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Abstract

Detailed phylogenetic relationships, evolutionary histories and phylogeographical hypotheses are still quite rare for the many genera of Apiaceae. One of the reasons for this is that traditional generic circumscriptions based on morphology and anatomy largely failed to recognize the group as monophyletic. This is also true for apioid members of Apiaceae with c. 400 genera. Here we focus on Grammosciadium s.l. centred in Turkey and unravel its evolutionary history. Based on three loci from the plastid genome and nuclear-encoded internal transcribed spacer regions 1 and 2 from the nuclear encoded ribosomal RNA operon, we show that Grammosciadium in its current circumscription is not monophyletic and represents an assemblage of species from four genera (Carum, Chamaesciadium, Fuernrohria, Grammosciadium). Diversification of this group started c. 7 Mya in the late Miocene. Most present-day species arose during the Pliocene with most of the intra-species diversification occurring during the Pleistocene. Plastome type variation does not resolve any of the main clades and there is little spatial structure of the distribution of high plastid genetic variation. The majority of species of pre-Quarternary origin are found east of the Anatolian Diagonal, running diagonally across central and eastern Turkey. This might indicate that the Diagonal acted as a barrier to gene flow and migration during the Pliocene and that eastern Anatolia may have served as a cradle for Pleistocene diversification and speciation processes in Turkey. We also provide a key to the species of the revised genus and introduce several taxonomic changes.

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... The monotypic Fuernrohria, and the allied genera Grammosciadium, Caropodium Stapf & Wettst. ex Stapf (1886: 317) and Vinogradovia Bani, D.A.German & M.A.Koch (2017: 184) were studied in a recent phylogenetic work (Koch et al. 2017). Those latter authors (Koch et al. 2017) corroborated the close relationships of Fuernrohria to taxa of Grammosciadium and Caropodium, as it had already been shown in several previous studies (Katz -Downie et al. 1999, 2001, Valiejo-Roman et al. 2006, Koch et al. 2017 Hedge et al. (1971: 75)) in the molecular phylogenies, it is obvious that Fuernrohria and Vinogradovia are the closest relatives, and they are sister to each other within the Grammosciadium clade (Koch et al. 2017). ...
... ex Stapf (1886: 317) and Vinogradovia Bani, D.A.German & M.A.Koch (2017: 184) were studied in a recent phylogenetic work (Koch et al. 2017). Those latter authors (Koch et al. 2017) corroborated the close relationships of Fuernrohria to taxa of Grammosciadium and Caropodium, as it had already been shown in several previous studies (Katz -Downie et al. 1999, 2001, Valiejo-Roman et al. 2006, Koch et al. 2017 Hedge et al. (1971: 75)) in the molecular phylogenies, it is obvious that Fuernrohria and Vinogradovia are the closest relatives, and they are sister to each other within the Grammosciadium clade (Koch et al. 2017). According to Ulusoy et al. (2017) and Koch et al. (2017), some morphological and anatomical characters such as the presence of sclerenchyma tissue or stone cells in transverse sections of roots, and also the shape and number of vascular bundles in transverse sections of fruits, support the phylogenetic relationships outlined above. ...
... ex Stapf (1886: 317) and Vinogradovia Bani, D.A.German & M.A.Koch (2017: 184) were studied in a recent phylogenetic work (Koch et al. 2017). Those latter authors (Koch et al. 2017) corroborated the close relationships of Fuernrohria to taxa of Grammosciadium and Caropodium, as it had already been shown in several previous studies (Katz -Downie et al. 1999, 2001, Valiejo-Roman et al. 2006, Koch et al. 2017 Hedge et al. (1971: 75)) in the molecular phylogenies, it is obvious that Fuernrohria and Vinogradovia are the closest relatives, and they are sister to each other within the Grammosciadium clade (Koch et al. 2017). According to Ulusoy et al. (2017) and Koch et al. (2017), some morphological and anatomical characters such as the presence of sclerenchyma tissue or stone cells in transverse sections of roots, and also the shape and number of vascular bundles in transverse sections of fruits, support the phylogenetic relationships outlined above. ...
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In this study, morphological and anatomical data (including fruit micromorphological characters) of the genus Fuernrohria were studied and compared with the related genera Grammosciadium, Caropodium and Vinogradovia. The morphological description of Fuernrohria setifolia, which is the only known species in the genus, is expanded. Photos of inflorescences, infructescences and leaves of the species taken from the wild are provided. Leaf segment and mericarps are illustrated, and also the distribution area of the genus is mapped. Anatomical features of root, stem, leaf sheath, leaf segment and fruit are presented with photographs of cross sections for each of them. Vegetative anatomical characteristics of the species and micromorphological description of fruit are given and exhibited for the first time in the present study. The results show that two important diagnostic characters for Fuernrohria are determined for the first time as “shape and size of commissural vittae in mericarps” and “number of vascular bundles in leaf cross section”.
... nezaketiae B.Bani and G. cornutum (Nábělek) C.C.Towns. distributed in Turkey [15]. Based on the importance of the distinctive property of fruit anatomical characters in the Apiaceae family, detailed fruit anatomy of these 4 taxa belonging to the genus Grammosciadium is given in this study. ...
... These findings support the previous studies that justified the large vascular bundles of the fruits of G. macrodon subsp. nezaketiae [15,25]. ...
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... Flora 263 (2020) 151536 (continued on next page) N. Friesen, et al. Flora 263 (2020) 151536 ITS-2) was amplified using primers ITS-A (Blattner, 1999) and ITS-25 r (5'-GGGTAATCCCGCCTGACCTGG-3', developed by K. Mummenhoff or/and M. Koch in the nineties, but the right sequence was first published in Koch et al. 2017). The PCR conditions were identical, as described in Friesen et al. (2006). ...
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... In this pattern, the lineages of Apathya between the Anatolian and Iranian groups diverged into two clusters around 13.66 mya in the middle Miocene, and a vicariant event creating an east and west sea barrier between the southeastern Anatolia and Iran led to the split of genus Apathya in this area (Kapli et al., 2013). This vicariant event not only affected speciation of animals in the area, diversification in plant genus Grammosciadium began in Serkan Gül approximately the Miocene 7.5 Mya, and G. cornutum and G. scabridium are restricted to along the glacier mountains of Hakkari (Koch et al., 2017). ...
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... Drude (1898) and Heywood (1971) also included the genus to Scandiceae-Scandicinae. Koso-Poljansky (1916) listed the genus among genera related to Oenantheae. Molecular studies attributed Rhabdosciadium and Grammosciadium to the clade Careae (Downie et al. 2000;Valiejo-Roman et al. 2006;Koch et al. 2017), together with Aegopodium Linnaeus (1753: 265), Caropodium Stapf & Wettstein ex Stapf (1886: 317), Carum Linnaeus (1753: 263), Chamaesciadium Meyer (1831: 122), Falcaria Fabricius (1759: 34), Fuernrohria Karl Koch (1842: 356), Gongylosciadium Rechinger (1987: 308), Grammosciadium s.s., Hladnikia Wilhelm Koch (1835: 320) and Olymposciadium Hermann Wolff (1922: 132). Such a position of Rhabdosciadium and Grammosciadium is rather distant from the other Scandiceae. ...
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Unlabelled: CONSEL is a program to assess the confidence of the tree selection by giving the p-values for the trees. The main thrust of the program is to calculate the p-value of the Approximately Unbiased (AU) test using the multi-scale bootstrap technique. This p-value is less biased than the other conventional p-values such as the Bootstrap Probability (BP), the Kishino-Hasegawa (KH) test, the Shimodaira-Hasegawa (SH) test, and the Weighted Shimodaira-Hasegawa (WSH) test. CONSEL calculates all these p-values from the output of the phylogeny program packages such as Molphy, PAML, and PAUP*. Furthermore, CONSEL is applicable to a wide class of problems where the BPs are available. Availability: The programs are written in C language. The source code for Unix and the executable binary for DOS are found at http://www.ism.ac.jp/~shimo/ Contact: shimo@ism.ac.jp
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Background and Aims Aubrieta is a taxonomically difficult genus from the Brassicaceae family with approximately 20 species centred in Turkey and Greece. Species boundaries and their evolutionary history are poorly understood. Therefore, we analysed bio- and phylogeographic relationships and evaluated morphological variation to study the evolution of this genus. Methods Phylogenetic analyses of DNA sequence variation of nuclear-encoded loci and plastid DNA were used to unravel phylogeographic patterns. Morphometric analyses were conducted to study species delimitation. DNA sequence-based mismatch distribution and climate-niche analyses were performed to explain various radiations in space and time during the last 2·5 million years. Key Results Species groups largely show non-overlapping distribution patterns in the eastern Mediterranean and Asia Minor. We recognized 20 species and provide evidence for overlooked species, thereby highlighting taxonomical difficulties but also demonstrating underexplored species diversity. The centre of origin of Aubrieta is probably Turkey, from which various clades expanded independently towards Asia Minor, south to Lebanon and west to Greece and the Balkans during the Pleistocene. Conclusions Pleistocene climatic fluctuations had a pronounced effect on Aubrieta speciation and radiation during the last 1·1 million years in the Eastern Mediterranean and Asia Minor. In contrast to many other Brassicaceae, speciation processes did not involve excessive formation of polyploids, but displayed formation of diploids with non-overlapping present-day distribution areas. Expansions from the Aubrieta centre of origin and primary centre of species diversity showed adaptation trends towards higher temperature and drier conditions. However, later expansion and diversification of taxa from within the second centre of species diversity in Greece started ∼0·19 Mya and were associated with a general transition of species adaptation towards milder temperatures and less dry conditions.
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Ever since the 19th century, the immense arid lands of the Orient, now called the Irano-Turanian (IT) floristic region, attracted the interest of European naturalists with their tremendous plant biodiversity. Covering approximately 30% of the surface of Eurasia (16000000 km2), the IT region is one of the largest floristic regions of the world. The IT region represents one of the hotspots of evolutionary and biological diversity in the Old World, and serves as a source of xerophytic taxa for neighbouring regions. Moreover, it is the cradle of the numerous species domesticated in the Fertile Crescent. Over the last 200 years, naturalists outlined different borders for the IT region. Yet, the delimitation and evolutionary history of this area remain one of the least well-understood fields of global biogeography, even though it is crucial to explaining the distribution of life in Eurasia. No comprehensive review of the biogeographical delimitations nor of the role of geological and climatic changes in the evolution of the IT region is currently available. After considering the key role of floristic regions in biogeography, we review the history of evolving concepts about the borders and composition of the IT region over the past 200 years and outline a tentative circumscription for it. We also summarise current knowledge on the geological and climatic history of the IT region. We then use this knowledge to generate specific evolutionary hypotheses to explain how different geological, palaeoclimatic, and ecological factors contributed to range expansion and contraction, thus shaping patterns of speciation in the IT region over time and space. Both historical and ecological biogeography should be applied to understand better the floristic diversification of the region. This will ultimately require evolutionary comparative analyses based on integrative phylogenetic, geological, climatic, ecological, and species distribution studies on the region. Furthermore, an understanding of evolutionary and ecological processes will play a major role in regional planning for protecting biodiversity of the IT region in facing climatic change. With this review, we aim to introduce the IT floristic region to a broader audience of evolutionary, ecological and systematic biologists, thus promoting cutting-edge research on this area and raising awareness of this vast and diverse, yet understudied, part of the world.
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Aim: To estimate the most likely pathways of expansion of the first diverging lineages of the angiosperm family Apiaceae across the Southern Hemisphere and to Eurasia by reconstructing the biogeographical history of the family through space and time. Location: Southern Hemisphere, with emphasis on Africa. Methods: Divergence times were assessed under a penalized-likelihood method (r8s) and a data set of 129 cpDNA rps16 intron sequences. Confidence intervals were estimated using ABCq, BCa, bootstrap-t and standard normal methods. Biogeographical distributions were reconstructed using DEC analyses over rps16 intron and/or nrDNA ITS trees. Results: Crown Apiaceae likely originated by the Late Cretaceous in Australasia. Apiaceae subfamilies diverged between 45.9 and 71.2 Ma in the Southern Hemisphere, specifically, Mackinlayoideae in Australasia, Azorelloideae in South America and Apioideae and Saniculoideae in southern Africa. From the Palaeocene to Oligocene, Africa showed connections via transoceanic dispersals as a sink continent with Australasia and as a source continent with South America and Eurasia. These dispersals explain the present intercontinental disjunctions of the subfamilies. The first diverging lineages of Apioideae and Saniculoideae likely originated in Africa and diversified in situ since the Palaeocene, with no input from newcomers until the Miocene. Subsequently, several dispersals mainly from Eurasia are estimated back to northern and eastern Africa. Main conclusions: The Southern Hemisphere has played a key role in the origin and early diversification of Apiaceae, currently a mostly north temperate family. African Apiaceae was likely assembled by Palaeocene lineages that diversified in situ and are now restricted mainly to southern Africa and post Miocene newcomers mostly restricted to northern and eastern Africa by effective environmental barriers.
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The genus Grammosciadium as circumscribed by Boissier (1844, 1872) should not be divided by splitting of Caropodium and Stenodiptera. A review of the nomenclature of the seven species is given, followed by a key to the four sections.
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Apiaceae (Umbelliferae) is a large angiosperm family that includes many medicinally important species. The ability to identify these species and their adulterants is important, yet difficult to do so because of their subtle fruit morphological differences and often lack of diagnostic features in preserved specimens. Moreover, dried roots are often the official medical organs, making visual identification to species almost impossible. DNA barcoding has been proposed as a powerful taxonomic tool for species identification. The Consortium for the Barcode of Life (CBOL) Plant Working Group has recommended the combination of rbcL+matK as the core plant barcode. Recently, the China Plant BOL Group proposed that the nuclear ribosomal DNA internal transcribed spacer (ITS), as well as a subset of this marker (ITS2), be incorporated alongside rbcL+matK into the core barcode for seed plants, particularly angiosperms. In this study, we assess the effectiveness of these four markers plus psbA-trnH as Apiaceae barcodes. A total of 6,032 sequences representing 1,957 species in 385 diverse genera were sampled, of which 211 sequences from 50 individuals (representing 7 species) were newly obtained. Of these five markers, ITS and ITS2 showed superior results in intra- and interspecific divergence and DNA barcoding gap assessments. For the matched dataset (173 samples representing 45 species in 5 genera), the ITS locus had the highest identification efficiency (73.3%), yet ITS2 also performed relatively well with 66.7% identification efficiency. The identification efficiency increased to 82.2% when using an ITS+psbA-trnH marker combination (ITS2+psbA-trnH was 80%), which was significantly higher than that of rbcL+matK (40%). For the full sample dataset (3,052 ITS sequences, 3,732 ITS2 sequences, 1,011 psbA-trnH sequences, 567 matK sequences, and 566 rbcL sequences), ITS, ITS2, psbA-trnH, matK, and rbcL had 70.0%, 64.3%, 49.5%, 38.6%, and 32.1% discrimination abilities, respectively. These results confirm that ITS or its subset ITS2 be incorporated into the core barcode for Apiaceae and that the combination of ITS/ITS2+psbA-trnH has much potential value as a powerful, standard DNA barcode for Apiaceae identification.This article is protected by copyright. All rights reserved.
Article
AimLong-distance dispersal (LDD) by migratory birds is often invoked to explain the broad and disjunct distributions of many aquatic plants. Such distributions may also be achieved by recurrent short-distance dispersal (SDD) to adjacent areas and extinctions in connecting areas. To test the relative importance of LDD and SDD in shaping the distributions of hydrophytes, we examined different dispersal models for a clade of hydrophytic umbellifers (Apiaceae tribe Oenantheae).LocationWorldwide, with emphasis on Eurasian–North American disjunctions.MethodsA dated phylogeny of the group was obtained with Bayesian methods using nrDNA internal transcribed spacer (ITS) and cpDNA rps16–trnK sequences from 100 species and infraspecific taxa of Oenantheae and two outgroup species. Ancestral habits were inferred using maximum likelihood (R package ape). Six connectivity models were compared using a maximum-likelihood-based method (Lagrange), four with symmetrical dispersal rate matrices and two assuming asymmetrical exchanges between Eurasia and North America, with or without spatial and temporal constraints on LDD.ResultsThe age of the crown node of Oenantheae was estimated at 26.3 Ma, and western Europe was reconstructed as its most likely ancestral area. The divergence between Peucedanum sandwicense, an endemic of Hawaii, and its sister Oenanthe occurred 17.2 Ma, pre-dating the emergence of the Hawaiian Islands. Throughout the phylogeny, the hydrophytic habit (including helophytes and amphiphytes) dominated. Of the six connectivity models considered, the model restricting all intracontinental and transoceanic LDDs and assuming an almost unidirectional dispersal from Eurasia to North America received the highest likelihood score. This model was also characterized by the highest dispersal rate. A stratified model assuming a higher probability for dispersals between Eurasia and North America when these continents were connected with land bridges received a lower likelihood score.Main conclusionsThe results suggest that the broad and often disjunct distribution of Oenantheae hydrophytes is mostly achieved through an increased dispersal rate and recurrent SDD rather than frequent LDD. Our data confirm the asymmetry of the floristic exchange between Eurasia and North America and do not support the increase of this exchange when the continents were connected with land bridges.
Article
To analyse spatial and temporal patterns of dispersal events in the euapioids (Apiaceae subfamily Apioideae). Worldwide, with an emphasis on the Northern Hemisphere. A phylogeny of euapioids was inferred from 1194 nuclear ribosomal DNA internal transcribed spacer (nrDNA ITS) sequences using Bayesian methods. The reconstruction of ancestral areas was performed simultaneously with phylogenetic inference using a Markov discrete phylogeographical model with Bayesian stochastic search variable selection (BSSVS). Routes with significant non-zero migration rates were identified using Bayes factors. For each significant track and each tree, the distribution of dispersals in time was scored and the asymmetry of dispersals was evaluated. The root of the euapioid umbellifers was reconstructed at c. 44.51 Ma (95% highest posterior density interval: 39.11–51.55 Ma). The Eastern Asiatic Region was reconstructed as the most probable ancestral area for the root of the tree. Seventeen migration routes have significant non-zero migration rates. Five of these tracks represent long-distance transoceanic routes: (1) western Eurasia–North America, (2) Eastern Asiatic Region–North America, (3) Australia–Neotropical Kingdom, (4) Australia–Neantarctic, and (5) Neotropical Kingdom–sub-Saharan Africa. Most dispersal events occurred among the areas that comprise the major diversification centres of apioid umbellifers: western Eurasia and the Mediterranean, Irano-Turanian and Eastern Asiatic regions. The floristic exchange among these regions was more or less symmetrical in all directions and continuous in time. The exchange between North America and the Eastern Asiatic Region was asymmetrical; the asymmetry of dispersals between western Eurasia and North America was less pronounced and not statistically significant. Floristic exchange was significantly asymmetrical for the Nearctic–Neotropical Kingdom, Nearctic–Neantarctic, western Eurasia–sub-Saharan Africa, and western Eurasia–Siberia migration tracks. The observed dispersal pattern – intense and symmetrical within the same climatic zone versus scarce and often unidirectional between climatic zones or along long-distance transoceanic tracks – suggests that both the availability of suitable habitats and geographical barriers have played crucial roles in determining the present distribution of euapioid umbellifers.
Article
Fossil pollen of the Umbelliferae occur frequently and with considerable diversity in Eocene sediments in France. We have been able to recognise 43 forms belonging to 23 genera. The Thanetian II, the Sparnacian, the Marinesian and the Ludian are characterised by particular assemblages. While in the Thanetian II pollen of a very archaic character predominate, the Ludian shows the greatest diversification of the family.With respect to climate, the Sparnacian warming is confirmed by the appearance of Steganotaenia, a taxon characteristic of the tropical savannah. A drier climatic phase developed in the Marinesian, indicated by the development of Hohenackeria, a mediterranean taxon of xerophilous affinities. The appearance and extension of modern temperate taxa in the Ludian confirm the climatic change at the end of the Eocene.RésuméLes pollens fossiles d'Ombellifères sont fréquents et diversifiés dans les sédiments de l'Eocène français. Nous avons pu distinguer 43 formes rapportées à 23 genres. Le Thanétien II, le Sparnacien, le Marinésien et le Ludien sont caractérisés par des ensembles particuliers. Si dans le Thanétien II les pollens à caractère très archaïque dominent, on constate, au Ludien, la grande diversification de cette famille.Du point de vue climatique, le réchauffement du Sparnacien est confirmé par l'apparition de Steganotaenia, taxon de savane tropicale. Une phase climatique plus sèche semble se developper au Marinésien, comme en témoigne le développemment d'Hohenackeria, taxon méditerranéen assez xérophile. Au Ludien, l'apparition ou l'extension de taxons actuellement tempérés confirme le changement climatique de la fin de l'Eocène.
Article
Synopsis An analysis of the 8 published volumes of the Flora of Turkey was made in order to determine if there was a factual basis for the concept of an Anatolian Diagonal affecting plant distribution in Turkey. The data accumulated indicated that a substantial number of species ( c. 2600) have distributions associated with the Diagonal this total accounts for c . 32% of the number of species in the 8 volumes: 5% are ± restricted to the Diagonal; 14% are absent from the W of it and 13% from the E. Some suggestions are made for the reasons for this floral break.
Article
The majority of endemic and subendemic Iranian Umbelliferae taxa have never been investigated using molecular approachs to phylogeny. 36 species from 28 genera of the Umbelliferae from this region were included for the first time in analysis using ITS sequence data to clarify their systematics. For some newly investigated (Leiotulus, Semenovia, Ducrosia, Echinophora, Pycnocycla, Turgenia lisaeoides, Prangos, Alococarpum, Bilacunaria, Elaeosticta, Johreniopsis, Pseudotrachydium) taxa the molecular data correspond to, or at least do not contradict with, their traditional, morphologically grounded taxonomy. Leutea was shown closely related to Ferula (not Peucedanum), and Opsicarpium – to Pimpinella. A taxonomically isolated position of Mozaffariania has been demonstrated. Haussknechtia appeared to be the closest relative of Demavendia and Zeravschania. It was shown that Calyptrosciadium belongs to Komarovia -clade, having long-distant geographical disjunction. Close affinity of Heptaptera and Prangos was not supported. Similarly, Thecocarpus appeared to be distant from other Echinophoreae, putting monophyly of the tribe in question. As a result, new molecular data have cleared up systematic questions in some taxonomically uncertain Iranian Umbelliferae. (© 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)
Article
Aim The angiosperm genus Cryptotaenia (family Apiaceae, tribe Oenantheae) exhibits an anomalous distribution pattern, with five of its eight species being narrow endemics geographically isolated from their presumed relatives. We examined the monophyly of the genus and ascertained the phylogenetic placements of its constituent members in order to explain their distribution patterns. Location Eastern North America, eastern Asia, the Caucasus, southern Italy, Macaronesia and Africa. Methods In total, 173 accessions were examined for nuclear rDNA ITS sequence variation, representing nearly all major lineages of Apiaceae subfamily Apioideae and seven species of Cryptotaenia. Sampling of tribes Oenantheae, Scandiceae and Pimpinelleae was comprehensive. Phylogenetic analyses included Bayesian, maximum parsimony and neighbour-joining methods; biogeographical scenarios were inferred using dispersal–vicariance analysis (diva). Results Cryptotaenia is polyphyletic and includes three distant lineages. (1) Cryptotaenia sensu stricto (C. canadensis, C. japonica, C. flahaultii and C. thomasii) is maintained within tribe Oenantheae; C. canadensis and C. japonica, representing an eastern North American–eastern Asian disjunction pattern, are confirmed to be sister species. (2) Cryptotaenia elegans, endemic to the Canary Islands, is placed within Scandiceae subtribe Daucinae along with two woody endemics of Madeira, Monizia edulis and Melanoselinum decipiens. The phylogeny of these Canarian and Madeiran endemics is unresolved. Either they constitute a monophyletic sister group to a clade comprising some Mediterranean and African species of Daucus and their relatives, or they are paraphyletic to this clade. The herbaceous/woody genus Tornabenea from Cape Verde, once included in Melanoselinum, is not closely related to the other Macaronesian endemics but to Daucus carota. (3) The African members of Cryptotaenia (C. africana, C. calycina and possibly C. polygama) comprise a clade with some African and Madagascan umbellifers; this entire clade is sister group to Eurasian Pimpinella. Main conclusions Elucidating the phylogeny of the biogeographically anomalous Cryptotaenia sensu lato enabled hypotheses on the biogeography of its constituent lineages. Cryptotaenia sensu stricto exhibits a holarctic distribution pattern, with its members occurring in regions that were important glacial refugia. The genus probably originated in eastern Asia and from there dispersed to Europe and North America. For the Macaronesian endemic species –C. elegans, M. edulis and M. decipiens–diva reconstructs either a single dispersal event to Macaronesia from the Mediterranean/African region, or a single dispersal followed by a back-dispersal to the mainland. The radiation of Tornabenea from Cape Verde followed a second dispersal of Daucinae to Macaronesia. Woodiness in Melanoselinum/Monizia and Tornabenea, therefore, is a derived and independently acquired trait. The African members of Cryptotaenia are derived from an ancestor arriving from the Middle East.
Article
Premise of the study: The high mountains in southern Anatolia and the eastern Mediterranean are assumed to play a major role as a primary center of genetic diversity and species richness in Eurasia. We tested this hypothesis by focusing on the widespread perennial arctic-alpine Arabis alpina and its sympatrically distributed closest relatives in the eastern Mediterranean. Methods: Plastid (trnL intron, trnL-F intergenic spacer) and nuclear (ITS) DNA sequence analysis was used for phylogenetic reconstruction. Broad-scale plastid haplotype analyses were conducted to infer ancestral biogeographic patterns. Key results: Five Arabis species, identified from the eastern Mediterranean (Turkey mainland and Cyprus), evolved directly and independently from A. alpina, leaving Arabis alpina as a paraphyletic taxon. These species are not affected by hybridization or introgression, and species divergence took place at the diploid level during the Pleistocene. Conclusions: Pleistocene climate fluctuations produced local altitudinal range-shifts among mountain glacial survival areas, resulting not only in the accumulation of intraspecific genotype diversity but also in the formation of five local species. We also show that the closest sister group of Arabis alpina consists exclusively of annuals/winter annuals and diverged prior to Pleistocene climatic fluctuations during the colonization of the lowland Mediterranean landscape. These findings highlight that Anatolia is not only a center of species richness but also a center for life-history diversification.
Article
Arabis drummondii, A. holboellii and their hybrid A. x divaricarpa are widespread perennials of open habitats in North America. A phylogenetic analysis based on noncoding chloroplast DNA sequences (trnL intron and trnL/F intergenic spacer) resolved A. drummondii as a monophyletic taxon, but found A. holboellii to bear chloroplast haplotypes from highly diverged evolutionary lineages. This raised the question of a possible polyphyletic origin of A. holboellii. Arabis x divaricarpa was found to be of recent and polytopic origin, a result consistent with its presumed hybrid origin. One hundred and three chloroplast haplotypes were detected within 719 Arabis accessions investigated. The majority of chloroplast-types were estimated to have arisen prior to the Wisconsin glaciation. Phylogeographical analysis using nested clade analysis, suggested for A. holboellii (i). past fragmentation events, partitioning genetic variation in several instances between the Sierra Nevada, the Southern Rocky Mountains and the Colorado Plateau on the one hand and the Central to Northern Rockies of the United States and adjacent Cascades on the other; and for both parental species (ii). recolonization of major areas formerly covered by the Wisconsin glaciation by three haplotypes; and (iii). restricted gene flow indicating isolation by distance in areas south of the last glacial maximum. Arabis x divaricarpa is closely codistributed with its parental species and resampled their haplotypes. The highest genetic diversity was found in the Rocky Mountains from Idaho and Montana south to Utah and Colorado. This area was further hypothesized to have played a major role in the origin of both parental species and probably represented an important glacial refugium. However, evidence for glacial refugia was also found in arctic and boreal regions of Alaska and near the Great Lakes. In comparison to nuclear ribosomal internal transcribed spacer data, chloroplast DNA divergence was very high and evidently predated the origin of A. drummondii and possibly A. holboellii. Divergence of major chloroplast lineages dates back to the middle of the Pleistocene at least. Extensive hybridization is the most likely evolutionary factor working on A. holboellii to explain the revealed discrepancy in nuclear DNA and chloroplast DNA diversification.
Article
The evolutionary history of a set of taxa is usually represented by a phylogenetic tree, and this model has greatly facilitated the discussion and testing of hypotheses. However, it is well known that more complex evolutionary scenarios are poorly described by such models. Further, even when evolution proceeds in a tree-like manner, analysis of the data may not be best served by using methods that enforce a tree structure but rather by a richer visualization of the data to evaluate its properties, at least as an essential first step. Thus, phylogenetic networks should be employed when reticulate events such as hybridization, horizontal gene transfer, recombination, or gene duplication and loss are believed to be involved, and, even in the absence of such events, phylogenetic networks have a useful role to play. This article reviews the terminology used for phylogenetic networks and covers both split networks and reticulate networks, how they are defined, and how they can be interpreted. Additionally, the article outlines the beginnings of a comprehensive statistical framework for applying split network methods. We show how split networks can represent confidence sets of trees and introduce a conservative statistical test for whether the conflicting signal in a network is treelike. Finally, this article describes a new program, SplitsTree4, an interactive and comprehensive tool for inferring different types of phylogenetic networks from sequences, distances, and trees.
Article
Continental shelf island systems, created by rising sea levels, provide a premier setting for studying the effects of past fragmentation, dispersal, and genetic drift on taxon diversification. We used phylogeographical (nested clade) and population genetic analyses to elucidate the relative roles of these processes in the evolutionary history of the Aegean Nigella arvensis alliance (= 'coenospecies'). We surveyed chloroplast DNA (cpDNA) variation in 455 individuals from 47 populations (nine taxa) of the alliance throughout its core range in the Aegean Archipelago and surrounding mainland areas of Greece and Turkey. The study revealed the presence of three major lineages, with largely nonoverlapping distributions in the Western, Central, and Eastern Aegean. There is evidence supporting the idea that these major lineages evolved in situ from a widespread (pan-Aegean) ancestral stock as a result of multiple fragmentation events, possibly due to the influence of post-Messinian sea flooding, Pleistocene eustatic changes and corresponding climate fluctuations. Over-sea dispersal and founder events appear to have played a rather insignificant role in the group's history. Rather, all analytical approaches identified the alliance as an organism group with poor seed dispersal capabilities and a susceptibility to genetic drift. In particular, we inferred that the observed level of cpDNA differentiation between Kikladian island populations of Nigella degenii largely reflects population history, (viz. Holocene island fragmentation) and genetic drift in the near absence of seed flow since their time of common ancestry. Overall, our cpDNA data for the N. arvensis alliance in general, and N. degenii in particular, indicate that historical events were important in determining the phylogeographical patterns seen, and that genetic drift has historically been relatively more influential on population structure than has cytoplasmic gene flow.