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Noccaea birolmutlui, a New Crucifer Species from South West Anatolia, Turkey


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Noccaea birolmutlui Özgişi & Özüdoğru is described as a new species from Kızıldağ National Park, Isparta, in SW Anatolia. It resembles N. ochroleuca but can be readily distinguished by several morphological characters including shape and size of basal/stem leaves, fruit shape, style length, ovule number etc. Nuclear-encoded ribosomal internal transcribed spacer (ITS) region based phylogeny proved that this species clearly differs than N. ochroleuca and these two species belong to different clades in Noccaea s.str. Diagnostic characters, a description, detailed photos and taxonomic comments on the species are given. Morphological affinities and ITS based phylogenetic relationships of new species are discussed. Furthermore, the IUCN Red List category of N. birolmutlui is proposed as critically endangered (CR) due to its limited distribution.
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Phytotaxa 345 (1): 059–067
Copyright © 2018 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Karol Marhold:22 Feb. 2018; published: 20 Mar. 2018
Noccaea birolmutlui, a New Crucifer Species from South West Anatolia, Turkey
Noccaea birolmutlui Özgişi & Özüdoğru is described as a new species from Kızıldağ National Park, Isparta, in SW Anatolia.
It resembles N. ochroleuca but can be readily distinguished by several morphological characters including shape and size of
basal/stem leaves, fruit shape, style length, ovule number etc. Nuclear-encoded ribosomal internal transcribed spacer (ITS)
region based phylogeny proved that this species clearly differs than N. ochroleuca and these two species belong to different
clades in Noccaea s.str. Diagnostic characters, a description, detailed photos and taxonomic comments on the species are
given. Morphological affinities and ITS based phylogenetic relationships of new species are discussed. Furthermore, the
IUCN Red List category of N. birolmutlui is proposed as critically endangered (CR) due to its limited distribution.
Keywords: Brassicaceae, Coluteocarpeae, Cruciferae, Noccaea, Turkey
The genus Noccaea Moench (1802: 89) is one of the most taxonomically problematic genera among crucifers and
its generic circumscription varies according to different concepts (Al-Shehbaz 2014; https://brassibase.cos.uni-; accessed 01 December 2017). Although Noccaea was described for the first time by Moench (1802),
species of within the genus were subsumed under  Linnaeus (1753: 645) (Prantl 1891; Schulz 1936) until
Meyer (1973; 1979) divided  into 12 segregate genera. Meyer’s concept was generally supported by early
molecular studies (Mummenhoff & Zunk 1991, Mummenhoff & Koch 1994, Mummenhoff et al. 1997a,b). However,
subsequent studies (Koch & Mummenhoff 2001, Koch & Al-Shehbaz 2004) showed that this concept was artificial.
Because Meyer’s concept was impractical for taxonomist, his treatment was not followed by some authors and generic
limits of segregates are still controversial (Greuter & Raus 1983, Greuter et al. 1986, Al-Shehbaz, 1986, 2014, Ali et
al. 2016). Based on both morphological and molecular studies, Al-Shehbaz (2014) united all of Meyer’s (1973; 1979)
segregates in a more inclusive Noccaea in the tribe Coluteocarpeae whereas database for Brassicaceae (BrassiBase, presented an alternative view including 14 genera.
In the first volume of  Hedge (1965) recognized 25  species
and two doubtful records. After this account, 26 species were added to  or to others of Meyer’s segregates
including Callothlaspi Meyer (1973: 457), Microthlaspi Meyer (1973: 452), Noccaea, etc. (Davis et al. 1988, Gemici
& Leblebici 1995, Yıldırımlı 2000, Aytaç et al. 2006, Fırat et al. 2014). Based on both morphological and molecular
studies, Al-Shehbaz (2014) subsumed some Eunomia Candolle (1821:555) species and all of Meyer’s segregates under
the genus Noccaea s.l. in the tribe Coluteocarpeae except  s.str. and Noccidium Meyer (1973: 456). Following
this treatment the current number of Noccaea species in Turkey is 50.
During the herbarium studies for “The Revision of Turkish Noccaea Speciesproject an interesting voucher
was found in the Herbarium of Biology Department, Hacettepe University (HUB), which was collected by Dr. Birol
Mutlu at Kızıldağ National Park, Isparta. This Noccaea specimen could not be identified with the Flora of Turkey and
related references (Hedge 1965; Meyer 2006; Al-Shehbaz 2014). Later, specimens in flowering and fruiting stage were
collected from the original location by the authors and were used in the current study.
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60 Phytotaxa 345 (1) © 2018 Magnolia Press
Material and Methods
Flowering and fruiting specimens of Noccaea ochroleuca (Boissier & Heldreich 1849:39) Meyer (1973:461) and
new species were collected in the field. All morphological analyses were undertaken on both living and herbarium
specimens. Herbarium specimens of the new species were compared with its morphologically closely related species N.
ochroleuca based on relevant literature (Hedge 1965, Meyer 2006, Al-Shehbaz 2014). There are some taxonomically
ambiguous populations around Isparta, Konya and Antalya border which may or may not belong to N. ochroleuca.
Therefore, we only used N. ochroleuca samples from Davraz Mountains following Meyer’s treatment (Meyer 2006).
For scanning electron microscopy (SEM) of seeds, dry seeds were mounted directly on stubs and coated with gold in
a sputter coater. SEM examination was carried out using a Jeol 5600 LV-SEM microscope at Eskişehir Osmangazi
University. Photographs of living material were taken with a Nikon D90 digital camera.
Internal transcribed spacer regions (including ITS1 and ITS2) and the 5.8S gene of nuclear ribosomal DNA
(hereafter ITS) were sequenced for the new species, N. ochroleuca and N. papillosa (Boissier 1867:330) Meyer
(1973:460). Additionally, ITS sequences of 32 available Coluteocarpeae members representing all Meyer’s segregate
plus Linnaeus (1753: 646) as an outgroup were taken from Genbank. (
DNA extraction, amplification, and sequencing:—Total genomic DNA was isolated using DNeasy Plant Mini
Kit (Qiagen, Hilden, Germany) following the manufacturer’s instructions. ITS region was amplified using primers
ITS1 and ITS 4 (White et al. 1990). Amplification of ITS followed the protocol in Warwick et al. (2004). Purification
and sequencing were performed by MedSanTek (İstanbul, Turkey).
Phylogenetic analysis:—Sequences were edited with Codon Code Aligner (CodonCode Corporation) and aligned
with MUSCLE v.3.6 (Edgar 2004). Alignment file was provided as supplementary data. The best fitting nucleotide
substutition model was selected by the Akaike information criterion (AIC) implemented in MEGA v.6 (Tamura et
al. 2013). The GTR + G model was selected for Bayesian analysis and a Yule Model process of speciation was used
as the tree prior. Bayesian analyses were implemented in BEAST v.1.8 (Drummond et al. 2012). Two independent
MCMC runs were conducted with chain lengths of 10 million and sampled every 1000 generations. Each run was
checked using Tracer v.1.5 (Rambaut & Drummond, 2009), with the first 10% of samples discarded as burn-in, and
then combined in LogCombiner (Drummond et al. 2012).
The aligned ITS data set included 38 sequences that belong to 34 species and 648 bp positions were available for
phylogenetic analysis long, of which 228 were variable and 123 were potentially parsimony informative.
Noccaea birolmutlui was placed in a clade with some representatives of Noccaea s.str including N. jankae (Kerner
1867:35) Meyer (1973:462), N. montana (Linnaeus 1753: 647) Meyer (1973:462), N. densiflora (Boissier & Kotschy
(Boissier 1867:328) Meyer (1973:462), N. macrantha (Lipsky 1894:271) Meyer (1973:464) and Raparia bulbosa
(Spruner 1843:74) Meyer (1973:458), whereas its morphologically closely relative N. ochroleuca is found in different
clade along with N. papillosa (Fig. 1).
Noccaea birolmutlui
Type:—TURKEY. B3 Isparta: Şarkikaraağaç; Kızıldağ National Park, 1400–1500 m, clearings in Cedrus libanii forest, 09.06.2017,
 5017 &  (holotype: HUB!, isotypes: OUFE!, GAZI!, EGE!, INU!). Paratypes: B3 Isparta: Şarkikaraağaç; Kızıldağ
National Park, ormanevleri Hastane inşaatı arası, Cedrus libani-Quercus karışık ormanı, 1150–1250 m, 26.05.1994, Mutlu 443
(HUB!); Isparta, Şarkikaraağaç, Kızıldağ Milli Parkı, Piknik Alanı Çevresi, Cedrus libani orman açıklığı 38° 02’ 25”K, 31° 21’
54”K, 1400 m, 26.06.2016,  1211 (ANK!, HUB!, OUFE!, NGBB!)
 Phytotaxa 345 (1) © 2018 Magnolia Press 61
 Phylogenetic tree of tribe Coluteocarpae based on the Bayesian analysis of the ITS data. Posterior probability
values > 0.5 are shown at the nodes. Taxon names follow concept presented in BrassiBase (https://brassibase.cos.uni-
Noccaea birolmutlui differs from the related N. ochroleuca by having long petiolate (up to 28 mm) and entire (vs.
short petiolate (up to 14 mm) and dentate) basal leaves, oblong to ovate (vs. lanceolate) cauline leaves, obdeltoid (vs.
obcordate) siliculae, 5-6 ovules (vs. 4) per locule.
Glabrous, glaucescent perennial herbs 15–40 cm tall; stems single or rarely several from the base, terete. Basal leaves
rosette forming, green, spatulate, tapering into a long petiole, 10–72 × 4–14 (16) mm (inc. petiole), apex rounded,
margins entire; cauline leaves subcoriaceous, glaucus, oblong to ovate, ca. 6–17 (19) × 5–13 (15) mm, apex obtuse
to acute, margins entire, amplexicaul or auriculate, sometimes overlapping, auricles obtuse. Inflorescence simple
or compound raceme (up to 6 racemes), racemes 25–107-flowered, congested in flower, elongating in fruit, lateral
racemes shorter than terminal in fruiting time. Pedicel 1.5–4.5 mm in flower; horizontally spreading up to 15 mm in
fruit. Sepals 2.1–2.2 × 0.9–1 mm. Petals 5.2–5.5 × 1.5–2 mm, white. Siliculae glaucus, pruinose, obdeltoid, 2.5–8.5 ×
1.8 (2.1) –6 (6.2) mm with a apical notch. Tips of notch retuse (Notch sometimes absent). Valves of siliculae winged up
to 1.8 mm at apex, narrower at base. Style 0.8–2.1 mm long in fruit, always exceeding sinus. Stigma capitate. Ovules
(5)6 per locule. Seeds orange to orange-brown, tuberculate, 2(–5) per locule, ca. 1.3 (1.5) × 0.7 mm.
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 Phytotaxa 345 (1) © 2018 Magnolia Press
 General view of Noccaea birolmutlui and N. ochroleuca. N. birolmutlui  N. ochroleuca
 Phytotaxa 345 (1) © 2018 Magnolia Press 
 Noccaea birolmutlui: Basal leaves;  Cauline leaves;  Fruits. N. ochroleuca:  Basal leaves;  Cauline
leaves; Fruits.
Habitat: Cedar forest.
 : May–June.
 The specific epithet is named after Dr. Birol Mutlu, who is the first collector of new species. The
Turkish name of this species is given as “Kızıldağ Dağarcığı” according to the guidelines of Menemen et al. (2013).
Seed morphology: Seeds of N. birolmutlui are orange to orange-brown, tuberculate, ca. 1.3 (1.5) × 0.7 mm and
2(–5) per locule (Fig. 4). Seed coat is colliculate, lacking the mucilage and has epicuticular wax only. Epidermal
cells are irregularly pentagonal to elliptic. Cell walls are evenly thickened, the boundaries are below the cell centers.
Anticlinal walls are flat, periclinal walls are obviously convex. Surface ornamentation is ruminate.
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 Phytotaxa 345 (1) © 2018 Magnolia Press
 Seed of Noccaea birolmutlui  General appearance,  Seed surface
  N.biromutlui is a local endemic restricted to the part of Cedar forest in Kızıldağ
National Park (Isparta, Turkey) (Fig. 4). The occupancy area (AOO) of new species was calculated as ca. 1 km2 in
which about <1000 individuals were estimated to occur. Although N. birolmutlui occurs in a protected area (National
Park), it has very small distribution area with low population size and therefore, in accordance with the criteria of the
IUCN (2017), it is here assessed as “Critically Endangered”
Examined specimens:—N. ochroleuca: —TURKEY. C3 Isparta: Davraz mountain, 1800–1900 m, 09.06.2017,
 5009 & (HUB!), Mons Davros dagh Pisidiae, ad nives, 05.1845, Heldreich (isolectotype B photo!
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 Distribution of Noccaea birolmutlui ( ) and N. ochroleuca ( ).
Noccaea birolmutlui is morphologically similar to N. ochroleuca but it is distinguished by several morphological
characters including basal and cauline leaves, fruit shape and ovule numbers.
These characters are typically used in species delimitation of Noccaea (Meyer, 2006; Al-Shehbaz, 2014 etc.).
Of these, fruit shape can be extremely variable for Noccaea species in the different stages of plant development
as indicated by Özgişi et al. (unpublished data). But as our collections represent different ontogenetic stages of N.
birolmutlui, fruit shape can be used as a diagnostic character.
Bayesian based molecular phylogenetic analysis of ITS data clearly show that N. birolmutlui and N. ochroleuca
are distinct species placed in different clades (Fig. 1). Raparia bulbosa is the phylogenetically closest species to N.
birolmutlui, but morphologically it is distinguished from latter by having napiform root and large rose-lilac petals.
However, to reveal the true phylogenetic position of N. birolmutlui, more representatives of Noccaea (128 species
according to Al-Shehbaz 2014) and additional molecular markers should be included in a phylogenetic analysis.
In his  account for Flora of Turkey, Hedge (1965) used broad species delimitation for N. ochroleuca and
apart from type specimen from Davraz Mountain (Isparta, Turkey), he included some additional accessions which are
treated as N. phyrgia (Bornmueller 1929: 57) Meyer (1973: 465), N. edinensium Meyer (1973: 460) and 
(Stojanov & Kitanov 1943: 97) Meyer (2006: 116) at the present time. Of them, N. phyrgia has a spotted distribution
in Anatolia and reported from Ankara, Bolu, Bursa and Konya provinces, whereas N. edinensium and  are
restricted to Murat Mountain (Kütahya, north-east Aegen Part of Turkey) and Uludağ (Çanakkale, north-west Turkey).
This would indicate that these species are allopatric. Finally, taxonomically ambiguous populations that outlined in
Material and Methods chapter, needs further evaluations using comprehensive morphometric and also genetic analyses.
Thus, species boundary of N.ochroleuca can be outlined correctly.
 Morphological comparison of Noccaea birolmutlui and its morphologically closest relative N. ochroleuca.
Characters Noccaea birolmutlui Noccaea ochroleuca
Basal leaves
Petiole (mm) 4.2–28 × 0.5–2.3 4–14 × 0.5–3.1
Size (mm) 10–72 × 4–14 (16) 8–52 × 2–9
Margin Entire Somewhat dentate
Cauline leaves
Shape Oblong to ovate Lanceolate
Size (mm) 6–17 (19) × 5–13 (15) 3–7 × 6–31
Apex Obtuse to acute Acute
Inflorescence A simple raceme or a compound raceme (up to 6
A simple raceme or compound raceme (up to 4
Racemes 25–107 flowered 17–47 (50) flowered
Shape Obdeltoid Obcordate
Size (mm) 2.5– 8.5 × 1.8 (2.1)–6 (6,2) 5–10 (12) × 2–7
Margins Winged up to 1.8 mm at apex, green Winged up to 1.2 mm at apex, often purplish
Style 0.8–2.1 mm Up to 4.7 (5.0) mm
Ovules per locule (5) 6 4
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66 Phytotaxa 345 (1) © 2018 Magnolia Press
This study was supported by Eskişehir Osmangazi University BAP Coordinatorship (project number: 2015/681) and
Hacettepe University Research Project Coordination Unit (project number: FBA-2017-12879). We thank Yelda Güzel
for helping us to describe seed surface of new species and Dmitry German for providing a reference.
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... The tribal classification of Al-Shehbaz (2012) was accepted by some researchers (Ali et al., 2016a;German, 2017;Özgişi et al., 2018a;Özgişi et al., 2018b;, accessed 27 August 2019). ...
... However, the generic classification of Coluteocarpeae remains controversial. While some researchers (Özgişi et al., 2018a;Özgişi et al., 2018b;Güzel et al., 2018;Özüdoğru, 2018) have accepted the concept of Al-Shehbaz (2014), others (Ali et al., 2016a;2016b;Karaismailoğlu and Erol, 2018;; https://brassibase., ...
... In addition to the debates (Greuter and Raus, 1983;Greuter et al., 1986;Artelari, 2002;Aytaç et al., 2006;Ali et al., 2016a;German, 2017;Özgişi et al., 2018a;Özgişi et al., 2018b;Karaismailoğlu and Erol, 2018) regarding the generic concept by Meyer (1973;1979), variations among the morphological features that were used for species delimitation resulted in misevaluation of the species (Özgişi, 2018a). Because of these variations, the morphology-based keys to identify specimens, which was proposed by Meyer (1973;1979; and Al-Shehbaz (2014) have been inadequate in some cases (i.e. for distinguishing N. densiflora (Boiss. ...
... Konvergensinin yanı sıra teşhiste kullanılan morfolojik karakterlerdeki varyasyon da yanlış teşhislere neden olmaktadır (Özgişi vd., 2018a;2018b;Güzel vd., 2018;Özgişi, 2020a). Bu durum cins sistematiği zaten tartışmalı olan bu grup içinde tür sayısında da bir tutarsızlığın oluşmasına neden olmaktadır. ...
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Yaklaşık 75 taksonla temsil edilen Thlaspi L. cinsinin sistematiği, 351 Brassicaceae cinsi arasında en tartışmalı olanlardan biridir. Schulz'dan etkilenen Meyer, Thlaspi cinsini 11 farklı cinse ayırmıştır. Bu cinslerden biri olan Thlaspicerasas F. K. Meyer, daha sonra Noccaea cinsi altında bir tür kompleksi olarak kabul edilmiştir. Bu tür kompleksi dokuzu Türkeye’ye endemik olan toplam 11 türden oluşmaktadır. Sınıflandırmada sıklıkla kullanılan meyve karakterindeki konvergensin ve varyasyonun varlığı toplam tür sayısı hakkında tartışmalara neden olmaktadır. Bu çalışmada Bayesian İstatistiksel Metoduna dayalı BPP tür sınırlandırma programı tür sınırlandırma çalışmaları için kullanılmıştır. Çekirdek DNA’sına ait Internal Transcribed Spacers (ITS) ile plastid DNA’sına ait trnL-F ve trnQ-5'rps16 moleküler belirteçleri kullanılarak tür sınırlandırma çalışmaları üç farklı senaryo (Küçük, orta ve büyük atasal popülasyon büyüklüğü) altında incelenmiştir. Ayrıca biyoklimatik veriler (19 biyoklimatik verinin yanı sıra aylık en düşük, ortalama, en yüksek sıcaklık değerleri ve aylık ortalama yağış miktarları) kullanılarak türlerin biyoklimatik tercihleri tespit edilmiştir. Tür sınırlandırma çalışmalarında küçük atasal popülasyon büyüklüğü modeli altında gerçekleştirilen çalışmalara göre 0.63 ardıl olasılık değeri ile tür kompleksi altı türden oluşmaktadır. Orta atasal popülasyon büyüklüğü modeline göre ise 0.78 ardıl olasılık değeri ile üç farklı türün varlığı tespit edilmiştir. Büyük atasal popülasyon büyüklüğü modeli ise 1.00 ardıl olasılık değeri ile kompleksin sadece tek bir türden oluştuğunu göstermiştir. Türlerin biyoklimatik isteklerini belirlemeye yönelik çalışmada ise Noccaea huber-morathii (F.K.Meyer) Al-Shehbaz türünün daha soğuk ve nemli alanları tercih ederken Noccaea meyerii (F.K.Meyer) Al-Shehbaz türünün daha kuru ve sıcak alanları tercih ettiği tespit edilmiştir.
... Coluteocarpeae V.I. Dorof. is a tribe of Brassicaceae (Cruciferae) which is being in focus of intensive ongoing taxonomic and phylogenetic studies resulted in raise of novel (and contrasting) generic concepts, discovery of new species, clarification of various nomenclatural items and distribution patterns, etc. (Dorofeyev 2013, Al-Shehbaz 2014, Fırat et al. 2014, Ali et al. 2016a, b, German 2017, Güzel et al. 2018, Özgişi et al. 2018a, b, Özüdoğru 2018a to name some recent publications). In course of the study of Noccaea Moench (1802: 89) s. l., the biggest genus of the tribe, in Turkey, by one of us (B. ...
The status of three names related to the genus Pseudosempervivum is reconsidered. As a result, Noccaea crassiuscula and Pseudosempervivum amanum are reduced to synonymy of P. sempervivum while P. gurulkanii is found conspecific with N. venusta. Problems in delimitation of P. sempervivum and P. aucheri are also briefly considered. The name Cochlearia sempervivum is lectotypified.
... The first revision of Thlaspi in Turkey was made by Hedge (1965), who recognized 25 species and 2 doubtful records for Turkish flora. Since then, 26 new Thlaspi taxa (or members of Meyer's other segregates, depending on treatments and authors), including subspecies, were added to the total number (Davis et al., 1988;Gemici and Leblebici, 1995;Yıldırımlı, 2000a;Aytaç et al., 2006;Özgişi et al., 2018a), some of which were synonymized by Al-Shehbaz (2014) in his comprehensive synopsis. ...
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Noccaea ali-atahanii, a new Brassicaceae species from southern Anatolia, is described herein. It is mostly similar to N. aghrica, but can be readily distinguished by the presence of apical placentation and orbiculate fruit with wings extending almost evenly from the base to the apex. Its phylogenetic relationships within the genus are provided based on the nuclear-encoded ribosomal internal transcribed spacer region. Carpological and ecological features of N. ali-atahanii and related species are also discussed.
Thlaspi harungalipii sp. nova (Brassicaceae) is described and illustrated from Southern of Turkey. This taxon is similar to T. violascens, from which it clearly differs in flower and fruit characters. In this study, the seed micromorphological and anatomical characters, and phylogenetic relations within the genus based on the nuclear-encoded ribosomal internal transcribed spacer region of the new species are determined. Furthermore, the IUCN category of the new species is discussed, and its distribution is mapped.
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Circumscribing species boundries is necessary in systematic plant biology. Even a mistake in delimiting taxa may lead to incorrect scientific interpretations. Draba rimarum (Rech.f.) A.R. Khosravi & A. Eslami-Farouji is an endemic Iranian species with a narrow geographic distribution, and is genetically close to D. aucheri. The present study provided a phylogenetic review, time divergence, and planar network of both species to unravel the distinct position of both species along with the prediction of any conflicting or ambiguous signals. Regarding this purpose, here we represent that phylogenetic trees may fail to show reliable results toward the distinct position of genetically close species.
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This study was conducted for extensive and systemic investigation of anatomical, palynological and seed morphological properties of two endemic Noccaea species, naturally growing in Turkey. Independent sample T-test and box plot were carried out using quantitative characters of the studied species. The anatomical results showed that the species had similar characteristics, though there were significant differences in root cortex cells and trachea; stem epidermis, cortex cell and endodermis; leaf upper and lower epidermis, lower cuticle, mesophyll and palisade parenchyma. Pollen grains of two endemic species were observed as radially symmetric, isopolar, with tricolpate aperture, prolate pollen shape and had small size. Pollen surface ornamentation was micro-reticulate in both species. Considering palynological characters, equatorial axis, AMB exine and intine have taxonomic importance. Seeds of N. birolmutlui were ovate to orbicular in shape and orange-brown in color, with colliculate ornamentation; however, the seeds of N. camlikensis were ovate to oblong in shape and brown and shiny in color, with colliculate-reticulate ornamentation.
The accurate identification of an organism is the basis of all biological disciplines. Although there have been many different species concepts and methods proposed, researchers generally choose the most appropriate concept according to their interests. However, each species concept has both advantages and disadvantages. In such cases, an integrated concept based on evidence obtained from different species concepts and methods is suitable for the accurate delimitation of a species. The biggest dilemma of integrated species delimitation methods is the inconsistency between species concepts and methods. Herein, the congruency of the different concepts and methods was tested to reveal the relationship of the Noccaea densiflora species complex. nrITS, chloroplast trnL-F, and trnQ-5'rps16 regions were used for species delimitation using the multispecies coalescent model (MSC) as implemented in two developed Bayesian Methods. ITS and trnL-F regions were also used for TCS haplotype network analyses. In addition to morphological measurements from different populations and vouchers, specific bioclimatic data values were also used for the principal component analysis. Based on all of the analyses, it was determined that only one population of N. densiflora and and one of N. microstyla were clearly distinct, whereas the rest of the specimens remain taxonomically uncertain.
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The genus Thlaspi has been variously subdivided since its description by Linnaeus in 1753, but due to similarities in fruit shape several segregates have still not gained broad recognition, despite the fact that they are not directly related to Thlaspi. This applies especially to segregates now considered to belong to the tribe Coluteocarpeae, which includes several well-studied taxa, e.g., Noccaea caerulescens (syn. Thlaspi caerulescens), and the widespread Microthlaspi perfoliatum (syn. Thlaspi perfoliatum). The taxonomy of this tribe is still debated, as a series of detailed monographs on Coluteocarpeae was not published in English and a lack of phylogenetic resolution within this tribe was found in previous studies. The current study presents detailed phylogenetic investigations and a critical review of morphological features, with focus on taxa previously placed in Microthlaspi. Based on one nuclear (ITS) and two chloroplast (matK, trnL-F) loci, four strongly supported major groups were recovered among the Coluteocarpeae genera included, corresponding to Ihsanalshehbazia gen. nov., Friedrichkarlmeyeria gen. nov., Microthlaspi s.str., and Noccaea s.l. In addition, two new species of Microthlaspi, M. sylvarum-cedri sp. nov. and M. mediterraneo-orientale sp. nov., were discovered, which are well supported by both morphological and molecular data. Furthermore, M. erraticum comb. nov. (diploid) and M. perfoliatum s.str. (polyploid) were shown to be distinct species, phylogenetically widely separate, but with some overlap in several morphological characters. Detailed descriptions, notes on taxonomy, geographical distribution, and line drawings for the new species and each species previously included in Microthlaspi are provided. In addition, the current taxonomic state of the tribe Coluteocarpeae is briefly discussed and it is concluded that while several annual taxa are clearly distinct from Noccaea, many perennial taxa, after thorough phylogenetic and morphological investigations, may have to be merged with this genus.
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Using sequence data from the ITS region (internal transcribed spacers ITS1 and ITS2 of nuclear ribosomal DNA and the 5.8S rRNA gene) and chloroplast DNA sequence data from the trnL intron. we examined the evolutionary relationships of Old and New World Smelowskia species with its putatively related genera (Brassicaceae). ITS and trnL sequence data were obtained from all 13 Smelowskia species and from 13 taxa of the related genera Descurainia, Ermania, Gorodkovia, Hedinia, Polyctenium, Redowskia. Sinosophiopsis, and Sophiopsis. Results from maximum parsimony analyses showed a polyphyletic origin for Smelowskia, with the taxa split into two major clades. Smelowskia taxa plus Ermania, Gorodkovia, Hedinia, Redowskia, Sinosophiopsis, and Sophiopsis formed one clade (Smelowskia s.l. clade), whereas the other clade included Smelowskia holmgrenii (Holmgrenii subclade) and the genus Polyctenium (Polyctenium subclade). Descurainia formed a separate clade. ITS and trnL molecular data, along with a comparison of key morphological traits for each genus, support recognition of a single genus that would include Smelowskia, the three monotypic genera Ermania. Gorodkovia, and Redowskia, as well as the genera Hedinia, Sinosphiopsis, and Sophiopsis (although not all species were examined in these genera). The molecular data are consistent with an Asian origin for Smelowskia s. str., followed by diversification in Asia and North America, subsequent to migration into arctic-alpine regions of western North America. ITS sequence divergence estimates ranged from 0 to 4.1% within Smelowskia s. Str., and up to 6.45% within the Smelowskia s.l. clade; whereas trnL sequence divergence estimates ranged from 0 to 1.85% within Smelowskia s. str., and up to 3.71% within the Smelowskia s.l. clade, indicating pleistocenic speciation within Smelowskia as a whole.
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A general review of the taxonomic status of Thlaspi past and present is given, and a critical evaluations of its segregates based on both morphological and molecular data are presented. ITS molecular phylogenetic study of Thlaspi aghricum and related species, as well as seed-coat morphology and anatomy strongly support the placement of the species in Noccaea. The new combination N. aghrica is proposed, and detailed description and distribution of the species are given.
The controversial limits of Thlaspi are discussed, and all of its segregates proposed or revised during the past four decades are critically evaluated. As a result, the majority of specific and infraspecific names previously assigned to Thlaspi are currently placed in Noccaea, a genus the limits of which are expanded to include almost all of Thlaspi segregates. A detailed description of Noccaea, a synoptic account of its species, along with full bibliography and typification, and key to all 128 species are presented. The new names N. germanii, N. iranica, and N. meyeri and the following 36 new combinations are proposed: N. abchasica, N. antitaurica, N. atlantica, N. bornmuelleri; N. bovis, N. bulbosa, N. campylophylla, N. cappadocica, N. capricornuta, N. cilicica, N. cornuticarpa, N. crassiuscula, N. dolichocarpa, N. eigii, N. elegans, N. huber-morathii, N. jaubertii, N. kurdica, N. lilacina, N. maassoumii, N. natolica, N. orbiculata, N. oxyceras, N. perfoliata, N. platycarpa, N. pulvinata, N. rechingeri, N. rostrata, N. rosularis, N. stenocarpa, N. szowitsiana, N. triangularis, N. umbellata, N. vesicaria, N. vesicaria subsp. boissieri, and N. zangezurica. The genus Coluteocarpus is reduced to synonymy of Noccaea. Brossardia papyracea var. kotschyi, Coluteocarpus reticulatus var. boissieri, Hutschinsia trinervia, Iberidella andersonii, Thlaspi affine, T. japonicum, T. kovatsii, T. szowitsianum, and Vesicaria reticulata are lectotypified.
Mummenhoff, K. & Zunk, K.: Should Thlaspi (Brassicaceae) be split? Preliminary evidence from isoelectric focusing analysis of Rubisco. ‐ Taxon 40: 427–434. 1991. ‐ ISSN 0040‐0262. Relationships among taxa of Thlaspi sensu lato were tested by isoelectric focusing (IEF) analysis of Ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco). 14 species were studied, belonging to 3 of the genera previously segregated by Meyer on the basis of seed anatomy. Differences in Rubisco IEF patterns resulted in a grouping of the species into three clusters which are consistent with the respective genera of Meyer.
Continuing a series of miscellaneous contributions, by various authors, where hitherto unpublished data relevant to the Med-Checklist project are presented, this instalment deals with the families Bignoniaceae, Boraginaceae, Callitrichaceae, Capparaceae, Chenopodiaceae, Compositae, Euphorbiaceae, Fagaceae, Leguminosae, Malvaceae, Nyctaginaceae, Orobanchaceae, Plantaginaceae, Ranunculaceae, Resedaceae, Rosaceae, Scrophulariaceae; Commelinaceae, Cyperaceae and Gramineae. It includes new country and area records, taxonomic and distributional considerations. New taxa are described in Capparis, Eragrostis, Plantago and Ventenata; new combinations and names at new rank are proposed in Aegonychon, Alcea, Amelanchier, Anemonastrum, Centaurea, Cerinthe, Chamaecytisus, Cytisus, Digitalis, Iranecio, Malva, Mattiastrum, Paracynoglossum, Pilosella, Sorbus and Trigonella.