS1 and ITS4 primers (White et al. 1990). The new sequence was added to a sequence matrix, based on published sequences of Doostmohammadi et al. (2022). All sequences were aligned using MAFFT v. 6.0 (Katoh and Toh 2010) and edited manually using PhyDE v. 0.9971 (Müller et al. 2010). ...
... (Fig. 6), by far the most species-rich subgenus in Iran, with many perennial, mountainous species. It has been demonstrated that this subgenus has probably originated in the Iranian plateau, with several relict and morphologically isolated species distributed at present along the Zagros and Alborz Mountains (Doostmohammadi et al. 2022). ...
Veronica kurdistanica is described and illustrated as a new species. It grows on calcareous rocks in a few sites in an alpine area in western Iran (Kurdistan province). The new species belongs to V. kurdica species group and is putatively closely related to V. daranica, with which it is com-pared. Veronica kurdistanica is distinguished from V. daranica particularly by its glandular in-dumentum. Molecular phylogenetic analysis of nrDNA (ITS) region confirms this relationship as it is assembled in a sub-clade including V. daranica, V. khorassanica and V. kurdica.
... However, several new taxa were added afterwards (Saeidi-Mehrvarz et al. 2001;Saeidi-Mehrvarz and Assadi 2003;Saeidi-Mehrvarz 2005, 2011 and the genus includes 61 species with 18 species endemic to the country according to the latest "Flora of Iran" (Saeidi-Mehrvarz 2011). At least two species have since been added to the flora of Iran (Doostmohammadi et al. 2021(Doostmohammadi et al. , 2022. Among these, most species occur in the Zagros and Alborz (also Elburz) Mountains of Iran, which are important centers of diversity of Veronica. ...
... 12808 HKS) was generated using ITS1 and ITS4 primers (White et al. 1990). The new sequence was added to a sequence matrix, based on published sequences of Doostmohammadi et al. (2022). All sequences were aligned using MAFFT v. 6.0 (Katoh and Toh 2010) and edited manually using PhyDE v. 0.9971 (Müller et al. 2010). ...
... (Fig. 6), by far the most species-rich subgenus in Iran, with many perennial, mountainous species. It has been demonstrated that this subgenus has probably originated in the Iranian plateau, with several relict and morphologically isolated species distributed at present along the Zagros and Alborz Mountains (Doostmohammadi et al. 2022). ...
A new species, Veronica kurdistanica (Plantaginaceae), is described and illustrated. It grows on limestone cliffs in mountainous alpine areas of western Iran (Kurdistan province). The new species belongs to the species group of V. kurdica and is considered to be closely related to V. daranica , V. khorassanica and V. kurdica , with which the new species is compared. Molecular phylogenetic analysis of nrDNA (ITS) region confirms this relationship. Veronica kurdistanica is distinguished from the mentioned species by its glandular indumentum, length and shape of leaves and bracts, number of flowers per raceme, length and width of calyx and corolla, and size of capsules and seeds.
... Nevertheless, even the 14 species that have been described since 2000 are mostly based on a morphological/phenetic species concept. And even those species that were established based on phylogenetic evidence in a DNA-based phylogenetic analysis were previously recognized at lower ranks and therefore not their recognition but only their rank is based on genetic divergence (Doostmohammadi & al. 2022), with the possible exception of V. dalmatica (Padilla-García & al. 2018). Thus, the question remains whether the morphological/phenetic species concept is sufficient, with other kind of data mostly aiding in deciding on the taxonomic rank. ...
... In essence, the biosystematic species concept of Fischer has simply been updated and expanded by Albach (e. g., Albach 2007, Doostmohammadi & al. 2022) by using DNA-characters and phylogenetic information to assess species rank, similar to how Fischer used ploidy information (Fischer 1967). In fact, the biosystematic species concept of Fischer does not differ considerably from the unified species concept of De Queiroz (2005, 2007 with species being "separately evolving metapopulation lineage segments" (De Queiroz 2007) with morphology, karyology, and DNA-based phylogenetic information as diagnostic properties. ...
... One example is dense, interwoven indumentum. This has, based on DNA sequence data, separately evolved in Veronica bombycina and V. bolkardaghensis, which were considered conspecific but have recently been separated based on DNA sequence data (Doostmohammadi & al. 2022). However, this step has not (yet) been taken for V. incana, which also appears to consist of at least two separate lineages (Nehrke & Albach,in prep.;Fig. ...
The species is the central category of organizing the diversity of life. Yet, many different species concepts, such as the morphological, biological, ecological, and phylogenetic concepts , have been used by biologists, preventing clear comparability. Based on the history of species description in the genus Veronica, I demonstrate that the biosystematics approach by Manfred A. Fischer from the 1960s onwards has started to renew our understanding of what a species is in this genus. Detailed investigations of morphology, ecology, reproductive barriers, and, later, genetic cohesion and testing of species limits have continued from his earliest analyses of the V. hederi-folia-complex. We now recognize 455 species in the genus Veronica but still only few species have been rigorously analyzed, leaving certainly a lot of work for future systematists working on this beautiful genus. K e y w o r d s : biosystematics; history; phylogeny; polyploidy; species concept Z u s a m m e n f a s s u n g : Was ist eine Art in Veronica? Reflexionen anläßlich des 80. Geburtstages von Manfred A. Fischer Die Art ist die zentrale Kategorie für die Organisation der Vielfalt des Lebens. Dennoch wurden von Biologen viele verschiedene Artkonzepte wie das morphologische, biologische, ökologische und phylogenetische Konzept verwendet, was eine klare Vergleichbarkeit von Arten verhindert. Anhand der Geschichte der Artbeschreibung in der Gattung Veronica zeige ich, dass der biosystematische Ansatz von Manfred A. Fischer seit den 1960er-Jahren dazu führte, dass sich unser Verständnis, was eine Art in dieser Gattung ist, komplett erneuerte. Detaillierte Untersuchungen der Morpholo-gie, der Ökologie, der Fortpflanzungsbarrieren und-später-der genetischen Kohäsion sowie die Prüfung der Artgrenzen wurden seit seinen ersten Analysen des V. hederifolia-Komplexes fort-gesetzt. Heute sind 455 Arten in der Gattung Veronica bekannt, aber nur wenige wurden gründ-lich analysiert, so dass es für künftige Systematiker, die sich mit dieser schönen Gattung befassen, sicherlich noch viel zu tun gibt.
... are largely consistent with previous publications. According to our and previously published data on molecular-phylogenetic analyses of the Veronica species [1,2,6,8,10,28,54,55], the Figure 5. Dendrograms of Euclidean distances using Ward's method based on chemical constituents obtained by microwave-assisted water extractions in an amount equal to or greater than 2%. 1-hexahydrofarnesyl acetone; 2-phytol/caryophyllene oxide/hexahydrofarnesyl aceton; 3-phytol. ...
... are largely consistent with previous publications. According to our and previously published data on molecular-phylogenetic analyses of the Veronica species [1,2,6,8,10,28,54,55], the species from Croatia selected for this research belong to eight subgenera: Comparing the two clusters created based on the free volatile compounds (Figures 4 and 5) and based on the molecular data for ITS regions (Figure 2), it can be said that clustering based on the volatile compounds from microwave extraction resulted in clusters that better fit the molecular clusters. The first cluster ( Figure 5) consists of species belonging to Chamaedrys, Pocilla, Stenocarpon and Beccabunga subgenera. ...
(1) Background: The purpose of this study was to compare the free volatile compounds of 18 Veronica species (Plantaginaceae), as previously analyzed by gas chromatography coupled with mass spectrometry, with their DNA sequences for internal transcribed spacers ITS2 and ITS1-5.8S-ITS2 of the nuclear ribosomal DNA. (2) Methods: Two sets of DNA sequence data were generated and used for phylogenetic analysis: ITS2 sequences (~360 bp) obtained by next-generation sequencing and ITS1-5.8S-ITS2 sequences (~580 bp) sequenced by the Sanger sequencing method. Clustering from previously analyzed free volatile compounds was performed by Ward’s method. (3) Results: Both sets of DNA sequence data showed that the 18 analyzed Veronica species were grouped into eight main groups corresponding to the following subgenera: Pentasepalae, Pocilla, Chamaedrys, Veronica, Beccabunga, Cochlidiosperma, Stenocarpon and Pseudolysimachium. Results of the clustering analysis of free volatile compounds showed better clustering when using microwave-extracted volatiles. Three clusters were detected with the following main compounds: hexahydrofarnesyl acetone, hexadecanoic acid, phytol, caryophyllene oxide and (E)-caryophyllene. (4) Conclusion: The phylogenetic analysis of ITS2 data obtained by NGS technology and ITS1-5.8S-ITS2 data obtained by Sanger sequencing resulted in the grouping of 18 Veronica species into eight subgenera, which is in accordance with the existing classification. Statistical testing showed that there was no correlation between such clustering of Veronica species and clustering that was based on free volatile compounds. The achieved results can be viewed in the light of parallel evolution among some of the species of the Veronica genus as well as the fact that volatile compound composition can be influenced by environmental factors or epigenetic modifications.
... Stereothrix (Fabaceae, Fabales, eudicots) [24] and Veronica subg. Pentasepalae (Plantaginaceae, Lamiales, eudicots) [25] from Eurasia were addressed. Whole plastome comparison revealed phylogenetic relationships in Crassula (Crassulaceae, Saxifragales, eudicots) [26] and in the family Magnoliaceae (Magnoliales, early branching angiosperms) [27]. ...
Systematics and taxonomy are basic sciences and are crucial for all applications dealing with living organisms [...]
Cryptic species are black boxes of taxonomy and need to be addressed with an integrative taxonomic approach. Simulium (Diptera: Simuliidae) may contain a substantial amount of cryptic species diversity due to its large cross-continental distribution and habitat-based canalization in taxonomic characters. Except for Simulium sergenti, the following six species Simulium equinum, Simulium paraequinum, Simulium pseudequinum, Simulium turgaicum, Simulium balcanicum, and Simulium lineatum are the most widely distributed species of the subgenus Wilhelmia in the Palearctic region. However, the conspecific diversity and evolutionary history of these species are not yet well understood. In this study, therefore, we used mitochondrial COI and nuclear ITS1-2 sequences to perform species delimitation tests, and demographic and phylogeographic analyses to understand candidate species and evolutionary history of the seven species. While our results did not support the species status of S. balcanicum, S. turgaicum, and S. lineatum, it did suggest several candidate species within S. equinum and S. pseudequinum. Molecular clock analysis revealed that significant branching events occurred in the Pliocene, and demographic fluctuations in the populations occurred intensely during the Pleistocene. Overall, the results indicated that Anatolia has served as a significant refugium for ancestral haplotypes, and it is an important dispersal corridor between the eastern and western Palearctic for Wilhelmia species.
تاب «فلور ایران: تیره گل میمون (Scrophulariaceae Juss.»، شماره 68ام از مجموعه کتابهای فلور ایران به زبان فارسی است که همسو با اجرای پروژه ملی فلور ایران (مجری: دکتر مصطفی اسدی)، تهیه و توسط مؤسسه تحقیقات جنگلها و مراتع کشور در سال 1390 منتشر شد. این جلد توسط مجموعهای از نویسندگان شامل شهریار سعیدیمهرورز، فریده عطار، سیدمحمدمهدی حمدی، فریبا شریفنیا، مصطفی اسدی، صدیقه یوسفنعنایی و ایرج مهرگان نوشته شده است و ویراستاران آن مصطفی اسدی، علیاصغر معصومی، پرویز باباخانلو و ولیاله مظفریان هستند. این کتاب در 483 صفحه تألیف شده است، طرح روی جلد آن مشابه با فلورهای پیشین (فلور ایران، اسدی 1367) است که بهصورت رنگی و گلاسه و صفحات آن با کاغذ مناسب چاپ شده است.
