Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України
The objective of this study was to carry out macro- and micromorphological studies of fruits and seeds of representatives of the genus Lonicera of the flora of Ukraine (both natural and introduced), to clarify their features, to provide additional characteristics and to evaluate the possibility of u...
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| Дата: | 2020 |
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2020
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Plant Introduction| _version_ | 1860145086896013312 |
|---|---|
| author | Tsarenko, O.M. Bulakh, O.V. Kolesnichenko, O.V. Hrysiuk, S.M. |
| author_facet | Tsarenko, O.M. Bulakh, O.V. Kolesnichenko, O.V. Hrysiuk, S.M. |
| author_sort | Tsarenko, O.M. |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2023-08-26T20:39:45Z |
| description | The objective of this study was to carry out macro- and micromorphological studies of fruits and seeds of representatives of the genus Lonicera of the flora of Ukraine (both natural and introduced), to clarify their features, to provide additional characteristics and to evaluate the possibility of using carpological features for the diagnosis of taxa of the genus.
Material and methods. Materials of the National Herbarium of the M.G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine (KW) and Herbarium of the M.M. Gryshko National Botanical Garden of the National Academy of Sciences of Ukraine (KWHA) were used, as well as samples from living plants from the collections of the M.M. Gryshko National Botanical Garden and O.V. Fomin Botanical Garden. The citations are given by the original text of the labels. The fruits and seeds of seven species of the genus (four are introduced, and the other are species of natural flora), which are listed in “Vascular plants of Ukraine. A nomenclatural checklist”, have been studied. Morphological features of fruits and seeds were studied under the light microscope (LM, MBS-9) and scanning electron microscope (SEM, JSM-6060 LA). Samples were sprayed with a layer of gold according to the standard method. The study was conducted at magnification from ×30 to ×3000. Descriptions of fruits and seeds were performed using terminology, summarized in specialized papers. Fruit and seed sizes were measured using Axio Vision Rel. 4.8.
Results. The fruits of Lonicera species are free or adnate at the base, surrounded by, for the most part, free bracteoles. A pseudocarp of L. caerulea is formed of the bracteoles of paired flowers concrescent in a tubular cupule, which completely and tightly wraps free paired fruits. The L. nigra has an uneven concrescent of the bracteoles that freely surround the paired fruits, which have concrescent only at the base. The fruits of the species Lonicera are black and dark blue, or red and yellow. The fruits of subgenus Periclymenum (L. caprifolium, L. etrusca, L. periclymenum) are free and spherical. The subgenus Chamaecerasus species’ fruits are mainly spherical, but in some species the fruits are also hemispherical, elliptical and elongated (L. caerulea, L. tatarica). Fruits in this subgenus are free (L. caerulea, L. tatarica) or coalescent at the base (L. nigra, L. xylosteum). All the studied species are characterized by glabrous fruits, except fruits of L. xylosteum that are pubescent with glandular and simple hairs. The seeds are elliptical, ovoid, nearly spherical, flat, concave on one side and convex on the other side, with a recess on the periphery and a crest in the center, at the base they are mainly narrow-rounded. All species of the section Eucaprifolium and one species of the section Coeloxylosteum have truncated seeds. The surface of the seeds is almost similar and pitted; in L. xylosteum it is shallow-pitted; in L. nigra it is tubercular-pitted. In some species, the seed surface is pitted, and also flat cells are present.
Conclusions. Based on the results of a critical review of literature materials and analysis of own data, the detailed characteristics of fruits and seeds of the representatives of the genus Lonicera within the flora of Ukraine have been compiled. These characteristics are useful for more accurate identification of fruiting Lonicera plants. The species of the subgenus Periclymenum have exclusively free fruits, while the fruits in the subgenus Chamaecerasus are free or coalescent at the base. The features of color and shape of the fruits and seeds, the peculiarities of pubescence of the fruits, are suggested to be used to distinguish L. caerulea, L. caprifolium, L. etrusca, L. nigra, L. periclymenum, L. tatarica, and L. xylosteum. Analysis of the microstructure of the surface of the seeds revealed that there no features that can be used as diagnostic at the level of subsections, sections, and subgenera. However, seeds microstructure showed some differences at the species level. |
| doi_str_mv | 10.46341/PI2020012 |
| first_indexed | 2025-07-17T12:53:40Z |
| format | Article |
| fulltext |
Plant Introduction, 85/86, 109–123 (2020)
© The Authors. This content is provided under CC BY 4.0 license.
RESEARCH ARTICLE
Carpological features of Lonicera L. (Caprifoliaceae Juss.) of the flora of
Ukraine
O.M. Tsarenko 1*, O.V. Bulakh 1, O.V. Kolesnichenko 2, S.M. Hrysiuk 2
1 M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Tereschenkivska str. 2, 01004 Kyiv, Ukraine;
* tsarenko_olga@ukr.net
2 National University of Life and Environmental Sciences of Ukraine, Heroiv oborony str. 15, 03041 Kуiv, Ukraine
Received: 03.03.2020 | Accepted: 25.04.2020 | Published: 30.06.2020
Abstract
The objective of this study was to carry out macro- and micromorphological studies of fruits and seeds of
representatives of the genus Lonicera of the flora of Ukraine (both natural and introduced), to clarify their
features, to provide additional characteristics and to evaluate the possibility of using carpological features
for the diagnosis of taxa of the genus.
Material and methods. Materials of the National Herbarium of the M.G. Kholodny Institute of Botany of
the National Academy of Sciences of Ukraine (KW) and Herbarium of the M.M. Gryshko National Botanical
Garden of the National Academy of Sciences of Ukraine (KWHA) were used, as well as samples from living
plants from the collections of the M.M. Gryshko National Botanical Garden and O.V. Fomin Botanical
Garden. The citations are given by the original text of the labels. The fruits and seeds of seven species of the
genus (four are introduced, and the other are species of natural flora), which are listed in “Vascular plants
of Ukraine. A nomenclatural checklist”, have been studied. Morphological features of fruits and seeds were
studied under the light microscope (LM, MBS-9) and scanning electron microscope (SEM, JSM-6060 LA).
Samples were sprayed with a layer of gold according to the standard method. The study was conducted
at magnification from ×30 to ×3000. Descriptions of fruits and seeds were performed using terminology,
summarized in specialized papers. Fruit and seed sizes were measured using Axio Vision Rel. 4.8.
Results. The fruits of Lonicera species are free or adnate at the base, surrounded by, for the most part,
free bracteoles. A pseudocarp of L. caerulea is formed of the bracteoles of paired flowers concrescent
in a tubular cupule, which completely and tightly wraps free paired fruits. The L. nigra has an uneven
concrescent of the bracteoles that freely surround the paired fruits, which have concrescent only at the
base. The fruits of the species Lonicera are black and dark blue, or red and yellow. The fruits of subgenus
Periclymenum (L. caprifolium, L. etrusca, L. periclymenum) are free and spherical. The subgenus Chamaecerasus
species’ fruits are mainly spherical, but in some species the fruits are also hemispherical, elliptical and
elongated (L. caerulea, L. tatarica). Fruits in this subgenus are free (L. caerulea, L. tatarica) or coalescent at
the base (L. nigra, L. xylosteum). All the studied species are characterized by glabrous fruits, except fruits
of L. xylosteum that are pubescent with glandular and simple hairs. The seeds are elliptical, ovoid, nearly
spherical, flat, concave on one side and convex on the other side, with a recess on the periphery and a
crest in the center, at the base they are mainly narrow-rounded. All species of the section Eucaprifolium
and one species of the section Coeloxylosteum have truncated seeds. The surface of the seeds is almost
similar and pitted; in L. xylosteum it is shallow-pitted; in L. nigra it is tubercular-pitted. In some species, the
seed surface is pitted, and also flat cells are present.
Conclusions. Based on the results of a critical review of literature materials and analysis of own data, the
detailed characteristics of fruits and seeds of the representatives of the genus Lonicera within the flora of
Ukraine have been compiled. These characteristics are useful for more accurate identification of fruiting
Lonicera plants. The species of the subgenus Periclymenum have exclusively free fruits, while the fruits in the
subgenus Chamaecerasus are free or coalescent at the base. The features of color and shape of the fruits
https://doi.org/10.46341/PI2020012
UDC 581.471/581.48/582.973
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0002-1216-6628
https://orcid.org/0000-0001-8471-3750
https://orcid.org/0000-0003-4767-6844
https://orcid.org/0000-0002-7523-427X
110 Plant Introduction • 85/86
O.M. Tsarenko, O.V. Bulakh, O.V. Kolesnichenko, S.M. Hrysiuk
Introduction
The genus Lonicera is represented in the world
flora by almost 200 species (Theis et al., 2008;
Takhtajan, 2009). The species are distributed
in all areas of the Northern Hemisphere (in
temperate and subtropical regions of Europe,
North and Central America, North Africa, and
Asia). South-East Asia is considered to be the
primary center of origin for this genus. Some
representatives of this genus are decorative
plants with beautiful flowers and fruits and
were introduced into the culture (Browicz,
1976; Poyarkova, 1958; Zaytsev, 1962).
Mosyakin & Fedorochuk (1999) have listed
seven Lonicera species for the flora of Ukraine
– four are species of natural flora (L. caerulea L.,
L. etrusca Santi, L. nigra L., L. xylosteum L.),
and three species (L. caprifolium L.,
L. periclymenum L., L. tatarica L.) are
introduced from other countries and are widely
used in botanical gardens and parks of Ukraine
(Barbarych, 1961; Zaytsev, 1962; Glukhov et al.,
2002; Lavrinenko, 2016).
According to the APG IV (APG, 2016) and
the results of recent phylogenetic studies of
Dipsacales Juss. ex Bercht. et J. Presl, the genus
Lonicera belongs to the tribe Caprifolieae of
the family Caprifoliaceae Juss. (Theis et al.,
2008; Takhtajan, 2009; Reveal, 2012). Also, the
features of fruits and seeds of Lonicera are well
discussed in the literature and listed among
the morphological features confirming recent
molecular-phylogenetic data on Dipsacales
(Bell et al., 2001, 2005; Donoghue, 2003;
Jacobs et al., 2008; Theis et al., 2008; Göktürk
& Sümbül, 2014).
Carpological characteristics are essential
for the identification of these plants in
the fruiting stage. The information on the
morphological features of seeds is also
important for specialists in botanical gardens
and arboretums, who directly work with seeds
during the seed exchange programs.
It is also important to note, most of the
authors of the monographic elaboration of
the genus Lonicera in the regional Floras and
Identification Keys usually used features of the
perianth, a number of flowers on the shoot
apex, shoot pubescence, and shape of leaves to
delimit taxa. They only briefly focused on the
characteristics of fruits (color, size) and seeds as
supporting features (Rehder, 1909; Poyarkova,
1958; Barbarych, 1961; Zaytsev, 1962). Despite
this, we believe, that carpological features
(including micromorphological peculiarities)
play an essential role in diagnostics and
identification of species in the genus Lonicera,
in particular those represented in the flora of
Ukraine, during the fruiting stage.
The fruits of Lonicera are inferior syncarpous
succulent multi-seeded berries with various
colors – yellow, dark-red, blue, and black. They
are arranged in pairs, free or coalescent at
the base, often one of the paired berries may
be underdeveloped. The ovary contains 2- or
3-4 locules with a double-rowed location of
numerous anatropous ovules with one massive
integument (Plysko, 2000). The species of
Lonicera are characterized by different levels
of coalescence of adjacent fruits. Free fruits
were found in L. syringantha Maxim.; partial
fusion was found in L. canadensis Bartram
ex Marshall and L. tatarica, and the highest
coalescence of adjacent fruits is typical for
L. caerulea (Artiushenko & Fedorov, 1986;
Plysko, 2000).
Malinkina (2001) found that L. tatarica
and L. xylosteum have a pair of inferior
syncarpous 3-locular ovaries, which unified
on a single pedicel with two free bracts. At
the base of each ovary, there are two small
round bracteoles, which are partially fused
with the ovary. Some species have coalescent
bracteoles, which form a cupule enclosing the
ovaries (Poyarkova, 1958).
Zaitseva (1998, 1999, 2006) studied the
organogenesis of the Lonicera fruit in detail.
and seeds, the peculiarities of pubescence of the fruits, are suggested to be used to distinguish L. caerulea,
L. caprifolium, L. etrusca, L. nigra, L. periclymenum, L. tatarica, and L. xylosteum. Analysis of the microstructure
of the surface of the seeds revealed that there no features that can be used as diagnostic at the level of
subsections, sections, and subgenera. However, seeds microstructure showed some differences at the
species level.
Keywords: Lonicera, fruit, seed, macromorphology, micromorphology, surface microstructure of seed, flora of Ukraine
Plant Introduction • 85/86 111
Carpological features of Lonicera of the flora of Ukraine
At the beginning of fruit development,
2–3 layers of cells separate from the outer
epidermis and together form the exocarp.
Between these layers there is a layer of
parenchyma cells – mesocarp. The pericarp
of the fruit of Lonicera is formed from similar
parenchymal cells.
The cells of the exocarp are polygonal
in most species in cross-section. The cells
of mesocarp are round and thin-walled,
containing chromoplasts and drops of oil. The
inner part of the mesocarp is rich in calcium
oxalate. Endocarp is usually reduced to a
single unsclerified layer of cells (Artiushenko &
Fedorov, 1986; Plysko, 2000; Jacobs et al., 2009).
The seeds are early separated from the
pericarp. Developed seeds have a small
(up to 1/6–1/4 length of seed), straight,
well-differentiated embryo. The embryo
is surrounded by a well-differentiated
endosperm and a multilayered spermoderm
with lignified exotesta cells (Malinkina, 2001;
Jacobs et al., 2009; Lavrinenko, 2012). The
microstructure of the seed surface is specified
by the features of the walls of exotesta cells.
Anticlinal and periclinal walls of exotesta are
thickened, lignified, and have pores. The hilum
is small, submerged, located at the base of the
seed (Jacobs et al., 2009; Lavrinenko, 2012).
Analyzing research articles (Rehder, 1909;
Barbarych, 1961; Zaytsev, 1962; Artiushenko
& Fedorov, 1986; Zaitseva, 1998, 1999, 2006;
Glukhov et al., 2002; Jacobs et al., 2009;
Lavrinenko, 2012, 2016), we found that a
number of questions regarding carpological
characteristics of the genus Lonicera remained
outside the attention of researchers (i.e.,
features of the fruits pubescence, surface
microstructure of the seeds, etc.). This fact
motivated us to investigate macro- and
micromorphology of the fruits and seeds
of Lonicera of the flora of Ukraine. These
studies were aimed to provide additional
characteristics of the fruits and seeds and
to evaluate the possibility of application of
carpological features for the identification of
Lonicera specimens.
Material and methods
Specimens preserved in the National
herbarium of M.G. Kholodny Institute of
Botany of the National Academy of Sciences
of Ukraine (KW) and the Herbarium of
M.M. Gryshko National Botanical Garden of
the National Academy of Sciences of Ukraine
(KWHA), and also selected specimens from
living collections of M.M. Gryshko National
Botanical Garden and O.V. Fomin Botanical
Garden were used during the research.
The fruits and seeds of all seven species
listed in the “Vascular plants of Ukraine.
A nomenclatural checklist” (Mosyakin &
Fedoronchuk, 1999) were investigated.
Morphological features of the seeds and
fruits were studied using a light microscope
(LM, MBS-9) and scanning electron
microscope (SEM, JSM-6060 LA). The study
was conducted at magnification from ×30 to
×3000. Descriptions of fruits and seeds were
performed using terminology, summarized
in a number of relevant specialized papers
(Artiushenko & Fedorov, 1986; Plysko, 2000;
Ziman et al., 2004, 2011). Fruit and seed sizes
were measured using Axio Vision Rel. 4.8.
Results
Investigated species belong to two subgenera:
Chamaecerasus (L.) Rehder (sections Isika
(Adanson) Rehder and Coeloxylosteum Rehder)
and Periclymenum (L.) Rehder (section
Eucaprifolium (Spach) Pojarkova). Based on
published materials and obtained results,
their characteristics have been compiled.
The species are listed below according to the
system of the genus Lonicera represented in
the “Flora of the USSR” (Poyarkova, 1958).
Genus Lonicera L.
Subgenus Chamaecerasus (L.) Rehder
Section Isika (Adanson) Rehder
Subsection Caeruleae (Rehder) Nakai
Lonicera caerulea L. (Fig. 1; Tables 1 & 2)
Two fruits are free. A pseudocarp is formed
by bracteoles of two paired flowers, which are
connate in a common tubular cupule. Cupule
completely and tightly wraps free paired fruits.
The fruits are dark blue, with bluish cover,
fleshy, spherical, hemispherical or elliptical,
8–11 mm in diameter, containing up to 10–20
(–30) seeds.
The seeds are light- or dark-brown, broadly
elliptical or rounded, 1.5–2.3 mm long and
1.4–2.1 mm wide, flat, slightly concave on one
112 Plant Introduction • 85/86
O.M. Tsarenko, O.V. Bulakh, O.V. Kolesnichenko, S.M. Hrysiuk
side, convex on the other, with a cavity on
the periphery and a crest in the center, at the
base narrow-rounded, with a small notch. The
hilum is small, submerged, located at the base
of the seed.
The surface of the seed is mostly pitted.
Exotesta cells are polygonal, with concave
periclinal and distinctly visible anticlinal walls.
The outer periclinal walls have ribs, which
become visible at the magnification over ×500.
Ribs were without clear orientation.
Samples examined: 1. Transcarpathian region, Rakhiv
district, around village Kvasy, the north-western slope of
Mt. Petros, among stones covered with sphagnum moss,
occurs in the form of small thickets, 24.VIII.1957. Leg.,
det. [SM?] Kazakevich (KW); 2. Living plants from the
collection of the M.M. Gryshko National Botanical Garden
NAS of Ukraine, 26.IX.2017. Leg., det. O.M. Tsarenko;
3. Living plants from the collection of the O.V. Fomin
Botanical Garden, 20.VII.2017. Leg., det. N.M. Belemets,
O.M. Tsarenko; 4. Carpathians, Chornohora, Mt. Petros,
1850 m, on the rocky placers of the south-western slope
of the mountain between the krummholz (Pinus mugo),
9.VIII.1954. Leg., det. K. Malinovski (KWHA).
Subsection Rhodanthae (Maximowicz) Rehder
Lonicera nigra L. (Fig. 2; Tables 1 & 2)
The infructescence consists of two fruits
connate at the base. Four bracteoles are
irregularly and incompletely connate into a
goblet-shaped cupule, which freely wraps
the fruit at its base. The fruits are black, with
bluish cover, fleshy, spherical, 6–10 mm in
diameter, connate in pairs only at the base.
The seeds are light- or dark-brown,
elliptical, 2.2–2.5 mm long and 1.8–2.4 mm
wide, flat, on one side slightly concave, on
the other side convex, with a cavity on the
periphery and a crest in the center, at the base
narrowly rounded, with a small notch. The
hilum is small, submerged, located at the base
of the seed.
The surface of the seed is tuberculate-
pitted. The cells of the exotesta are polygonal,
with concave or convex periclinal and
distinctly visible anticlinal walls. The outer
periclinal walls are ribbed or wrinkled (seen at
magnification over ×500).
BA
C D
Figure 1. The seed of Lonicera caerulea: A – general view; B–D – surface at different magnification.
Plant Introduction • 85/86 113
Carpological features of Lonicera of the flora of Ukraine
Samples examined: 1. Transcarpathian region, Rakhiv
district, climb to Hoverla, slopes to r. Hoverla at an altitude
of 700 m. 27.VII.1947. Leg., det. M. Klokov (KW); 2. Chernivtsi
region, Putilsky district, village Selyatin, spruce forest,
Urochysche Ropachevske, on the edge of the forest.
28.VI.1947. Leg., det. N. Kosets (KW); 3. Transcarpathian
region, Rakhiv district, village Bogdan, Urochysche Turkul,
spruce forest, 7.VI.1953. Leg., det. Komendar (KWHA).
Section Coeloxylosteum Rehder
Subsection Tataricae Rehder emend. Nedoluzhko
Lonicera tatarica L. (Fig. 3; Tables 1 & 2)
Two free fruits, bracteoles are also free and
cover the fruits to the middle. The fruits are
red or yellow, fleshy, spherical, sometimes
elliptical, 6–8 mm in diameter.
The seeds are light- or dark-brown,
elliptical, 3.0–3.5 mm long and 1.5–2.7 mm
wide, somewhat flat, slightly concave on one
side, convex on the other, with a cavity on the
periphery and a crest in the center, at the base
narrowly rounded, somewhat truncated. The
hilum is small, submerged, located at the base
of the seed.
The surface of the seed is pitted. The cells
of the exotesta are polygonal, with concave
periclinal and distinctly visible anticlinal walls.
The outer periclinal walls with ribs, which are
visible at magnification over ×500. Ribs do not
have a strict orientation.
Samples examined: 1. Odessa region, Hadzhibey,
a swamp near the village Maryanovka, 3.VII.2003. Leg.,
det. E.Y. Bondarenko, 124485 (KW); 2. Zhytomyr region,
Korostynskyi district, around village Ushtun, forest
park, quarter No. 8, 4.VI.1951, Leg., det. Barbarich (KW);
3. Kharkiv city, Zoological Garden, cultural, 30.V.1921,
Leg., det. Kozlov (KW); 4. Sumy region, Middle-Stara
Buta district, village Stara Buta, Desniansko-Starobutsky
NNP, outskirts of pine forest, 9.VI.1997, 069875. Leg., det.
Panchenko (KW).
Subsection Ochranthae Zabel emend. Rehder
Lonicera xylosteum L. (Fig. 4; Tables 1 & 2)
The infructescence consists of two fruits,
which are connate only at the base. Bracteoles
are free. The fruits are dark-red, yellow, fleshy,
spherical, 6–9 mm in diameter. The surface
of the fruits is sparsely pubescent with two-
BA
C D
Figure 2. The seed of Lonicera nigra: A – general view; B–D – surface at different magnification.
114 Plant Introduction • 85/86
O.M. Tsarenko, O.V. Bulakh, O.V. Kolesnichenko, S.M. Hrysiuk
celled glandular hairs with a spherical head
(up to 100 µm in length) and simple solitary
hairs (up to 450 µm in length). The surface of
simple hairs is verrucose. The quantity of hairs
increases at the apex of the fruit.
The seeds are light- or dark-brown,
elliptical-rounded or asymmetrical, slightly
curved, angular, 2.5–3.0 mm long and 2.0–
2.7 mm wide, flat, slightly concave on one
side, convex on the other, with a cavity on the
periphery and a hardly noticeable crest in the
center, narrow-rounded at the base. The hilum
is small, submerged, located at the base of the
seed.
The surface of the seed is shallowly pitted.
The cells of the exotesta are polygonal, with
concave periclinal and indistinct anticlinal
walls. Outer periclinal walls are covered with
a tortuous net visible at the magnification over
×500.
Samples examined: 1. Living plants from the collection
of the O.V. Fomin Botanical Garden, 20.VII.2017. Leg., det.
N.M. Belemets, O.M. Tsarenko; 2. Sumy region, Sumy
district, village Kijnytsa, Kijnitsky Arboretum, 26.V.1970,
Leg., det. Litvinenko (KWHA); 3. Transcarpathian region,
Rakhiv district, village Bogdan-Lusy (Tyschora), rocky
places, 20.V.1953, Leg., det. Komendar (KW).
Subgenus Periclymenum (L.) Rehder
Section Eucaprifolium (Spach) Pojarkova
Lonicera caprifolium L. (Fig. 5; Tables 1 & 2)
Two fruits are free. Bracteoles are also free,
small or absent. The fruits are bright-red,
fleshy, spherical or elliptic, 5–8 mm long,
4– mm wide, with 1–5 seeds. The fruits are
located on short pedicels as if they were
coalescent to the leaves.
The seeds are light- or dark-brown, broad-
elliptical, sometimes rounded, often curved,
3–6 mm long and 2–3 mm wide, somewhat
flat, slightly concave on one side, convex on
the other, with a cavity on the periphery and a
crest in the center, somewhat truncated at the
base, with a small notch. The hilum is small,
submerged, located at the base of the seed.
BA
C D
Figure 3. The seed of Lonicera tatarica: A – general view; B, C – surface; D – perforations on the internal
periclinal and anticlinal cell walls of the testa.
Plant Introduction • 85/86 115
Carpological features of Lonicera of the flora of Ukraine
The surface of the seeds is diversiform: in
some areas of the seeds, it is pitted. In others
– the cells of the exotesta are almost flat,
with tortuous anticlinal walls, the walls of the
cells are clearly visible. The outer periclinal
walls are ribbed and wrinkled (seen at the
magnification over ×1000).
Figure 4. The seed (A, B) and the fruit (C–F) of Lonicera xylosteum: A – general view of the seed; B – surface
of the seed; C, D – simple hair on the fruit; E, F – glandular hair on the fruit.
BA
C
E
D
F
116 Plant Introduction • 85/86
O.M. Tsarenko, O.V. Bulakh, O.V. Kolesnichenko, S.M. Hrysiuk
Samples examined: 1. Yalta district, near Alushta, in the
bushes of the park, 12.VIII.1959. Leg., det. Klokov (KW);
2. Living plants from the collection of the M.M. Gryshko
National Botanical Garden of NAS of Ukraine, 26.IX.2017.
Leg., det. O.M. Tsarenko; 3. Chernihiv, Agrobiostation,
Arboretum, 19.V.2010, 097764. Leg., det. Pototskaya (KW).
Lonicera periclymenum L. (Fig. 6; Tables 1 & 2)
Two free fruits, bracteoles are free too. The
fruits are red, fleshy, spherical, up to 6 mm in
diameter.
The seeds are light-brown, elliptical, ovoid,
2.0–3.5 mm long and 1.5–2.0 mm wide, flat,
slightly concave on one side, convex on the
other, with a cavity on the periphery and a
crest in the center, narrow-rounded at the
base, somewhat truncated. The hilum is small,
submerged, located at the base of the seed.
The surface of the seeds is pitted in some
areas of the seeds, and the cells of the exotesta
are almost flat, with concave or more or less
BA
C
E
D
F
Figure 5. The seed of Lonicera caprifolium: A – general view; B–F – the surface of different seed sections.
Plant Introduction • 85/86 117
Carpological features of Lonicera of the flora of Ukraine
curved periclinal and tortuous anticline walls,
the walls of the cell are distinctly visible. The
outer periclinal walls are ribbed and wrinkled
(seen at the magnification over ×1000).
Samples examined: 1. Living plants from the collection
of the M.M. Gryshko National Botanical Garden of
NAS of Ukraine, 26.IX.2017. Leg., det. O.M. Tsarenko;
2. Crimea, Suuk-su, hotel park, 11.VI.1915. Leg. Tutkovsky,
det. Mosyakin 17.I.1975 (KW).
Lonicera etrusca Santi (Fig. 7; Tables 1 & 2)
Several fruits are free, some of which are much
smaller or undeveloped. The bracteoles are
free too. The fruits are red, fleshy, spherical,
up to 8–10 mm in diameter.
The seeds are brown, broadly elliptical or
ovoid, somewhat angular, 3.0–4.0 mm long and
2.7–3.5 mm wide, flattened-convex, slightly
concave on one side, convex on the other, with
BA
C
E
D
F
Figure 6. The seed of Lonicera periclymenum: A – general view; B – hilum; C–F – surface.
118 Plant Introduction • 85/86
O.M. Tsarenko, O.V. Bulakh, O.V. Kolesnichenko, S.M. Hrysiuk
a cavity on the periphery and a crest in the
center, asymmetrically truncated at the base.
The hilum is small, submerged, located at the
base of the seed.
The seed surface is pitted. The cells of
exotesta are polygonal, with distinct borders,
the cells have equally thickened anticlinal
and concave periclinal walls. On the outer
periclinal walls, ribs are visible at magnification
over ×200.
Samples examined: Living plants from the collection of
the O.V. Fomin Botanical Garden, 20.VII.2017. Leg., det.
N.M. Belemets, O.M. Tsarenko.
Discussion
According to our results, we did not find any
characteristics that could be diagnostic at
the level of subsections or higher. We have
only found that there are some species-level
differences in the microstructure of the seed
surface. The surface of the seeds of all the
species studied is almost identical. It is mostly
pitted due to concave external periclinal walls
of the exotesta. The surface of L. xylosteum
seeds is shallowly pitted, as reported by Jacobs
et al. (2009). However, the seed surface in
L. nigra is tubercular-pitted. The seed surface
BA
C D
Figure 7. The seed of Lonicera etrusca: A – general view; B–D – surface at different magnification.
Plant Introduction • 85/86 119
Carpological features of Lonicera of the flora of Ukraine
in L. caprifolium and L. periclymenum is pitted,
but almost flat cells of the exotesta are also
present.
Our research confirmed the feasibility of
the fruit microcarpological features for more
accurate identification of species-level taxa
in the genus Lonicera. Only a few publications
devoted to microcarpology of the genus
Lonicera (Plysko, 2000; Jacobs et al., 2009;
Lavrinenko, 2012). Several micromorphological
characteristics of Lonicera species have been
applied to confirm the molecular phylogeny of
the genus (Jacobs et al., 2009). These features
also can be successfully applied for resolving
controversial questions of taxonomy, what
has been shown for Dipsacales (Tsymbalyuk
et al., 2018; Tsarenko et al., 2019) and some
rare Caryophyllales Juss. ex Bercht. et J. Presl
(Amini et al., 2011; Martynyuk et al., 2015a,
2015b, 2016, 2018; Shykhaleyeva et al., 2018).
The fruits of studied species, excepting
L. xylosteum, are reported as glabrous
(Poyarkova, 1958; Zaytsev, 1962; Browicz,
1976; Artiushenko & Fedorov, 1986; Jacobs
et al., 2009). Our observation confirmed
the absence of trichomes on the fruits of
L. caerulea, L. nigra, L. tatarica, L. caprifolium,
L. periclymenum, and L. etrusca. Only fruits of
L. xylosteum have been sparsely pubescent by
glandular two-celled headed and simple (with
Taxa Construction Color Size Shape Pubescence
Subgenus Chamaecerasus
Section Isika
Subsection Caeruleae
L. caerulea
two free fruits; of
bracteoles of paired
flowers merged in a
tubular cupule
dark blue
with bluish
cover
8–11 mm in
diameter
spherical,
hemispherical,
elliptical
glabrouse
Subsection Rhodanthae
L. nigra
two fruits connate at
the base; bracteoles
incompletely connate into
a goblet-shaped cupule
black with
bluish cover
6–10 mm in
diameter spherical glabrouse
Section Coeloxylosteum
Subsection Tataricaе
L. tatarica two free fruits, bracteoles
are free red, yellow 6–8 mm in
diameter
spherical,
sometimes
elliptical
glabrouse
Subsection Ochranthae
L. xylosteum
two connate only at the
base fruits; bracteoles are
free
dark-red,
yellow
6–9 mm in
diameter spherical
sparsely pubescent
with glandular hairs
and solitary simple
hairs
Subgenus Periclymenum
Section Eucaprifolium
L. caprifolium two free fruits; bracteoles
are free, small or absent bright-red
5–8 mm
long, 4–5
mm wide
spherical or
elliptic glabrouse
L. etrusca
several free fruits, some of
which are much smaller or
undeveloped; bracteoles
are free
red 8–10 mm in
diameter spherical glabrouse
L. periclymenum free fruits; bracteoles are
free red 6 mm in
diameter spherical glabrouse
Table 1. Morphological features of the fruit of Lonicera of the flora of Ukraine.
120 Plant Introduction • 85/86
O.M. Tsarenko, O.V. Bulakh, O.V. Kolesnichenko, S.M. Hrysiuk
Taxa Color Size Shape Surface
Subgenus Chamaecerasus
Section Isika
Subsection Caeruleae
L. caerulea light- or
dark-brown
1.5–2.3 mm long and
1.4–2.1 mm wide broadly elliptical or rounded mostly pitted
Subsection Rhodanthae
L. nigra light- or
dark-brown
2.2–2.5 mm long and
1.8–2.4 mm wide elliptical tuberculate-pitted
Section Coeloxylosteum
Subsection Tataricaе
L. tatarica light- or
dark-brown
3.0–3.5 mm long and
1.5–2.7 mm wide elliptical pitted
Subsection Ochranthae
L. xylosteum light- or
dark-brown
2.5–3.0 mm long and
2.0–2.7 mm wide
elliptical-rounded, asymmetrical,
slightly curved, angular pitted
Subgenus Periclymenum
Section Eucaprifolium
L. caprifolium light- or
dark-brown
3.0–6.0 mm long and
2.0–3.0 mm wide
broad-elliptical, sometimes
rounded, often curved
pitted and the cells of
the exotesta almost flat
L. etrusca brown 3.0–4.0 mm long and
2,7–3,5 mm wide
broadly elliptical, ovoid,
somewhat angular pitted
L. periclymenum light-brown 2.0–3.5 mm long and
1.5–2.0 mm wide elliptical, ovoid pitted and the cells of
the exotesta almost flat
Table 2. Macro- and micromorphological features of the seed of Lonicera of the flora of Ukraine.
a finely warty surface) hairs. Other observed
morphological features of the studies species
also corresponded to descriptions reported
before (Rehder, 1909; Poyarkova, 1958;
Barbarych, 1961; Zaytsev, 1962; Browicz, 1976;
Jacobs et al., 2009; Lavrinenko, 2016).
Conclusions
Based on the results of the analysis of literary
sources and our own data of macro- and
micromorphological studies of the fruits
and seeds of representatives of the genus
Lonicera, their detailed characteristics were
prepared. It should facilitate the more precise
identification of the species in the fruiting
state. A carpological feature that is specific to
representatives of the subgenus Periclymenum
is the presence of exclusively free fruits,
unlike representatives of the subgenus
Chamaecerasus with both free and connate at
the base fruits.
At the species level, we suggest using
the following features of fruits and
seeds: differences in fruits coloration of
representatives of the genus; the presence of
pseudocarp in the L. caerulea; the difference
in shape (i.e., for L. caerulea and L. tatarica
with the spherical, elliptical shape of the fruits
was also noted); the presence of glandular
and simple hairs on fruits of L. xylosteum; the
difference in the shape of the seeds (elliptic
or ovoid in L. etrusca, L. nigra, and L. tatarica,
and broadly elliptic or almost rounded in
L. caerulea, L. periclymenum, and L. xylosteum).
There are no characteristics of the
microstructure of the surface of the seed that
can be diagnostic at the level of subsections
or higher. The seeds have only some
differences at the species level. In most of
the studied species, the seed surface is pitted
– seeds of L. nigra are tubercular-pitted and
seeds of L. xylosteum with shallow-pitted
surfaces, while seeds of L. caprifolium and
L. periclymenum have almost flat exotesta cells.
Plant Introduction • 85/86 121
Carpological features of Lonicera of the flora of Ukraine
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Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України
О.М. Царенко 1*, О.В. Булах 1, О.В. Колесніченко 2, С.М. Грисюк 2
1 Інститут ботаніки імені М.Г. Холодного НАН України, вул. Терещенківська, 2, м. Київ, 01004, Україна;
* tsarenko_olga@ukr.net; anemone@ukr.net
2 Національний університет біоресурсів і природокористування України, вул. Героїв оборони, 15, м.
Київ, 03041, Україна
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Plant Introduction • 85/86 123
Carpological features of Lonicera of the flora of Ukraine
Національного ботанічного саду імені М.М. Гришка та Ботанічного саду імені О.В. Фоміна. Цитування
опрацьованих зразків представлено відповідно до оригінального тексту етикеток. Загалом було
досліджено морфологічні ознаки плодів та насінин семи видів роду (чотири з яких є інтродуцентами,
а решта – види природної флори), що вказані у “Vascular plants of Ukraine. A nomenclatural checklist”.
Для досліджень використовували світловий (МБС-9) та сканувальний електронний (JSM-6060
LA) мікроскопи. Зразки напилювали шаром золота за стандартною методикою. Дослідження
проводили при збільшенні від ×30 до ×3000. Описи плодів і насінин здійснювали з використанням
загальноприйнятої ботанічної термінології. Виміри розмірів плодів та насінин виконували з
використанням програми Axio Vision Rel. 4.8.
Результати. Представники досліджених видів мають вільні або зрослі при основі плоди, оточені
обгорткою з переважно вільних приквітничків. Псевдокарпій L. caerulea сформований бракеолями
парних квіток, які зростаються у трубчасту обгортку, що повністю і щільно обгортає вільні парні
плоди. Для L. nigra відмічено нерівномірне зростання приквітничків між собою, які вільно оточують
парні плоди, що зрослися лише при основі. Плоди досліджених видів за забарвленням чорні,
темно-сині, червоні та жовті. Для підроду Periclymenum (L. caprifolium, L. etrusca, L. periclymenum)
характерні вільні та кулясті за формою плоди. У видів підроду Chamaecerasus плоди переважно
кулясті, але у деяких представників разом з кулястими трапляються також напівкулясті, еліптичні
та видовжені (L. caerulea, L. tatarica) плоди, які можуть бути як вільними (L. caerulea, L. tatarica), так й
зрослими при основі (L. nigra, L. xylosteum). Для всіх досліджених видів, за виключенням L. xylosteum,
характерні голі плоди. Поверхня плодів L. xylosteum опушена залозистими та простими волосками.
Насінини досліджених видів за формою еліптичні, яйцеподібні, майже округлі, сплюснуті, з одного
боку ввігнуті та з іншого опуклі, з заглибленням по периферії та гребенем по центру, переважно
зі звужено-заокругленою основною. Досліджені представники секції Eucaprifolium та один вид з
секції Coeloxylosteum мають усічені при основі насінини. Поверхня насінин всіх досліджених видів
майже однакова. Вона здебільшого ямкувата, але у L. xylosteum – вона неглибоко-ямкувата, у L. nigra
– горбкувато–ямкувата, а у деяких видів відмічена не лише ямкувата структура поверхні, а також
присутність майже пласких клітин екзотести.
Висновки. За результатами опрацювання літературних джерел та на підставі аналізу власних даних
макро- та мікроморфологічних досліджень плодів та насінин представників роду Lonicera в межах
флори України складено їхні детальні характеристики, що сприятиме точнішому визначенню цих
видів у стані плодоношення. Відмічено, що у представників підроду Periclymenum плоди виключно
вільні, на відміну від підроду Chamaecerasus, у представників якого плоди можуть бути як вільними
так і зрослими при основі. Особливості забарвлення плодів, форми плодів та насінин, а також
особливості опушення плодів запропоновано використовувати для визначення видів L. caerulea,
L. caprifolium, L. etrusca, L. nigra, L. periclymenum, L. tatarica та L. xylosteum. У результаті досліджень не
виявлено ознак мікроструктури поверхні насінин, за якими можна діагностувати представників на
рівні підсекцій, секцій та підродів, проте встановлено, що вони можуть частково використовуватися
для розмежування таксонів на видовому рівні.
Ключові слова: Lonicera, плід, насінина, макроморфологічні особливості, мікроморфологічні особливості, мікроструктура
поверхні насінини, флора України
|
| id | oai:ojs2.plantintroduction.org:article-1558 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:53:40Z |
| publishDate | 2020 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/c9/98523646acc6a73eac8a2de688fccdc9.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-15582023-08-26T20:39:45Z Carpological features of Lonicera L. (Caprifoliaceae Juss.) of the flora of Ukraine Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України Tsarenko, O.M. Bulakh, O.V. Kolesnichenko, O.V. Hrysiuk, S.M. The objective of this study was to carry out macro- and micromorphological studies of fruits and seeds of representatives of the genus Lonicera of the flora of Ukraine (both natural and introduced), to clarify their features, to provide additional characteristics and to evaluate the possibility of using carpological features for the diagnosis of taxa of the genus. Material and methods. Materials of the National Herbarium of the M.G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine (KW) and Herbarium of the M.M. Gryshko National Botanical Garden of the National Academy of Sciences of Ukraine (KWHA) were used, as well as samples from living plants from the collections of the M.M. Gryshko National Botanical Garden and O.V. Fomin Botanical Garden. The citations are given by the original text of the labels. The fruits and seeds of seven species of the genus (four are introduced, and the other are species of natural flora), which are listed in “Vascular plants of Ukraine. A nomenclatural checklist”, have been studied. Morphological features of fruits and seeds were studied under the light microscope (LM, MBS-9) and scanning electron microscope (SEM, JSM-6060 LA). Samples were sprayed with a layer of gold according to the standard method. The study was conducted at magnification from ×30 to ×3000. Descriptions of fruits and seeds were performed using terminology, summarized in specialized papers. Fruit and seed sizes were measured using Axio Vision Rel. 4.8. Results. The fruits of Lonicera species are free or adnate at the base, surrounded by, for the most part, free bracteoles. A pseudocarp of L. caerulea is formed of the bracteoles of paired flowers concrescent in a tubular cupule, which completely and tightly wraps free paired fruits. The L. nigra has an uneven concrescent of the bracteoles that freely surround the paired fruits, which have concrescent only at the base. The fruits of the species Lonicera are black and dark blue, or red and yellow. The fruits of subgenus Periclymenum (L. caprifolium, L. etrusca, L. periclymenum) are free and spherical. The subgenus Chamaecerasus species’ fruits are mainly spherical, but in some species the fruits are also hemispherical, elliptical and elongated (L. caerulea, L. tatarica). Fruits in this subgenus are free (L. caerulea, L. tatarica) or coalescent at the base (L. nigra, L. xylosteum). All the studied species are characterized by glabrous fruits, except fruits of L. xylosteum that are pubescent with glandular and simple hairs. The seeds are elliptical, ovoid, nearly spherical, flat, concave on one side and convex on the other side, with a recess on the periphery and a crest in the center, at the base they are mainly narrow-rounded. All species of the section Eucaprifolium and one species of the section Coeloxylosteum have truncated seeds. The surface of the seeds is almost similar and pitted; in L. xylosteum it is shallow-pitted; in L. nigra it is tubercular-pitted. In some species, the seed surface is pitted, and also flat cells are present. Conclusions. Based on the results of a critical review of literature materials and analysis of own data, the detailed characteristics of fruits and seeds of the representatives of the genus Lonicera within the flora of Ukraine have been compiled. These characteristics are useful for more accurate identification of fruiting Lonicera plants. The species of the subgenus Periclymenum have exclusively free fruits, while the fruits in the subgenus Chamaecerasus are free or coalescent at the base. The features of color and shape of the fruits and seeds, the peculiarities of pubescence of the fruits, are suggested to be used to distinguish L. caerulea, L. caprifolium, L. etrusca, L. nigra, L. periclymenum, L. tatarica, and L. xylosteum. Analysis of the microstructure of the surface of the seeds revealed that there no features that can be used as diagnostic at the level of subsections, sections, and subgenera. However, seeds microstructure showed some differences at the species level. Мета – провести макро- та мікроморфологічні дослідження плодів та насінин природних й інтродукованих представників роду Lonicera флори України, уточнити їхні особливості, надати доповнену характеристику та оцінити можливість використання карпологічних ознак для діагностики таксонів цього роду. Матеріал і методи. Для досліджень було залучено матеріал з Національного гербарію Інституту ботаніки імені М.Г. Холодного НАН України (KW) і гербарію Національного ботанічного саду імені М.М. Гришка НАН України (KWHA), а також відібрано зразки з живих рослин у колекціях Національного ботанічного саду імені М.М. Гришка та Ботанічного саду імені О.В. Фоміна. Цитування опрацьованих зразків представлено відповідно до оригінального тексту етикеток. Загалом було досліджено морфологічні ознаки плодів та насінин семи видів роду (чотири з яких є інтродуцентами, а решта – види природної флори), що вказані у “Vascular plants of Ukraine. A nomenclatural checklist”. Для досліджень використовували світловий (МБС-9) та сканувальний електронний (JSM-6060 LA) мікроскопи. Зразки напилювали шаром золота за стандартною методикою. Дослідження проводили при збільшенні від ×30 до ×3000. Описи плодів і насінин здійснювали з використанням загальноприйнятої ботанічної термінології. Виміри розмірів плодів та насінин виконували з використанням програми Axio Vision Rel. 4.8. Результати. Представники досліджених видів мають вільні або зрослі при основі плоди, оточені обгорткою з переважно вільних приквітничків. Псевдокарпій L. caerulea сформований бракеолями парних квіток, які зростаються у трубчасту обгортку, що повністю і щільно обгортає вільні парні плоди. Для L. nigra відмічено нерівномірне зростання приквітничків між собою, які вільно оточують парні плоди, що зрослися лише при основі. Плоди досліджених видів за забарвленням чорні, темно-сині, червоні та жовті. Для підроду Periclymenum (L. caprifolium, L. etrusca, L. periclymenum) характерні вільні та кулясті за формою плоди. У видів підроду Chamaecerasus плоди переважно кулясті, але у деяких представників разом з кулястими трапляються також напівкулясті, еліптичні та видовжені (L. caerulea, L. tatarica) плоди, які можуть бути як вільними (L. caerulea, L. tatarica), так й зрослими при основі (L. nigra, L. xylosteum). Для всіх досліджених видів, за виключенням L. xylosteum, характерні голі плоди. Поверхня плодів L. xylosteum опушена залозистими та простими волосками. Насінини досліджених видів за формою еліптичні, яйцеподібні, майже округлі, сплюснуті, з одного боку ввігнуті та з іншого опуклі, з заглибленням по периферії та гребенем по центру, переважно зі звужено-заокругленою основною. Досліджені представники секції Eucaprifolium та один вид з секції Coeloxylosteum мають усічені при основі насінини. Поверхня насінин всіх досліджених видів майже однакова. Вона здебільшого ямкувата, але у L. xylosteum – вона неглибоко-ямкувата, у L. nigra – горбкувато–ямкувата, а у деяких видів відмічена не лише ямкувата структура поверхні, а також присутність майже пласких клітин екзотести. Висновки. За результатами опрацювання літературних джерел та на підставі аналізу власних даних макро- та мікроморфологічних досліджень плодів та насінин представників роду Lonicera в межах флори України складено їхні детальні характеристики, що сприятиме точнішому визначенню цих видів у стані плодоношення. Відмічено, що у представників підроду Periclymenum плоди виключно вільні, на відміну від підроду Chamaecerasus, у представників якого плоди можуть бути як вільними так і зрослими при основі. Особливості забарвлення плодів, форми плодів та насінин, а також особливості опушення плодів запропоновано використовувати для визначення видів L. caerulea, L. caprifolium, L. etrusca, L. nigra, L. periclymenum, L. tatarica та L. xylosteum. У результаті досліджень не виявлено ознак мікроструктури поверхні насінин, за якими можна діагностувати представників на рівні підсекцій, секцій та підродів, проте встановлено, що вони можуть частково використовуватися для розмежування таксонів на видовому рівні. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2020-06-30 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1558 10.46341/PI2020012 Plant Introduction; No 85/86 (2020); 109-123 Інтродукція Рослин; № 85/86 (2020); 109-123 2663-290X 1605-6574 10.46341/PI85-86 en https://www.plantintroduction.org/index.php/pi/article/view/1558/1491 Copyright (c) 2020 O.M. Tsarenko, O.V. Bulakh, O.V. Kolesnichenko, S.M. Hrysiuk http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Tsarenko, O.M. Bulakh, O.V. Kolesnichenko, O.V. Hrysiuk, S.M. Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України |
| title | Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України |
| title_alt | Carpological features of Lonicera L. (Caprifoliaceae Juss.) of the flora of Ukraine |
| title_full | Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України |
| title_fullStr | Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України |
| title_full_unstemmed | Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України |
| title_short | Карпологічні особливості Lonicera L. (Caprifoliaceae Juss.) флори України |
| title_sort | карпологічні особливості lonicera l. (caprifoliaceae juss.) флори україни |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1558 |
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