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NORDIC JOURNAL OF
BOTANY
Nordic Journal of Botany
1
––––––––––––––––––––––––––––––––––––––––
© 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
Introduction
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
AbdurrahmanSefali
A. Sefali (https://orcid.org/0000-0002-0092-0857) ✉ (asef4petal@gmail.com), Dept of Primary Education, Faculty of Education, Bayburt Univ.,
Bayburt, Turkey.
Research
2
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 ( ).
3
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.
4
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°30′N, 40°30′E,
1 July 2020, A.Sefalı 507 (holotype: ISTE 117267; isotypes:
ANK 60610, BIN 9405, EGE 43195).
Etymology
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).
Description
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.
5
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.
Phenology
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.
6
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
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
entire
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
margin
silky hairy towards the
apex and margin
Scape
indumentum
only long simple (silky) hairs with short, branched hairs and
shorter glandular hairs
short, much branched,
dendroid hairs
only long simple (silky)
hairs
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
annulus
white with yellow to
rosy-pink
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
7
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
E00024849!)
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:
<http://dx.doi.org/10.5061/dryad.4f4qrc4> (Sefali 2021).
References
Anderberg, A. A. and Kelso, S. 1996. Phylogenetic implications of
endosperm cell wall morphology in Douglasia, Androsace and
Vitaliana (Primulaceae). – Nord. J. Bot. 16(5): 481–486.
APG IV 2016. An update of the Angiosperm Phylogeny Group
classification for the orders and families of flowering plants:
APG IV. – Bot. J. Linn. Soc. 181: 1–20.
Baytop, A. 1998. İngilizce-Türkçe botanik kılavuzu. – İstanbul
Üniversitesi Basımevi ve Film Merkezi Üniversite Yayın (4058).
Boucher, F. C. et al. 2012. Reconstructing the origins of high-alpine
niches and cushion life form in the genus Androsace S.L.
(Primulaceae). – Evolution 66: 1255–1268.
Christenhusz, M. J. M. and Byng, J. W. 2016. e number of
known plants species in the world and its annual increase. –
Phytotaxa 261: 201–217.
Dentant, C. 2018. e highest vascular plants on Earth. – Alp. Bot.
128: 97–106.
Dentant, C. et al. 2018. Taxonomic revision of West-Alpine cush-
ion plant species belonging to Androsace subsect. Aretia. – Bot.
Lett. 165: 337–351.
Erdtman, G. 1969. Handbook of palynology, morphology, taxon-
omy and ecology. – Munksgaard, Copenhagen, Denmark.
Grossheim, A. A. 1967. Androsace L. – In: Flora of the Caucasus.
7. Izdatelstvo Nauka, Leningrad, pp. 167–174.
Jacquemoud, F. and Jordan, D. 2020. Androsace albimontana (Prim-
ulaceae): une nouvelle espèce des Alpes (France, Suisse, Italie)
à distinguer de A. pubescens. – Candollea 75: 149–155.
Lamond, J. 1978. Androsace L. – In: Davis, P. H. (ed.), Flora of
Turkey and the East Aegean Islands, Vol. 6. Edinb. Univ. Press,
pp. 122–127.
Mabberley, D. J. 2008. Mabberley’s plant book: a portable diction-
ary of plants, their classifications and uses. – Cambridge
Univ. Press.
Menemen, Y. et al. 2016. Türkçe Bilimsel Bitki Adları Yönergesi.
– Bağbahçe Bilim Derg. 3: 1–3.
Pax, F. and Knuth, R. 1905. Androsace L. (Primulaceae). – In:
Engler, A. (ed.), Das Pflanzenreich. Regni vegetabilis conspec-
tus IV. Wilhelm Englermann, Leipzig, pp. 172–220.
Punt, W. et al. 2007. Glossary of pollen and spore terminology. –
Rev. Palaeobot. Palynol. 143: 1–81.
Roquet, C. et al. 2013. Replicated radiations of the alpine genus
Androsace (Primulaceae) driven by range expansion and conver-
gent key innovations. – J. Biogeogr. 40: 1874–1886.
Schönswetter, P. and Schneeweiss, G. M. 2009. Androsace komoven-
sis sp. nov., a long mistaken local endemic from the southern
Balkan Peninsula with biogeographic links to the eastern Alps.
– Taxon 58: 544–549.
Schönswetter, P. et al. 2015. Androsace halleri subsp. nuria Schönsw.
& Schneew. (Primulaceae), a new taxon from the eastern Pyr-
enees (Spain, France). – Phytotaxa 201: 227–232.
8
Sefali, A. 2021. Data from: Androsace azizsancarii sp. nov. (Primu-
laceae): a new species from northeastern Anatolia, Turkey. –
Dryad Digital Repository, <http://dx.doi.org/10.5061/
dryad.4f4qrc4>.
Shishkin, B. K. and Bobrov, E. G. 1952. Androsace L. – In: Shishkin,
B. K. and Bobrov, E. G. (eds), Flora SSSR XVIII. Izdateľstvo
Akademii Nauk Belopusskoi SSR, Moskva, Leningrad,
pp. 217–242.
Smith, G. and Lowe, D. 1997. e genus Androsace. – Alpine Gar-
den Society, Avon Bank, UK.
Stevanovıć, V. et al. 2005. Androsace septentrionalis (Primulaceae),
a new species for the Balkan flora. – Ann. Bot. Fenn. 42: 35–39.
e Plant List 2013. Ver. 1.1. Published on the internet. – <www.
theplantlist.org/>, accessed 1 May 2021.
Xu, Y. et al. 2016. Pollen morphology of Androsace (Primulaceae)
and its systematic implications. – J. Syst. Evol. 54: 48–64.