Морфологічні типи плодів у базальних покритонасінних рослин флори України
Basal angiosperms is a paraphyletic group of plants before branching eudicots and monocots. This group of angiosperms is of great scientific interest because it is the most ancient representative of the flowering plants retained for today. In Ukraine, both ANA-grade and magnoliids are presented. The...
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| description | Basal angiosperms is a paraphyletic group of plants before branching eudicots and monocots. This group of angiosperms is of great scientific interest because it is the most ancient representative of the flowering plants retained for today. In Ukraine, both ANA-grade and magnoliids are presented. The species diversity of basal angiosperms is about half a percent of the flora of vascular plants of Ukraine, which caused a low research activity on this subject. The objectives of the present study were to elucidate the morphological diversity of fruits within basal angiosperms inhabiting Ukraine and estimate their evolutionary level and adaptations. The morphological diversity of fruit, according to our concept, comprises the structure of the gynoecium, ovary insertion, placentation, ovule number (pre-anthetic features), as well as principal fruit type and morphological adaptations for dispersal (post-anthetic features). The principal fruit types were distinguished according to the principles established previously for classifying fruits of monocots and rosids of the flora of Ukraine. Within the basal angiosperms of the flora of Ukraine, four principal fruit types were established: aggregate fruit, capsule, berry, and one-seeded fruit. Among them, the aggregate fruits are the most numerous. In Nymphaeaceae and Aristolochiaceae, which belong to the native flora of Ukraine, fruits are syncarpous, multi-seeded and inferior, while in the families cultivated in Ukraine (Annonaceae, Magnoliaceae, Calycanthaceae, Schisandraceae, and Lauraceae) fruits are apocarpous, superior and often have low seed number in a carpel. Each family has peculiar gynoecium and fruit features, which can be advanced as well as ancestral. In this article, the carpological spectrum was created to reveal the distribution of principal fruit types in families, genera, and species of the basal angiosperms of the flora of Ukraine. In these spectra, aggregate fruits are the most numerous; however, they occur in families of cultivated flora only. Most species have adaptation for dispersal by animals (birds or ants). The conclusion was made that in the native flora of Ukraine, the most derived members of basal angiosperms are retained. |
| doi_str_mv | 10.46341/PI2024006 |
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Plant Introduction, 101/102, 57–65 (2024)
© The Authors. This content is provided under CC BY 4.0 license.
RESEARCH ARTICLE
Morphological fruit types of basal angiosperms in the flora of Ukraine
Anastasiya Odintsova
Ivan Franko National University of Lviv, Hrushevskyi str. 4, 79005 Lviv, Ukraine; anastasiya.odintsova@lnu.edu.ua
Received: 22.08.2024 | Accepted: 25.09.2024 | Published online: 26.09.2024
Abstract
Basal angiosperms is a paraphyletic group of plants before branching eudicots and monocots. This group
of angiosperms is of great scientific interest because it is the most ancient representative of the flowering
plants retained for today. In Ukraine, both ANA-grade and magnoliids are presented. The species diversity
of basal angiosperms is about half a percent of the flora of vascular plants of Ukraine, which caused a low
research activity on this subject. The objectives of the present study were to elucidate the morphological
diversity of fruits within basal angiosperms inhabiting Ukraine and estimate their evolutionary level and
adaptations. The morphological diversity of fruit, according to our concept, comprises the structure of the
gynoecium, ovary insertion, placentation, ovule number (pre-anthetic features), as well as principal fruit
type and morphological adaptations for dispersal (post-anthetic features). The principal fruit types were
distinguished according to the principles established previously for classifying fruits of monocots and
rosids of the flora of Ukraine. Within the basal angiosperms of the flora of Ukraine, four principal fruit
types were established: aggregate fruit, capsule, berry, and one-seeded fruit. Among them, the aggregate
fruits are the most numerous. In Nymphaeaceae and Aristolochiaceae, which belong to the native flora
of Ukraine, fruits are syncarpous, multi-seeded and inferior, while in the families cultivated in Ukraine
(Annonaceae, Magnoliaceae, Calycanthaceae, Schisandraceae, and Lauraceae) fruits are apocarpous,
superior and often have low seed number in a carpel. Each family has peculiar gynoecium and fruit
features, which can be advanced as well as ancestral. In this article, the carpological spectrum was created
to reveal the distribution of principal fruit types in families, genera, and species of the basal angiosperms
of the flora of Ukraine. In these spectra, aggregate fruits are the most numerous; however, they occur in
families of cultivated flora only. Most species have adaptation for dispersal by animals (birds or ants). The
conclusion was made that in the native flora of Ukraine, the most derived members of basal angiosperms
are retained.
Keywords: Magnoliidae, Nymphaeideae, evolution, aggregate fruit, gynoecium, fruiting ovary, seed
https://doi.org/10.46341/PI2024006
UDC 581.47 : 582.65 + 7
Funding: There are no declared financial sources for this research.
Competing Interests: The author has declared no conflict of interest.
Introduction
Basal angiosperms is a paraphyletic group
of plants before branching eudicots and
monocots (APG, 2016). Within this group,
ANA-grade and magnoliids clade are
recognized. The basal angiosperms in the
flora of Ukraine include seven families
belonging to two subclasses – Nymphaeidae
J.W.Walker ex Takht. and Magnoliidae Novak
ex Takht. (Mosyakin, 2013). According to the
checklist of the flora of Ukraine (Mosyakin
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0001-7645-3311
58 Plant Introduction • 101/102
Odintsova
& Fedоronchuk, 1999), 13 genera and about
33 species are presented within basal
angiosperms. In the native flora of Ukraine,
only members of the families Nymphaeaceae
Salisb. and Aristolochiaceae Juss. occur, while
the cultivated Laurus nobilis L. (Lauraceae)
sometimes escaped and naturalized in the
Crimea (Mosyakin, 2013). The part of the
species diversity is about 0.55 % of the flora
of vascular plants of Ukraine. This is the
lowest number of species diversity within
the main clades of angiosperms (along with
monocots and eudicots), which caused a low
research activity on this subject; however,
this group has great scientific value as
the most ancient representatives of the
flowering plants inhabiting the territory of
the study. The objectives of the study were
to elucidate the morphological diversity of
fruits within basal angiosperms inhabiting
Ukraine and estimate their evolutionary level
and adaptations.
Material and methods
The morphological analysis of fruits in early
angiosperms was carried out to compile a
spectrum of principal types of fruits and to
identify their features in comparison with
monocots and eudicots. The morphological
diversity of fruit, according to our concept
(Odintsova, 2022), comprises the pre-anthetic
features (structure of the gynoecium, ovary
insertion, placentation, ovule number) as
well as post-anthetic features (fruit type,
dehiscence and morphological adaptations
for dispersal). The annotated list of fruit
characteristics was compiled using the
sources which are provided in the results
section. The number or genera and species
within families of early angiosperms are
given according to the taxonomic synopses
(Mosyakin & Fedoronchuk, 1999; Kokhno &
Parkhomenko, 2002).
The principal fruit types were distinguished
according to the principles established for the
classification of fruits of monocots (Odintsova
et al., 2021, 2022) and rosids of the flora of
Ukraine (Odintsova, 2023). Four principal fruit
types were established for this group in the
flora of Ukraine:
Aggregate fruit – polymerous apocarpous
fruit composed of numerous or several free
carpels with one to few seeds inside each
carpel. Fruiting carpels can be follicles, berries
or achenes.
Capsule – syncarpous multi-seeded
dehiscent fruit. In early angiosperms, the
capsule is found to be inferior and hexamerous.
Berry-like fruit – indehiscent syncarpous
fruit with fleshy fruit wall.
One-seeded fruit – unilocular indehiscent
fruit with a solitary seed. It can be berry-like
or drupaceous and monomerous in the studied
material.
Results
Subclassis Nymphaeidae
Nymphaeaceae (2 genera / 3 species) –
fruit is a polymerous multi-seeded spongy
berry, carpels are numerous; placentation is
laminar (Fig. 1 A) (Schneider & Williamson,
1993). In Nymphaea L., ovary is partly inferior
(petal and stamens are adnated to the fruit
wall at different levels); carpels are free
above the syncarpous ovary, forming carpel
appendages encircling a stigmatic disc; floral
axis is projected between carpels. Fruits of
Nymphaea rupture irregularly underwater due
to the enlargement of seeds and the volume of
mucilage. Seeds are arillate, with mucilaginous
arillus.
In Nuphar Sm., the ovary is superior,
apically constricted and bear concave disc
with stigmatic rays, carpels 5–30, fused. Fruit
is ovoid, dehisces irregularly (or from the base
to tip) releasing floating aerenchymatous
white floating packets of seeds and filled by
mucilage. Seeds are nonarillate, dispersed by
water after the decay of aerenchyma (Smits
et al., 1989; Hart & Cox, 1995; Padgett, 2007;
Shiga, 2007).
According to Spjut (1994, p. 84), fruit
in Nymphaea is hesperidium, “a simple
indehiscent fruit with axile placentae and
a pericarp that is internally fleshy and
externally of a leathery rind”. The fruit in
Nuphar is ceratium, “fruit ... like a berry but
is distinguished from it by the bursting of the
outer stout layer of the pericarp; the inner
lining of each locule detaches, and floats on
the water like a sac containing the seeds”
(Spjut, 1994, p. 57). Ceratium is a capsular fruit
which opens by separation of pericarpium
layers, with persistent placentas.
Plant Introduction • 101/102 59
Morphological fruit types of basal angiosperms in the flora of Ukraine
BA
ov
cc
ov
cc
ov
DC
Figure 1. Peculiarities of fruits of early angiosperms in the flora of Ukraine: А – syncarpous polymerous
berry with laminar placentation in Nuphar lutea, preanthetic stage; B – apocarpous polymerous berry
in Schisandra chinensis; C & D – fleshy capsule in Asarum europaeum, general view (C) and longitudinally
dissected one (D). cc – central column; ov – ovule. Scale bars: 1 mm.
In Nymphaeaceae, the carpels are united
without forming an internal compitum, by the
deformation of the basically ascidiate carpels.
The floral axis expands between carpels which
appear to be plicate. Stigma is sessile, in the
stigma center there is a protrusion formed by
the alongated floral apex; ovules numerous (up
to 400 per carpel in Nymphaea) (Igersheim &
Endress, 1998).
Both genera have unlignified fruit wall
which dehisces to release seeds. In this reason,
the definition of the fruit as berry is debatable,
60 Plant Introduction • 101/102
Odintsova
like this: “The fruit of Nuphar lutea (L.) Sm. is
a fleshy capsule (or dehiscent berry)...” (Hart &
Cox, 1995). Sometimes, fruit in Nymphaeaceae
is regadred as a coenocarpous fleshy berry-
like multi-follicle, dispersed by water or
endozoochorously (Didukh et al., 2004).
Schisandraceae (1 genus / 1 species –
Schisandra chinensis (Turcz.) Baill.) – fruit is
aggregate, composed of numerous globose
1–5-seeded carpels arranged spirally on
elongated receptacle, a torus (Keng, 1993).
Fruit is baccetum – apocarpous berry (Spjut,
1994). Carpels are ascidiate, ovules 2–5
(Igersheim & Endress, 1997). The receptacle
elongates after flowering, resulting in raceme-
like aggregate fruit, with small individual
carpels, red at maturity, widely separated on
the torus, and contain 1–2(–3) seeds (Saunders,
2000). Carpels 14–40 per flower, however,
Saunders (2000) in the fig. 37 F just as in the
studied material (Lviv city, Olena Pchilka
str., cultivated), indicated some carpels as
underdeveloped in the pendent fruit (Fig. 1 B).
In the conditions of Kyiv, S. chinensis pruduces
13 to 34 red fruiting carpels per fruit with 1–2
seeds inside (Skrypchenko & Slyusar, 2020);
ovules two per carpel (Sliusar, 2018).
Subclassis Magnoliidae
Annonaceae (1 genus / 1 species – Asimina
triloba (L.) Dun.) – fruit is an apocarpous
polymerous berry composed of 3–15 carpels;
ovules numerous, biseriate; fruits dispersed
by birds or bats (Kessler, 1993). Fruit is a
baccetum, with fleshy indehiscent fruit wall,
a sarcocarp (Spjut, 1994). Carpels are plicate
(Igersheim & Endress, 1997).
Aristolochiaceae (2 genera / 2 species)
– syncarpous hexamerous capsule. It is
semi-inferior, fleshy, rupturing irregularly
at ripening (Asarum europaeum L.). In
Aristolochia clematitis L. the capsule is fully
inferior, dry, dehisces septicidal-ventrally.
Capsule septicidal or rupturing, placentation
is axile (Huber, 1993). Spjut (1994) designated
the fruit in Aristolochia L. species as ceratium
or septicidal capsule. Ovary has synascidiate
and symplicate zones in Asarum, and almost
completely symplicate in Aristolochia
(Igersheim & Endress, 1998). In Aristolochia
stamens are adnated to the style forming
a gynostemium. Ovules 2–10 per carpel,
placentation partly axile in Asarum L.; and more
that 12 ovules per carpel, placentation parietal
in Aristolochia (Igersheim & Endress, 1998). In
the studied material (Lviv city, Holosko forest),
fruits of Asarum europaeum are crowned with
three tepals, ovary is inferior, and seeds are
not numerous, bear elaiosoma (Fig. 1 C, D).
Seeds are dispersed by ants in Asarum and
by wind in Aristolochia (Didukh et al., 2004).
The research of seed dispersal of Asarum
europaeum and other spring geophytes in
beech-dominated forests in Central Europe
demonstrated that slugs Arion rufus L. (red
slug) and Limax cinereo-niger Wolf (ash-grey
slug) are common consumers of elaiosomes
or entire seeds of ant-dispersed plants
(Türke et al., 2010). The terrestrial gastropods
can generally act as seed dispersers of
myrmecochorous plants and even substitute
myrmecochory, especially where ants are
absent or uncommon (Türke et al., 2012).
It was also revealed that diaspore removal
rates in Asarum europaeum are low probably
due to the small size of ants relative to heavy
diaspores. Ant-plant relationships in that case
are not mutualistic but rather neutral/slightly
negative, because the plants do not obtain
any benefits from their interactions with ants
(Prokop et al., 2022).
Calycanthaceae (2 genera / 5 species
– Calycanthus L. with 3 species and
Chimonanthus Lindl. with 2 species) – fruit is
indehiscent, dry, the carpels few to many, free,
spirally arranged, uni-ovulate, enclosed in a
cup-shaped receptacle (a pseudocarp); ovule
basal (Kubitzki, 1993). In Calycanthus, achenes
15–35; in Chimonanthus achenes 5–15. In the
conditions of the North Ukraine, Calycanthus
species have fruits with 3–13 carpels / seeds
inside (Gavrylyuk, 2015).
Seeds are toxic for mammals, so they
germinate after dacay of external fruit
coverings in the fallen fruits. Spjut (1994)
designated the fruit as pometum – an
apocarpous fruit of carpels embedded in a
hypanthium with one cavity (like in Rosa L.).
Carpels numerous (up to 35), free, spirally
arranged, ovary plicate; ovules two, but only
one of them (the lowest) develop seed (Endress
& Igersheim, 1997). Carpellodes are situated on
the upper rim of the floral cup, and persistent
in fruit forming a woody crown around the
orifice that includes the fruitlets.
Plant Introduction • 101/102 61
Morphological fruit types of basal angiosperms in the flora of Ukraine
Lauraceae (3 genera / 3 species – Lindera
Thunb., Cinnamomum Schaeff., Laurus L.) –
fruit is a one-seeded berry or a drupe with
a weaky developed endocarp; carpel one,
uni-ovulate, ovule pendulous; receptacle
and peduncle of fruit enlarged and forming
a cupule at the base of fruit (Rohwer, 1993).
Fruits are dispersed by frugivorous birds. In
Laurus nobilis L., endocarp is lignified, with
“wavy walls” (Roth, 1977). For Cinnamomum
litsaeifolium Thwaites, Spjut (1994) used
the name of fruit – glans (ukr. – zholud) due
to the cupule, a woody floral cup; and for
C. macrocarpum Hook. fil., he uses the name
amphisarcum, a berry-like indehiscent fruit
with dry crust exocarp. Carpel one, ovary is
ascidiate; ovule one, median, pendent (Endress
& Igersheim, 1997).
Magnoliaceae (2 genera / 17 species) –
fruits apocarpous, dry, woody; carpels are
numerous, free, arranged spirally on the
elongated receptacle; ovules two per carpel;
fruiting carpels dehisce dorsally (in Magnolia
Plum. ex L.) or indehiscent caducous
samaroids (ukr. – krylatyj bahatohorishok) in
Liriodendron L. In Magnolia, seeds are 1–2 per
carpel, hanged on the elongated vessels of
funiculi from the dehiscent, or opened fruit;
testa has arilloid (Nooteboom, 1993). Carpels
are plicate with short ascidiate zone; ovules
mostly two in a carpel (Igersheim & Endress,
1997). According to Spjut (1994, p. 61), fruit in
Magnolia is a coccetum, “a multiple fruit with
dehiscent fruitlets opening along the dorsal
and ventral sutures”. In M. tripetala (L.) L.,
instead, Spjut (1994, p. 76) adopted the name
fruit-follicetum, “a multiple fruit of dehiscing
apocarps that open only along one suture,
dorsally or ventrally”. For Liriodendron, Spjut
(1994, p. 108) recognized a samaretum, “a
multiple fruit of winged indehiscent apocarps,
the wing of each apocarp exceeding the
length of the seed”. The German botanists
have a tradition to designate the fruit in
Magnolia as legumes-fruit (germ. – die Hülse;
die Hülsenfrucht; ukr. – bib). In this fruit,
the fruiting carpels are legumes, they open
by both ventral and dorsal dehiscense lines
(Leins & Erbar, 2010; Kadereit et al., 2014).
Discussion
In basal angiosperms of the flora of Ukraine,
both aggregate and simple fruits were found
to be superior or inferior, syncarpous or
monomerous apocarpous. Aggregate fruits
occur in Schisandraceae, Annonaceae,
Magnoliaceae, Calycanthaceae (Table 1). All
these families are cultivated in Ukraine and
have woody habits (Kokhno & Parkhomenko,
2002). Aggregate fruits are fleshy, berry-like
(Schisandra chinensis, Asimina triloba), dry
follicles with sarcotestal seeds (Magnolia),
or multi-achene born on the conical
receptacle (Liriodendron) or the invaginated
receptacle (Calycanthaceae). Carpels are
numerous and arranged spirally (Endress &
Igersheim, 1997; Igersheim & Endress, 1997).
In Nymphaeaceae and Aristolochiaceae the
gynoecium is congenitally syncarpous, with
a tendency to inferiority of the ovary. Fruits
are fleshy untypical berries (Nymphaeaceae)
or fleshy capsules (Asarum), or dry capsules
(Aristolochia). These two families represent
a high level of morphological progress in
the female reproductive sphere. The high
number of carpels in Nymphaeaceae may be
equivocally a result of a secondary increase as
well as a plesiomorphic condition (De Craene
et al., 2003). In Aristolochiaceae, two trimerous
condensed carpel whorls are presented. In
Lauraceae the gynoecium is apocarpous and
Basic fruit type Subclasses and families
Aggregate fruit Magnoliidae: Annonaceae *, Magnoliaceae *, Calycanthaceae *
Nymphaeidae: Schisandraceae *
Capsule Magnoliidae: Aristolochiaceae
Berry Nymphaeidae: Nymphaeaceae
One-seeded fruit Magnoliidae: Lauraceae*
Table 1. Principal fruit types in the families of basal angiosperms of the flora of Ukraine.
Note. * – cultivated taxa.
62 Plant Introduction • 101/102
Odintsova
monomerous; ovule solitary. In this family,
the terminal stage of the gynoecium and fruit
evolution is observed resulting in extremely
reduced carpels’ and ovules’ number
(De Craene et al., 2003).
Seed number in a carpel varies from
many (Nymphaeaceae, Annonaceae) to
few (Schisandraceae, Aristolochiaceae,
Magnoliaceae) and one (Calycanthaceae,
Lauraceae). In Nymphaeaceae, the highest
number of ovules within basal angiosperms
is regarded as a derived character (Endress &
Igersheim, 2000). Placentation is axile, parietal
or laminar in the gynoecium with numerous
ovules, and basal or apical in monovulate
carpels. Laminar placentation is very
uncommon for angiosperms; it is regarded as a
derived condition from marginal placentation
(Shivaprakash & Bawa, 2022).
Within studied families, somewhat
different adaptations to dispersal are evident
which cause different fruit names of the
representatives given by Spjut (1994). Namely,
in Nymphaeaceae, fruit in Nymphaea is
defined as hesperidium, and fruit in Nuphar
– as ceratium. Fruit in Magnolia is defined
as a coccetum, and fruit in Liriodendron,
as a samaretum. In Asarum europaeum and
Aristolochia clematitis capsules open by
different lines and seeds are dispersed by
different agents (wind or animals).
Within basal angiosperms of the flora of
Ukraine, there were no monomerous multi-
seeded fruits, nor schizocarps and multi-
seeded pyrenaria found. Contrary to the
monocots and rosids of the flora of Ukraine
(Odintsova et al., 2021; Odintsova, 2023), there
were no fruit with ambiguous gynoecium
type found (syncarpous or apocarpous;
monomerous or pseudomonomerous).
All studied families of basal angiosperns
have peculiar and unique features of the fruit.
In Nymphaeaceae, fruit has deformation of the
gynoecium structure during the development
and laminar placentation. In Schisandraceae,
the fruit turn to be raceme-like inflorescence
due to the elongation of the fruiting receptacle.
In Schisandraceae, like in Annonaceae, a rare
fruit type occurs, a polymerous apocarpous
berry. In Aristolochia, the gynostemium is
formed. In Calycanthaceae and Lauraceae, a
hypanthium is formed enclosing whole carpels
or only base of carpel. In Magnolia, the fruit is
polymerous legume-like (ukr. – bahatobib).
Figure 2. Distribution of principal fruit types in the
families (A), genera (B), and species (C) of basal
angiosperms in the flora of Ukraine. * – cultivated
taxa.
Plant Introduction • 101/102 63
Morphological fruit types of basal angiosperms in the flora of Ukraine
Despite the fact that basal angiosperms are
the most ancient group of flowering plants,
not only the most primitive apocarpous
fruits, but also a syncarpous inferior fruits
have been described in its members. It is
an evidence for early diversification of the
morphological structure of the gynoecium
in angiosperms. The polymerous gynoecium
in some basal angiosperms may be a reversal
derived from a whorled arrangement of
several carpels. The cyclization and reduction
of the gynoecium are common trends of the
gynoecium evolution in basal angiosperms
(De Craene et al., 2003).
In Schisandraceae, Nymphaeaceae,
Annonaceae and Lauraceae fruit wall is
fleshy or partly fleshy that is an adaptation to
dispersal by birds or water (Nymphaeaceae).
The spectra of the distribution of principal
fruit types in families, genera and species
of basal angiosperms in the flora of Ukraine
demonstrated, that the aggregate fruits are
the most numerous (Fig. 2). Capsules, berries
and one-seeded fruits are almost equally
presented. In is notable that the most derived
syncarpous and inferior fruits are revealed
only in the taxa of the native flora of Ukraine,
while aggregate and one-seeded superior
fruits available only in cultivated taxa.
Conclusions
The present research revealed four principal
fruit types within basal angiosperms of the
flora of Ukraine. The most common are
the aggregate fruits of numerous spirally
arranged berries, follicles or achenes. In
Lauraceae, the fruit is one-seeded and
one-carpellate (monomerous apocarp). In
the families of the native flora of Ukraine,
Nymphaeaceae and Aristolochiaceae, the
most derived characters of fruit occur
(syncarpous gynoecium and inferior ovary).
It could be evidence that only the most
derived representatives of the ancient group
spread on the moderate climate of Central
Europe by natural ways. The adaptation
for dispersal as well as morphological and
morphogenetic features of fruits in basal
angiosperms are revealed to be very diverse
and should be more precisely examined in
the Ukrainian flora.
Acknowledgements
This article is dedicated to my good friend
Yaroslav Khomei, who suspended his research
due to the need to defend his country from
invaders.
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Plant Introduction • 101/102 65
Morphological fruit types of basal angiosperms in the flora of Ukraine
Морфологічні типи плодів у базальних покритонасінних рослин флори України
Анастасія Одінцова
Львівський національний університет імені Івана Франка, вул. Грушевського, 4, Львів, 79005,
Україна; anastasiya.odintsova@lnu.edu.ua
Базальні покритонасінні – це парафілетична група рослин до відокремлення еудікотів та монокотів.
Ця група має велике наукове значення, оскільки охоплює найбільш давніх представників квіткових
рослин, які збереглися до сьогодні. В Україні представлені як ANA-града, так і клада магноліїд.
Видове різноманіття базальних покритонасінних становить лише приблизно півпроцента від усіх
судинних рослин України, що зумовлює незначну кількість досліджень цієї групи. Метою цього
дослідження є висвітлення морфологічної різноманітності плодів серед базальних покритонасінних
флори України та оцінка їхнього еволюційного рівня і адаптацій. Згідно з нашою концепцією,
морфологічна різноманітність плодів охоплює структуру гінецею, положення зав’язі, плацентацію,
кількість насінних зачатків (пре-антетичні ознаки) та тип плоду і морфологічні адаптації до
рознесення (пост-антетичні ознаки). Основні типи плодів виділялися згідно з попередньо
встановленими принципами, застосованими для однодольних рослин та розидів флори України.
У межах базальних покритонасінних флори України було виділено чотири основні типи плодів:
збірний плід, коробочка, ягода, однонасінний плід. Серед них збірні плоди були найбільш поширені.
У Nymphaeaceae та Aristolochiaceae, які належать до природної флори України, плоди синкарпні,
нижні, багатонасінні, позаяк у родинах з культурної флори України (Annonaceae, Magnoliaceae,
Calycanthaceae, Schisandraceae та Lauraceae) плоди апокарпні, верхні та з малонасінними плодиками.
У кожній родині були виявлені особливі риси гінецею і плоду, серед них були і просунуті, і примітивні
ознаки. У статті представлені карпологічні спектри розподілу основних типів плодів серед родин,
родів і видів базальних покритонасінних флори України. У цих спектрах збірні плоди переважають,
однак, вони виявлені виключно в родинах культурної флори. Більшість видів мають адаптації до
рознесення тваринами (птахами або мурахами). Зроблений висновок про переважання в природній
флорі України найбільш просунутих представників базальних покритонасінних.
Ключові слова: Magnoliidae, Nymphaeideae, еволюція, збірний плід, гінецей, зав’язь, насінина
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Sliusar, G. (2018). Peculiarities of Schisandra chinensis
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| id | oai:ojs2.plantintroduction.org:article-1642 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2026-03-12T15:49:12Z |
| publishDate | 2024 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/31/934a9124160b4ddea6982e119a190031.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-16422026-03-05T18:42:33Z Morphological fruit types of basal angiosperms in the flora of Ukraine Морфологічні типи плодів у базальних покритонасінних рослин флори України Odintsova, Anastasiya Basal angiosperms is a paraphyletic group of plants before branching eudicots and monocots. This group of angiosperms is of great scientific interest because it is the most ancient representative of the flowering plants retained for today. In Ukraine, both ANA-grade and magnoliids are presented. The species diversity of basal angiosperms is about half a percent of the flora of vascular plants of Ukraine, which caused a low research activity on this subject. The objectives of the present study were to elucidate the morphological diversity of fruits within basal angiosperms inhabiting Ukraine and estimate their evolutionary level and adaptations. The morphological diversity of fruit, according to our concept, comprises the structure of the gynoecium, ovary insertion, placentation, ovule number (pre-anthetic features), as well as principal fruit type and morphological adaptations for dispersal (post-anthetic features). The principal fruit types were distinguished according to the principles established previously for classifying fruits of monocots and rosids of the flora of Ukraine. Within the basal angiosperms of the flora of Ukraine, four principal fruit types were established: aggregate fruit, capsule, berry, and one-seeded fruit. Among them, the aggregate fruits are the most numerous. In Nymphaeaceae and Aristolochiaceae, which belong to the native flora of Ukraine, fruits are syncarpous, multi-seeded and inferior, while in the families cultivated in Ukraine (Annonaceae, Magnoliaceae, Calycanthaceae, Schisandraceae, and Lauraceae) fruits are apocarpous, superior and often have low seed number in a carpel. Each family has peculiar gynoecium and fruit features, which can be advanced as well as ancestral. In this article, the carpological spectrum was created to reveal the distribution of principal fruit types in families, genera, and species of the basal angiosperms of the flora of Ukraine. In these spectra, aggregate fruits are the most numerous; however, they occur in families of cultivated flora only. Most species have adaptation for dispersal by animals (birds or ants). The conclusion was made that in the native flora of Ukraine, the most derived members of basal angiosperms are retained. Базальні покритонасінні – це парафілетична група рослин до відокремлення еудікотів та монокотів. Ця група має велике наукове значення, оскільки охоплює найбільш давніх представників квіткових рослин, які збереглися до сьогодні. В Україні представлені як ANA-града, так і клада магноліїд. Видове різноманіття базальних покритонасінних становить лише приблизно півпроцента від усіх судинних рослин України, що зумовлює незначну кількість досліджень цієї групи. Метою цього дослідження є висвітлення морфологічної різноманітності плодів серед базальних покритонасінних флори України та оцінка їхнього еволюційного рівня і адаптацій. Згідно з нашою концепцією, морфологічна різноманітність плодів охоплює структуру гінецею, положення зав’язі, плацентацію, кількість насінних зачатків (пре-антетичні ознаки) та тип плоду і морфологічні адаптації до рознесення (пост-антетичні ознаки). Основні типи плодів виділялися згідно з попередньо встановленими принципами, застосованими для однодольних рослин та розидів флори України. У межах базальних покритонасінних флори України було виділено чотири основні типи плодів: збірний плід, коробочка, ягода, однонасінний плід. Серед них збірні плоди були найбільш поширені. У Nymphaeaceae та Aristolochiaceae, які належать до природної флори України, плоди синкарпні, нижні, багатонасінні, позаяк у родинах з культурної флори України (Annonaceae, Magnoliaceae, Calycanthaceae, Schisandraceae та Lauraceae) плоди апокарпні, верхні та з малонасінними плодиками. У кожній родині були виявлені особливі риси гінецею і плоду, серед них були і просунуті, і примітивні ознаки. У статті представлені карпологічні спектри розподілу основних типів плодів серед родин, родів і видів базальних покритонасінних флори України. У цих спектрах збірні плоди переважають, однак, вони виявлені виключно в родинах культурної флори. Більшість видів мають адаптації до рознесення тваринами (птахами або мурахами). Зроблений висновок про переважання в природній флорі України найбільш просунутих представників базальних покритонасінних. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2024-09-26 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1642 10.46341/PI2024006 Plant Introduction; No 101/102 (2024); 57-65 Інтродукція Рослин; № 101/102 (2024); 57-65 2663-290X 1605-6574 10.46341/PI101-102 en https://www.plantintroduction.org/index.php/pi/article/view/1642/1554 Copyright (c) 2024 Anastasiya Odintsova http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Odintsova, Anastasiya Морфологічні типи плодів у базальних покритонасінних рослин флори України |
| title | Морфологічні типи плодів у базальних покритонасінних рослин флори України |
| title_alt | Morphological fruit types of basal angiosperms in the flora of Ukraine |
| title_full | Морфологічні типи плодів у базальних покритонасінних рослин флори України |
| title_fullStr | Морфологічні типи плодів у базальних покритонасінних рослин флори України |
| title_full_unstemmed | Морфологічні типи плодів у базальних покритонасінних рослин флори України |
| title_short | Морфологічні типи плодів у базальних покритонасінних рослин флори України |
| title_sort | морфологічні типи плодів у базальних покритонасінних рослин флори україни |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1642 |
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