Notes on Crocus L. Series Flavi Mathew (Iridaceae) and a new species with unique corm structure

Abstract

Crocus asymmetricus (Iridaceae) is described as a new species endemic to the southern part of the Anatolian diagonal in Turkey. It is phylogenetically related to C. vitellinus and morphologically to C. antalyensis, but differs from these species in showing an asymmetric corm and a single point of root emergence. Both C. antalyensis and C. asymmetricus are illustrated and compared in this paper. A Bayesian phylogenetic tree of the nuclear rDNA ITS region confirms the affiliation of C. asymmetricus to C. ser. Flavi, and its close relationship to C. vitellinus. A new identification key to the species of C. ser. Flavi occurring in Turkey is also presented.

Full text

Phytotaxa 438 (2): 065–079
https://www.mapress.com/j/pt/
Copyright © 2020 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Giovanni Astuti: 13 Mar. 2020; published: 6 Apr. 2020
https://doi.org/10.11646/phytotaxa.438.2.1
65
Notes on Crocus L. Series Flavi Mathew (Iridaceae) and a new species with unique
corm structure
ALMILA ÇIFTÇI1,4, DOERTE HARPKE2,5, RACHEL MOLLMAN1,6, HASAN YILDIRIM3,7 & OSMAN EROL1,8*
1 
2 Leibniz Institute of Plant Genetics and Crop Research (IPK), 06466 Gatersleben, Germany.
3 
4
�
almila.ciftci@istanbul.edu.tr; https://orcid.org/0000-0002-3406-3064
5
�
e; https://orcid.org/0000-0003-1667-2912
6
�
zmollman@gmail.com; https://orcid.org/0000-0002-6873-6212
7
�
hasanyldrm@gmail.com; https://orcid.org/0000-0003-3951-4343
8
�
erol@istanbul.edu.tr; https://orcid.org/0000-0002-6310-1402
*Author for correspondence
Abstract
Crocus asymmetricus (Iridaceae) is described as a new species endemic to the southern part of the Anatolian diagonal in
Turkey. It is phylogenetically related to C. vitellinus and morphologically to C. antalyensis, but differs from these species in
showing an asymmetric corm and a single point of root emergence. Both C. antalyensis and C. asymmetricus are illustrated
and compared in this paper. A Bayesian phylogenetic tree of the nuclear rDNA ITS region confirms the affiliation of C.
asymmetricus to C. ser. Flavi, and its close relationship to C. vitellinus. A new identification key to the species of C. ser. Flavi
occurring in Turkey is also presented.
Introduction
Crocus Linnaeus (1753: 36) (Iridaceae) is one of the largest genera in the flora of Turkey. This genus is represented by
over 135 species according to Rukšāns (2017). Some species are taxonomically doubtful, because of unclear locality
data, inadequate type specimens, and lack of clear diagnoses. Despite this, Turkey is the centre of diversity of the genus
and scientists still discover new taxa even near heavily trafficked roads and human settlements. Most of the Turkish
Crocus taxa belong to C. sect. Nudiscapus Mathew (1982: 61), which is further divided into series. In several cases,
a detailed examination of corm tunics allows the placement of a taxon in a series or even identification at the species
level (Kerndorff et al. 2015). Therefore, the corm still represents a useful part of the plant for diagnostic purpose.
Within C. sect. Nudiscapus, C. ser. Flavi Mathew (1982: 84) is characterized by membranous (sometimes papery)
parallel-fibrillated tunics and multifid stigma branches (Mathew 1982), with the exceptions of C. adanensis Baytop
& Mathew (1975: 245) and C. paschei Kerndorff (1994: 76), both of which have a three-branched stigma and poorly
developed rings at the base of the corm. Phylogenetic studies (Petersen et al. 2008, Harpke et al. 2013) placed the latter
two taxa as sister to all the other C. ser. Flavi members.
In the current circumscription, C. ser. Flavi includes 16 taxa: C. adanensis, C. antalyensis Mathew (1972: 327)
subsp. antalyensis, C. antalyensis subsp. striatus Erol & Koçyiğit (2010: 187), C. antalyensis subsp. gemicii Sik
& Erol (2011: 282), C. antalyensioides Rukšans (2015: 6), C. balansae J.Gay ex Baker (1879: 234), C. candidus
Clarke (1812: 145), C. mouradi Whittall (1889: 473), C. flavus Weston (1771: 237), C. graveolens Boissier & Reuter
(1882:107), C. hyemalis Boissier (1859: 93), C. istanbulensis Mathew (1982: 99) Rukšâns (2017: 271), C. olivieri Gay
(1831: 319), C. paschei Kerndorff (1994:76), C. sarichinarensis Rukšans (2010: 157) Rukšans (2017: 442), and C.
vitellinus Wahlenberg (1828: 1000).
An amateur scientist, Mehmet Çelik, found a Crocus population near the locus classicus of C. adanensis in
Maraş province, Turkey. The individuals of this population are characterized by asymmetric corms, a shape unique
among all species of the genus, thus suggesting that they represent a new species. Therefore, we carried out a thorough
morphological investigation and used the rDNA internal transcribed spacer (ITS) region to confirm its rank as a
putative new species of C. ser. Flavi and to identify its closest relatives.

ÇIFTÇI 
66 • Phytotaxa 438 (2) © 2020 Magnolia Press
FIGURE 1. Crocus asymmetricus from its locus classicus: habit (A, B); peeled corm (C); corms with tunics (D).
The most interesting feature of Crocus asymmetricus sp. nov. is the organization of its corm tunics. Its flower
morphology is similar to C. antalyensis, but it differs by its asymmetrically oriented corm and tunics, as well as by its
lack of persistent cataphylls (Figs. 1, 2, 3).

NOTES ON CROCUS L. SERIES FLAVI MATHEW Phytotaxa 438 (2) © 2020 Magnolia Press • 67
Material and Methods
Morphological investigations:—Standard deviations (SD) were calculated for each quantitative morphological
parameter. The number of individuals measured (indicated by n) for each morphological character is given in the
description. Photographs of the plants were taken with a Canon 5Ds digital camera (60 and 100 mm macro-lenses).
Macromorphological features were examined under a stereo-binocular microscope. We compared the morphology
of the undescribed specimens to the relevant species descriptions available in the taxonomic literature (Maw 1886,
Bowles 1952, Mathew 1982, Mathew 1984, Rukšāns 2017) and herbarium material conserved in EGE, ISTE, ISTF, K
(acronyms follow Thiers 2019).
FIGURE 2. Illustration of C. asymmetricus: A. Habitus; B. Flower dissection; C. Leaf cross section; D. Corm: D1. General view, D2.
Young (above) and mature corm (below); E. Corm tunics (E1 and E2); F. Basal tunic. Drawing made from specimen ISTF 41370.

ÇIFTÇI 
68 • Phytotaxa 438 (2) © 2020 Magnolia Press
FIGURE 3. Illustration of Crocus antalyensis: A. Habitus; B. Flower dissection; C. Leaf cross section; D. Corm: D1. General view, D2.
Young (above) and mature (below) corm; E. Tunics (E1, E2, E3). Drawing made from specimen ISTF 41144.
Molecular methods:—The extraction of genomic DNA and amplification of the nuclear ribosomal region
ITS of five individuals were performed according to Harpke et al. (2013). Both strands of the PCR products were
directly Sanger sequenced with Applied Biosystems BigDye Terminator technology on an ABI3730xl automatic DNA
sequencer using the primers from PCR amplifications.

NOTES ON CROCUS L. SERIES FLAVI MATHEW Phytotaxa 438 (2) © 2020 Magnolia Press • 69
FIGURE 4. Phylogenetic tree obtained by Bayesian inference of the nuclear rDNA ITS. Numbers along branches are posterior
probabilities.

ÇIFTÇI 
70 • Phytotaxa 438 (2) © 2020 Magnolia Press
TABLE 1. Morphological comparison of Crocus asymmetricus and its related species (abbreviations written in bold and a list of all abbreviations given at the end of the table in
alphabetical order).*
Species/
Characters
C. asymmetricus
C. antalyensis
C. adanensis
C. graveolens
C. vitellinus
C. hyemalis
C. olivieri
C. balansae
C. istanbulensis
C. flavus
C. mouradi
C. candidus
C. antalyensioides
C. paschei
C. sarichinarensis
Corm shape asym sym sym sym sym sym sym sym sym sym sym sym sym sym sym
Emergence
of root one full two
– several full full full full full full full full full full full full
Tunics
c/m
heads
towards
one side
(asym) spl
into narrow
bands, or
triangles
from base
p, spl into
narrow bands
or prl fibrils
from base
m, spl into
stripes or
prl fibrils
m/c, spl at
base into prl
fibrils
m/c
spl at base
into narrow
strips or
fibrils
m
spl length
ways into
prl strips at
base
m
spl at base
into coarse,
prl fibres or
triangular
teeth
m
spl at base
into prl
fibrils
c
coarsely
fibrous,
fibrils prl,
weakly
reticulate at
apex
m with
fibrous
points at
apex, spl at
the base into
coarse prl
fibres
m, distinctly
fibrous with
prl fibrils
m, spl into
prl fibrils
p, spl at
base into
many prl
fibrils
p, with
occasional
splits at
base
prl fibrous,
outer tunics
split strap-
like at base
Tunic neck
narrow,
sharp,
triangular
teeth
formed
by old
cataphylls
short, p
fibrous
points of
main tunic
fibrous
points of
main tunic
poorly
developed,
soft fibrous
points
sharp
triangular
fibrous
points
narrow
sharp
fibrous
splits
bristly
formed
by old
cataphylls
formed
by old
cataphylls
narrow
sharp
tunic splits
formed
by old
cataphylls
short, p
formed
by old
cataphylls
Basal rings ab ab poorly
developed ab ab ab ab ab ab ab ab ab ab ab ab
Flower
segment
shape
el; ob to sac el to obl; ob
to sac
obov to sla;
ob to sac
wai; obl or
el; ac to ob
nd wai; obl
to obov; ob
to sac
obl to ov;
sac to ob
el, obl to
obov; ob to
sac
ov; sac el to obl;
sac
obl or ov; ob
to sac
obl
or ov; ob,
sac to a
el; ob ov to sla;
ob to sac
ov; ob to
sac
obov to
obl; ob to
sac
......continued on the next page

NOTES ON CROCUS L. SERIES FLAVI MATHEW Phytotaxa 438 (2) © 2020 Magnolia Press • 71
TABLE 1. (Continued)
Species/
Characters
C. asymmetricus
C. antalyensis
C. adanensis
C. graveolens
C. vitellinus
C. hyemalis
C. olivieri
C. balansae
C. istanbulensis
C. flavus
C. mouradi
C. candidus
C.
antalyensioides
C. paschei
C.
sarichinarensis
Colour of
outer segments
dull li-wh
with three
distinct pur
stripes and
featherings
pl or deep
li bl, or
wh with
bl staining
on base,
sometimes
buff-colored
with vio
stripes or
flecks
dull li-wh
y o usually
with 3
major
br-ish-vio
stripes
bright y
to o y,
sometimes
with br-
ish-vio
stripes or
speckles
wh with
pur
staining or
speckling,
sometimes
with
central
band
of vio
reaching
apex
usually
uniform
bright
o y or
sometimes
pl y, rarely
with br-ish
striping or
staining
y to o y,
sometimes
with
translucent
striping,
very rarely
pl y
uniform y
to o
pl y to
deep o y,
sometimes
striped or
stained br,
vio or gr at
base
pl to dark
y, same
color inside
and out,
sometimes
with gr-ish
staining at
base
spotted or
suffused
gr-ish-bl or
pur
uniform
bl or with
dkr vio-bl
shading
along
midrib, gre-
ish gr blotch
at base
outside sil to
buff-colored
with a
distinct gr-
ish speckled
basal blotch
or only dkr
speckled at
the base,
maybe
extending
up in three
indistinct
lines
y, occ wh
or creamy,
always
suffused,
speckled or
diffusely
striped gr-
ish to br-ish
Colour of
inner segments
pur,
sometimes
with one
inconspicuous
vio stripe
pl or deep
li bl, or
wh with bl
staining on
base
pl li y obright y
to yo
wh, with
pointed
gr-ish
basal
blotch
bright o y
y to o y
with gre-
ish, br-ish
or gr-ish
basal
blotch
y to o,
same
colour
inside
and out,
without
marking
pl y to
deep o ypl to dk ywh with y
base
Same as
outer
Same as
outer, but sl
dkr
y, with a
dk gr-ish
or br-ish
speckled,
well-defined
basal blotch
Throat bright y, gla y, pub
usually wh,
sometimes
with sl y
center, gla
deep y,
gla or pub
o to y, gla
or pub y, gla y, gla or
pub
y, gla or
pub
deep y to
o, gla
y, gla or
pub y, gla or pub bright y,
gla y, gla pl to deep
y, gla
pl to deep
y, usually
with a
distinct wh
zone above
throat, gla
......continued on the next page

ÇIFTÇI 
72 • Phytotaxa 438 (2) © 2020 Magnolia Press
TABLE 1. (Continued)
Species/
Characters
C. asymmetricus
C. antalyensis
C. adanensis
C. graveolens
C. vitellinus
C. hyemalis
C. olivieri
C. balansae
C. istanbulensis
C. flavus
C. mouradi
C. candidus
C. antalyensioides
C. paschei
C. sarichinarensis
Outer flower
segment size
(mm)
19–30 ×
4–9
20–35 ×
6–11 20–25 × 4–7 18–28 × 4–720–30 ×
6–9
24–42 ×
6–15
15–35 ×
4–12
25–32 ×
7–9
20–27 ×
7–11
20–35 × (4–)
6–12
15–30 ×
5–10
20–23 ×
6–10
35–40 ×
17–20
28–38 ×
9–15
21–34 ×
10–16
Inner flower
segment size
(mm)
20–28 ×
5–10
20–28 ×
7–9 20–25 × 4–718–28 × 4–7 20–30 ×
6–9
23–42 ×
6–15
15–35 ×
4–12
25–32 ×
7–9
20–27 ×
7–11
20–35 ×
4–12
15–30 ×
5–10
20–23 ×
6–10
35–40 ×
17–20
19–30 ×
6–12
21–34 ×
10–16
Colour of
anthers pl y pl y y y y rarely
bla
bla-m,
rarely y y y to o y y deep y y y y y
Length of
anthers (mm) 9–14 10–12 7–9 9–13 9–13 9–13 6–15 12–15 9–11 8–15 8–15 9–10 10–20 7–10 10–17
Style mul
many bra
mul
6–12 sle
bra
tri
sometimes
with very
shallow
secondary
division, but
not widening
near the tips
mul
many cr
bra
mul
many bra
mul
deeply
divided
into 8–15
sle bra
mul
6 sle bra
mul
(8–)12–15
bra
mul
6 sle bra
mul
primarily
divided into
3 at the apex,
divided into
secondary
branches
mul
6 or more
distinct,
sle bra
mul 6 sle
bra
mul 8–15
bra with
short
subdivisions
tri, sl
expanded
and fringed
at apex
mul primarily
divided into
3 at the apex,
divided into
secondary
branches
......continued on the next page

NOTES ON CROCUS L. SERIES FLAVI MATHEW Phytotaxa 438 (2) © 2020 Magnolia Press • 73
TABLE 1. (Continued)
Species/
Characters
C. asymmetricus
C. antalyensis
C. adanensis
C. graveolens
C. vitellinus
C. hyemalis
C. olivieri
C. balansae
C. istanbulensis
C. flavus
C. mouradi
C. candidus
C. antalyensioides
C. paschei
C. sarichinarensis
Colour of
style oo or y o to rdeep y
or odeep y or oo
y to o, s
wh-ish at
tip
y to o y to o y to oy to o, occ
wh-ish y to o
wh-ish to
creamy
rarely y
deep o to red y
Length
of style
according to
stamens
always
lon mostly sl sh lon sh or sl
lon
usually sh
or eq sh or eq sl sh to lon sl sh or lon sl sh rarely eq
or sh
usually sh /
sometimes
sl lon
eq or just
exceeding
usually eq
rarely long
mostly lon
rarely eq sh, rarely eq
Flowering
time Feb Feb–Mar Feb–Mar (Jan–)
Feb–Apr
Nov–Jan S;
Feb–Mar(–
Apr) N
Nov–
Dec(–Jan)Feb–Apr Jan–Mar Feb Mar–Apr Mar Feb–Mar Feb–Mar Mar Mar
Corm: asym = asymmetric, full = full perimeter of basal node, one = one side of basal node, several = from 1-3 points of basal node, sym = symmetric; two = two sides of the basal
node
Tunic: ab = absent, c = coriaceous, m = membranous, p = papery, pr = present, prl = parallel spl = splitting
Flower: ac = acute, el = elliptic, gla = glabrous, nd wai = non-distinct waist, ob = obtuse, obl = oblanceolate, ov = obovate, pub = pubescent, sac = subacute, sla = sublanceolate, wai
= waisted
Color: bl = blue, bla = blackish, br = brown, dk = dark, dkr= darker, gr = grey, gre = green, li = lilac, lt = light, ltr = lighter, m = maroon, o = orange, occ = occasionally, pl = pale, pur
= purple, r = red, sil =silvery, vio = violet, wh = white, y = yellow
Style: bra = branches, cr bra = curved branches, d = deeply, eq = equal, lon = longer; mul = multifid, sh = shorter, sl = slightly, sle = slender, tri = trifid
Anthers: col. anth. = colour of anthers, l. anth. = length of anthers
Flowering time: Apr = April, Dec = December, Feb = February, Jan = January, Mar = March, N = in Northern part, Nov = November, S = in Southern part
* The table was prepared according to previous literature (Bowles, 1952; Mathew, 1982, 1984; Maw, 1886; Ruksans, 2017) and specimens from herbaria (GAT, B, E, ISTE, ISTF,
GB) listed in Appendix 2.

ÇIFTÇI 
74 • Phytotaxa 438 (2) © 2020 Magnolia Press
FIGURE 5. Mitotic metaphase plate with 2n = 8 chromosomes (a), and idiogram (b) of C. asymmetricus. Scale bar = 10 µm.
Phylogenetic analysis:—Information about the investigated material and accessions numbers are given in the
Appendix. For newly obtained sequences, forward and reverse strands were checked and manually edited where
necessary, after which they were combined in consensus sequences for each locus and individual. Representatives of
other groups of section Nudiscapus where used as outgroups (C. almehensis C.D.Brickell & B.Mathew, C. aleppicus
Baker, C. fleischeri J.Gay, C. tournefortii J.Gay). Sequences were submitted to ENA (European Nucleotide Archive)
and are accessible through accession number ERZ1300521. Sequences were aligned manually, retaining only one
sequence in the case of identical sequences shared by the individuals within a population. The alignment was subjected
to phylogenetic analyses using Bayesian phylogenetic inference (BI) with MrBayes 3.2 (Ronquist et al. 2011). For
BI, 2 times 4 chains were run for 1 million generations under the appropriate GTR+Γ+I model of sequence evolution,
sampling a tree every 100 generations. Converging log-likelihoods, potential scale reduction factors for each parameter
and inspection of tabulated model parameters in MrBayes suggested that stationary had been reached in all analyses.
The first 25% of trees of each run were discarded as burn-in. Two independent runs of BI analysis were performed to
confirm that separate analyses converged on the same result. Each of the two analyses resulted in the same topology
and similar posterior probabilities (pp) for nodal support (Fig. 4).
Karyological investigations:—Living corms were collected from the type locality on 11 February 2019. The
root tips were then fixed with Clarke’s solution (1:3 acetic acid:alcohol) for 3–4 hours, washed in distilled water,
hydrolyzed and stained with Carmine. The mitotic metaphase squash preparations (20 slides) were examined under a
Zeiss AxioScope A1 microscope, and Kameram Software v3.1.0.0.
Description of the new species
Crocus asymmetricus Erol sp. nov.
Crocus asymmetricus is similar to C. antalyensis. It is easily distinguished from C. antalyensis by its asymmetric corm and lack of long,
persistent neck of old cataphylls at the apex.
Type:—TURKEY. Maraş: border of Maraş – Osmaniye, Ceyhan Valley 850–920m a.s.l., 6 March 2019,  (holotype ISTF
41370!, isotype: EGE!)
Corm ovoid-globose, asymmetric when mature, 8–18 mm in width (mean 12.28 ± 2.5; n = 17) × 11–19 mm in length
(mean 15.52 ± 2.6; n = 17); roots emerge from one side. Corm tunics coriaceous to membranous, split into narrow
bands or triangles from base, rings absent; neck narrow sharp triangular teeth, 1.79–4.52 mm long (mean 3.17 ± 0.9;
n = 17). Cataphylls 2-3, white, dried cataphylls white. Prophyll absent. Bract and bracteole present, conspicuous,
silvery white. Leaves synanthous, 3–8, usually 5 (mean 4.83 ± 1.09; n = 18), green, elongate lanceolate, 0.84–3.06 mm
in width at broadest (mean 1.8 ± 0.5; n = 18), white stripe usually 1/2 of leaf diameter, 0.33–1.07 mm (mean 0.76 ±
0.2; n = 18), without ribs, glabrous. Perianth tube creamy white (yellow on herbarium specimens) with purple stripes.
Flowers star-shaped, perigone segments elliptic, obtuse to subacute, outer segments 19–30 mm long (mean 25.31 ±
3.4; n = 13), 4–9 mm wide (mean 7.4 ± 1.4; n = 13). Inner segments 20–28 mm long (mean 23.9 ± 2.3; n = 13) and 5–10

NOTES ON CROCUS L. SERIES FLAVI MATHEW Phytotaxa 438 (2) © 2020 Magnolia Press • 75
mm wide (mean 7.47 ± 1.9; n = 13). Segments purple on inside, outer segments dull lilac-white on abaxial side with
three distinct purple stripes and featherings, inner segments purple, sometimes with one inconspicuous purple stripe
and featherings. Throat bright yellow, glabrous. Filaments creamy, 3–6 mm long (mean 4.67 ± 1.05; n = 13), glabrous,
anthers 9–14 mm long (mean 10.84 ± 1.5; n = 13), pale yellow, connective white. Styles orange, many branched, with
long branches 5–10 mm long (mean 7.66 ± 0.71; n = 13). Styles always longer than stamens (Fig. 3). Mature capsule
and seed not observed. Flowering February. Chromosome number 2n = 2x = 8 (Fig. 5).
Etymology:—The species epithet is derived from the asymmetric corm of the new species. The Turkish name given
to the species is “Karacaoğlan çiğdemi” according to the guidelines laid out by Menemen et al. (2013). Karacaoğlan
was a folk poet who lived near the type locality of C. asymmetricus in 1700s.
Taxonomic relationships:—The asymmetric corm of Crocus asymmetricus is unique within the genus. Other
corm tunic characters and the multifid stigma branches suggest that this species belongs to C. ser. Flavi. Prior to the
discovery of this new species, the only species in series C. ser. Flavi showing lilac-blue flowers and multi-branched
stigma were C. antalyensis and C. antalyensioides. These two closely related taxa are characterized by long, brown,
persistent neck of old cataphylls at the apex (Fig. 3, Table 1), whereas the new species has deep lilac-blue flowers
without persistent cataphylls (Fig. 1, 2, Table 1). Besides its slightly asymmetric corms (Figs. 2, 3), C. asymmetricus
shows another unique features, i.e. the arrangement of root emergence, where roots emerge from only one point rather
than from around the basal node as in other species. Since its original description published in 1975 no one paid enough
attention to the corm of C. adanensis. Indeed, after a careful observation of peeled corms (free from tunics) of this
species we found that roots only emerge from the base of the corm in two or three bundles (Figs. 6a, b). This feature is
shared with C. asymmetricus discovered in the Ceyhan valley. The ITS sequences of individuals of this population are
also unique within the genus: eight substitutions and two indels distinguish it from the closest species C. vitellinus. Our
phylogenetic analysis places the new species in a clade comprising C. vitellinus, C. graveolens, and C. hyemalis (pp =
1) (Fig. 4). Morphologically, C. asymmetricus can be distinguished from the latter four species by its flowers color in
addition to the unique corm shape and root emergence (Table 1, Fig. 1).
All species of C. ser. Flavi with the exception of C. hyemalis are part of the Turkish flora. A new identification
key for species occurring in Turkey is presented below (the two subspecies of C. antalyensis are excluded, but a key
for them was provided by Erol et al. 2011).
Identification key to Crocus ser. Flavi in Turkey
1. Corm asymmetric, roots emerge from one side ................................................................................................. Crocus asymmetricus
- Corm symmetric, roots emerge from around basal node...................................................................................................................2.
2. Cataphylls persistent, old cataphylls form a persistent brown sheath ...............................................................................................3.
- Cataphylls not persistent, white .........................................................................................................................................................7.
3. Flowers lilac-blue ..............................................................................................................................................................................4.
- Flowers yellow or pale yellow ..........................................................................................................................................................5.
4. Style yellow-orange, perigone segments narrow, with a length to width ratio of 3.3–3.8 ....................................Crocus antalyensis
- Style white-cream, rarely yellow, perigone segments are ovate-sublanceolate, with a length to width ratio of 2.0–2.5 .....................
........................................................................................................................................................................ Crocus antalyensioides
5. Leaves 1.0–1.5 mm wide, outside of outer flower segments always striped .................................................. Crocus sarichinarensis
- Leaves 2.5–4 mm wide, outside of outer flower segments usually unstriped ...................................................................................6.
6. Style has 3 branches, each splitting into shorter indistinct lobes, and always shorter than anthers ...............................Crocus flavus
- Style has 6–15 branches, reaching the same height as anthers or slightly longer ......................................................Crocus mouradi
7. Style with 3 distinct branches, corm base splits weakly into basal rings ..........................................................................................8.
- Style with more than 3 branches, basal rings absent on base of corm, with parallel fibrils ..............................................................9.
8. Throat white ............................................................................................................................................................. Crocus adanensis
- Throat yellow with a white zone upper side ................................................................................................................ Crocus paschei
9. Corm tunic fibrils coarse, fibrils slightly reticulate at apex ............................................................................... Crocus istanbulensis
- Corm tunic membranous, splitting into narrow triangular strips or fibrils parallel to base ............................................................10.
10. Style has ca. 6 branches, outside of outer perigone segments usually unmarked, perianth tube may be violet .............................11.
- Style with 12–15 branches, outside of outer perigone segments usually with brown-violet stripes or markings ...........................12.
11. Leaves 1–4(–5) per corm, width (1.5–)2–5(–7) mm, flowers yellow ..........................................................................Crocus olivieri
- Leaves 1–2(–4) per corm, width 4.5–9 mm, flowers white with yellow base ..........................................................Crocus candidus
12. Leaf width 4–6 mm, filaments glabrous rarely pubescent ........................................................................................Crocus balansae
- Leaf width 2–4 mm, filaments pubescent-papillose ........................................................................................................................13.
13. Leaves 2–4, 1.5–3 mm wide ......................................................................................................................................Crocus vitellinus
- Leaves 5–8, 0.5–1.5 mm wide ................................................................................................................................Crocus graveolens

ÇIFTÇI 
76 • Phytotaxa 438 (2) © 2020 Magnolia Press
FIGURE 6. Corm of C. adanensis: basal view (a), side view (b).

NOTES ON CROCUS L. SERIES FLAVI MATHEW Phytotaxa 438 (2) © 2020 Magnolia Press • 77
Conclusions
Based on our morphological and molecular analyses, C. asymmetricus represents a distinct, new species. Future studies
will be aimed to add further details, as other characters, such as those of mature capsules and seeds, are considered of
diagnostic importance within the genus (Karaismailoğlu et al. 2018, Kerndorff et al. 2015). Characterization of the
karyotypes of this series in Turkey should also be interesting. Indeed, although the variation in chromosome numbers
(2n = 6, 8, 14) is comparatively low in this group of crocuses, Karamplianis & Constantinides (2019) reported that
karyotypes and number of B chromosomes differ among the four C. ser. Flavi taxa growing in Greece and possessing
the same chromosome number. Further studies involving the karyotypes of the other ten C. ser. Flavi taxa could be
useful in order to gain detailed insights into the karyotype evolution of this group.
Acknowledgments
We would especially like to thank Mehmet Çelik for collecting and sending us specimens of C. asymmetricus and
Kadir Terzioğlu for his kind help in the field trips. This work was supported by the Research Fund of Istanbul
University, Istanbul, Turkey (project number 27545 and 24390). We also thank Nezahat Gökyiğit Botanik Bahçesi
(ANG Foundation) for funding visits to K, E and GAT herbaria.
References
Baytop, T., Mathew, B. & Brighton, C. (1975) Four new taxa in Turkish Crocus (Iridaceae). Kew Bulletin 30: 241–246.
https://doi.org/10.2307/4103158
Boissier, P.E. & Blanche, C.I. (1859) Diagnoses Plantarum Orientalium novarum ser. 2. ser. 2, 4: 93 Lipsiae [Leipzig] pp. 148.
Boissier, P.E. (1882) Flora Orientalis sive enumeratio plantarum in Oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum
5 (1). H. Georg, Basileae & Genève, 428 pp.
Bowles, E.A. (1952)  Edn. 2. pp. 222.
Clarke, E.D. (1812) . London, 145 pp.
Erol, O., Koçyiğit, M., Sik, L., Ozhatay, N. & Kucuker, O. (2010) Crocus antalyensis subsp. striatus subsp. nov. (Iridaceae) from southwest
Anatolia, Nordic Journal of Botany 28: 186–188.
https://doi.org/10.1111/j.1756-1051.2009.00447.x
Erol, O., Kaya, H.B., Şik, L., Tuna, M., Can, L. & Tanyolac, M.B. (2014) The genus Crocus, series Crocus (Iridaceae) in Turkey and 2
East Aegean islands: a genetic approach.  38: 48–62.
https://doi.org/10.3906/biy-1305-14
Erol, O., Şik, L., Kaya, H.B., Tanyolaç, B. & Küçüker, O. (2011) Genetic diversity of Crocus antalyensis B.Mathew (Iridaceae) and a new
subspecies from southern Anatolia. Plant Systematics and Evolution 294: 281–287.
https://doi.org/10.1007/s00606-011-0465-8
Gay, J. (1879) Notes on New Croci. Gardeners’ Chronicle New series 11: 234. [pp. 828]
Gay, J. (1831) Nouvelles Especes de Crocus. Bulletin des Sciences Naturelles et de Géologie 25: 319. [pp. 368]
Harpke, D., Carta, A., Tomović, G., Ranšelović, V., Ranšelović, N., Blattner, F.R. & Peruzzi, L. (2015) Phylogeny, karyotype evolution
and taxonomy of Crocus series Verni (Iridaceae). Plant Systematics and Evolution 301: 309–325.
https://doi.org/10.1007/s00606-014-1074-0
Harpke, D., Meng, S., Rutten, T., Kerndorff, H. & Blattner, F. (2013) Phylogeny of Crocus (Iridaceae) based on one chloroplast and two
nuclear loci: ancient hybridization and chromosome number evolution, Molecular Phylogenetics and Evolution 66: 617–627.
https://doi.org/10.1016/j.ympev.2012.10.007
Harpke, D., Peruzzi, L., Kerndorff, H., Karamplianis, T., Constantinidis, T., Randelovic, V., Randelovic, N., Juskovıc, M. & Blattner, F.R.
(2014) Phylogeny, geographic distribution, and new taxonomic circumscription of the Crocus reticulatus species group (Iridaceae).
 38: 1182–1198.
https://doi.org/10.3906/bot-1405-60
Karaismailoğlu, M.C., Şik, L., Gemicioğlu, A. & Erol, O. (2018) Seed structure of some taxa of the genus Crocus L.(Iridaceae) series
Crocus.  42: 722–731.
https://doi.org/10.3906/bot-1712-17

ÇIFTÇI 
78 • Phytotaxa 438 (2) © 2020 Magnolia Press
Karamplianis, T. & Constantinides, T. (2019) The taxa of Crocus ser. Flavi (Iridaceae) in Greece: a taxonomic and karyomorphometric
study.  22: 171–194.
Kerndorff, H. (1994) Two new taxa in Turkish Crocus. Herbertia 49: 76–86.
Kerndorff, H., Pasche, E. & Harpke, D. (2015) The genus Crocus (Liliiflorae, Iridaceae): life-cycle, morphology, phenotypic characteristics,
and taxonomical relevant parameters. Stapfia 103: 27–65.
Kerndorff, H. (1994) Two New Taxa in Turkish Crocus (Iridaceae), Herbertia 49: 76–78.
Mathew, B. (1982) . Timber Press, London, 224 pp.
Mathew, B. (1984) Crocus L. In: Davis, H. (Ed.)  8. Edinburgh University Press, Edinburgh,
pp. 413–438.
Mathew, B. & Brighton, C.A. (1972) A new Crocus from Turkey, Kew Bulletin 27: 327–329.
https://doi.org/10.2307/4109462
Mathew, B., Petersen, G. & Seberg, O. (2009) A reassessment of Crocus based on molecular analysis. Plantsman 8 (1): 50–57.
Maw, G. (1886) A Monograph of the Genus Crocus. Dulau & co., London, 54 pp.
Menemen, Y., Aytaç, Z. & Kandemir, A. (2013) Türkçe bilimsel bitki adları yönergesi. Bağbahçe Dergisi 47: 28–31. [in Turkish]
Rukšâns, J. (2015) Some New Crocus Taxa (Iridaceae) from Western Turkey and East Aegean Islands,  64:
1–38.
Rukšāns, J. (2010) Crocuses. A complete guide to the genus. Timber Press, Portland, 157 pp.
Rukšāns, J. (2017) . The Latvian Academy of Sciences, Riga, 442 pp.
Thiers, B. (2019) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual
Herbarium. Available from: http://sweetgum.nybg.org/ih/ (accessed 12 September 2019)
Wahlenberg, G. (1828) Gewächse im Morgenland gesammelt von J. Berggren, und näher bestimmt von Goran Wahlenberg. Isis (Oken)
21: 972–1006.
Weston, R. (1771) Botanicus Universalis et Hortulanus 2: 237. [pp. 384]
Whittall, E. (1889) Notes from Smyrna.  35: 473.

NOTES ON CROCUS L. SERIES FLAVI MATHEW Phytotaxa 438 (2) © 2020 Magnolia Press • 79
Appendix 1. This appendix lists the source of each sequence used for our phylogenetic analysis and its EMBL/NCBI
accession number (taxon, herbarium vouchers, collection number, origin, EMBL accession nrITS region. Missing data
is indicated with “-“.
Crocus adanensis T.Baytop & B.Mathew, GAT7148, -, Turkey,HE663988, Crocus aleppicus Baker, IABH18357,
-, Jordan, HE801175; Crocus almehensis C.D.Brickell & B.Mathew, TARI69170, -, Iran, HE801162; Crocus
antalyensis B.Mathew, HKEP9511, GAT7237, Turkey, HE664015; Crocus antalyensis, HKEP1006a, GAT7186,
Turkey, HE663990; Crocus asymmetricus Erol, ISTF 41370, Crocus sp. DH-2020, Turkey, LR761644; Crocus
candidus E.D.Clarke, GAT7137, HKEP 0907, Turkey, HE66398; C. flavus Weston, GAT7140, HKEP0915, Turkey,
HE663984; Crocus fleischeri Baker, GAT7139, HKEP0914, Turkey, HE663983; Crocus graveolens Boiss. & Reut.,
GAT7457, HKEP1111, Turkey, HE664010; Crocus hyemalis Boiss. & C.I.Blanche, GAT7118, -, Jordan ,HE801057;
Crocus olivieri J.Gay, GAT23020, HKEP1342a, Serbia, LR778179, Crocus vitellinus Wahlenb., -, -,Turkey, LR778178;
Crocus paschei Kerndorff,-,-, Turkey, LR778177; Crocus tournefortii J.Gay, GAT7202, -, Greece, HE801123;
Appendix 2. Voucher numbers of herbarium specimens used for the morphological investigations.
C. adanensis K802478, ISTE23969, ISTE12468, ISTE95577, GAT7148; C. antalyensioides ISTF41127, GAT56933,
GB0152368; C. antalyensis ISTF40145, ISTF40143, ISTF40142, ISTF40168, ISTF40079, ISTF41127, ISTF41126,
ISTF41117, ISTF41116, ISTF41113, ISTF41109, ISTF41105, ISTF41103, ISTF41144, ISTF41149, ISTF41141,
K802490, E333389, E333390, ISTE36684, ISTE31337, ISTE31340, ISTE9825, ISTE9826, ISTE60001, ISTE12482,
ISTE23980, ISTE21323, ISTE93252, ISTE93289, ISTE99292, GAT7186, GAT7237, GAT7458; C. balansae
K802441, K802440, K802442, ISTF41122, E333325, E333326, E333327, E333328, E333329, E333330, ISTE21342,
ISTE95469, ISTE93253, ISTE19651, ISTE36513, ISTE38935, ISTE23806; C. candidus ISTF41161, E346204,
E346205, E333324, ISTE79127, ISTE65012, ISTE62669, ISTE64999, ISTE65002, ISTE64993, ISTE106596,
ISTE23812, ISTE79119, ISTE65009, ISTE65015, ISTE99290, ISTE96004, GAT7137; C. flavus ISTF41130,
ISTF41128, ISTF41151, ISTF41150, K802444, K802400, B100151920, B100151921, B100151922, B100151923,
B100210241, B100510434, GAT23031, GAT23032, GAT23034, GAT31633, GAT33768, GAT33769, GAT33775,
GAT33780, GAT33781, ISTE27221, ISTE27225, ISTE27239, ISTE27281, ISTE27286, ISTE27294, ISTE27296,
ISTE27212, ISTE27214, ISTE27217, ISTE95468, ISTE99247, ISTE21333, ISTE108652, ISTE108651, ISTE10804,
ISTE31535, ISTE23990, ISTE10667, ISTE10672, ISTE95467, ISTE31341, ISTE31346, ISTE31352, ISTE31306,
ISTE31316, ISTE23750, ISTE23807, ISTE23808, ISTE23815, ISTE23816, ISTE23817, ISTE27257, ISTE31320,
ISTE96668, ISTE99289, ISTE60018, ISTE66649, ISTE65008, ISTE10790, ISTE12446, ISTE21283, ISTE21289,
ISTE21321, ISTE34806, ISTE3482, ISTE3498, ISTE66248, ISTE80850, ISTE82998, ISTE83516, ISTE96168,
ISTE99266; C. graveolens E333392, E333393, E333394, E333395, E333396, E333397, E333398, E333399,
E333400, E333401, E333402, E333403, E333404, E333405, E333406, E333407, E333408, ISTE96002, ISTE99287,
ISTE99237, ISTE99240, ISTE99250, ISTE99248, ISTE96031, ISTE78351, ISTE12460, ISTE12469, ISTE4173,
ISTE65127, ISTE93255, GAT7144, GAT7256, GAT7457, GAT30297, GAT56985; C. hyemalis K318284, K802431,
K802425, K802428, K318285, K802429, K802426, K802430, K802427, E333294, E333295, E333297, E333298,
E333299, E333300, E333301, E333302, E333303, GAT7218; C. istanbulensis K802399, ISTE46030, ISTE69709,
ISTE99254, ISTE96169, GAT7399; C. mouradii ISTF36901, K400300, ISTE23390; C. olivieri ISTF34678,
ISTF41119, ISTF41120, ISTF41287, K802443, K341684, K99742, E333311, E333310, E333312, E333313, E333314,
E333315, E333316, E333317, E333318, E333319, E333320, E333321, E333322, E333323, ISTE69768, ISTE1162,
ISTE1365, ISTE10247, ISTE21255, ISTE21313, ISTE21326, ISTE99249, ISTE99291, ISTE99279, ISTE99246,
ISTE96024, ISTE95484, ISTE95479, ISTE96669, ISTE95478, ISTE21364, ISTE21370, ISTE27231, ISTE24197,
ISTE25002, ISTE25979, ISTE27251, ISTE31556, ISTE34411, ISTE34416, ISTE34560, ISTE34573, ISTE48464,
ISTE50018, ISTE52903, ISTE66244, ISTE66255, ISTE7919, ISTE80846, ISTE88431, ISTE95817, GAT7135,
GAT23020, GAT33772, GAT47240; C. paschei ISTF41083, ISTF41085, ISTF41086; C. sarichinarensis ISTF41130;
C. vitellinus ISTF13965, ISTF13974, ISTF41094, ISTF1443, ISTF1444, ISTF1469, ISTF1470, ISTF12118, ISTF820,
K802437, K802439, K802436, K802438, E333309, B100355309, B100355310, B100355311, ISTE99282, ISTE99283,
ISTE77725, ISTE77731, ISTE5129, ISTE10243, ISTE23824, ISTE23825, ISTE23828, ISTE23967, ISTE23972,
ISTE23981, ISTE23982, ISTE23989, ISTE31390, ISTE31400, ISTE5788, ISTE23983;