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Androsace azizsancarii sp. nov. (Primulaceae): a new species from northeastern Anatolia, Turkey


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Androsace azizsancarii (Primulaceae) is described and illustrated as a new species from Bayburt Province in northeastern Anatolia, Turkey. Diagnostic morphological characteristics, a full description and a distribution map are provided. The new species is morphologically closest to Androsace albana, A. multiscapa and A. villosa, but it is easily distinguished from these species by indumentum, leaf, flower and seed characters.
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Nordic Journal of Botany
© 2021 Nordic Society Oikos. Published by John Wiley & Sons Ltd
Subject Editor: Panayiotis Trigas
Editor-in-Chief: Torbjörn Tyler
Accepted 25 May 2021
2021: e03208
doi: 10.1111/njb.03208
2021 e03208
Published 24 July 2021
Androsace azizsancarii (Primulaceae) is described and illustrated as a new species
from Bayburt Province in northeastern Anatolia, Turkey. Diagnostic morphological
characteristics, a full description and a distribution map are provided. e new species
is morphologically closest to Androsace albana, A. multiscapa and A. villosa, but it is
easily distinguished from these species by indumentum, leaf, flower and seed characters.
Keywords: Bayburt, Flora of Turkey, new species, taxonomy
Primulaceae is one of the 22 families in the order Ericales (APG IV 2016, Christenhusz
and Byng 2016). e family Primulaceae consists of about 53 genera with approximately
2790 species in the broad sense, including the former families Myrsinaceae and
eophrastaceae, which are mainly distributed in the north temperate zone (APG IV
2016, Christenhusz and Byng 2016).
e genus Androsace L. (Primulaceae) comprises about 156 species (Mabberley 2008,
Jacquemoud and Jordan 2020) that are mainly distributed in the extratropical mountain
ranges of the northern hemisphere. ey have experienced some periods of rapid
diversification (Anderberg and Kelso 1996, Roquet et al. 2013, Schönswetter et al. 2015)
as most Androsace species tend to occur in generally cold environments (Boucher et al.
2012, Roquet et al. 2013). Ancestors of the genus (short-lived) probably had to adapt
to the conditions on the cold steppes, and although these ancestral species spread across
the mountains, their insufficient dispersal ability prevented them from migrating back
to their original habitats, and instead forced them to adapt to new habitats to survive
(Boucher et al. 2012, Roquet et al. 2013). e cold climatic conditions influenced the
enhanced diversification of Androsace species, especially in Alpine areas (Boucher et al.
2012, Roquet et al. 2013). ere has been ongoing discovery of new Androsace taxa in the
Eurasian mountains, which shows that despite some taxonomic novelties, we have little
knowledge about plant diversity on the mountain summits (Dentant 2018). e genus
Androsace has been divided into six quite distinct sections, comprising Pseudoprimula
Pax., Chamaejasme Koch, Aretia (L.) Duby, Andraspis (Duby) Koch, Douglasia (Gray)
Wendelbo and Aizoidium Hand.-Mazz. Androsace sect. Andraspis comprises approximately
20 species, mostly annuals and biennials, or short-lived perennials, distributed over the
Androsace azizsancarii sp. nov. (Primulaceae): a new species
from northeastern Anatolia, Turkey
A. Sefali ( (, Dept of Primary Education, Faculty of Education, Bayburt Univ.,
Bayburt, Turkey.
whole Holarctic, from the Arctic regions of Eurasia and North
America, extending towards the warm and arid regions of the
Mediterranean, Asia Minor, Iran and Afghanistan (Pax and
Knuth 1905, Smith and Lowe 1997, Stevanovıć et al. 2005).
e diversity center of A. sect. Andraspis is the Caucasus area,
mainly Armenia (Shishkin and Bobrov 1952, Grossheim
1967, Stevanovıć et al. 2005). According to Smith and Lowe
(1997), A. sect. Andraspis includes two distinct species groups:
the A. septentrionalis and A. albana (A. armeniaca) groups. e
A. albana group is distributed over the Caucasus and Turkey
and includes five species (Smith and Lowe 1997). Overall,
eight Androsace species (nine taxa) are distributed in Turkey,
including four species of the A. albana group, i.e. Androsace
albana Steven, A. armeniaca Duby, A. intermedia Ledeb. and
A. multiscapa Duby (Lamond 1978).
During field studies in July 2020, an unusual Androsace
population was discovered on the southern part of Anzer
Mountain, which is a part of the Soğanlı Mountain chain,
in the Bayburt Province of Turkey. is species had mostly
simple hairs and a few small glandular hairs, but no forked
or branched hairs. It resembled A. albana and other members
of the A. albana group, especially A. multiscapa, as well as A.
villosa L. e latter, is a caespitose perennial species, forming
dense cushions or lax mats, and it is distributed in Europe and
the Mediterranean region. A detailed morphological study of
the specimens collected from Anzer Mountain revealed that
they are distinct from all related species, and they belong to
a new, undescribed species. is species is described here as
Androsace azizsancarii.
Material and methods
Androsace azizsancarii specimens were collected from
Bayburt Province (near the Rize border) in the northeastern
Anatolia region, in 2020 (Fig. 1). e holotype specimen
was deposited in the herbarium of İstanbul University
(ISTE), and isotypes were deposited at Ege University
(EGE), Ankara University (ANK) and the Biology
Dept at Bingöl University. All morphological measure-
ments were made on dried specimens using a millimetric
ruler. A stereo-binocular microscope was used to exam-
ine the gross morphology of the new species. Plant spe-
cies nomenclature follows e Plant List (2013) website.
e Androsace specimens were compared with the relevant
taxonomic literature (Pax and Knuth 1905, Grossheim
1967, Lamond 1978, Smith and Lowe 1997, Mabberley
2008, Schönswetter and Schneeweiss 2009, Boucher et al.
2012, Xu et al. 2016, Dentant et al. 2018, Jacquemoud
and Jordan 2020) and material in the herbaria of ISTE,
Selçuk University (KNYA), EGE, Biology Dept of Bingöl
University, Nigde Ömer Halisdemir University, Çukurova
University (CUEF), ANK and Bolu Abant İzzet Baysal
University (AIBU), and some digital herbarium materials
were examined from the the E, G and K herbaria. Detailed
morphological measurements were performed on the
new species and it was compared with the closely related
species Androsace albana and A. multiscapa, using a stereo-
binocular microscope.
A total of 20 specimens of the new species were exam-
ined. At least 50 pollen grains and 50 mature seeds were
measured using a light microscope and a scanning elec-
tron microscope (SEM). e pollen terminology was
adopted from Punt et al. (2007), and classifications using
the P/E ratio and shape followed Erdtman (1969). During
the field studies, photographs of the living material of the
new species and its related taxa were taken using a digital
camera. e general terminology used by Baytop (1998)
was adopted.
Figure 1. Distribution map for A. azizsancarii ( ), A. albana ( ), A. multiscapa ( ).
Figure 2. Androsace azizsancarii sp. nov. (A–D) habitus in flowering time, (E–F) indumentum of late period, (F) habitus in early fruiting
time, (G–H) nonflowering rosettes, (I) basal leaves, (J) leaf indumentum, (K) habitat.
Results and discussion
Androsace azizsancarii Sefalı sp. nov. (Fig. 2–4)
A species related to A. albana, A. multiscapa and A. villosa.
It differs from A. albana by its shorter scapes up to 4.5 cm
long (versus up to 28 cm long), the silky, simple hairs (ver-
sus branched and simple hairs), pink corolla (versus white)
and larger seeds 3.0–3.1 × 2.0 mm (versus 1.4 × 0.8–1.0
mm). It differs from A. multiscapa by its simple hairs (versus
dendroid), leaf margin with 1(–2) pairs of teeth (versus 0–1)
and the pink corolla (versus white). It also differs from A.
villosa by its larger leaves (5–)6–18(–20) × 3–4 mm (versus
2–7(–9) × 1–2 mm), the pink, campanulate corolla (versus
white, rotate) and the dark brown to black seeds 3.0–3.1 ×
2.0 mm (versus pale brown to yellow, 1.7–2.2 × ca 1.2 mm).
Type: Turkey, Bayburt: Soganlı Mountains, south of Anzer
Mountain, on moraines, 2831 m a.s.l., 40°30N, 40°30E,
1 July 2020, A.Sefalı 507 (holotype: ISTE 117267; isotypes:
ANK 60610, BIN 9405, EGE 43195).
Androsace azizsancarii was named in honor of Prof. Dr.
Aziz Sancar, who is the Nobel Laureate in Chemistry 2015.
e Turkish name of this species was chosen as ‘Sancarınca
(Menemen et al. 2016).
Perennial plant with simple hairs, 1.0–6.5 cm tall, usually
forming a single rosette 1.4–4.0 cm in diameter. Leaves in
hemispherical basal rosettes, somewhat fleshy, narrowly
oblong to linear-lanceolate or linear, incurved, (5–)6–18
(–20) × 3–4 mm; adaxial surface often hairy, especially towards
the apex, with long soft hairs 0.2–1.3 mm long towards the
margins; margin with 1(–2) pairs of teeth towards the apex.
Scapes (5–)10–30, 1–3 cm long during anthesis, elongated
to 1.3–4.5 cm long when fruiting, covered with long, soft,
simple, hairs; median scape usually present and stout. Bracts
linear to narrowly lanceolate, generally equaling pedicel dur-
ing anthesis and fruiting, 2.0–7.0 × 1.4 mm, densely vil-
lous. Inflorescence with (1–)2–5(–8) flowers. Pedicel 1–7
mm long at anthesis, elongated up to 12 mm when fruiting.
Figure 3. SEM photographs: Androsace azizsancarii sp. nov. (A–B) pollen grain and pollen surface, (D–E) seed and seed surface, (G–H)
indumentum, A. albana: (C) pollen grain, (F) seed surface, (I) indumentum.
Calyx 6.0–6.8(–8.2) mm, with long silky hairs and scattered
short glandular hairs; calyx teeth ovate to triangular up to
3.2 mm long. Corolla campanulate, 8–10 mm in diam-
eter, pink with yellow centre; tube ± equaling calyx; lobes
oblong to obovoid; throat constricted and annulate. Capsule
5–6 × 3–4 mm, ovoid. Seeds dark brown to black, ovoid,
3.0–3.1 × 2.0 mm.
Flowering from June to July; fruiting in August.
Figure 4. Morphological related species to Androsace azizsancarii: (A–D) A. albana, (D) branched hairs, (E–F) A. multiscapa (photos: Ahmet
Savran), (F) dendroid hairs, (G) A. villosa.
Pollen morphology and seed surface
Pollen grains of A. azizsancarii are tricolporate and isopolar.
e polar axis (P) is 14.07 ± 1.19 µm; equatorial axis (E) is
11.42 ± 1.63 µm; P/E ratio is 1.35 µm and pollen shape is
prolate. e exine is 0.44 ± 0.32 µm, and the intine is 0.24 ±
0.16 µm. e exine ornamentation is microreticulate.
Pollen grains of A. albana are tricolporate and isopolar.
e polar axis (P) is 12.30 ± 1.53 µm; equatorial axis (E)
is 9.04 ± 1.47 µm; P/E ratio is 1.36 µm and pollen shape is
prolate. e exine is 0.42 ± 0.30 µm, and the intine is 0.25 ±
0.27 µm. e exine ornamentation of pollen grains is micro-
reticulate (Table 1, Fig. 2).
Distribution and ecology
Androsace azizsancarii is a local endemic species restricted to
the southern slope of Anzer Mountain in Bayburt Province,
northeastern Anatolia. It is an element belonging to the
Hyrcano-Euxine (mt.) phytogeographical region. e new
species colonizes moraines, between 2820 and 2835 m a.s.l.
Species growing in the near vicinity include Allium balan-
sae Boiss., Androsace albana, Bupleurum falcatum L. subsp.
persicum (Boiss.) Koso-Pol., Campanula aucheri A.DC.,
Helichrysum pallasii (Spreng.) Ledeb., Psephellus appendici-
gerus (K.Koch) Wagenitz, P. pulcherrimus (Willd.) Wagenitz
and Oxytropis lazica Boiss.
Taxonomic relationships
e sections of Androsace have been distinguished by life
form, leaf shape, perianth and fruit characteristics. Androsace
azizsancarii belongs to A. sect. Andraspis and to the informal
A. albana-group. It includes characterized by annuals, bien-
nials or somewhat short-lived perennials (especially the mem-
bers of A. albana-group) (Smith and Lowe 1997). It has been
noticed that indumentum features are especially important in
the taxonomy within A. sect. Andraspis (Shishkin and Bobrov
1952, Grossheim 1967, Lamond 1978, Smith and Lowe
1997). Androsace azizsancarii stands out in terms of its long
silky indumentum. is feature makes A. azizsancarii clearly
different from all other members of A. sect. Andraspis and A.
albana-group. When looking at the members of A. albana-
group, this new species is closely similar to A. albana in
terms of its indumentum. However, A. albana has only long
simple hairs in the calyx region, not throughout the whole
plant. Moreover, A. albana has branched hairs, mostly in the
scape region.
Androsace bidentata K. Koch, is an imperfectly known spe-
cies from Turkey. is species is similar to A. albana and has
branched hairs, however, it was observed that it has larger
flowers and fewer-flowered heads. Another closely related
species is A. multiscapa, which has a completely different hair
type, known as dendroid hairs (Fig. 4, Table 2). Androsace
Table 1. Comparison of the pollen of Androsace azizsancarii with
A. albana.
Characters A. azizsancarii A. albana
Polar axis (P) (µm) 14.07 ± 1.19 12.30 ± 1.53
Equatorial axis (E) (µm) 11.42 ± 1.63 9.04 ± 1.47
P/E and shape 1.35–prolate 1.36–prolate
Ornamentation Microreticulate Microreticulate
Aperture type Tricolporate Tricolporate
Colpus length (Clg) (µm) 11.44 ± 0.95 9.45 ± 0.94
Colpus width (Clt) (µm) 2.13 ± 0.19 1.95 ± 0.24
Por (µm) 3.07 ± 0.36 2.64 ± 0.42
Exine thickness (µm) 0.44 ± 0.32 0.42 ± 0.30
Intine thickness (µm) 0.24 ± 0.16 0.25 ± 0.27
Table 2. Morphological comparison of A. azizsancarii with A. albana, A. multiscapa and A. villosa.
Characters A. azizsancarii A. albana A. multiscapa A. villosa
Plant height 1.0–6.5 cm 3–22(–28) cm 1.5–5.0 cm 1–2 cm
Indumentum long simple (silky) hairs and
scattered short glandular hairs
long simple, long branched and
scattered short glandular hairs
short, much branched,
dendroid hairs
long simple (silky)
hairs and scattered
short glandular hairs
Leaf size (5–)6–18(–20) × 3–4 mm 5.0–27.0(–40.0) × 1.5–8.0 mm 6–15 × 1–3 mm 2–7(–9) × 1–2 mm
Leaf shape linear-lanceolate or linear to
spatulate, incurved
oblanceolate, oblong-spathulate
or obovate
narrowly elliptic to
linear to elliptic or
narrowly ovate
mostly obtuse
Leaf margin with 1(–2) pair of teeth towards
the apex
entire to ± deeply and bluntly
toothed in upper 1/2
entire or with a pair of
teeth near apex
Leaf indumentum silky hairy towards the apex
and margin
with short simple hairs at margin
and apex
short simple and
dendroid hairy
towards the apex and
silky hairy towards the
apex and margin
only long simple (silky) hairs with short, branched hairs and
shorter glandular hairs
short, much branched,
dendroid hairs
only long simple (silky)
Calyx 6.0–6.8(–8.2) mm long, silky
hairy with scattered short
glandular hairs
4–5 mm long, with simple hairs,
short glandular hairs also
occasionally present
3–6 mm long, dendroid
hairs with scattered
short glandular hairs
4–5 mm long, pillose
to silky hairy with
scattered short
glandular hairs
Corolla color pink with yellow annulus white to rose with yellow annulus white with yellow
white with yellow to
Seed dark brown to black, ovoid,
3.0–3.1 × 2.0 mm
dark brown to black, ovoid-
orbicular, 1.4 × 0.8–1.0 mm
unknown pale brown to yellow,
1.7–2.2 × ca 1.2 mm
azizsancarii was also compared with other A. albana-group
species, and it was then observed that A. armeniaca Duby has
branched, dendroid or at least forked hairs, while A. azizsan-
carii does not have branched hairs. Another species, A. inter-
media Ledeb. is biennial and typically has markedly unequal
pedicels (A. azizsancarii is a short-lived perennial, without
markedly unequal pedicels). When the new species was com-
pared to other Turkish Androsace species (beyond A. sect.
Andraspis), it was observed that it superficially resembles A.
villosa, which has a silky indumentum, but its general habit is
mat-forming, with a shorter scape and white flowers.
As a result, it was concluded that A. azizsancarii is a dis-
tinct new species, clearly different from all other members of
A. albana group and easily distinguished from the widespread
A. villosa.
Selected specimens examined
Androsace albana – Turkey, Kastamonu, Ilgaz Mountain, 12
Jun 1885, E. Wiedemann, (E 00024846!); Artvin: Yusufeli,
Altıparmak Village, Kaçkar Mountains footage, moist places,
19 Jul 1989, Z. Aytaç 1991 (GAZI!); Bayburt; south side
of Anzer Mountain, 2860 m a.s.l., 5 Jul 2020, A. Sefali 494
(ISTE 117268!); Rize: Çamlıhemşin, Ortayayla Village,
Verçembek Mountain, above lake of Atmeydan, stony places,
3100 m a.s.l., 20 Aug 1982, A. Güner 4501 (HUB 09014!);
İkizdere, Cimil, above of Cermaniman highland, alpinic
steppe, 2300 m a.s.l., 23 Jul 1984, A. Güner 6000 (HUB
09012!); Kars: Posof, Centrum, Gümüşkonak, Riversides,
28 Jun 1986, Demirkuş 3645 (HUB 09013!), Posof, Ulgar
Mountain, Çamyazı Place to Alköy Village, 1586–2730 m
a.s.l., 16–20 Jun 1986, Demirkuş 3556 (HUB 09015!)
Androsace multiscapa – Lebanon, 1837, P. M. R.
Aucher-Eloy 2607 (Syntype G 00139871!); 1835, P. M. R.
Aucher-Eloy 320 (Syntype G 00139868!); Turkey, Konya,
Ereğli, Aydos Mountain, Yazıgöl highland, alpinic steppe,
2700 m a.s.l., 16 Jul 1981, S. Erik 2319 (HUB 09044!);
Niğde, Ulukışla, Çiftehan, Horoz Village, volcanic gravel,
1750–2900 m a.s.l., 31 Jun 1984, Ş. Yıldırımlı 7222 (HUB
09043!); Ulukışla, Maden Village, Bolkar Mountains,
Kızıl Tepe, Bozkaya Tepe, 2700–2950 m a.s.l., 2 Jul 2015,
K. Gurbanov & A. Savran 79 (Niğde University!)
Androsace villosa – Turkey, Bayburt, Kop Mountains,
Bahtlı Mountain, stony places, 2800 m a.s.l., 20 Jun 2018,
A. Sefalı 394 (BIN!); Erzurum, İspir, above Noryah, hills,
stony places, 2200 m a.s.l., 27 Jun 1981, O. Beyazoğlu (SÜ
11635!); Erzurum, Şenkaya, Allahuekber Mountain, Köroğlu
and Sarıgöl Plateau, meadows, 2850 m a.s.l., 16 Jun 1982,
A. Tatlı 6546 (SÜ 11634!); Konya, Little Geyik Mountain,
steppe, 2300 m a.s.l., 14 Jun 1967, R. Çetik & E. Yurdakul
43 (SÜ 11636!); Konya, Phrygia, Sultandagh, in jugis alpinis
supra Tschai, 9 Jul 1899, Bornmüller & Nicolaus 5502 (E
Androsace azizsancarii (paratypes) – Bayburt; Soganlı
Mountains, south of Anzer Mountain, on moraines, 2831
m a.s.l., 1 July 2020, A. Sefalı 507 (holotype: ISTE 117267!;
isotypes: ANK 60610!, BIN 9405!, EGE 43195!).
Acknowledgements – e author wishes to thank researcher Yakup
Yapar (Bingöl University), who provided the SEM photographs and
measured the pollen grains.
Data availability statement
Data are available from the Dryad Digital Repository:
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... Moreover, the presence of nectar is supported by the different color shade of the corolla mouth observed in different flowers within the same cushion, which may give a signal about the quality and quantity of floral rewards throughout the flowering period for potential pollinators (Melendez-Ackerman et al., 1997;Aragón and Ackerman, 2004). Color changes of the corolla mouth during anthesis were observed also in other Androsace species (Polunin, 1969;Zhang, 1982;Polunin and Stainton, 1984;Stainton and Polunin, 1988;Weiss, 1995;Sefali, 2021) as well as in E. nanum. The latter species, in particular, presents small (Ø =~6-7 mm) blue flowers with five yellow epipetalous fornices that become visible when anthers begin to shed pollen and when nectaries at the base of the ovary start to produce fair amounts of nectar. ...
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Androsace brevis (Hegetschw.) Ces. (Primulaceae) is a narrow endemic plant of the Southern Central European Alps that lives only above 2000 m asl and flowers immediately after snowmelt for a very short period during which the plant must guarantee successful sexual reproduction. Despite the vulnerability of this species, mainly due to increasing temperature, competition with plant communities shifting to higher altitudes, and possible mismatches with pollinators, nothing is known about its pollen and floral morphology. In this study we aim to provide a detailed description of these traits and to investigate how they might be related to possible pollination strategies and vectors. Pollen and flower samples were collected from the Lepontine and Orobic Alps (Northern Italy) populations. The pollen grains are small (polar axis: 19.17 μm ± 0.09; equatorial axis: 10.84 μm ± 0.1) with a prolate shape. The morphology of the flower, with the location of both stamens and pistil inside the corolla tube, and the coloured corolla mouths, ranging from yellow to purple, suggest that A. brevis requires insect-mediated pollination. Moreover, floral morphology does not show a particular insect-selective pattern in that the presence of both stamens and pistil less than 1 mm below the corolla mouth (Ø: 0.76 mm ± 0.05) might allow many high-altitude flower-visiting insects to reach both nectar and pollen. A generalist pollination strategy could be fundamental in contests characterized by harsh environmental conditions as it could counteract potential plant-pollinator mismatches due to increase in temperatures.
Androsace artvinensis (Primulaceae) from Artvin province (northeastern Anatolia) has been described as a new species to science. The new species is morphologically similar to A. armeniaca var. macrantha but differs from it by several morphological characters, such as bracts shape, calyx (calyx length and lobes), corolla (corolla length, lobes, and indumentum), hairs, leaves, pollen features, and habitat preference. Diagnostic morphological characteristics, a full description, and a distribution map are provided.
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Mountaineering, since the beginning of its history, has played an inconspicuous but key role in the collection of species samples at the highest elevations. During two historical expeditions undertaken to reach the summit of Mount Everest in 1935 and 1952, mountaineers collected five species of vascular plants from both the north and south sides of the mountain, at ca. 6400 m a.s.l. Only one of these specimens was determined immediately following the expedition (Saussurea gnaphalodes), and the remaining four were not identified until quite recently. In 2000, the second specimen from the 1935 expedition was described as a new species for science (Lepidostemon everestianus), endemic to Tibet. In this paper, the remaining three specimens from the 1952 Everest expedition are reviewed and analysed, bringing the number of species sharing the title of “highest known vascular plant” from two to five. I identify one of the 1952 specimens as Arenaria bryophylla, and describe two novel taxa based on analysis of the herbarium records: Saxifraga lychnitis var. everestianus and Androsace khumbuensis. Although elevation records on their own do not inform us about the ecological conditions and physiological capacity of plants at the upper limit of their distribution, this taxonomic investigation contributes to our knowledge of the biogeography of Himalayan flora and opens the way for future field-based investigations of mechanisms limiting plant growth on the roof of the world.
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High-altitude rockjasmines (genus Androsace) are a paramount example of evolutionary radiations in temperate mountains of the Northern Hemisphere. Yet, we show here that their taxonomy is incomplete and has been subject to many historical mistakes, probably due to the lack of exploration of mountains by the classical botanists who described these species. Here we wish to clarify the application of names with regard to four defined morphotypes, typical of high-elevation zones of the Western Alps, in order to set a definitive basis for morphological delimitation of a likely new species discovered in the Mont Blanc range (species not described in this work). To do so, we review the historical taxonomic treatments and positions, carefully reconsider types and species names for these morphotypes, and designate lectotypes and epitypes for each of them. In particular, we confirm the validity of names commonly used to refer to the taxa Androsace alpina, Androsace helvetica, and Androsace pubescens. We show that Androsace vandellii is an invalid name and that Androsace argentea should be used instead. Our work illustrates the utility of historical herbaria to clarify the taxonomy of complex groups of plants growing in inaccessible environments such as high-altitude regions.
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We have counted the currently known, described and accepted number of plant species as ca 374,000, of which approxi-mately 308,312 are vascular plants, with 295,383 flowering plants (angiosperms; monocots: 74,273; eudicots: 210,008). Global numbers of smaller plant groups are as follows: algae ca 44,000, liverworts ca 9,000, hornworts ca 225, mosses 12,700, lycopods 1,290, ferns 10,560 and gymnosperms 1,079. Phytotaxa is currently contributing more than a quarter of the ca 2000 species that are described every year, showing that it has become a major contributor to the dissemination of new species discovery. However, the rate of discovery is slowing down, due to reduction in financial and scientific support for fundamental natural history studies.
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In the course of molecular phylogeographical investigations in the Aretia group of Androsace (Primulaceae), a previously unrecognised entity from the eastern Pyrenees (Spain/France) was identified as genetically distinct lineage. The entity is here morphologically characterised and described as new subspecies, Androsace halleri subsp. nuria Schönsw. & Schneew.
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A close relationship of Androsace and related genera (Douglasia, Vitaliana and Pomatosace) has long been recognized. Recent molecular studies have provided abundant evidence that Douglasia, Vitaliana and Pomatosace are nested within Androsace and together constitute the monophyletic “Androsace group”. We investigated pollen morphology of 80 taxa representing all sections of Androsace s.s. as traditionally construed, as well as Douglasia, Vitaliana and Pomatosace, to see whether they are congruent with phylogenetic relationships. We uncovered subtle variation in pollen morphology within the group. The shape of pollen grains ranges from spheroidal to perprolate. Pollen size ranges from 9.37 μm in Androsace sect. Samuelia to 20.68 μm in Douglasia. Exine ornamentation includes microreticulate, microechinate, perforate and rugulate types. The polar view varies from circular, triangular and planaperturate, to triangular and angulaperturate. Various pollen morphological characters support the monophyly of major clades, including /Septentrionalis, /Pomatosace, /Orthocaulon and /Megista, which were recognized previously based on molecular evidence.
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We still have limited understanding of the contingent and deterministic factors that have fostered the evolutionary success of some species lineages over others. We investigated how the interplay of intercontinental migration and key innovations promoted diversification of the genus Androsace. Mountain ranges and cold steppes of the Northern Hemisphere. We reconstructed ancestral biogeographical ranges at regional and continental scales by means of a dispersal-extinction-cladogenesis analysis using dated Bayesian phylogenies and contrasting two migration scenarios. Based on diversification analyses under two frameworks, we tested the influence of life form on speciation rates and whether diversification has been diversity-dependent. We found that three radiations occurred in this genus, at different periods and on different continents, and that life form played a critical role in the history of Androsace. Short-lived ancestors first facilitated the expansion of the genus' range from Asia to Europe, while cushions, which appeared independently in Asia and Europe, enhanced species diversification in alpine regions. One long-distance dispersal event from Europe to North America led to the diversification of the nested genus Douglasia. We found support for a model in which speciation of the North American-European clade is diversity-dependent and close to its carrying capacity, and that the diversification dynamics of the North American subclade are uncoupled from this and follow a pure birth process. The contingency of past biogeographical connections combined with the evolutionary determinism of convergent key innovations may have led to replicated radiations of Androsace in three mountain regions of the world. The repeated emergence of the cushion life form was a convergent key innovation that fostered radiation into alpine habitats. Given the large ecological similarity of Androsace species, allopatry may have been the main mode of speciation.
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Androsace septentrionalis L. (Primulaceae) is recorded as a new species for the Balkan flora on the basis of herbarium material collected several times from Mt. Prokletije, situated near the borders of Serbia (Metochia and Kosovo province), Montenegro and Albania. The locality marks the southernmost limit of the species' range in Europe. The existence of A. septentrionalis in the Balkans may be the result of migration of the tundra-steppe flora from central and East Europe towards the mountains of the peninsula during the Ice Age.
Jacquemoud, F. & D. Jordan (2020). Androsace albimontana (Primulaceae): a new species from the Alps (France, Switzerland, Italy) to be distinguished from A. pubescens. Candollea 75: 149–155. In French, English and French abstracts. A new alpine species of the genus Androsace L. (Primulaceae), Androsace albimontana D. Jord. & Jacquemoud is described. Specimens attributed to the new taxon were until now considered by floras as being part of the high morphological and ecological variability of Androsace pubescens DC. (bi- to trifurcate vs simple trichomes, corolla mostly pink to rarely white vs white; acid, siliceous vs alkaline, calcareous substrate), a chasmophytic species growing exclusively on limestone cliffs. Morphology, habitat and ecology of the new taxon were determined in the siliceous mountain ranges of Aiguilles Rouges and Mont-Blanc (Haute-Savoie, France), and in Valais (Switzerland). Data from Italy (Monte Rosa, Valle d'Aosta, Piemonte) only provided by floras or botanical literature, will have to be subject of further field and herbarium studies. Distribution data are given, relationships with other species of Androsace subsect. Aretia (L.) Kress and the putative origin of the new taxon are discussed.
Nodules harboring nitrogen-fixing rhizobia are a well-known trait of legumes, but nodules also occur in other plant lineages, with rhizobia or the actinomycete Frankia as microsymbiont. It is generally assumed that nodulation evolved independently multiple times. However, molecular-genetic support for this hypothesis is lacking, as the genetic changes underlying nodule evolution remain elusive. We conducted genetic and comparative genomics studies by using Parasponia species (Cannabaceae), the only nonlegumes that can establish nitrogen-fixing nodules with rhizobium. Intergeneric crosses between Parasponia andersonii and its nonnodulating relative Trema tomentosa demonstrated that nodule organogenesis, but not intracellular infection, is a dominant genetic trait. Comparative transcriptomics of P. andersonii and the legume Medicago truncatula revealed utilization of at least 290 orthologous symbiosis genes in nodules. Among these are key genes that, in legumes, are essential for nodulation, including NODULE INCEPTION (NIN) and RHIZOBIUM-DIRECTED POLAR GROWTH (RPG). Comparative analysis of genomes from three Parasponia species and related nonnodulating plant species show evidence of parallel loss in nonnodulating species of putative orthologs of NIN, RPG, and NOD FACTOR PERCEPTION. Parallel loss of these symbiosis genes indicates that these nonnodulating lineages lost the potential to nodulate. Taken together, our results challenge the view that nodulation evolved in parallel and raises the possibility that nodulation originated ∼100 Mya in a common ancestor of all nodulating plant species, but was subsequently lost in many descendant lineages. This will have profound implications for translational approaches aimed at engineering nitrogen-fixing nodules in crop plants. symbiosis | biological nitrogen fixation | evolution | comparative genomics | copy number variation N itrogen sources such as nitrate or ammonia are key nutrients for plant growth, but their availability is frequently limited. Some plant species in the related orders Fabales, Fagales, Rosales, and Cucurbitales-collectively known as the nitrogen-fixing clade-can overcome this limitation by establishing a nitrogen-fixing endosymbi-osis with Frankia or rhizobium bacteria (1). These symbioses require specialized root organs, known as nodules, that provide optimal physiological conditions for nitrogen fixation (2). For example, nod-ules of legumes (Fabaceae, order Fabales) contain a high concentration of hemoglobin that is essential to control oxygen homeostasis and protect the rhizobial nitrogenase enzyme complex from oxidation (2, 3). Legumes, such as soybean (Glycine max), common bean (Phaseolus vulgaris), and peanut (Arachis hypogaea), represent the only crops that possess nitrogen-fixing nodules, and engineering this trait in other crop plants is a long-term vision in sustainable agriculture (4, 5). Nodulating plants represent ∼10 related clades that diverged >100 Mya, supporting a shared evolutionary origin of the underlying capacity for this trait (1). Nevertheless, these nodulating clades are interspersed with many nonnodulating lineages. This has led to two hypotheses explaining the evolution of nodulation (1). The first is that nodulation has a single origin in the root of the nitrogen-fixation clade, followed by multiple independent losses. Significance Fixed nitrogen is essential for plant growth. Some plants, such as legumes, can host nitrogen-fixing bacteria within cells in root organs called nodules. Nodules are considered to have evolved in parallel in different lineages, but the genetic changes underlying this evolution remain unknown. Based on gene expression in the nitrogen-fixing nonlegume Parasponia andersonii and the legume Medicago truncatula, we find that nodules in these different lineages may share a single origin. Comparison of the genomes of Parasponia with those of related nonnodulating plants reveals evidence of parallel loss of genes that, in legumes, are essential for nodulation. Taken together, this raises the possibility that nodu-lation originated only once and was subsequently lost in many descendant lineages.
An update of the Angiosperm Phylogeny Group (APG) classification of the orders and families of angiosperms is presented. Several new orders are recognized: Boraginales, Dilleniales, Icacinales, Metteniusiales and Vahliales. This brings the total number of orders and families recognized in the APG system to 64 and 416, respectively. We propose two additional informal major clades, superrosids and superasterids, that each comprise the additional orders that are included in the larger clades dominated by the rosids and asterids. Families that made up potentially monofamilial orders, Dasypogonaceae and Sabiaceae, are instead referred to Arecales and Proteales, respectively. Two parasitic families formerly of uncertain positions are now placed: Cynomoriaceae in Saxifragales and Apodanthaceae in Cucurbitales. Although there is evidence that some families recognized in APG III are not monophyletic, we make no changes in Dioscoreales and Santalales relative to APG III and leave some genera in Lamiales unplaced (e.g. Peltanthera). These changes in familial circumscription and recognition have all resulted from new results published since APG III, except for some changes simply due to nomenclatural issues, which include substituting Asphodelaceae for Xanthorrhoeaceae (Asparagales) and Francoaceae for Melianthaceae (Geraniales); however, in Francoaceae we also include Bersamaceae, Ledocarpaceae, Rhynchothecaceae and Vivianiaceae. Other changes to family limits are not drastic or numerous and are mostly focused on some members of the lamiids, especially the former Icacinaceae that have long been problematic with several genera moved to the formerly monogeneric Metteniusaceae, but minor changes in circumscription include Aristolochiaceae (now including Lactoridaceae and Hydnoraceae; Aristolochiales), Maundiaceae (removed from Juncaginaceae; Alismatales), Restionaceae (now re-including Anarthriaceae and Centrolepidaceae; Poales), Buxaceae (now including Haptanthaceae; Buxales), Peraceae (split from Euphorbiaceae; Malpighiales), recognition of Petenaeaceae (Huerteales), Kewaceae, Limeaceae, Macarthuriaceae and Microteaceae (all Caryophyllales), Petiveriaceae split from Phytolaccaceae (Caryophyllales), changes to the generic composition of Ixonanthaceae and Irvingiaceae (with transfer of Allantospermum from the former to the latter; Malpighiales), transfer of Pakaraimaea (formerly Dipterocarpaceae) to Cistaceae (Malvales), transfer of Borthwickia, Forchhammeria, Stixis and Tirania (formerly all Capparaceae) to Resedaceae (Brassicales), Nyssaceae split from Cornaceae (Cornales), Pteleocarpa moved to Gelsemiaceae (Gentianales), changes to the generic composition of Gesneriaceae (Sanango moved from Loganiaceae) and Orobanchaceae (now including Lindenbergiaceae and Rehmanniaceae) and recognition of Mazaceae distinct from Phrymaceae (all Lamiales).