Plant Syst. Evol. 248: 143–169 (2004)
DOI 10.1007/s00606-004-0171-x
A re-evaluation of morphological characters in European Gentianella
section Gentianella (Gentianaceae)
J. Greimler1, B. Hermanowski2, and C.-G. Jang1
1
Department of Systematics and Evolution of Higher Plants, Institute of Botany, University
of Vienna, Vienna, Austria
2
Department of Ultrastructure Research and Palynology, Institute of Botany, University
of Vienna, Vienna, Austria
Received September 8, 2003; accepted March 9, 2004
Published online: July 20, 2004
Springer-Verlag 2004
Abstract. Morphological traits were investigated in
Gentianella section Gentianella by morphometrics
and Scanning Electronic Microscopy (SEM). Variation in vegetative measures and in calyx, corolla,
ovary, and gynophor was analyzed in populational
samples and herbarium material. Special emphasis
was given to micro-morphology and variation of
papillae on the calyx using SEM. Three types of
papillae on the calyx lobes were found: (A) short
conical (G. amarella-group, G. insubrica, G. germanica); (B) long conical often curved (G. campestris,
G. anisodonta, G. engadinsesis, and G. liburnica; (C)
long cylindrical (G. aspera and G. pilosa).
G. austriaca, G. caucasea, G. crispata, G. fatrae,
and G. ramosa usually lack any papillae. Plants
without and with short conical papillae were found
in G. bulgarica and G. lutescens as well as in the
intermediate taxa G. bohemica and G. stiriaca. The
different types of papillae together with other calyx
characters (sinus, shape and margin of lobe) are of
high systematic importance and provide more stable
characters than morphometric flower measures.
Principal component and correlation analysis revealed a strong response of nearly all morphometric
traits to the environmental variable altitude. Adaptive and historical causes of morphological variation as well as taxonomical consequences are
discussed and a determination key is provided.
Key words: Gentianella section Gentianella, calyx,
papillae, morphology, SEM, morphometrics,
determination key.
The genus Gentianella Moench comprising ca.
260 species (von Hagen and Kadereit 2001) is a
member of the large subtribe Swertiinae of the
primarily temperate-alpine Gentianeae (Struwe
et al. 2002). Two taxa, segregated from
Gentianella on the basis of morphological
characters, Gentianopsis (Ma 1951) and
Comastoma (Toyokuni 1961), are clearly confirmed by molecular data (Yuan and Küpfer
1995, von Hagen and Kadereit 2001) as genera
that do not belong to the monophyletic uninectariate and biennial (occasionally annual)
Gentianella s. str. With one exception (G. aurea)
all European taxa of Gentianella s. str. are
members of the fimbriate group which corresponds to Gentiana sect. Endotricha (Wettstein
1896) or Gentianella section Gentianella
(Pritchard and Tutin 1972).
Section Gentianella comprises 16 species
plus five uncertain taxa (because they are
weakly differentiated narrow endemics or
intermediates or recently not observed) in
144
J. Greimler et al.: Morphology in European Gentianella
Europe (Appendix 1) according to taxonomy
applied in Pritchard and Tutin (1972) which is
based essentially on the framework given by
Kerner and Kerner (1882), and Wettstein
(1892, 1896). Geographical variants raised to
specific rank: G. fatrae (Holub 1983) and G.
stiriaca (Maurer 1998) we treat as separate
entities as well as G. germanica samples from
the Alps (= G. rhaetica, Kerner and Kerner
1882) and outside. Except three taxa all with
unique calyx morphology, G. caucasea (falcate
calyx lobes), G. crispata (crispate calyx lobes)
and G. columnae (4 calyx lobes, larger not
enclosing smaller ones), all taxa of the section
can be assigned to one of the three groups of
G. amarella, G. campestris and G. germanica.
High character differentation and species diversity within the G. germanica group (large 5merous flowers, mostly stalked ovary, with 9
species plus 4 intermediate or uncertain taxa) is
contrasted by little differentiation in the groups
of G. amarella (small 5-merous flowers, sessile
ovary, with 3 species) and G. campestris (large
4-merous flowers, two larger enclosing smaller
ones, with 2 species plus 1 uncertain taxon).
However, taxonomy in these two groups has
been questioned in recent studies (Renobales
Scheifler 2003, Winfield et al. 2003).
High morphological variation essentially in
vegetative characters within and among species
in section Gentianella is complicated by seasonal dimorphism (Wettstein 1896) or ecotypic
polymorphism (Zopfi 1991). The high variation in these characters being more or less
correlated with flowering time and ‘‘the intersection of these morphological gradients with
other more definite characters’’ (Pritchard and
Tutin 1972, p. 64) is a major reason for the
confusing taxonomy in European floras. As
can be judged from the investigated herbaria
populations of the early flowering variants
(aestival forms) probably always have been
rare in Middle Europe at least in the Alps and
close to them. Certainly these populations
have declined rapidly in the last decades (Zopfi
1991, Rosenbauer 1996, Skalicky pers. com.,
Kirschner pers. com.) apparently due to
impact from human activities.
Complications from seasonal dimorphism
or ecotypic polymorphism in vegetative characters are usually critical in delimitation of
intraspecific taxa. High polymorphism,
however, is also found in floral characters.
Problems of specific delimitation can be summarized as follows: (1) The exact distribution
of many species in Europe is yet unclear as
characters used for taxonomic limitation show
high intraspecific variation and little differentiation among taxa (Holub 1983, Dostal 1989,
Zopfi 1991, Petanidou et al. 1998). (2) Some of
the endemics (including ‘‘seasonal vicariant
taxa’’) that have been described (Samuelsson
1922, Kunz 1940, Ritter-Studnicka 1955,
Mayer 1969, Skalicky 1969, Holub 1983,
Renobales et al. 2002) exhibit only minor
differences from their closest relatives.
(3) There is confusion about specific characters namely, ciliae and papillae on the calyx
(e.g. Hess et al. 1972, Pritchard and Tutin
1972, Pignatti 1983).
In this study we investigate variation and
differentation of morphological features in
European Gentianella section Gentianella.
Applying a nested sampling with two datasets
(populations and taxa) we focus on: (1) variation in specific characters of the flower: calyx,
corolla, gynophor; (2) micro-morphology and
variation of papillae located on the calyx; (3)
relationships among morphological traits and
environmental/geographical variables; and (4)
the systematic relevance of these characters.
Materials and methods
Plant material. Population samples of Gentianella
section Gentianella were taken during field excursions between 1996 and 2003. Details on the 75
samples (including accessions provided by other
collectors) and sites are given in Appendix 2.
Specimens are kept in WU. Variation of critical
characters was also studied in specimens from the
herbaria GZU, LI, M, NMGW, PRC, RUEB,
TSB, UDM, W, and WU to complete results from
our population samples and to overcome bias
caused by the preponderance of central European
populations in our sampling. Nomenclature follows
Pritchard and Tutin (1972) except for taxa raised to
J. Greimler et al.: Morphology in European Gentianella
specific rank since then (Holub 1983, Maurer 1998)
and the two taxa within G. germanica (Kerner and
Kerner 1882).
Preparation. Collections made on the field
trips were immediately prepared for morphological
analysis. Flat preparations of calyx and corolla
were dried with the vouchers. In the lab these
preparations and vouchers were photographed
using the digital equipment Olympus E-10. Photographs have been modified in Adobe-Photoshop
6.0 and then displayed directly or used as templates
for drawings of morphological details.
Scannning Electron Microscopy (SEM).
Freshly collected flowers were preserved directly
in FAA (formaldehyde : acetic acid : distilled water:
70% ethanol; 2 : 1 : 7 : 10) or in 70 % ethanol which
was later replaced by FAA. Calices were prepared
for analysis of epidermal structures with SEM: (A)
Calyx lobes of flowers fixed in FAA were cut off,
transferred to fresh FAA and cleaned in the
Ultrasonic Cleaner B-32 (Branson) for max. 45
minutes. (B) The cleaned lobes were dehydrated in
formaldehyde dimethyl acetal (FDA) and then
critical-point-dried in the CPD 030 Critical Point
Dryer (Balzers). The lobes of calices fixed in 70%
ethanol were transferred to fresh 70% ethanol and
cleaned in an ultrasonic cleaner. Then they were
dehydrated in acetone for 24 hours and criticalpoint-dried. The dried lobes were sputtered with
gold for 5 minutes. Adaxial (C) and abaxial (D)
surfaces of lobes were examined with the Jeol JSM35CF Scanning Microscope and digitally documented using the Digital Image Processing System
Version 2.4.5.8. (Point electronic GmbH, Halle/
Saale, Germany). Digital images were reworked in
Adobe Photoshop 6.0.
Morphometry, statistics. Morphometric data
(counts, lengths) were gathered from 75 population
samples by measuring plant size, number of
internodes (above the rosette-like agglomerated
leaves at the stem base), number of flowers, pedicel,
total calyx (base of tube to apex longest lobe), calyx
tube, total corolla, corolla tube, gynophor
(= carpophor, the stalk of the ovary), and ovary.
Measures of pedicel and floral traits in full anthesis
were taken either from terminal flowers of the main
stem or from the subterminal nodes (which are
most similar in all dimensions) when the former
was missing or not accessible (occasionally in
attached vouchers). Between four and 14 individuals per population were measured (single charac-
145
ters occasionally could be measured in only two or
three individuals). From these data the individual
lengths of calyx lobes and corolla lobes and the
ratio lobe/tube for calyx and corolla, corolla/calyx,
and pedicel/corolla were calculated. Data distribution and total character variation were investigated
using the software SPSS 10: histograms (normal
distribution fit) and box-plots whereby a box
includes 50% of cases and median (horizontal
line), bar ranges from smallest to largest observed
value, that is not an outlier (more than 1.5 box
lengths from 25th or 75th percentile) or extreme
(more than three box lengths from 25th or 75th
percentile). Median, quartiles, minimum and maximum, were calculated for each of the 75 population samples for data set 1. Mean, standard
deviation, minimum, maximum were calculated
from all samples per taxon for data set 2, i. e. the
75 population samples plus additional data (2–3
individuals/voucher) from herbaria collected for
the above mentioned characters except number of
flowers, ratio corolla lobe/tube and ovary length,
which turned out negligible after analysis of data
set 1.
Four qualitative characters were scored on an
ordinal scale: (i) Calyx papillae: 1 absent, 2 shortconical, 3 long-conical, 4 long-cylindrical (scored
with a 10–16 fold lense); (ii) Calyx sinus between
two lobes: 1 obtuse, 2 acute; (iii) Shape of calyx
lobes: 1 linear, 2 lanceolate; (iv) Margin of calyx
lobes: 1 flat, 2 slightly revolute, 3 strongly revolute
(so that the margins, [nearly] touch each other
[Fig. 6d]). Measuring the proper lobes is important
in samples with very unequal ones as there is a
size hierarchy among them as can bee seen also in
± equal (= subequal) ones ( Fig. 1a, b). Calyx
sinus and shape was scored between/on the smaller
(inner) lobes: sinus between lobe 3 and 5 (not
always feasable in attached herbarium specimens)
and shape on lobe 3, 4 or 5; margins on the larger
(outer) lobes 1 or 2. Means calculated from the
individual assignments were used as population
scores in subsequent multivariate analyses (dataset
1). For the taxon samples (dataset 2) the percentage
of occurrence of each character (character state)
was calculated.
One way ANOVA (analysis of variance) was
applied on population samples to calculate variance components within and among populations
in each of the larger samples (N = 5 or more populations) i. e. G. anisodonta, G. aspera, G. austriaca,
146
J. Greimler et al.: Morphology in European Gentianella
Fig. 1. Calyx and corolla (flat
preparations). Basic calyx
types: a G. campestris, b
G. germanica. c Corolla: f =
fimbriae,
g = gynophor;
bar = 1 cm
G. germanica (Alps), and G. stiriaca. Principal
components analyses (PCA with correlation
matrix, unrotated solution and Varimax rotation)
were performed on the medians and scores of the
populations (labeled with taxon acronyms) using (i)
all samples with complete data (72 populations of
data set 1) and (ii) a subset (46 populations)
comprising the Alps and the adjacent north eastern
lowlands. Correlations among the medians were
investigated calculating the Pearson productmoment correlation coefficient. Number of flowers
and plant size were log-transformed prior to these
analyses. Altitude as a surrogate of environmental
variables and geographical longitude, latitude
(potential indicators of biogeographic differentiation) were included in these analyses as well as in a
final PCA of the subsample with the characters as
OTUs. For statistical analyses the software Splus
6.1 and SPSS 10 was used.
Results
Vegetative characters. Plant size, number of
internodes and number of flowers showed
higher variation among populations than within in all taxa sampled across a wide range of
habitats and elevations. ANOVAs were significant (F = 3.9 to 64.7; p < 0.005) in all cases.
This causes high variation within species
(Table 1). For length of pedicel we found
significantly higher variation among populations (F = 3.5; p < 0.005) only in G. germanica (Alps). Number of flowers ranged from 1
to 175 (n = 591) with an extreme left-sided
distribution [median 8 (quartiles 5/16)] caused
by a similar pattern in nearly all taxa. Investigating patterns of variation among taxa
revealed (Table 1): (1) Small plant sizes combined with low variation in G. bulgarica and
G. liburnica. (2) Lower numbers of internodes
in G. uliginosa than in G. amarella. (3a)
Shortest pedicels in G. liburnica, however, to a
high proportion (ca. 50%) within the lower
range of G. anisodonta and G. engadinensis. (3b)
An overlapping range of variation in length of
pedicel for G. bulgarica and G. ramosa.
Calyx. Two basic calyx types are shown in
Fig. 1a,b. The tetramerous type A with flat
lobes is restricted to G. campestris: two ovoidlanceolate larger sepals of equal size at least
partly cover two smaller narrowly lanceolate
or nearly linear ones (Fig. 1a) and G. columnae: two larger triangular-lanceolate subequal
sepals usually not covering the smaller narrowly lanceolate or nearly linear ones. The
reduced number of sepals, although rarely, can
be found in single flowers of most usually fivemerous taxa. Within the pentamerous type,
which is typical for the G. germanica group
J. Greimler et al.: Morphology in European Gentianella
147
Table 1. Metric data (min.-max., mean, standard deviation, number of individuals) for length of calyx,
pedicel, gynophore (mm), stem (cm); ratios of pedicel/corolla and calyx/corolla; number of internodes
Taxa
Calyx
Pedicel
R Pe/Co
R Ca/Co Gynophor
Stem
N
Internodes
amare
7–15
10.3±1.8
35
8–20
13.6±3.0
35
13–27
19.9±3.4
34
10–36
19.5±4.4
77
11–20
15.9±2.4
26
11–23
16.3±3.1
40
10–25
16.4±3.6
29
11–21
15.4±2.7
12
9–19
13.4±2.5
24
8–15
11.0±2.0
28
12–26
18.4±3.8
14
11–24
15.9±2.8
75
6–21
12.7±2.7
54
12–27
18.4±4.8
22
6–10
8.0±1.3
7
5–32
12.6±5.4
22
2–25
10.0±5.5
25
13–43
23.4±8.2
26
5–25
16.0±4.8
35
6–23
14.2±5.4
17
3–23
10.0±6.2
22
6–30
17.2±6.8
23
5–40
19.1±9.1
13
5–25
16.3±7.7
9
3–15
6.7±3.0
17
3–23
12.3±6.1
12
6–25
14.1±5.6
48
7–35
16.4±7.5
27
7–29
17.5±7.0
12
1–5
3.0±1.4
7
0.3–2.0
0.7±0.3
22
0.2–1.0
0.4±0.2
24
0.4–1.2
0.7±0.2
26
0.2–0.8
0.5±0.2
35
0.2–1.1
0.5±0.2
17
0.1–1.0
0.4±0.3
20
0.3–1.1
0.6±0.2
23
0.2–1.7
0.8±0.4
10
0.3–1.1
0.7±0.3
9
0.2–0.8
0.4±0.2
17
0.1–0.8
0.4±0.2
11
0.2–1.0
0.5±0.2
48
0.2–0.9
0.6±0.2
23
0.3–1.2
0.7±0.3
11
0.1–0.3
0.2±0.1
7
1.2–1.9
1.7±0.2
22
1.3–2.3
1.8±0.3
24
1.5–2.3
1.8±0.2
26
1.2–2.6
1.7±0.3
38
1.3–2.1
1.8±0.2
17
0.9–1.8
1.5±0.2
28
1.3–2.2
1.7±0.2
28
1.3–2.0
1.7±0.3
10
1.3–1.7
1.4±0.1
10
1.3–2.5
1.6±0.4
18
1.4–2.3
1.8±0.3
12
1.1–2.3
1.7±0.2
48
1.6–3.5
2.2±0.4
44
1.0–1.9
1.5±0.3
20
1.6–2.2
1.9±0.2
7
9–43
25.3±10.3
21
2–23
7.6±4.5
67
8–53
22.8±10.8
48
8–40
19.3±8.2
78
9–51
20.3±9.0
22
3–10
5.7±2.0
30
5–25
13.5±4.5
34
9–22
16.2±3.8
15
3–14
8.6±4.1
16
2–18
5.6±3.6
48
7–27
14.9±4.9
15
4–41
13.0±7.2
88
4–44
17.7±10.8
37
8–22
14.6±3.7
18
3–7
3.6±1.5
7
6–13
9.1±2.4
21
2–8
4.4±1.4
67
3–15
8.0±3.1
48
4–14
8.3±2.3
57
5–12
8.7±2.2
19
3–7
4.5±1.0
30
3–7
4.8±0.9
34
3–7
5.1±1.2
14
2–6
4.2±1.3
15
2–7
3.5±1.3
42
5–9
6.5±1.0
16
3–12
5.9±1.6
82
3–13
8.5±2.5
38
4–8
5.8±1.0
14
2–6
3.4±1.3
7
Min.–Max.
Mean±Std
N
aniso
Min.–Max.
Mean±Std
N
asper
Min.–Max.
Mean±Std
N
austri
Min.–Max.
Mean±Std
N
bohem Min.–Max.
Mean±Std
N
bulga
Min.–Max.
Mean±Std
N
camp
Min.–Max.
Mean±Std
N
caucas Min.–Max.
Mean±Std
N
colum
Min.–Max.
Mean±Std
N
engad
Min.–Max.
Mean±Std
N
fatrae
Min.–Max.
Mean±Std
N
germ_A Min.–Max.
Mean±Std
N
germ_O Min.–Max.
Mean±Std
N
insub
Min.–Max.
Mean±Std
N
liburn
Min.–Max.
Mean±Std
N
0–2
0.5±0.6
20
0–4
1.7±1.2
18
0–7
2.9±2.4
19
1–10
4.3±2.3
52
1–6
3.9±1.4
11
0–10
3.7±2.9
21
1–4
2.7±1.1
12
2–6
3.9±1.1
11
0–2
0.6±0.9
5
0–3
1.4±0.9
14
1–5
3.1±1.6
13
0–6
2.2±1.1
49
2–7
4.5±1.5
22
0–4
2.1±1.1
12
1–2
1.3±0.5
4
148
J. Greimler et al.: Morphology in European Gentianella
Table 1. (continued)
Taxa
lutesc
pilosa
ramos
styria
uligin
Total
Min.–Max.
Mean±Std
N
Min.–Max.
Mean±Std
N
Min.–Max.
Mean±Std
N
Min.–Max.
Mean±Std
N
Min.–Max.
Mean±Std
N
Min.–Max.
Mean±Std
N
Calyx
Pedicel
R Pe/Co
R Ca/Co Gynophor
Stem
N
Internodes
9–31
16.8±6.2
25
12–24
18.2±3.4
28
9–17
12.8±2.4
25
13–23
18.2±2.7
51
10–18
14.0±2.3
16
6–36
15.8±4.4
657
5–22
13.0±5.7
14
5–19
11.5±4.3
20
2–15
5.8±3.2
24
5–30
17.0±5.6
22
13–70
34.5±20.6
15
1–70
14.6±9.0
410
0.2–1.0
0.5±0.2
14
0.2–0.7
0.4±0.1
20
0.1–0.7
0.3±0.1
24
0.2–0.9
0.5±0.1
21
0.7–4.6
2.2±1.3
15
0.1–4.6
0.6±0.5
397
1.2–2.2
1.6±0.3
14
1.3–2.1
1.7±0.2
21
1.3–1.8
1.5±0.2
22
1.5–2.5
1.9±0.3
21
0.9–1.8
1.2±0.3
16
0.9–3.5
1.7±0.3
446
7–35
21.9±7.7
24
6–39
19.4±9.1
29
4–16
9.6±3.1
29
7–36
16.5±6.6
43
3–16
9.6±3.7
16
2–53
14.4±9.0
685
4–10
6.9±1.5
23
4–11
7.4±2.2
28
4–7
5.3±1.1
29
4–10
6.7±1.4
46
2–4
2.7±0.7
15
2–15
6.2±2.5
645
and G. amarella group (Fig. 1b: G. germanica,
Alps) there is a great variation concerning
proportions, margins, sinuses between adjacent lobes (Table 2), and the presence and
shape of papillae on the margin and midrib of
the lobes (Table 3). As a rule the sepals
decrease in size according to a 2/5 spiral. The
larger (outer) sepals occasionally have ‘‘wings’’
in their connate part forming the calyx tube.
Within the basic type B two calyx types
are unique: (1) crispate calyx lobes with often
blackish margins in Gentianella crispata; (2)
falcate and very delicate, narrowly linear lobes
and the tube often split down on one side in
G. caucasea. In the remaining taxa belonging to
the G. germanica group and G. amarella group
three major calyx types are found (Fig. 2):
(1) Lobes triangular-lanceolate to ovoidlanceolate, partly enfolding the inner ones,
extremly unequal in size, margins often conspicuously recurved (= lobe revolute), sinus
acute: G. anisodonta (Figs. 2a, 6d), G. engadinensis, and G. liburnica.
(2) Lobes (often narrowly) lanceolate, ±
equal in size, margins not or slightly recurved,
sinus acute: G. aspera (Fig. 2b), G. pilosa
0–4
2.7±1.5
15
2–5
3.5±1.0
11
1–5
3.1±1.1
15
1–8
4.5±1.8
32
0–1
0.5±0.6
4
0–10
3.0±2.0
360
(Fig. 2c), G. germanica, and G. ramosa
(Fig. 2d).
(3) Lobes linear, ± equal in size, margins
not or slightly recurved, sinus obtuse:
G. austriaca (Fig. 2e), G. bulgarica, G. fatrae
(Fig. 2f), G. lutescens (Fig. 2g), and often in
G. amarella (Fig. 2h).
However, in single individuals from populations of all taxa we found intermediate calyx
types due to occasionally high variation in the
characters sinus, lobe, margin. Such intermediate calyx types were often found in
G. bohemica, G. insubrica, and G. stiriaca but
also in G. amarella (Table 2).
We observed high metric variation of calyx
features: The ratio lobe/tube revealed significantly higher variation among populations
than within in all five largest taxon samples
(ANOVA, F = 2.6 to 7.8; p < 0.01). There is,
however, also high differentiation among taxa
(Fig. 3). Variation in total calyx length was
significantly higher among populations
(F = 3.2 to 8.2; p < 0.005) in G. aspera,
G. austriaca, and G. germanica (Alps).
Micromorphology of papillae on the calyx. In taxa with papillae, these occur on the
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24
40
70
30
61
34
5
38
42
22
100
71
29
18
82
11
16
85
46
37
48
15
27
43
57
68
32
29
36
18
100
97
3
100
46
78
31
17
57
26
23
52
48
92
8
72
28
49
51
53
36
11
89
18
100
10
90
29
91
9
23
52
48
31
86
14
78
6
94
47
100
63
81
19
Margin 1
2
3
N
N
69
31
42
Lobes
1
2
42
16
81
19
50
50
28
37
63
27
15
85
46
12
88
90
100
100
28
16
36
N
83
17
751
100
11
86
14
42
27
73
22
45
55
38
43
58
40
42
58
24
751
17
83
24
20
80
40
13
87
38
100
22
81
19
42
100
11
11
89
27
9
91
46
3
97
90
100
100
28
100
18
100
100
29
61
39
23
35
65
31
86
14
78
100
47
100
63
74
26
42
1
2
Sinus
St. amare aniso asper austri bohem bulga camp caucas colum engad fatrae germ_A germ_O insub liburn lutesc pilosa ramos styria uligin Tot.
Char.
Table 2. Discrete calyx character states (percent of observations given): Sinus between lobes: 1 obtuse, 2 acute; shape of lobes: 1 linear,
2 lanceolate; margin of lobes: 1 flat, 2 slightly revolute, 3 strongly revolute = margins (nearly) touch each other
J. Greimler et al.: Morphology in European Gentianella
149
margin of the leaves, gradually becoming more
pronounced towards the inflorescence. The
papillae are most conspicuous on the margins
of the calyx lobes from which they often run
down the tube along the fusion lines of the
sepals. They also occur (often decreased in
size) at least in part of the adaxial surface and
(in two taxa) also on their abaxial midrib or
the whole abaxial surface of the tube. Table 3
gives a quantiative summary about papillar
structures as shown in detail by the schematic
drawing (Fig. 4) and the SEM photographs
(Figs. 5–8):
We found three types of papillae:
(A) Short-conical (in herbario: short-triangular) papillae, about as long as broad (basal
diameter), sharply pointed, ± asymmetrical
and pointing slightly towards apex of sepal
were found in the G. amarella-group (Fig. 5ac), G. germanica (Fig. 5d-f), G. insubrica. As
the following types of papillae often become
shorter towards the apex, papillae approaching
type A could be found in all taxa near the very
apex.
(B) Long-conical (long-triangular), about
twice as long as broad, occasionally curved
papillae with sharp or rounded tip, ± pointing
towards apex were found typically in
G. campestris (Fig. 6a–c), G. anisodonta
(Fig. 6d–f), G. columnae, G. engadinensis, and
G. liburnica. These taxa, however, show some
variation in often having additionally (occasionally only) short type A papillae or also
longer papillae approaching type C but with a
broader base.
(C) These are long-cylindrical (long-linear)
with blunt or rounded tip, usually much more
than twice as long as broad, typically found in
G. pilosa (Fig. 7a–c) and G. aspera (Fig. 7d–f),
where they also occur (often shortened in G.
pilosa) on the abaxial midrib. However, in
both G. aspera occasionally and more often in
G. pilosa we found the midrib smooth and the
margins only short-papillose (approaching
types B and A) or nearly smooth, rarely
perfectly smooth.
Non-papillose taxa with smooth margins
of the calyx lobes (G. austriaca, Fig. 8a,b;
150
J. Greimler et al.: Morphology in European Gentianella
Table 3. Overview of occurrence and types of calyx papillae in European Gentianella sect. Gentianella.
Descriptions are based on freshly fixed flowers (see SEM-photographs) and herbarium observations by a
10–16 fold lense (in parentheses). N: number of individuals (1209 total) and percent of observed papillae
type per taxon (main type bold)
Taxon
N
Papillae
absent (%)
Papillae present (%)
(A) shortconical
(= shorttriangular)
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
G.
amarella
anisodonta
aspera
austriaca
bohemica
bulgarica
campestris
caucasea
columnae
crispata
engadinensis
fatrae
germanica Alps
germanica Out
insubrica
liburnica
lutescens
pilosa
ramosa
styriaca
uliginosa
108
66
195
98
46
39
51
15
24
30
38
16
109
107
20
11
25
36
88
44
29
11
98
37
36
89
9
15
2
63
64
27
Location
(B) longconical
(= longtriangular)
(C) longcylindrical
(=longlinear)
76
11
15
74
73
margin
lobe (%)
89
100
100
2
63
64
100
midrib
lobe (%)
84
100
87
88
11
3
60
3
98
18
20
25
13
8
12
89
95
100
18
40
19
2
82
80
G. fatrae, Fig. 8c; G. ramosa, Fig. 8d,e; G.
caucasea; G. crispata) occasionally have somewhat undulate-uneven (rarely very short papillose) margins towards the apex [the crispate
margin of G. crispata is a feature of the total
lobe and not of the epidermal cells].
High variation between the smooth and
short-papillose (type A) was found in G.
bohemica, G. bulgarica, G. lutescens, G. stiriaca
(Fig. 8f), and G. uliginosa .
Corolla and gynophor. The obconical or
nearly cylindrical corolla with four or five
lobes, which is fimbriate in the throat
(Fig. 1c), offers few characters: Corolla size,
color (could not be scored reliably from
herbaria) and the ratio lobe/tube (often difficult to see in old specimens). In corolla size
75
87
5
2
82
42
36
100
13
100
12
89
97
100
100
40
97
2
82
80
56
(Fig. 9) we found a common size range of
more than 75 % for the pairs G. engadinensis/
G. anisodonta and G. austriaca/G. lutescens.
This is due to high within species variation,
resulting from high differentiation among
populations (ANOVA for corolla size,
F = 8.1 to 13.8, p < 0.001) in G. anisodonta,
G. austriaca and G. germanica (Alps). The
ratio corolla lobe/tube calculated for the
population samples exhibited some geographical correlation (see below) due to high values
in G. stiriaca and clinal variation in
G. germanica towards the East and G. austriaca towards the West, respectively. Comparing
taxa, where this might be a useful additional
character, e.g. in G. aspera and G. anisodonta
we found a widely common range for this ratio
J. Greimler et al.: Morphology in European Gentianella
151
Fig. 2. Drawings after
flat preparations of the
calyx of (a) G. anisodonta (note the very
unequal
lobes),
(b)
G. aspera, (c) G. pilosa,
(d) G. ramosa – all with
acute sinuses; (e) G. austriaca, (f) G. fatrae, (g)
G. lutescens, (h) G. amarella – all with obtuse
sinuses; bar = 1 cm
5
Calyx, ratio lobe/tube
4
3
2
1
austri
bohem
bulga
camp
caucas
colum
14
67
41
19
23
26
22
47
16
uligin
asper
27
styria
24
ramos
12
pilosa
23
lutesc
30
insub
21
germ_O
77
germ_A
29
fatrae
29
engad
28
aniso
N=
amare
0
Taxa
Fig. 3. Variation in the ratio calyx lobe/tube within and among taxa (abbreviated epitheta) of Gentianella
section Gentianella. Horizontal line = median, circle = outlier, asterisk = extreme. germ_O: populations of
G. germanica outside the Alps; germ_A: in the Alps
152
J. Greimler et al.: Morphology in European Gentianella
Fig. 4. Schematic drawings of papillae in the middle
of the calyx lobes (abaxial view) with approximate
size relations among the three types: (a) short-conical,
about as long as broad, sharply pointed, ± asymmetrical and pointing slightly towards apex of sepal;
(b) long-conical, about twice as long as broad, often
curved with sharp or rounded tip, ± pointing
towards apex; (c) long-cylindrical (long-linear) with
blunted or rounded tip, usually more than twice as
long as broad. For exact size measures compare Figs.
5–7
(> 50%) in a boxplot view (not shown) despite
a statistically significant difference (t-test, p <
0.001) in the means.
High variation in length of gynophor was
found in nearly all taxa. Again ANOVA
revealed significantly higher varition among
than within populations (F = 2.1 to 31.1; p <
0.05) in the large samples except in G. stiriaca.
Sessile ovaries are not confined to the
G. amarella-group and G. engadinensis. We
found them in flowers of seven other
taxa (Table 1). On the other hand, gynophor
length in rare cases reaches up to 2 mm in
G. amarella and occasionally up to 3 mm in
G. engadinensis.
Correlations, PCA. Correlation analyses
performed on a subsample of dataset 1 (46
populations of the Alps and adjacent areas)
revealed the high importance of the environmental variable altitude which explained 57%
of total variation in number of internodes
(r = )0.758, p < 0.001), 36% in plant size
(r = )0.602, p < 0.001), and 17% in number
of flowers (r = )0.407, p = 0.005). Altitude
also explained high proportions of variation
in floral characters: 20% in calyx length
(r = )0.445, p = 0.002), 38% in corolla
length (r = )0.614, p < 0.001), 36% in ovary
length (r = )0.598, p < 0.001), and 29% in
gynophor length (r = )0.482, p = 0.001).
Both geographical variables longitude and
latitude (which were inversely related to altitude due to the west-northeast sampling along
the Alps) showed high positive correlations
with the above variables. For the ratio corolla
lobe/tube, which was not significantly associated with altitude we also found high correlations with both, longitude (r = 0.559, p <
0.001) and latitude (r = 0.515, p < 0.001).
Vegetative characters were highly correlated
with floral characters: number of internodes
with corolla length (r = 0.543, p < 0.001),
ovary length (r = 0.543, p < 0.001), and
gynophor length (r = 0.434, p = 0.003);
plant size with corolla length (r = 0.588,
p < 0.001), ovary length (r = 0.549,
p < 0.001), and gynophor length (r = 0.413,
p = 0.004).
A principle components analysis on all
populations of data set 1 did not reveal any
groups (not shown). Similar results were
obtained with the subsample as used for
correlation analysis (Varimax rotated solution
shown in Fig. 10a). Therefore the characters
were ordinated in another PCA together with
altitude of the sampling site and geographical
coordinates. This unrotated PCA underlined
the high importance of altitude (Fig.10b) in
J. Greimler et al.: Morphology in European Gentianella
153
Fig. 5. Short-conical
(type A) papillae (SEM
photographs of calyx
lobes, abaxial): G. amarella: (a) surface, (b)
margin, (c) midrib;
G. germanica: (d) surface,
(e) margin, (f) midrib.
Bar: 100 lm
constituting the first component, which explained 40% of the total variance. The correlations reported above were essentially
displayed on this first axis. Together with the
second (14%) and third component (11%)
65% of the total variance was explained.
Patterns, however, along the other axes did
not provide additional insights except demonstrating the high association of the ratio
corolla lobe/tube with geographical variables
in all dimensions.
As neither PCA with all characters
(Fig. 10a) nor ordination of selected metric
characters (Fig. 10c) did reveal groups in the
data we tested the ordinal characters alone.
The population scores of these characters
(character states see Table 2 and 3) did reveal
groups in the data. The best resolution was
obtained by including (1) papillae type, (2)
shape of lobe, and (3) margin. (Fig. 10d).
Discussion
Character variation. Focusing on micromorphology of epidermal structures on the calyx
revealed that there are three different types of
papillae and only few taxa with no papillae
(Figs. 4–8). The calyx lobes where they occur
have been discribed in the past by so many
subtle terms, such as ‘‘smooth, glabrous,
(somewhat) scabrid, (strongly) ciliate, shortly
hirsute, papillose, papillose-hirsute’’ (Pritchard
154
J. Greimler et al.: Morphology in European Gentianella
Fig. 6. Long-conical
(type B) papillae (SEM
photographs of calyx
lobes, abaxial): G. campestris: (a) surface, (b)
margin, (c) midrib;
G. anisodonta: (d) surface, (e) margin, (f) midrib. Bar: a, d 1 mm; b, c,
e, f 100 lm
and Tutin 1972 p. 63 ff) or ‘‘glatt, schwachrauh, knotig-rauh, kurz-rauhhaarig, rauh-gewimpert’’ (Jäger and Werner 2002 pp.
515–516). We propose a more straightforward
terminology by distinguishing between smooth
and papillose and defining the three types of
papillae and their occurence (Fig. 4, Table 3).
By this we hope to provide a more efficient tool
for identification and help reduce confusion
about these characters in many European
determination keys.
Calyx features (number of sepals, shapes,
sinuses between the lobes, ratio lobe/tube,
presence/absence of papillae) have always been
central diagnostic characters in section Gentianella since Kerner and Kerner (1882) and
Wettstein (1892, 1896). A biogeographical
pattern in these characters as suggested by
Wettstein (1896) and Skalicky (1969) is obvious in the G. germanica group (including
G. caucasea) although G. bulgarica and G.
ramosa do not fit well into this: (1) Smooth or
only shortly papillose often linear lobes and
obtuse calyx sinuses are typical for taxa of
eastern and south-eastern Europe (incl. the
Caucasus region). The very delicate calyx tubes
often split down on one side in G. caucasea are
occasionally found also in G. austriaca,
G. fatrae, and G. lutescens. (2) Long conical
papillae, combined with acute sinuses and
lanceolate lobes are found in the southern
Alps and Dinarids (G. anisodonta, G. engadin-
J. Greimler et al.: Morphology in European Gentianella
155
Fig. 7. Long-cylindrical
(type C) papillae (SEM
photographs of calyx
lobes, abaxial): G. pilosa
(a) surface, (b) margin,
(c) midrib. G. aspera:
(d) surface, (e) margin,
(f) midrib; Bar: d 1 mm;
a, b, c, e, f 100 lm
ensis, G. liburnica). (3) Long cylindrical papillae combined with lanceolate lobes and acute
sinuses are found only in a small area of the
southern eastern (G. pilosa) and along the
northern Alps extending to adjacent central
Europe (G. aspera). (4) Short–conical papillae
are typical for G. germanica of central and
north-western Europe and the southern Alps
narrow endemic G. insubrica.
These patterns of calyx variation, however,
are not always correlated with taxonomic
entities. Taxa with intermediate characters
such as G. bohemica (Skalicky 1969) and
G. stiriaca (Wettstein 1892, 1896; Hayek
1911–14; Maurer 1998) have been considered
transgressive. However, clinal variation can be
found in many Gentianella taxa. Such variation might be due to inter-/postglacial introgression events or even intergradation in
refugia during glaciation as supposed for
populations of Pritzelago alpina (Kropf et al.
2003). Molecular markers revealed high genetic similarities among populations across Gentianella taxa of the Eastern Alps (Jang and
Greimler, unpublished).
Characters of the corolla (color, size) and
gynophor length are of limited diagnostic
value although often used (Samuelsson 1922,
Mayer 1969, Pritchard and Tutin 1972). In
some cases unusual characters might point to
hybridization as e.g. the long gynophor found
in an Austrian population of G. amarella
156
J. Greimler et al.: Morphology in European Gentianella
Fig. 8. Smooth margins
(SEM photographs of
calyx lobes, abaxial): G.
austriaca: (a) surface,
(b) margin; G fatrae:
(c) margin; G. ramosa:
(d) surface, (e) margin;
G. styriaca: (f) margin.
Bar: 100 lm
(in this region sympatric with G. germanica).
Hybridization of G. amarella with taxa of the
G. germanica group is not uncommon (Pritchard 1961, Moravec and Vollrath 1967). We
have no explanation for the higher ratios lobe/
tube in the Nort-Eastern Alps in G. stiriaca
and populations of G. austriaca, G. germanica
and G. aspera occuring there. Floral similarities across taxa often reflect local adaptations
to attract similar pollinators as found in
orchids (Borba et al. 2002).
We found evidence that high proportions
of metric variation especially of vegetative
traits in Gentianella sect. Gentianella can be
explained by altitude used as a complex
environmental variable. In a detailed investi-
gation of G. austriaca (Greimler and Dobeš
2000) we also found high intraspecific variation in vegetative characters connected with
different habitats and different population
histories. This agrees with Zopfi (1991) who
favoured a model of ecotypic polymorphism in
contrast to Wettsteins (1896) simple seasonal
dimorphism model (aestival plants with fewer
and longer internodes compared to the autumnal plants). There are, however, clear patterns
of seasonal differentiation in populations outside the central European mountain ranges in
the G. amarella group and in G. campestris
(Lennartsson 1997, Rich et al. 1997). In the
central European mountain taxa of Gentianella, there are very few extant populations
J. Greimler et al.: Morphology in European Gentianella
157
60
50
Corolla length, mm
40
30
20
10
austri
bohem
bulga
camp
cauca
colum
crispa
13
96
102
21
8
21
23
23
47
23
uligin
asper
34
styria
16
ramos
21
pilosa
25
lutesc
30
liburn
26
insub
39
germ_O
71
germ_A
67
fatrae
42
engad
111
aniso
N=
amare
0
Taxa
Fig. 9. Variation in corolla length within and among taxa (abbreviated epitheta) of Gentianella section
Gentianella. Horizontal line = median, circle = outlier, asterisk = extreme. germ_O: populations of G.
germanica outside the Alps; germ_A: in the Alps
(Skalicky 1969 and pers. comm., Zopfi 1991,
Wagner and Mitterhofer 1998) showing seasonally and morphologically seperated cohorts.
Calyx features (sinus, shape, proportions)
as used traditionally are central for understanding systematic relationships in the section
and for distinguishing taxa looking beyond the
concept of Pritchard and Tutin (1972). Our
more detailed investigation of epidermal structures on the calyx, i.e. the presence/absence
and different shapes of papillae, provided
useful critical characters that so far have been
underestimated. As morphometric characters
of calyx, corolla and gynophor as well as
vegetative characters display a great withinspecies variation partly due to environmental
modification, such additional characters help
in systematic delimitation. However, all diagnostic characters can become less definitive
towards the edge of distribution of many
central European taxa. Morphologically this
has been interpreted as reflecting introgression
or perhaps ancient hybridization (Wettstein
1892; 1896; Hayek 1911–14; Skalicky 1969).
Conclusions towards a modified taxonomy. We here summarize various investigations and our observations on European
Gentianella section Gentianella with respect to
the determination key proposed below. Short
158
J. Greimler et al.: Morphology in European Gentianella
comments on taxa for which we have no
original data are given in Appendix 1.
We partly resolve the G. amarella group
in distinguishing between G. amarella and
G. uliginosa. However, due to some taxonomic
confusion and many missidentifications in the
G. amarella group we excluded many herbaria
samples of G. uliginosa which by far did not
agree with the pre-selective criteria as given in
Pritchard and Tutin (1972) and Holyoak
(1999). These characters have also been found
too variable for clearly identifying Dutch
populations of G. amarella and G. uliginosa
(Petanidou et al. 1998). Also we did not
observe the more outcurved calyx lobes (as in
British samples: Rich 1997, Holyoak 1999)
in Middle and Northern European samples of
G. uliginosa. Taking the characters given in
Pritchard and Tutin (1972) it is very difficult to
distinguish between G. uliginosa and G. anglica
due to the high variation of these characters
(upper leaves, calyx lobes) in what we consider
G. uliginosa. From the data given in Rich et al.
(1997), G. anglica well fits into the range of
G. uliginosa and therefore we include it in the
latter taxon.
Considering the poor genetic differentiation within the G. amarella-group (data in
Winfield et al. 2003) this rather reflects ecotypic variation of one species occuring with
various fully compatible variants in various
habitats than differentiation on the species
level (with subsequent hybridizations) as suggested by Rich et al. (1997) and Holyoak
(1999). Additionally there is some geographical differentiation. Because of high proportions of tetramerous flowers, often smooth
calyx margins and high variation in the gynophor length Spanish populations of G. ama-
rella were considered a different species
(G. hispanica) by Renobales et al. (2002).
However, recently (Renobales Scheifler 2003)
this taxon was included again in G. amarella.
G. engadinensis is a variant of G. anisodonta
with smaller and more reddish flowers and
nearly sessile ovaries as found by Samuelsson
(1922). However, there is some variation in
color within G. anisodonta. Moreover variants
with very small flowers and nearly sessile
ovaries we also found in other regions of the
large distribution area. Such small variants we
cannot distinguish from G. engadinensis, at
least not in the herbaria. G. liburnica, another
variant with small and dirty-violet flowers was
distinguished from G. engadinensis by more
narrow and acute leaves, smaller flowers and a
longer gynophor. Summarizing our observations and considering the diagnosis of
G. liburnica (Mayer 1969), none of these
characters holds. We cannot distinguish clearly
between all these variants of G. anisodonta and
therefore treat it as a group here. Apart from
problems of clearly characterizing all the
variants, this group requires further investigation focusing on patterns of differentiation in
context with the many refugial areas (Tribsch
and Schönswetter 2003) in the Southern Alps
and Illyric mountain ranges and possible
hybridization zones.
G. aspera shows some clinal variation. In
the Bavarian Prealps the taxon often lacks
long papillae in contrast to the densely longpapillose populations of the Alps. Towards the
western and northern border of its distribution
area this might also indicate introgression by
G. germanica. Samples of such probably intermediate variants were not included in the
statistics and key.
c
Fig. 10. (a) PCA 1 (varimax rotated) ordinating populations (labeled with first two characters of epitheta) does
not reveal groups. (b) PCA 2 shows (i) high negative relationships between the environmental variable altitude
(alt) and metric variables: lengths of calyx (cx), corolla (co), gynophor (gy), ovary (ov), plant size (sh); number of
internodes (ni) and flowers (nf); the ratios lobe/tube of calyx (rcx) and corolla (rco); and high association
between rco and the geographical variables longitude (lon) and latitude (lat). (c) Ordinating populations by
selected metric characters (gynophor, corolla, ratio lobe/tube of calyx) does not reveal structure. (d) Ordinating
populations by ordinal characters shows core groups in accordance with taxonomy despite high variation also
in these characters
J. Greimler et al.: Morphology in European Gentianella
159
160
J. Greimler et al.: Morphology in European Gentianella
G. austriaca includes G. fatrae as we
cannot distinguish the two entities by the
longer and narrower calyx lobes of the latter
(as given by Holub 1983) or by any other
character we have investigated. For the same
reason we have to include also G. lutescens.
This taxon, however, shows high variation in
all characters across its wide distribution area
and requires further investigation. For now
we include it into an informal G. austriacagroup.
In G. campestris there is no differentiation
of plants in the Pyrenees justifying a separate
species G. hypericifolia. Spanish authors have
included this taxon in G. campestris (Morales
et al. 1996, Renobales et al. 2002). According
to Renobales Scheifler (2003) the sepals in
these populations are mostly elliptical, ovate,
not obovate as given in Wettstein (1896: plate
4, fig. 2) and therefore not different from the
sepals in G. campestris.
We here propose to follow Kerner and
Kerner (1882) in assigning the alpine populations of G. germanica to a separate taxon
G. rhaetica although most differential characters given by the authors (relative leaf size and
shape, corolla size) are too variable to separate
the populations from the Alps and the regions
of northern Middle and Western Europe.
Another character given by the authors, the
length of gynophor in relation to calyx tube is
difficult to observe in herbaria. This character,
however, shows significant differentiation due
to the more delicate calyx and longer gynophor
in G. germanica s.str. as well as the total
relation of corolla to calyx (t-tests, all p <
0.001) which we scored additionally (Fig. 11).
Other characters as given in Lauber and
Wagner (2001): calyx lobes outcurved (some
regional differentiation within G. germanica
s.str.) and calyx winged do not show differentiation of taxonomic significance. As we have
evidence for high genetic differentiation between these two taxa (Jang et al., unpublished)
differentiation of all characters will be tested
again including further samples of other
regions of widespread G. germanica s.str. Due
to high intraspecific polymorphism, however,
morphometry often fails to provide clues for
taxonomy despite genetic differentiation (Woo
et al. 2002).
Finally, we have to deal with two obvious
intermediate taxa and one taxon combining
characters of other taxa. G. bohemica is a
morphological and geographical intermediate
between G. austriaca and G. germanica (Skalicky 1969). Intermediate populations between
G. austriaca, G. aspera, and G. rhaetica in the
Eastern Alps have been assigned to G. stiriaca
on various taxonomic ranks (Wettstein 1892,
1896; Hayek 1911–14). Recently this taxon has
been raised to specific rank again (Maurer
Fig. 11. Variation in calyx length (white),
ratio corolla/calyx (light grey), gynophor
length (grey), and ratio gynophor/calyx tube
(bricks) within and between G. germanica
(Alps) (= G. rhaetica) and G. germanica s.str.
(Out); all lengths in cm
J. Greimler et al.: Morphology in European Gentianella
1998: sub Gentiana). The case of G. insubrica is
less clear. Although Kunz (1940) empasized
that this is a distinct taxon to us it seems likely
that it is a southern isolated variant of
G. germanica or G. rhaetica with smaller
flowers. However, in G. insubrica the calyx
lobes are often very unequal (as in G. anisodonta) and elongated (as in G. ramosa). We
do not agree with the inclusion of this taxon
(Pignatti 1983) in G. anisodonta. Genetic
evidence has to be considered also in deciding wether these taxa should be included
elsewhere.
Determination key. This is a multiple
access key based on the investigated material.
Terms describing frequency refer to statistical
counts. Exclusive characters (>95% of all
observations) are given without adverb. In
other cases terms discribing frequencies are
used: usually (>70% of all observations),
often (>50%) frequently (>30%), occasionally (>5%), rarely (<5%). Measures include
minimum and maximum (in parentheses) and
the range covered by the standard deviation
(all rounded to next integer). The papillae
type can be judged with a ten-fold lense.
Data from Rich et al. (1997) were including
for the G. amarella group. Caveat: green or
hyaline epidermal cells alternating with darkpurplish cells can produce the impression
of short papillae which are actually not
present.
1 Calyx lobes with strongly crispate margin; margin usually strongly blackish and
without papillae
G. crispata
* Calyx lobes with flat or recurved, but not
crispate, margin
2
2 Flowers 4- (rarely 5-) merous; margin of
calyx lobes usually flat
3
* Flowers 5- (rarely 4-) merous; margin of
calyx lobes flat or recurved
5
3 Margin of calyx lobes usually with longconical, occasionally with short-conical papillae; calyx lobes usually very unequal
4
* Margin of calyx lobes smooth or with
short-conical papillae; calyx lobes subequal.
4-merous variants (incl. G. hispanica) of the
G. amarella-group 13
161
4 2 large ovate-lanceolate calyx lobes
(partly) enclosing the 2 smaller linear-lanceolate ones G. campestris (incl. G. hypericifolia)
* 2 large triangular-lanceolate calyx lobes
not enclosing the 2 smaller linear-lanceolate
ones
G. columnae
5 Calyx lobes falcate, linear; calyx tube
often split down one side
G. caucasea
* Calyx lobes not falcate
6
6 Margin of calyx lobes usually with longconical or long-cylindrical papillae
7
* Margin of calyx lobes smooth or with
short-conical papillae
9
7 Midrib of calyx lobes without papillae;
calyx lobes usually strongly revolute and very
unequal, margin usually with long-conical,
occasionally with short-conical or long-cylindrical papillae
G. anisodonta-group
* Midrib of calyx lobes often with papillae;
calyx lobes revolute or not, usually subequal,
margin with long-cylindrical or long-conical
papillae, occasionally with short-conical papillae
8
8 Calyx lobes usually broadly-lanceolate,
(1.1-)1.3-2.0(-2.6) · as long as tube; margin of
calyx lobes usually with long-cylindrical, occasionally with long-conical or short-conical
papillae
G. aspera
* Calyx lobes narrowly-lanceolate,
(1.4-)1.9-2.7(-3.0) · as long as tube; margin
of calyx lobes with long-conical or longcylindrical, occasionally with short-conical
papillae or rarely smooth
G. pilosa
9 Calyx sinus usually obtuse; calyx lobes
usually linear or ovate-lanceolate-widest towards apex
10
* Calyx sinus usually acute; calyx lobes
(triangular-) lanceolate
12
10 Calyx lobes usually strongly revolute, often ovate-lanceolate-widest towards
apex
G. bulgarica
* Calyx lobes flat or rarely slightly revolute, usually linear
11
11 Corolla (10-)14-19(-22) mm; gynophor
0-1(-2) mm; margin of calyx lobes usually with
short-conical papillae
G. amarella 15
* Corolla (21-)22-40(-51) mm; gynophor (0-)1-7(-10) mm; margin of calyx
162
J. Greimler et al.: Morphology in European Gentianella
lobes without (Central Europe) or frequently with short-conical papillae (Eastern
Europe)
G. austriaca-group
12 Calyx lobes (2.0-)2.1-3.8(-4.5) · as long
as tube; corolla usually much shorter than 25
mm
13
* Calyx lobes (0.8-) 1.1-1.6(-2.4) · as long
as tube
16
13 Tall plants, usually > 15 cm, > 7
internodes (autumnal variant); stems simple or
branched from above the base or smaller
plants with < 4 internodes and long ascending
branches and terminal internode plus pedicel
longer than leaves
G. amarella-group 15
Note: Rare(?) aestival variants of G. amarella with < 7 internodes can be distinguished
from the following taxa by at least 3 internodes
much longer than the leaves, the longest
internode 3-7 · as long as the leaves below.
* Small pants, < 15 cm, usually 4–7
internodes; stems usually densely branched
from the base; plants often of caespitose
cushion-like habit; terminal internode plus
pedicel shorter than leaves
14
14 Calyx lobes usually lanceolate, margin flat or slightly revolute, smooth; leaves
in middle of stem often ovate and (nearly)
blunt
G. ramosa
* Calyx lobes linear or lanceolate, margin
occasionally strongly revolute, often with
short-conical papillae; leaves in middle of stem
ovate and acute or lanceolate
G. bulgarica
15 Usually > 4 internodes; tall plants
(5-)15-80(-109) cm; terminal pedicel and
uppermost internode together forming 1030(-40)% heigth of plant; calyx often delicate
G. amarella
* Usually < 4 internodes; small plants
(4-)6-20(-55) cm; terminal pedicel and uppermost internode together forming (30-)50-90 %
heigth of plant; calyx often enlarged
G. uliginosa (incl. G. anglica)
16 Corolla (10-)14-19(-22) mm; gynophor
0-1(-2) mm
G. amarella-group 15
* Corolla (15-)22-33(-50) mm; gynophor
(0-)1-7 mm
G. germanica s.l. 17
17 Gynophor (0-)0.1-0.5(-0.8) · as long as
calyx tube; Corolla (1.1-) 1.5-2.0(-2.3) · as
long as calyx
G. rhaetica
* Gynophor (0.4-)0.6-1.1(-1.4) · as long as
calyx tube; Corolla (1.6-) 1.9-2.6(-3.5) · as
long as calyx
G. germanica s.str.
Additional key for taxa with intermediate
characters Plants without long-conical or longcylindrical papillae; the calyx lobes are often
narrowly-lanceolate, approaching linear; sinuses somewhat intermediate, more acute than
obtuse; often within one individual both
lanceolate and linear lobes as well as acute
and obtuse sinuses were found.
1 1 or 2 calyx lobes usually much larger
than the others, (1.1-)1.5-3.3(-3.5) · as long as
tube, with short-conical papillae G. insubrica
* Calyx lobes subequal, (1.0-)1.3-2.0(-2.4) ·
as long as tube, with short conical papillae or
smooth
2
2 Calyx lobes usually linear or narrowlylanceolate; corolla (21-)25-32(-35) mm
G. bohemica
* Calyx lobes often broadly-lanceolate;
corolla (22-)29-37(-47) mm
G. stiriaca
This research was supported by the
Austrian Science Fund (FWF, P15346-B03).
We further want to thank: A. Alegro, N.
Boscaiu, C. Dobeš, F. Gruber, M. Fischer,
M.A. Fischer, H. Niklfeld, M. Onete, F.
Prochazka, G. M. Schneeweiss, P. Schönswetter, L. Schratt-Ehrendorfer, E. Sinn, A.
Tribsch, and B. Wallnöfer for various help in
collecting; the herbaria GZU, LI, M,
NMGW, PRC, RUEB, TSB, UDM, W,
and WU for supplying vouchers via loan or
during visits; T.C.G. Rich for comments;
H. Rainer for technical support; A. Kästner
for the calyx drawings; T.F. Stuessy for
reading through an earlier version of the
manuscript.
J. Greimler et al.: Morphology in European Gentianella
163
Appendix 1. Synopsis of European taxa of Gentianella section Gentianella with short distribution notes and
comments on doubtful taxa. Bold letters indicate taxa that have been accepted or given for Europe
(G. caucasea see note) in Pritchard and Tutin (1972); numbers indicate taxa or informal groups, that can be
distinguished according to our observations
1
2
3
4
5
6
G. caucasea (Loddiges ex Sims)
J. Holub
G. columnae (Ten.) J. Holub
G. crispata (Vis.) J. Holub
G. amarella group
G. amarella (L.) Börner
incl. G. hispanica López Udias,
Fabregat & Renob.
G. uliginosa (Willd.) Börner
incl. G. anglica (Pugsley) E.F. Warburg
G. campestris group
G. campestris (L.) Börner
incl. G. hypericifolia (Murb.)
Pritchard
G. laevicalyx (Rohlena) Rohlena
G. germanica group
G. albanica (Jav.) J. Holub
7
8
9
10
11
12
13
14
15
16
17
G. anisodonta group
incl. G. anisodonta (Borbas) A.
& D. Löve
incl. G. liburnica E. Mayer &
H. Kunz
incl. G. engadinensis (Wettst.)
J. Holub
G. aspera (Hegetschw. & Heer) Dostal
ex Skalicky, Chrtek & Gill
G. austriaca group
incl. G. austriaca (A. & J. Kerner)
J. Holub
incl. G. fatrae (Borb) J. Holub
incl. G. lutescens (Velen.) J. Holub
G. bohemica Skalicky
G. bulgarica (Velen.) J. Holub
G. germanica (Willd.) E.F. Warburg
G. insubrica (H. Kunz) J. Holub
G. pilosa (Wettst.) J. Holub
G. ramosa (Hegetschw.) J. Holub
G. rhaetica (= Gentiana rhaetica
A. & J. Kerner, not combined
in Gentianella)
G. stiriaca (= Gentiana stiriaca Wettst.,
not combined in Gentianella)
Caucasus, ? (presence in European part of
Turkey questioned)
C. Appennini
Balkans, S. Italy
N., W. and C. Europe extending to S.E. Europe
C. Spain
N., N.C. and W. Europe
S. England
N. W. and C. Europe, Pyrenees
Pyrenees
?, no voucher seen, diagnosis not seen;
former Jugoslavia
?, only one voucher (two small plants) seen,
diagnosis not seen; Albania
S.C. and S.E. Alps, N.W. Dinarids
S.E. Alps, N.W. Dinarids
(E.)C. Alps
N. and E.C. Alps, C. Europe
N.E. Alps, E.C. Europe
W.Carpathians
E. Europe, N. Balkans
C. Europe
C. Balkans, S. Carpathians
N.C. and W. Europe
S.C. Alps
S.E. Alps
C. and S.W. Alps
E. Alps
N.E. Alps
164
J. Greimler et al.: Morphology in European Gentianella
Appendix 2. Vouchers of population samples with collection number (deposited in WU), location, altitude,
coordinates East/North, and collector. Additional vouchers (in italics) studied from public herbaria are
given with data base accession number (as numbers on vouchers are missing in many cases), collector, and
herbarium
G. albanica
Greece: 42, W. Greuter, M.
G. amarella
Austria: 347, A. Polatschek, W; 348, R. Seipka, W; 349, R. Seipka, W; 350, A. Polatschek, W; 351, A.
Polatschek, W; 352, A. Polatschek, W. Czech Republic: 28a, Southern Bohemia, Sudslavice N Vimperk,
600 m, 1347’, 4905’, J. Greimler; 86, Southern Bohemia, Sudslavice N Vimperk, 600 m, 1347’, 4905’, J.
Greimler; 91, Southern Bohemia, Jaroskov, 750 m, 1340’, 4907’, J. Greimler. 356, F. Kovar, W; 408, F.
Svestka, WU. Danmark: 342, N. Jakobsen & J. Svensden, W. Germany: 43, F.G. Dunkel, M; 44, O. Angerer,
M; 45, C. Correns, M; 46, P. Junge, M. Hungary: 407, V. Borbas, WU. Lettland: 354, K. Starcs, W; 398, K.
Starcs, WU; 403, K. Starcs, WU. Norway: 341, B. Federley, W; 344, P.Wendelbo & A. Roesvik, W. Russia:
393, T.A. Teploukhoff, WU. Slovac Republic: 45, Mala Fatra, Chata Vratna S Terchova, 760 m, 1902’,
4912’, J. Greimler & G.M. Schneeweiss. 353, F. Svestka, W; 357, J. Chrtek & B. Deylova, W. Sweden: 345,
K.H. Rechinger, W; 346, V. Samuelsson, W; 395, Enander, WU; 399, F. Vierhapper, WU. United
Kingdome: 47, S.L. Jury, M; 48, M. Weigend, M; 197, R.&M. Gulliver, NMGW; 200, W.A. Shoolbred,
NMGW; 201, W.A. Shoolbred, NMGW; 202, W.A. Shoolbred, NMGW; 203, F. Rose, NMGW; 206, E.
Vachell, NMGW; 207, A.E. Wade, NMGW; 209, A.H. Trow, NMGW; 211, H.J. Dawson, NMGW; 212, R.
Lewis, NMGW; 231, R.A. Boniface, NMGW; 232, C.E. Hubbard, NMGW; 233, T.C.G. Rich, A. McVeigh &
J. Carey, NMGW; 234, T.C.G. Rich, A. McVeigh & J. Carey, NMGW; 235, T.C.G. Rich, A. McVeigh,
NMGW; 355, E.K. Horwood, W; 400, F. Vierhapper, WU.
G. anisodonta
Austria: 53, Salzburg, Schaldminger Tauern, W Steirischer Kalkspitze, 2280 m, 1337’, 4717’, J. Greimler;
55, Salzburg, Radstädter Tauern, Zauchensee, 1500 m, 1327’, 4716’, J. Greimler; 81, Salzburg, Hohe
Tauern, Goldberggruppe, Frauenkogel, 2200 m, 1310’, 4710, G.M. Schneeweiß; 84, Salzburg, Hohe
Tauern, Goldberggruppe, Heukareck, 2100 m, 1310’, 4718’, G.M. Schneeweiß. Croatia: 59, Gorski-Kotar
mountains, Snjeznik, 1450 m, 1435’, 4526’, A. Alegro. Italy: 42, Sondrio, Alpi Orobie, Passo San MarcoMonte Azzarini, 2200, 0940’, 4603’, L. Schratt-Ehrendorfer. 459, H. Becker, LI; 468, L. Poldini, TSB;
469, L. Poldini, TSB; 470, L. Poldini, TSB.
G. aspera
Austria: 14, Oberösterreich, Totes Gebirge, Hinterstoder, 650 m, 1407’, 4742’, J. Greimler; 68, Steiermark, Totes Gebirge, Bad Aussee, Loser, 1550 m, 1347’, 4740’, J. Greimler; 82, Salzburg, Hohe Tauern,
Goldberggruppe, Plattenberg, 1650 m, 1257’, 4709’, G.M. Schneeweiß; 83 Salzburg, Hohe Tauern,
Glockner-Gruppe, Fusch, 1900 m, 1247’, 4713’, G.M. Schneeweiß. 50, Arnold, M; 51, F. Schuhwerk, M;
53, J. Sellmair, M; 54, J. Sellmair, M; 55, W. Freiberg, M; 56, E. Dörr, M; 57, J. Koch, M; 58, J. Koch, M.
Czech Republic: 17, Southern Bohemia, Kocelovice, 450 m, 1350’, 4928’, J. Greimler. Germany: 59,
Arnold, M; 60, F. Vollmann, M; 61, Arnold, M; 62, Arnold, M; 63, Sendtner, M; 64, C.J. Mayer, M; 65, E.
Hepp, M; 66, F. Vollmann, M; 67, G. Hegi, M; 68, Diehm, M; 69, H. Paul, M; 70, E. Dörr, M; 71, P.
Eggensberger, M; 72, P. Eggensberger, M; 73, R. Urban, M; 74, H. Merxmüller, M; 75, W. Lippert, M; 76,
R. Urban, M; 77, D. Lemp, M; 78, C. Niederbichler, M; 79, H. Roessler, M; 80, Eschelmüller & Dörr, M; 81,
E. Dörr, M; 82, F. Schuhwerk, M; 83, F. Eberlein, M; 84, F. Eberlein, M; 85, F. Eberlein, M; 86, A. Lang,
M; 87, F. Eberlein, M; 88, F. Eberlein, M; 89, F. Eberlein, M; 90, Dr. Wolley, M; 91, H. Löffelmann, M; 92,
W. Kortenhaus, M; 93, W. Kortenhaus, M; 94, W. Kortenhaus, M; 95, A. Mayer, M; 96, G. Weisenbeck, M;
97, W. Lippert, M; 98, R. Grützmann, M; 99, E. Dörr, M; 100, C. Niederbichler, M; 101, J. Mayer, M; 102,
R. Binsfeld, M; 103, F. Schuhwerk, M; 104, H.J. Tillich, M; 105, O. Angerer, M; 106, O. Mergenthaler, M;
107, G. Hegi, M; 108, F. Vollmann, M; 109, F. Vollmann, M; 110, Ernst, M; 111, Arnold, M; 112, H. Wild,
M; 113, R. Urban, M; 114, W. Kortenhaus, M; 115, N. Müller, M; 116, E. Dörr, M; 117, E. Dörr, M; 118, F.
Schuhwerk, M; 119, J. Mayer, M; 120, Dihm, M.
J. Greimler et al.: Morphology in European Gentianella
165
Appendix 2 (continued)
G. austriaca
Austria: 3, Niederösterreich, Raxplateau, 1610 m, 1545’, 4744’, J. Greimler; 4, Niederösterreich,
Schneeberg, 1100 m, 1550’, 4744’, J. Greimler; 5, Niederösterreich, Gutensteiner Alpen, Pernitz, 470 m,
1556’, 4753’, J. Greimler; 6, Niederösterreich, Wienerwald, Lindabrunn, 360 m, 1609’, 4755’, J.
Greimler; 7, Niederösterreich, Wienerwald, Gaaden, 350 m, 1613’, 4803’, J. Greimler; 8, Steiermark, Prein/
Rax, above Preiner Gscheid, 1350 m, 1543’, 4741’, J. Greimler; 9, Burgenland, Seewinkel, Weiden am See,
120 m, 1651’, 4754’, J. Greimler; 44, Niederösterreich, Wiener Becken, Moosbrunn, 180 m, 1626’, 4800’,
J. Greimler. Hungary: 58, Günser Gebirge, Bozsok SE Köszeg, 380 m, 1629’, 4720’, J. Greimler.
G. bohemica
Austria: 461, J. Kerner, GZU; 462, J. Kerner, GZU. Czech Republic: 15, Southern Bohemia, Onsovice, 650
m, 1346’, 4907’, J. Greimler; 16, Southern Bohemia, Javornik NW Vimperk, 900 m, 1339’, 4908’, J.
Greimler. 306, L. Krajcr, PRC; 307, M. Protiva, PRC; 308, R. Schreiber, PRC; 309, J. Simak, PRC; 310, A.
Matatko, PRC; 311, J. Diener, PRC; 312, B. Vopravil, PRC; 313, P. Hora, PRC; 314, Sourek-Sadova,
PRC; 315, M. Jerist, PRC; 316, S. Hejny, PRC; 318, J. Obdrzalek, PRC; 319, Rohlena,
PRC; 320, J. Rohlena, PRC; 321, J. Rohlena, PRC; 322, ?, PRC.
G. bulgarica
Albania: 421, Dimonie, WU; 423, I. Dörfler, WU. Bulgaria: 420, I.K. Urumoff, WU; 422, J. Wagner, WU;
424, J. Wagner, WU; 425, St. Gheorghieff, WU; 426, Velenovsky, WU. Romania: 76-79, Eastern Carpathians, Muntii Bucegi, Sinaia, 1950-2210 m, 2528’, 4524’, J. Greimler & C-G. Jang.
G. campestris
Austria: 127, F. Schuhwerk, M. France: 155, L. Giraudias, M. Spain: 128, P. Montserrat, M; 129, A.
Meebold, M; 130, P. Garin, M; 131, P. Montserrat, M. Switzerland: 25, Graubünden, Maloja SW St.
Moritz, 2400 m, 0940’, 4625’, J. Greimler; 30, Graubünden, Pass dal Fuorn (Ofenpaß), 2200 m, 1018’,
4639’, J. Greimler; 89, Luzern, Alpnachstad-Pilatus, 2050 m, 0816’, 4657’, J. Greimler. 126, H. Hertel,
M.
G. caucasea
Georgia: 60, Chewi, Cauacasus, Kasbegi, 2040, 4446’, 4234’, C.-G. Jang; 62, Chewi, Cauacasus, Mt.
Kasbek, 2430, 4437’, 4240’, C.-G. Jang; 99, Kartli, Trialetis kedi, Didgori, 1700 m, 4427’, 4145’, P.
Schönswetter & A. Tribsch; 100, Chewi, Cauacasus, Kasbegi, 1900 m, 4440’, 4239’, P. Schönswetter & A.
Tribsch.
G. columnae
Italy: 47, Abruzzo, ĹAquila-Gran Sasso, 1760 m, 1340’, 4225’, J. Greimler & B. Wallnöfer; 96, Umbria,
Perugia, Monte Sibillini, NE Casteluccio, 1250 m, 1315’, 4251’, J. Greimler & B. Wallnöfer; A3, Abruzzo,
Gran Sasso, Mte. Tremoggia, 2100 m, 1342’, 4226’, G.M. Schneeweiß & P. Schönswetter.
G. crispata
Bosnia: 446, E. Brandis, WU; 447, F. Vierhapper, WU; 448, K. Maly, WU; 449, H. Raap, WU; 450, F.
Fiala, WU; 451, T. Pichler, WU; 452, F. Fiala & G. Beck, WU; 453, F. Fiala, WU; 454, H. Handel-Mazzetti,
WU; 455, S. Murbeck, WU; 456, H. Handel-Mazzetti & E. Janchen, WU; 457, H. Handel-Mazzetti & E.
Janchen, WU.
G. engadinensis
Italy: 29, Alto Adige, Ortler-Alpen, Sulden, 1950 m, 1035’, 4631’, J. Greimler; 51, Alto Adige, 1900 m,
Ortler-Alpen, Trafoi, 1032’, 4632’, J. Greimler; 97, Sondrio, Alpi Retiche, Lago di S. Giacomo, 1950 m,
1015’, 4633’, L. Schratt-Ehrendorfer. Switzerland: 98, Graubünden, Pass dal Fuorn (Ofenpaß), 2100 m,
1018’, 4640’, P. Schönswetter.
G. fatrae
Slovac Republic: 43, Mala Fatra, Mala Fatra, Velky Krivan, 1670 m, 1902’, 4911’, J. Greimler & G.M.
Schneeweiß; 38, Velka Fatra, Velka Fatra, Tlsta E Blatnica, 1350 m, 1858’, 4856’, J. Greimler & G.M.
Schneeweiß.
166
J. Greimler et al.: Morphology in European Gentianella
Appendix 2 (continued)
G. germanica (Alps)
Austria: 22, Tirol, Igls-Patscherkofel, 2000 m, 1127’, 4713’, J. Greimler & C. Dobes; 40, Tweng, 1250 m,
1335’, 4712’, J. Greimler; 49, Steiermark, Rottenmanner Tauern, Scheibel-Alm, 1650 m, 1426’, 4726’, J.
Greimler; 50, Steiermark, Rottenmanner Tauern, Großer Bösenstein, 1950 m, 1425’, 4726’, J. Greimler;
54, Salzburg, Schladminger Tauern, Obersee, 1880 m, 1336’, 4717’, J. Greimler; 56, Steiermark, Wölzer
Tauern, Niederer Zinken - Kleiner Zinken, 2150 m, 1421’, 4716’, P. Schönswetter; 69, Salzburg,
Schladminger Tauern, Preber, 1600 m, 1352’, 4711’, C.-G. Jang; 70, Kärnten, Hohe Tauern, Goldberggruppe, Fragant, 1850 m, 1301’, 4657’, C.-G. Jang; 71, Tirol, Ötztaler Alpen, Vent, 2000 m, 1054’,
4651’, J. Greimler; 72, Tirol, Ötztaler Alpen, Rofen, 2450 m, 1053’, 4652’, J. Greimler; 121, Salzburg,
Hohe Tauern, Sportgastein, Nassfeld, 1600 m, 1303’, 4703’, F. Gruber. 458, H. Wittmann & P. Pilsl, LI;
471, A. Kerner, WU; 472, A. Pernhoffer, WU. Italy: 23, Alto Adige, Ötztaler Alpen, Plawenn, 2050 m,
1035’, 4646’, J. Greimler & C. Dobes. Switzerland: 24, Graubünden, Fuldera, 1620 m, 1022’, 4637’, J.
Greimler & C. Dobes.
G. germanica (outside Alps)
Belgium: 325, M. Crepin, PRC; 326, A. Hardy, PRC; 327, E. Peaubert, PRC; 409, A. Hardy, WU. Czech
Republic: 323, J. v. Sterneck, PRC; 324, K. Hosic, PRC. France: 258, F. Rose, NMGW; 328, A. Schmidely,
PRC; 384, H. Caron, W; 389, A. Huguenin, W; 391, Abbe F. Riguet, W; 392, C. Billot, W. Germany: A27,
Sachsen-Anhalt, Trebitz near Kloschwitz, 100 m, 1145’, 5132’, J. Greimler. 135, Dihm, M; 136, F.
Schuhwerk, M; 137, F. Schuhwerk, M; 138, E. Dörr, M; 141, W. Freiberg, M; 142, W. Freiberg, M; 143, W.
Freiberg, M; 144, W. Freiberg, M; 145, Ruppert, M; 146, W. Freiberg, M; 147, W. Freiberg, M; 148, W.
Freiberg, M; 149, J. Koch, M; 150, J. Koch, M; 151, W. Freiberg, M; 152, W. Freiberg, M; 153, W.
Freiberg, M; 366, A.Vocke, W; 367, J. Bornmüller, W; 368, J. Bornmüller, W; 369, Hausknecht, W; 370, T.
Vogel, W; 371, T. Vogel, W; 372, Westram, W; 373, W. Freiberg, W; 374, G. Sennholz, W; 375, Girth, W;
376, C.J. Mayer, W; 377, Girth, W; 378, H. Neumann, W; 380, G. Sennholz, W; 381, H. Schulz, W; 410, F.
Wirtgen, WU; 411, J. Bornmüller, WU; 412, A. Hayek, WU; 413, Sagorski, WU. Switzerland: 87, Jura
mountains, Dittingen S Basel, 350 m, 0731’, 4726’, J. Greimler; 88, Jura mountains, Diegten SE Basel,
550 m, 0749’, 4725’, J. Greimler. United Kingdome: 276, R.A. Boniface, NMGW; 277, W.H. Griffin,
NMGW; 280, E. Vachell, E. Knowling & J. Davy, NMGW; 281, J. E. Lousley, NMGW; 282, A. McVeigh &
J. Carey, NMGW; 283, T.C.G. Rich & A. McVeigh, NMGW; 292, A. McVeigh & J. Carey, NMGW; 294, A.
McVeigh & J. Carey, NMGW; 295, A. McVeigh & J. Carey, NMGW; 296, A. McVeigh & J. Carey,
NMGW.
G. insubrica
Italy: 27, Lombardia, Monte Generoso 1680 m, 0903’, 4556’, J. Greimler & C. Dobes. Switzerland: 463,
W. Koch, RUEB; 464, W. Lüdi, RUEB; 465, Jaquet, RUEB; 466, J. Coaz, RUEB; 467, W. Koch, RUEB.
G. liburnica
Slovenia: 337, G. Sauli, TSB.
G. lutescens
Poland: 32, Krakow, Vysokie Tatry, Tatry Zachodnie, 1400 m, 1959’, 4915’, P. Schönswetter & A.
Tribsch; 34, Krakow, Vysokie Tatry, Tatry Zachodnie, 1950 m, 1954’, 4914’, P. Schönswetter & A.
Tribsch. Romania: 73, Eastern Carpathians, Piatra Craiului, Zarnesti, 800 m, 2517’, 4534’, J. Greimler &
C-G. Jang; 75, Eastern Carpathians, Piatra Craiului mica, 1710 m, 2516’, 4533’, J. Greimler & C-G. Jang.
475, J. Römer, WU; 476, Porcius, WU; 477, A. Rehmann, WU; 478, A. Scherfel, WU.
G. pilosa
Italy: 18, Friuli-Venezia Giulia, Alpi Giulie, Cave del Predil, 980 m, 1334’, 4626’, J. Greimler & C-G.
Jang; A4, Veneto, Prealpi Carniche, Mte. Tiarfin, 1235’, 4628’, G.M. Schneeweiß & P. Schönswetter. 479,
I. Dörfler, WU; 480, O. Krebs, WU; 481, M. Statzer, WU; 482, Petter, WU; 483, Ressmann, WU; 484, I.
Dörfler, WU; 485, M.& I. Dörfler, WU; 486, S. Rizzardini, UDM; 487, E. Tomasi, UDM; 488, F. Martini,
UDM.
J. Greimler et al.: Morphology in European Gentianella
167
Appendix 2 (continued)
G. ramosa
Italy: 427, H. Handel-Mazzetti, WU; 431, G.B. Traverso, WU;436, Muret, WU; 439, R. Wettstein, WU;
440, R. Wettstein, WU. Switzerland: 90, Gotthardgebiet, Oberalp, 2040 m, 0846’, 4640’, J. Greimler; 26,
Graubünden, Maloja SW St. Moritz, 2100 m, 0941’, 4624’, J. Greimler. 428, H. Handel-Mazzetti, WU;
429, Wolf, WU; 430, H. Handel-Mazzetti, WU; 432, F.O. Wolf, WU; 433, R. Wettstein, WU; 434, K.
Ronniger, WU; 435, Favrat, WU; 437, F. Vierhapper, WU; 438, Favrat, WU; 441, H. Handel-Mazzetti,
WU; 442, H. Handel-Mazzetti, WU; 443, H. Handel-Mazzetti, WU; 444, H. Handel-Mazzetti, WU; 445, R.
Wettstein, WU.
G. stiriaca
Austria, all Steiermark: 12, Hochschwab, Seetal, 1050 m, 1513’, 4737’, J. Greimler; 13, Ennstaler Alpen,
Lugauer, 1530 m, 1442’, 4733’, J. Greimler; 48, Hochschwab, Aflenzer Staritzen, 1350 m, 1516’, 4738’,
J. Greimler; 52, Schneealpe, Neuberg, 1080 m, 1536’, 4740’, M.A. Fischer; 57, Ennstaler Alpen, Johnsbach, 700 m 1435’, 4733’, J. Greimler. 460, A. Lonsing, LI; 473, J. Nevole, WU; 474, A. Klammerth, WU.
G. uliginosa
Finnland: 414, H. Lindberg, WU; 415, H. Lindberg, WU; 417, H. Lindberg, WU. Germany: 418,
Schlechtendal, WU; 419, Ruthe, WU. Sweden: 164, H. Nilsson, NMGW; 416, K.F. Dusen, WU. United
Kingdome: 166, J.E. Lousley, NMGW; 168, R.F. May, NMGW; 169, I.M. Vaughan, T.A.W & D. Davies,
NMGW; 170, P. Saunders, NMGW.
References
Borba E. L., Shepherd G. J., van den Berg C.,
Semir J. (2002) Floral and vegetative morphometrics of five Pleurothallis (Orchidaceae) species: correlation with taxonomy, phylogeny,
genetic variability and pollination systems.
Ann. Bot. 90: 219–230.
Dostal J. (1989) Nova Květena CSSR. Vol. 2.
Academia, Praha.
Greimler J., Dobeš C. (2000) High genetic diversity
and differentiation in relict lowland populations
of Gentianella austriaca (A. and J. Kern.) Holub
(Gentianaceae). Plant Biol. 2: 628–637.
Hagen K. B. von, Kadereit J. W. (2001) The
phylogeny of Gentianella (Gentianaceae) and its
colonization of the southern hemisphere as
revealed by nuclear and chloroplast DNA
sequence variation. Org. Divers. Evol. 1: 61–79.
Hayek A. (1911–1914) Flora von Steiermark.
Zweiter Band – Erste Abteilung. Gebrüder
Borntraeger, Berlin.
Hess H. E., Landolt E., Hirzel R. (1972) Flora der
Schweiz. Band 3. Birkhäuser, Basel Stuttgart.
Holub J. (1983) A brief note on Slovak taxa of
Gentianella. Preslia 55: 371–373.
Holyoak D. T. (1999) Gentianella uliginosa (Willd.)
Börner (Gentianaceae) rediscovered in North
Devon. Watsonia 22: 428–429.
Jäger E. J., Werner K. (2002) Rothmaler: Exkursionsflora von Deutschland. Band 4. Gefäßpflanzen. Kritischer Band. 9. Auflage. Spektrum,
Heidelberg Berlin.
Kerner A., Kerner J. (1882) 649. Gentiana Rhaetica.
In: Kerner A., Kerner J. (eds.) Schedae ad
Floram exsiccatam Austro-Hungaricam 2. Frick,
Vindobonae, pp. 122–128.
Kropf M., Kadereit J. W., Comes H. P. (2003)
Differential cycles of range contraction and
expansion in European high mountain plants
during the Late Quaternary: insights from
Pritzelago alpina (L.) O. Kuntze (Brassicaceae).
Molec. Ecol. 12: 931–949.
Kunz H. (1940) Beitrag zur Revision einiger Gentianeen. Verh. naturforsch. Ges. Basel 51/2: 1–20.
Lauber K., Wagner G. (2001) Flora Helvetica. 3.
Aufl. Haupt, Bern Stuttgart Wien.
Lennartsson T. (1997) Seasonal differentiation, a
conservative reproductive barrier in two grassland Gentianella (Gentianaceae) species. Plant
Syst. Evol. 208: 45–69.
Ma Y. C. (1951) Gentianopsis – a new genus of
Chinese Gentianaceae. Acta Phytotax. 1: 5–19,
pls I–V.
Maurer W. (1998) Flora der Steiermark II/1. IHW,
München.
Mayer E. (1969) Zur Kenntnis der Gattung
Gentianella Moench in Jugoslawien. I. Der
168
J. Greimler et al.: Morphology in European Gentianella
G. anisodonta-Komplex. Österr. Bot. Z. 116:
393–399.
Morales R., Macı́a M. J., Dorda E., Garcı́a
Villaraco A. (1996) Archivos de Flora Iberica.
Núm. 7. Real Jardin Botanico-CSIC, Madrid.
Moravec J., Vollrath H. (1967) Gentianella ·
austroamarella hybr. spec. nov. Folia Geobot.
Phytotax. 3: 333–336.
Petanidou T., Ellis-Adam A. C., den Nijs J. C. M.,
Oostermeijer J. G. B. (1998) Pollination ecology
of Gentianella uliginosa, a rare annual of the
Dutch coastal dunes. Nord. J. Bot. 18: 537–548.
Pignatti S. (1983) Flora d’Italia. Vol. secondo.
Edagricole, Bologna.
Pritchard N. M. (1961) Gentianella in Britain. III.
Gentianella germanica (Willd.) Börner. Watsonia
4: 290–303.
Pritchard N. M., Tutin T. G. (1972) 6. Gentianella
Moench. In: Tutin T. G., Heywood V. H. (eds.)
Flora Europaea, Vol. 3. Cambridge University
Press, Cambridge, pp. 63–67.
Renobales G., Fabregat Llueca C., López Udias S.
(2002) Una nueva especie del genero Gentianella
(Gentianaceae) del sistema ı́berico. Anales Jard.
Bot. Madrid 59: 217–226.
Renobales Scheifler G. (2003) Notas acerca del
tratamiento de las Gentianaceae para ‘‘Flora
iberica’’. Anales Jard. Bot. Madrid 60: 461–
469.
Rich T. C. G. (1997) Gentianella uliginosa (Willd.)
Boerner (Gentianacaeae) present in England?
Watsonia 21: 208–209.
Rich T. C. G., Holyoak D. T., Margetts L. J.,
Murphy R. J. (1997) Hybridisation between
Gentianella amarella (L.) Boerner and G. anglica
(Pugsley) E. F. Warb. (Gentianacaeae). Watsonia 21: 313–325.
Ritter-Studnicka H. (1955) Eine neue Unterart von
Gentiana crispata Vis. aus den Karstfeldern
Westbosniens. Feddes Repert. 57: 203–208.
Rosenbauer A. (1996) Gentianaceae. In: Sebald O.,
Seybold S., Philippi G., Wörz A. (eds.) Die
Farn- und Blütenpflanzen Baden-Württembergs.
Spezieller Teil (Spermatophyta, Unterklasse Asteridae) Buddlejaceae bis Caprifoliaceae. E. Ulmer, Stuttgart, pp. 16–42.
Samuelsson G. (1922) Zur Kenntnis der Schweizer
Flora. Vierteljahrsschr. naturforsch. Ges. Zürich
67: 224–267.
Skalicky V. (1969) Die Sammelart Gentianella
germanica (Willd.) E. F. Warburg s. l. im
Böhmischen Massiv. Preslia 41: 140–147.
Struwe L., Kadereit J. W., Klackenberg J., Nilsson
S., Thiv M., von Hagen K. B., Albert V. A.
(2002) Systematics, character evolution, and
biogeography of Gentianaceae, including a new
tribal and subtribal classification. In: Struwe L.,
Albert V. A. (eds.) Gentianaceae. Cambridge
University Press, Cambridge, pp. 21–309.
Toyokuni H. (1961) Séparation de Comastoma,
genre nouveau d’avec Gentianella. Bot. Mag.
(Tokyo) 74: 198.
Tribsch A., Schönswetter P. (2003) Patterns of
endemism and comparative phylogeography
confirm paleo-environmental evidence for Pleistocene refugia in the eastern Alps. Taxon 52:
477–497.
Wagner J., Mitterhofer E. (1998) Phenology, seed
development, and reproductive success of an
alpine population of Gentianella germanica in
climatically varying years. Bot. Acta 111: 159–
166.
Wettstein R. (1892) Untersuchungen über Pflanzen
der österreichisch-ungarischen Monarchie. Die
Arten der Gattung Gentiana aus der Sektion
,,Endotricha‘‘ (Frœl.). Österr. Bot. Z. 42: 1–6, 40–
45, 84–88, 125–130, 156–161, 193–196, 229–235.
Wettstein R. (1896) Die Europäischen Arten der
Gattung Gentiana aus der Section Endotricha
Frœl. und ihr entwicklungsgeschichtlicher Zusammenhang. C. Gerold, Wien.
Winfield M. O., Wilson P. J., Labra M., Parker J.
S. (2003) A brief evolutionary excursion comes
to an end: the genetic relationship of British
species of Gentianella sect. Gentianella (Gentianaceae). Plant Syst. Evol. 237: 13 7–151.
Woo H.-K., Kim J.-H., Yeau S.-H., Lee N. S.
(2002) Morphological and isozyme divergence in
Korean Hepatica sensu stricto (Ranunculaceae).
Plant Syst. Evol. 236: 33–44.
Yuan Y. M., Küpfer P. (1995) Molecular phylogenetics of the subtribe Gentianinae (Gentianaceae) inferred from the sequences of internal
transcribed spacers (ITS) of nuclear ribosomal
DNA. Plant Syst. Evol. 196: 207–226.
Zopfi H. J. (1991) Aestival and autumnal vicariads
of Gentianella (Gentianaceae): a myth? Plant
Syst. Evol. 174: 139–158.
J. Greimler et al.: Morphology in European Gentianella
Address of the authors: Josef Greimler (e-mail:
josef.greimler@univie.ac.at), Chang-Gee Jang,
Department of Systematics and Evolution of
Higher Plants. Barbara Hermanowski, Department
169
of Ultrastructure Research and Palynology. All:
Institute of Botany, University of Vienna, Rennweg
14, A-1030 Vienna, Austria.