ArticlePDF Available

Abstract

As the result of surveying the relevant type specimens, together with macro‐ and micro‐morphological studies, chromosome counting and ITS sequencing, Astragalus trifoliastrum was found to be a species independent of A. laguriformis (with which it has peviously been synonymized). In contrast, A. wanensis , assumed to be a synonym of A. trifoliastrum , indeed appears to be identical with A. trifoliastrum . The diploid chromosome number of 2n = 16 is reported for the first time for A. trifoliastrum .
Astragalus trifoliastrum (Fabaceae), a revived species for the
ora of Turkey
Ali Bagheri , Seher Karaman Erkul , Ali Asghar Maassoumi , Mohammad Reza Rahiminejad
and Frank R. Blattner
A. Bagheri and M. R. Rahiminejad, Dept of Biology, Faculty of Sciences, Univ. of Isfahan, Isfahan, 73441-81746, Iran. S. K. Erkul
(seherkaraman@yahoo.com), Dept of Biology, Faculty of Arts and Sciences, Aksaray Univ., PO Box 68100, Aksaray, Turkey. A. A. Maassoumi,
Dept of Botany, Research Inst. of Forests and Rangelands, PO Box 13185-116, Tehran, Iran. F. R. Blattner, Leibniz-Inst. of Plant Genetics and
Crop Research (IPK Gatersleben), Corrensstr. 3, DE-06466 Gatersleben, Germany. FRB also at: German Centre of Integrative Biodiversity
Research (iDiv) Halle-Jena-Leipzig, DE-04103 Leipzig, Germany.
As the result of surveying the relevant type specimens, together with macro- and micro-morphological studies, chromo-
some counting and ITS sequencing, Astragalus trifoliastrum was found to be a species independent of A. laguriformis
(with which it has peviously been synonymized) . In contrast, A. wanensis , assumed to be a synonym of A. trifoliastrum ,
indeed appears to be identical with A. trifoliastrum . e diploid chromosome number of 2n 16 is reported for the fi rst
time for A. trifoliastrum .
In Astragalus L., with more than 3000 species and one of the
largest genera of fl owering plants (Podlech and Zarre 2013),
478 species are currently recognized in Turkey (Ayta ç and
Ekici 2012), and 202 of them (42%) are endemic.
Astragalus trifoliastrum Hub.-Mor. & V.A. Matthews in
A. sect. Hymenostegis Bunge is a taxonomically problematic
taxon. It was described from Van in eastern Turkey based on
the Renz and Simon or Huber-Morath collections. In their
revision, Zarre and Podlech (1996), as well as Podlech and
Zarre (2013), treated Astragalus trifoliastrum as a synonym of
A. laguriformis Freyn, a species fi rst described from Erbil in
Iraqi Kurdistan. Another species occurring in the Van area
is Astragalus wanensis Bornm. ex Rech. f. that resembles
samples of A. trifoliastrum .
Considering the diff erent geographical distributions of
A. laguriformis and A. trifoliastrum we here aim to clarify
the veracity of Podlech and Zarre ' s (2013) treatment of
these taxa. For this purpose we analysed the type specimens,
morphology, seed structures, chromosome numbers, and
sequences of the nuclear rDNA internal transcribed spacer
(ITS) region. Moreover, we compared A. trifoliastrum
with A. wanensis to infer if they are conspecifi c or represent
diff erent species.
Material and methods
Specimens examined
is study was based on a number of herbarium vouchers
including:
1) Turkey: Van: G ü rp ı nar, between Hamurkesen-I ş ı kp ı nar,
steppe, 2155 m a.s.l., 13 Jul 2011, S. Karaman 2650
(Fig. 1); ibid., 11 Aug 2011, S. Karaman 2682; Van:
Pass of Kuruba ş , steppe, 1980 m a.s.l., 20 Jul 2013, S.
Karaman 2781; Van: Ho ş ap, dry mudstone hills, near
Ho ş ap Castle, 10 Jul 1968, E. M. Rix et al. 827.(E); Van:
Erek Mount., west slopes, steppe, 2000 m a.s.l., 10 Aug
1989, Z. Ayta ç 1991 and Metzger (GAZI); Van: Ö zalp,
Dambelk village, steppe, 2200 m a.s.l., 5 Jul 1997, F.
Ö zg ö k ç e 5780 (VANF); Van: 7 km from Van to Er ç ek,
igneous slopes, steppe, 1850 m a.s.l., 5 Jun 1966, Davis
44283 (K, E, W).
2) Iraq: Kurdistania, in montis Sefi nadpag. Schaklava (ditio-
nis Erbil), 1000 m a.s.l., 4 Jul (6) 1893, Bornmuller 1194
(holotype: BRNM, isotype: B, W, WU).
3) Iran: Kermanshah, Biston versus Songhor, Kuh-e
Dalekhani, 1800 m a.s.l., Hamzehee and Hatami 1096
(TARI).
SEM micromorphology of seeds
Seed photographs were taken from dry samples of
A. laguriformis and A. trifoliastrum by scanning electron
microscopy (SEM).  e samples were glued on aluminium
stubs with the help of double sided adhesive tape and vac-
uum coated with gold (P ı nar et al. 2009, Altuner et al. 2012,
Ç eter et al. 2013). e images were taken with a Jeol JSM
6060 scanning electron microscope in the SEM laboratory
of the Dept of Biology, Gazi University and TPAO (T ü rkiye
Petrolleri Anonim Ortakl ı ğ ı ).
© 2015  e Authors. Nordic Journal of Botany © 2015 Nordic Society Oikos
Subject Editor: Arne Strid. Editor-in-Chief: Torbj ö rn Tyler. Accepted 3 February 2015
Nordic Journal of Botany 000: 001–008, 2015
doi: 10.1111/njb.00831, ISSN 1756-1051
Early View (EV): 1-EV
... Our investigation and previous studies show that the seeds in most sections of this clade are non-flattened and non-globose, but reniform-suborbicular, reniformelliptic, and ovoid in outline. In the few species with flattened seeds, the shape is almost ovoid or finally with a sunken hilum that can be interpreted as reniform-ellipsoid, mitiform-ellipsoid, ellipsoid-ovoid, mitiform-ovoid (Bacchetta & Brullo, 2010;Erkul & Aytaç, 2013;Bagheri, Erkul, Maassoumi, Rahiminejad & Blattner, 2015;Abd El-Ghani et al., 2021). Oblong seeds (named rectangular-reniform by Ranjbar, 2009) were found only in A. paradoxus . ...
... Oblong seeds (named rectangular-reniform by Ranjbar, 2009) were found only in A. paradoxus . The studied seeds of this clade are 2.1-4 mm long and are classified as medium-sized (Bacchetta & Brullo, 2010;Vural &Şapçı, 2012;Erkul & Aytaç, 2013;Bagheri et al., 2015;Atasagun, Aksoy & Martin, 2018;Kaya, 2023). Interestingly, the largest seeds in this clade are reported in A. dipodurus of sect.Macrophyllium , which are known otherwise to have a large habit (Zarre, 2000), big pollen grains (Ranjbar, Assadi & Karamian, 2012), and high ploidy level (dodecaploid in A. oleifolius : 2n=12x=96: Samad et al. (2014). ...
... Seed surface with narrow to wide pit-like depressions were seen especially in sect. Hymenostegis ,Leucocercis , Microphysa , Poterion andPterophorus (Ranjbar, 2009;Bacchetta & Brullo, 2010;Vural &Şapçı, 2012;Erkul & Aytaç, 2013;Bagheri et al., 2015;Atasagun et al., 2018;Abd El-Ghani et al., 2021;Kaya, 2023). ...
Preprint
In big plant genera such as Astragalus (Fabaceae), the evaluation of morphological evidence from reliable resources is a prerequisite for understanding the evolutionary history, relationships, and taxonomic placements. The present study is the most comprehensive effort to assess the seed morphological characters in a large sample of species. Seed characteristics including size, color, shape, and testa ornamentation of 494 seed samples representing 348 species and 83 sections and all 11 main clades recognized in Astragalus and five species of related genera were investigated using light and Scanning Electron Microscopy. We provide here a standard terminology for all examined characters to suggest a consensus for future seed morphological studies. Accordingly, seed morphological features show considerable variability, making them useful in some taxonomic aspects, but they may also be heterogeneous, which sometimes limits their application. Based on our results none of the natural groups known as section or clades in Astragalus can be defined based on one character, but rather using a combination of distinctive character states. Considering the currently available phylogenetic trees of Astragalus , the present seed morphological survey allows us to suggest some evolutionary trends. In the course of evolution in most clades, the seed size tends to reduce, reddish seeds tend to be frequent over other colors, black dots or spots appear or distribute more densely on the surface, oblong and square as well as semi-oblong-elliptic shapes become abundant, the pit-like depressions on the seed surface tend to increase, and the multiple reticulate-rugulate testa ornamentation appears more frequently.
... Based [30] chromosome numbers for the taxa in A. sect. Hymenostegis are unknown, except for a few species [31][32][33][34]. Taking into account the large number of Astragalus species and their worldwide distribution, genome size analyses are still rare [26,[35][36][37][38]. ...
... Hymenostegis are provided in Table 1, Figures 1 and 2. This number is lower than the number of individuals we initially collected, as not in all cases (enough) seeds germinated to reliably infer karyotypes. Until now only for 12 taxa of section Hymenostegis chromosome numbers have been determined [31][32][33][34]39]. We here add new data for 15 species and confirm earlier chromosome counts for two species. ...
Article
Full-text available
Astragalus section Hymenostegis is one of the important characteristic elements of thorn-cushion formations in the Irano-Turanian floristic region. In this paper, we examined the chromosome number of 17 species (15 new reports) and provide estimates of genome size for 62 individuals belonging to 38 taxa of A. sect. Hymenostegis, some species outside this section, plus two Oxytropis species. Based on chromosome counts 11 species were found to be diploid (2n = 16), four species tetraploid (2n = 32) and two taxa hexaploid (2n = 48). From genome size measurements on silica-gel dried material, three ploidy levels (2x, 4x and 6x) were inferred, with a majority of species being diploid. The 2C values reach from 2.07 pg in diploid Astragalus zohrabi to 7.16 pg in hexaploid A. rubrostriatus. We found indications that species might occur with different cytotypes. A phylogenetic framework using nrDNA ITS sequences was constructed to understand the evolution of ploidy changes and genome sizes. It showed that genome size values among the studied taxa differ only slightly within ploidy levels and are nearly constant within most species and groups of closely related taxa within the genus Astragalus. The results of this study show that there is a rather strong correlation between genome sizes and chromosome numbers in sect. Hymenostegis. The resolution of the ITS-based phylogenetic tree is too low to infer evolutionary or environmental correlations of genome size differences. Polyploidization seems to contribute to the high species number in Astragalus, however, in sect. Hymenostegis it is not the main driver of speciation.
... Additionally, we observed that the indumentum of bracts may vary from glabrous to densely hairy in contrast to the description provided by Podlech , 1996). However, A. trifoliastrum is not actually synonymous; it is reclassified and recorded again (Bagheri et al., 2015). Morphologically, A. trifoliastrum can be distinguished from A. laguriformis by oblong to elliptic leaflets (not narrowly elliptic), ca. ...
... 2.0–3.5 mm), with upper side glabrous and lower side sparsely covered with very short adpressed hairs (not both sides densely covered with adpressed hairs), ovoid raceme (not globose to ovoid), ca. 4–7 cm long (not 2–5 cm), purplish calyx (not yellowish) with ascending to almost spreading hairs (not rather densely covered with short, spreading and with ascending to nearly spreading hairs) (Bagheri et al., 2015). Astragalus zohrabi was given as a synonym of A. lagopodioides in the Flora of Turkey (Chamberlain and Matthews, 1970). ...
Article
Full-text available
A synopsis of the sect. Hymenocoleus Bunge, sect. Hymenostegis Bunge, and sect. Macrophyllium Boiss. belonging to Astragalus L. in Turkey is given based on a revisional study carried out in Turkey. As a result of this study, an account of 19 species including keys and general distribution as well as their conservation status in Turkey is given. It is determined that A. narmanicus Karaman & Aytaç is new to science and A. expetitus Maassoumi, A. pereshkhoranicus Maassoumi & F.Ghahrem., and A. marivanensis Maassoumi & Podlech do not exist in Turkey. Additionally, the name of the sect. Hymenocoleus is preserved.
Book
Full-text available
This is a new version of Check list of Astragalus in the world. Several specific names were added into the new version.
Book
Full-text available
This is a checklist of the genus Astragalus in the World with special references. User can make a copy from this important work and give as following references. Maassoumi, A. A., 2020 : A Checklist of Astragalus in the world: New Grouping, New Changes and Additional species with Augmented data. 560 pages
Article
Full-text available
The taxa of Astragalus section Hymenostegis are an important element of mountainous and steppe habitats in Southwest Asia. A phylogenetic hypothesis of sect. Hymenostegis has been obtained from nuclear ribosomal DNA internal transcribed spacer (ITS) and plastid ycf1 sequences of up to 303 individuals from 106 species, including all 89 taxa currently assigned to sect. Hymenostegis, 14 species of other Astragalus sections, and two species of Oxytropis and one Biserrula designated as outgroups. Bayesian phylogenetic inference and parsimony analyses reveal that three species from two other closely related sections group within sect. Hymenostegis, making the section paraphyletic. DNA sequence diversity is generally very low among Hymenostegis taxa, which is consistent with recent diversification of the section. We estimate that diversification in sect. Hymenostegis occurred in the middle to late Pleistocene, with many species arising only during the last one million years, when environmental conditions in the mountain regions of Southwest and Central Asia cycled repeatedly between dry and more humid conditions.
Article
Full-text available
Pollen and seed morphology were examined in 35 specimens representing 25 Turkish species of the taxonomically complex genus Hesperis L. by light and scanning electron microscopes. Three main types and 2 subtypes were recognized based on the seed coat surface, pollen shape, and exine sculpturing. The seed coat ornamentation of Type I was ocellate, and pollen shape was oblate-spheroidal. While Type II was represented by reticulate seed coat and prolate-spheroidal pollen shape, Type III had tuberculate seed coat. This study revealed that both palynological and seed morphological characters are of significant importance in the taxonomy of the genus.
Article
Full-text available
Pollen morphology of four Matricaria species and 28 Tripleurospermum species was investigated with light microscopies (LM) and scanning electron microscopies (SEM). Pollen slides were prepared using Wodehouse technique. Measurements were based on 20 or more pollen grains per specimen. For SEM studies, dried pollen grains were transferred on aluminum stubs and coated with gold for 4 min in a sputter-coater. The pollen grains of Matricaria and Tripleurospermum are radially symmetric and isopolar. The pollen grains of the Matricaria are oblate-spheroidal with the polar axes 16.6–31.2 μm and the equatorial axes 18.7–23.9 μm. Tripleurospermum is oblate-spheroidal, suboblate and prolate-spheroidal with the polar axes 15.6–32.2 μm and the equatorial axes 17.7-38.5 μm. The pollen grains of Tripleurospermum are operculate and tricolporate. Matricaria is operculate and usually tricolporate or rarely syncolporate, tricolpate and tetracolporate. The pollen grain of both taxa shows echinate ornamentation. The spines are commonly conical with a broadened base and a tapered apical portion. The spine length varies between 1.8–4 μm in Tripleurospermum and 2.3–3.3 μm in Matricaria. The width of spines varies between 2.8–4.6 μm in Tripleurospermum and 2.4–3.6 μm in Matricaria. Inter-spinal area shows granulate–perforate, reticulate–perforate, rugulate–perforate ornamentations and the tectum surrounding the spine base is micro perforate. Overall exine thickness ranges from 2.8 to 4.8 μm in Tripleurospermum, 3.6 to 5.2 μm in Matricaria. Intine is thicker under pores in Tripleurospermum (0.3–0.62 μm) than in Matricaria (0.6–0.8 μm). Inter-spinal ornamentations, pollen shape and the numbers of perforations at the spin base have been observed as important morphological characters.
Article
Full-text available
Even a single peptide that is present in the pollen wall and cytoplasm could cause pollen allergy. To produce skin-prick test kits, the first step is the extraction of these molecules. In this study, Cedrus atlantica pollens were subjected to 220 and 330 MPa for 10 and 30 min in order to extract these molecules. After high hydrostatic pressure processing (HHPP), the total amounts of proteins (TAPs) are measured and compared with the results of the conventional extraction method (CEM). As a result, the TAPs extracted by HHPP is 18.0210 μ g/mL at 220 MPa for 10 min, 22.5770 μ g/mL at 220 MPa for 30 min, 23.3810 μ g/mL at 330 MPa for 10 min and 25.9270 μ g/mL at 330 MPa for 30 min, while this is 1.9460 μ g/mL in 24 h by the CEM. In addition to these results, visual pollen deformation and eruption, pollen wall and surface damage have also been observed.
Article
Full-text available
The program MODELTEST uses log likelihood scores to establish the model of DNA evolution that best fits the data. AVAILABILITY: The MODELTEST package, including the source code and some documentation is available at http://bioag.byu. edu/zoology/crandall_lab/modeltest.html.
Article
Full-text available
Sequences of the internal transcribed spacers (ITS) of the nuclear ribosomal DNA are important molecular markers in phylogenetic analyses. To obtain sequences from herbarium material in which DNA often is severely degraded, the ITS region has to be amplified in two steps. Two methods that reduce bench time and reagents used are described. (i) Separately amplified preparations of subunits ITS-1 and ITS-2 are combined before purification. The presence of two fragments in the sequencing reaction does not impair the quality of sequences. (ii) Newly designed internal primers amplify partly overlapping regions of the two subunits. A combination of these internal primers with the external primers in one PCR allows the amplification of the entire ITS region even when degraded DNAs are used. This recombinant PCR approach, taking into account the +A bases added by several Taq DNA polymerases, will also be useful with other marker regions used in molecular phylogenetics.
Article
Full-text available
MrBayes 3 performs Bayesian phylogenetic analysis combining information from different data partitions or subsets evolving under different stochastic evolutionary models. This allows the user to analyze heterogeneous data sets consisting of different data types—e.g. morphological, nucleotide, and protein—and to explore a wide variety of structured models mixing partition-unique and shared parameters. The program employs MPI to parallelize Metropolis coupling on Macintosh or UNIX clusters. Availability: http://morphbank.ebc.uu.se/mrbayes Contact: fredrik.ronquist@ebc.uu.se * To whom correspondence should be addressed.
Article
N the inorphologic identification of chromosomes, the location of the I centromere is the most useful landmark, and one which is characterized by great constancy. It would seem that not much could be added to the definitions by E. B. WILSON (1928) of the locations on the chromosome of the centrornere or, in the terminology of that time, the spindle attachment: “Attachment of the chromosome to the spindle is commonly limited to a small area, and is of two general types, namely: (1) terminnl or telomitic and (2) non-ferminal or atelomitic, being in the former case at one end, and in the latter at some other point or points. Non-terminal attachment may be at the middle point (median) or at an intermediate point (submedian, sub-terminal). All gradations exist between these various cases;” (I.c., p. 130-131). In the acconipanying picture (l.c., Fig. 56, p. 132), here reprinted as Fig. l., the four locations of median, submedian, subterminal and terminal are represented, and, in addition, “lateral”, which corresponds to the modern term “diffuse centromere”. Nevertheless, at the present time, with the immense increase in research activity in mammalian cytology, the terminology of the centromeric position has become burdened by much obscurity and confusion. One cause of confusion is that different authors, and even the same author on different occasions, have used the terms median, submedian etc. with great amplitude, and it is often difficult to know in a specific case what each term signifies. Another cause of confusion is that a set of terms for chromosomes with specific centromeric positions, such as metacentric, acrocentric, telocentric, have come into wide usage without being clearly defined in relation to the positional terms median, submedian, subterminal and terminal. During the spring of 1963 the present writers exchanged epistolary
Article
Mitochondrial control region sequences from European populations of the blue tit Parus caeruleus were used to reveal the Pleistocene history and the post-glacial recolonization of Europe by the species. The southern subspecies, P. c. ogliastrae was found to represent a stable population with isolation-by-distance structure harboring a lot of genetic variation, and the northern subspecies P. c. caeruleus a recently bottlenecked and expanded population. We suggest that after the last Ice Ages, the subspecies have colonized Europe from two different southern refuges following previously proposed general recolonization routes from the Balkans to northern and Central Europe, and from the Iberian Peninsula north- and eastwards. The two subspecies form a wide secondary contact zone extending from southern Spain to southern France.