Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція
The morphostructure of Oxalis purpurea bulbs during the growing season, with an emphasis on the number and functional purpose of their scales, has been investigated. It was found that two protective membranous scales, two leathery scales, and three fleshy scales are constantly present in the investi...
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2026
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Plant Introduction| _version_ | 1860145182084694016 |
|---|---|
| author | Zhila, Alla Tymchenko, Olga Vakulenko, Tetyana |
| author_facet | Zhila, Alla Tymchenko, Olga Vakulenko, Tetyana |
| author_sort | Zhila, Alla |
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| datestamp_date | 2026-01-15T02:04:21Z |
| description | The morphostructure of Oxalis purpurea bulbs during the growing season, with an emphasis on the number and functional purpose of their scales, has been investigated. It was found that two protective membranous scales, two leathery scales, and three fleshy scales are constantly present in the investigated species. The first fleshy scale has a narrow hyaline edge. Additionally, the phenomenon of metatopy in a representative of the Oxalidaceae family and recaulescence in a bulbous plant are described for the first time. |
| doi_str_mv | 10.46341/PI2025017 |
| first_indexed | 2026-02-08T08:11:58Z |
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© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 109, 14–19 (2026) ISSN 1605-6574, e-ISSN 2663-290X
RESEARCH ARTICLE
Oxalis purpurea L. (Oxalidaceae): bulb morphostructure, recaulescence
Alla Zhila *, Olga Tymchenko, Tetyana Vakulenko
M.M. Gryshko National Botanical Garden of the NAS of Ukraine, Sadovo-Botanichna str. 1, 01103 Kyiv, Ukraine;
* allazhila58@gmail.com
Received: 08.10.2025 | Accepted: 25.11.2025 | Published: 14.01.2026
Abstract
The morphostructure of Oxalis purpurea bulbs during the growing season, with an emphasis on the
number and functional purpose of their scales, has been investigated. It was found that two protective
membranous scales, two leathery scales, and three fleshy scales are constantly present in the investigated
species. The first fleshy scale has a narrow hyaline edge. Additionally, the phenomenon of metatopy in a
representative of the Oxalidaceae family and recaulescence in a bulbous plant are described for the first
time.
Keywords: Oxalidaceae, Oxalis purpurea, bulbs, scales, metatopy, recaulescence
https://doi.org/10.46341/PI2025017
UDC 582.751.1 : 581.446.2
Authors’ contributions: Conceptualization – A. Zhila. Formal analysis – A. Zhila, O. Tymchenko. Methodology – A. Zhila, T. Vakulenko.
Software – T. Vakulenko. Validation – A. Zhila, O. Tymchenko, T. Vakulenko. Writing, original draft – A. Zhila. Writing, review & editing
– A. Zhila, O. Tymchenko, T. Vakulenko.
Funding: Research project Nr. 409-OR “Induction of phenotypic plasticity of tropical and subtropical plants: search for predictive
markers of resistance” (state registration number 0123U101383, budget program code 6541030).
Competing Interests: The authors declare no conflict of interest.
Introduction
Southern African bulbous representatives of
the genus Oxalis L. (Oxalidaceae) comprise
about 270 species, of which 122 originate
from the Cape Floristic Kingdom (Freiberg &
Manning, 2013). The species Oxalis purpurea L.
is natively distributed in the Western Cape,
a region in the winter-rainfall part of South
Africa (Goldblatt & Manning, 2000). In its
native range, this species is considered
endemic, classified as LC (SANBI, 2025).
However, O. purpurea is a highly decorative
groundcover plant and is therefore widely
cultivated in other parts of the world. This
species, characterized by the presence of true
bulbs, also occurs in Ukraine.
The taxonomic structure and nomenclatural
history of the genus Oxalis are complicated. In
particular, the taxonomic revision of Southern
African Oxalis based on morphological
characters (Salter, 1944) does not align with
palynological findings (Dreyer, 1996) and
recent phylogenetic studies (Oberlander
et al., 2004, 2009, 2011; Oberlander, 2009).
However, it is considered the most complete.
Considering Salter (1944), O. purpurea belongs
to the section Stictophyllae. Following the
POWO (2025) list, it has 34 heterotypic
and two homotypic synonyms. Several
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0003-1246-1376
https://orcid.org/0000-0002-5691-5264
https://orcid.org/0000-0002-6141-6689
Plant Introduction • 109 15
Oxalis purpurea L. (Oxalidaceae): bulb morphostructure, recaulescence
morphological synapomorphies support the
clade Purpurea, which combines Southern
African bulbous Oxalis, based on molecular
data (Gebregziabher, 2004; Jooste et al., 2016).
However, the discrepancy between molecular
phylogeny and morphological data (Oberlander
et al., 2011; Vaio et al., 2013) highlights the need
for searching for additional morphological
characters that allow distinguishing the taxa
within this clade.
Oxalis purpurea is a species with wide
edaphic tolerance (Haukka et al., 2013), a
high level of genetic variability (Becker et al.,
2022), an extremely high level of vegetative
reproduction, pronounced self-compatibility,
and substantial damage resistance (Zietsman
et al., 2008). In disturbed habitats, O. purpurea
often becomes a malicious weed, included
in the GCW (2007). It is a winter-growing
dwarf synanthous geophyte (Harrower, 2009)
with immensely shortened internodes in the
aboveground part of the shoot, with trifoliate
leaves collected in basal rosettes, and a short
rhizome-like underground part of the shoot
continuing into a bulb.
Descriptions of the bulb structure of
Southern African Oxalis differ even within
the same species, as they were carried out
at different ontogenetic stages (Zhila &
Tymchenko, 2020). According to Moodley
(1988), O. purpurea has one of the simplest
bulb structures, the seasonal growth of which
begins with the onset of winter rains and has
a developmental cycle typical for all bulbous
Oxalis species from this region.
Thus, the bulbs of Southern African Oxalis
consist of the following structures: a basal
plate, outer protective leathery scales, inner
nourishing fleshy scales, and a bud that will
develop in the next growing season (Salter,
1944; Moodley, 1988; Gebregziabher, 2004).
A sprouted bulb with an already formed
aboveground part has a similar structure.
Levyns (1974), who described the annual
development cycle of O. purpurea, reported
one more type of scale that forms on the basal
plate at the beginning of bulb development
– thin and short-lived, disappearing before
the bulb matures. When studying the
morphostructure of bulbs during growth,
another type of scale, membranous, is noted,
located on the underground part of the shoot,
often referred to as the ‘rhizome’ (Zhila &
Tymchenko, 2020).
This work aimed to comprehensively
analyze the features of shoot formation in
the underground part of O. purpurea plants,
including the type of bulb scales, their number,
and functional purpose, as well as the activity
of axillary meristems.
Material and methods
Oxalis purpurea plants were introduced into
the M.M. Gryshko National Botanical Garden
of the NAS of Ukraine. Observations were
conducted during the 2019 and 2023–2024
growing seasons. Plants were kept in pot
culture in a cool greenhouse. During the
growing season, three plants were dug up once
a month to establish the bulb morphostructure
by sequentially removing scales. The
morphostructures were described and drawn
using an Lomo MBS-9 light microscope.
The applied descriptive terminology follows
Gebregziabher (2004) with additions of
Zhila & Tymchenko (2020). Descriptions and
measurements of buds formed on fleshy scales,
in five replicates, were made using a Carl Zeiss
Stemmi 2000 microscope at magnifications
×10 to ×80.
Results and discussion
The bulb in O. purpurea is ovoid, approximately
1 cm in diameter. The outer covering scales are
blackish-brown and sticky. Leaves are petiolate
(up to 5 cm long, widened at the base), leaflets
are dark green, 1–3 cm long, 1–3 cm wide,
sparsely hairy on the abaxial side and along
the edges, without oxalate deposits. Leaflets
are flat, not folded. Flowers are pentamerous,
located solitary on a hairy peduncle up to 11
cm long. Sepals are up to 1 cm long, lanceolate,
and hairy. Petals up to 2.5 cm long, purple
but always yellow at the base. Under the test
conditions, the fruits did not form.
The bulb of O. purpurea is imbricate, with
few scales. In a state of relative dormancy,
the bulb consists of a shortened section of
the main shoot axis (basal plate) with scales
developed on it, and a terminal bud that
produces the main shoot axis (Pütz, 1994).
The terminal bud of the main shoot axis has
a monopodial growth pattern (Jeannoda-
Robinson, 1977). After a period of relative
16 Plant Introduction • 109
Zhila et al.
dormancy, thread-like roots (Fig. 1: 2) begin to
grow from the mother bulb around the basal
plate (Fig. 1: 3). The main shoot axis partially
remains in the bulb and partially became
located between the bulb and the soil surface
forming so-called ‘rhizome’, on which thread-
like roots also develop (Fig. 1: 12). The rest of
the main shoot axis develops above the soil
surface. Subsequently, one or two thread-
like roots around the basal plate thicken
and accrete with the basal plate, forming a
contractile taproot-like roots (Fig. 1: 1; Zhila &
Tymchenko, 2016).
The first two internodes of the main shoot
axis with membranous scales at the nodes
(Fig. 1: 9), in the axils of which buds (bulbils)
are formed (Fig. 1: 10), are called ‘threads’ (Pütz,
1994). They are shortened and can elongate
due to intercalary growth with simultaneous
contraction of the contractile root at the end
of the growing season. At the beginning of root
Figure 1. Schematic illustration of bulb and main
shoot axis structure in Oxalis purpurea: 1 – rachis-
like contractile basal root; 2 – basal thread-like
root; 3 – basal plate; 4 – thin evanescent scales;
5 – protective membranous scales on the basal
plate; 6 – leathery scale; 7 – pediculate bud on the
first fleshy scale; 8 – first fleshy scale with hyaline
edge; 9 – first membranous scale on the main axis
of the shoot (‘thread’); 10 – axillary bulb embedded
under the first membranous scale; 11 – first
elongated internode; 12 – thread-like adventitious
root on the stem.
contraction, the scales attached to the basal
plate of the bulb detach from it and remain
in their original position. The internode
continues to elongate further (Fig. 1: 11).
The elongation of the ‘threads’ and the
contraction of the root can reach 30 cm
in O. pes-caprae L. (Pütz, 1994). The buds
located on the ‘threads’ develop into renewal
bulbs and typically get the same size as the
mother bulb (Pütz, 1994). Axillary buds are
present at all bulb scales (Pütz, 1994; Zhila &
Tymchenko, 2020) and, during the contraction
of the contractile root, can be displaced to a
considerable distance from the mother bulb.
Gebregziabher (2004) investigated
O. purpurea bulbs from two localities when
the plants were in the floral state. The bulbs of
both specimens varied in terms of the length
of the so-called ‘rhizome’, their diameter
(a difference of 1.5 times), the length of the
beak of the covering scales, the number of
fleshy scales, and the presence of trichomes
on different parts. Common features for both
specimens were: a narrow basal part (basal
plate); the presence of two types of covering
scales – golden-brown membranous and
honey-brown leathery; sticky secretory cells
on the lower half of the outer surface of the
covering scales; a beak-like apex of the bulb;
the presence of a narrow membranous edge
on the largest fleshy scale.
In the bulbs of the studied specimens, the
presence of scales on the basal plate (Fig. 1: 3)
was established in the following sequence:
1) Thin, short-lived scales (Fig. 1: 4).
2) Two protective membranous scales
(Fig. 1: 5) with a smooth surface, where the
inner surface is sticky. The first scale is semi-
enclosing, the second is very wide, covering
almost the entire bulb.
3) Two firm leathery scales (Fig. 1: 6) with
pubescent beaks.
4) Three fleshy scales, different in width,
almost equal in length. The first scale (up to
12.5 mm long) is the longest and has a narrow
hyaline edge (Fig. 1: 8), on which pediculate
buds (Fig. 1: 7) are formed.
Commonly, seed plants are hemiaxillar –
their lateral shoots develop only from buds
in the axils of leaves (Esau, 1977). However,
some species have shoots (vegetative or
generative) that appear in a supra-axillary
position. Such displacement is referred to as
metatopy (Endress, 2010). To date, two types of
Plant Introduction • 109 17
Oxalis purpurea L. (Oxalidaceae): bulb morphostructure, recaulescence
metatopy are well known: axial displacement
(concrescence) and epiphyllous displacement
(recaulescence). Epiphyllous displacement
(recaulescence) is the development of a
lateral bud on its covering leaf (Troll, 1937).
Recaulescences usually occur in flowers and
inflorescences, for example, in Bougainvillea
glabra Choisy from Nyctaginaceae (Cooper,
1932), Phyllonoma integerrima (Turcz.) Loes.
from Phyllonomaceae (Dickinson & Sattler,
1974), and Helwingia japonica (Thunb.) F.Dietr.
from Helwingiaceae (Dickinson & Sattler, 1975).
In studied O. purpurea bulbs, in all three
fleshy scales, the axillary bud is located not
in the axil of the scale but on the midrib of
its adaxial surface, above the proximal end.
Hence, an epiphyllous displacement of the
lateral bud was observed (Fig. 2 A, B). The
displaced pediculate buds were conical in
shape, with pointed apices.
There are no known reports of metatopy in
Oxalidaceae, and in bulbous plants, it has not
been observed at all. Therefore, our report is
the first in this regard.
Conclusions
The phenomenon of recaulescence on the
fleshy scales of O. purpurea bulbs discovered
by us demonstrates the need for further
research on the structure of Oxalis bulbs,
which may have potential taxonomic
significance. This phenomenon, previously
unreported for Oxalidaceae and bulbous
plants, is documented for the first time here.
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Plant Introduction • 109 19
Oxalis purpurea L. (Oxalidaceae): bulb morphostructure, recaulescence
Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція
Алла Жила *, Ольга Тимченко, Тетяна Вакуленко
Національний ботанічний сад імені М.М. Гришка НАН України, вул. Садово-Ботанічна, 1, Київ, 01103,
Україна; * allazhila58@gmail.com
Досліджено морфоструктуру цибулин Oxalis purpurea упродовж вегетаційного періоду з акцентом на
оцінку кількісних показників та функціональне призначення лусок. Встановлено, що у досліджуваного
виду сталим є число лусок, що складається з двох захисних плівчастих лусок, двох шкірястих і трьох
м’ясистих лусок. Перша м’ясиста луска має вузький гіаліновий край. Крім того, вперше описано
явище метатопії у представника родини Oxalidaceae та рекаулесценції у цибулинної рослини.
Ключові слова: Oxalidacea, Oxalis purpurea, цибулина, луски, метатопія, рекаулесценція
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| resource_txt_mv | wwwplantintroductionorg/a5/36b9c80ad6eb433dc7f9fb2aaf4aaaa5.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-16802026-01-15T02:04:21Z Oxalis purpurea L. (Oxalidaceae): bulb morphostructure, recaulescence Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція Zhila, Alla Tymchenko, Olga Vakulenko, Tetyana The morphostructure of Oxalis purpurea bulbs during the growing season, with an emphasis on the number and functional purpose of their scales, has been investigated. It was found that two protective membranous scales, two leathery scales, and three fleshy scales are constantly present in the investigated species. The first fleshy scale has a narrow hyaline edge. Additionally, the phenomenon of metatopy in a representative of the Oxalidaceae family and recaulescence in a bulbous plant are described for the first time. Досліджено морфоструктуру цибулин Oxalis purpurea упродовж вегетаційного періоду з акцентом на оцінку кількісних показників та функціональне призначення лусок. Встановлено, що у досліджуваного виду сталим є число лусок, що складається з двох захисних плівчастих лусок, двох шкірястих і трьох м’ясистих лусок. Перша м’ясиста луска має вузький гіаліновий край. Крім того, вперше описано явище метатопії у представника родини Oxalidaceae та рекаулесценції у цибулинної рослини. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2026-01-14 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1680 10.46341/PI2025017 Plant Introduction; No 109 (2026): Early view; 14-19 Інтродукція Рослин; № 109 (2026): Early view; 14-19 2663-290X 1605-6574 en https://www.plantintroduction.org/index.php/pi/article/view/1680/1579 Copyright (c) 2026 Alla Zhila, Olga Tymchenko, Tetyana Vakulenko http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Zhila, Alla Tymchenko, Olga Vakulenko, Tetyana Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція |
| title | Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція |
| title_alt | Oxalis purpurea L. (Oxalidaceae): bulb morphostructure, recaulescence |
| title_full | Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція |
| title_fullStr | Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція |
| title_full_unstemmed | Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція |
| title_short | Oxalis purpurea L. (Oxalidaceae): морфоструктура цибулини, рекаулесценція |
| title_sort | oxalis purpurea l. (oxalidaceae): морфоструктура цибулини, рекаулесценція |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1680 |
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