Four multipurpose species of the genus Vitex (Lamiaceae) in the Democratic Republic of the Congo show different responses to propagation techniques for nursery production

Abstract

Citer l'article / To cite the article Numbi Mujike D., Meerts P., Mbinga Lokoto B., Ngoy Shutcha M., 2023. Four multipurpose species of the genus Vitex (Lamiaceae) in the Democratic Republic of the Congo show different responses to propagation techniques for nursery production. Bois et Forêts des Tropiques, 357 : 43-56. Doi : https://doi.

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TECHNIQUES DE MULTIPLICATION DE VITEX SP. / RECHERCHE
Bois et Forêts des Tropiques – ISSN : L-0006-579X
Volume 357 – 3e trimestre – octobre 2023 – p. 43-56 43
Désiré N M1, 2
Pierre M2, 3, 4
Boniface M L1
Mylor N S1
1 University of Lubumbashi
Faculty of Agronomic Sciences
Ecology, Ecological Restoration
and Landscape Research Unit
Route Kasapa, Campus universitaire
BP 1825, Lubumbashi
Democratic Republic of the Congo
2 Université libre de Bruxelles
Laboratory of Plant Ecology
and Biogeochemistry
Avenue F.D. Roosevelt 50
CP 244, Brussels
Belgium
3 Meise Botanic Garden
Nieuwelaan 38, Meise
Belgium
4 Service général de l’Enseignement
supérieur et de la Recherche
scientifique
Fédération Wallonie-Bruxelles, Brussels
Belgium
Auteur correspondant /
Corresponding author:
Désiré NUMBI MUJIKE –
desire.mujike@gmail.com
mujiken@unilu.ac.cd
:
https://orcid.org/0000-0001-8735-2153
Doi : 10.19182/bt2023.357.a36943 – Droit d’auteur © 2023, Bois et Forêts des Tropiques – © Cirad – Date de soumission : 8 avril 2022 ;
date d’acceptation : 25 avril 2023 ; date de publication : 1er octobre 2023.
Licence Creative Commons:
Attribution - 4.0 International.
Attribution-4.0 International (CC BY 4.0)
Citer l’article / To cite the article
Numbi Mujike D., Meerts P., Mbinga Lokoto B., Ngoy Shutcha M., 2023. Four
multipurpose species of the genus Vitex (Lamiaceae) in the Democratic
Republic of the Congo show dierent responses to propagation tech-
niques for nursery production. Bois et Forêts des Tropiques, 357 : 43-56.
Doi : https://doi.org/10.19182/bt2023.357.a36943
Four multipurpose species of the genus
Vitex (Lamiaceae) in the Democratic Republic
of the Congo show dierent responses
to propagation techniques for nursery production
Photos 1.
a. Young Vitex doniana plants in the shadehouse. b. Budding of
Vitex doniana roots in a propagation frame. c. A plantation of
species of the genus Vitex in the experimental garden of UNI-
LU’s Faculty of Agronomic Sciences. d. Rooted marcotte of Vitex
fischeri in Lubumbashi. e. Rooted root cutting of Vitex madiensis
in Lubumbashi. f. Rooted marcotte of Vitex madiensis in Lubu-
mbashi. g. Cutting roots of Vitex mombassae in a propagation
chassi in Lubumbashi. Fragment of a rooted root cutting of Vitex
doniana in a propagation frame in Lubumbashi. h. Fragment of
a rooted root cutting of Vitex doniana in a propagation frame in
Lubumbashi.
Photos D. Numbi Mujike, 2019.
a
c
e
g
b
d
f
h

RESEARCH / PROPAGATION TECHNIQUES FOR VITEX SP.
Bois et Forêts des Tropiques – ISSN: L-0006-579X
Volume 357 – 3rd quarter – October 2023 – p. 43-56
44
RÉSUMÉ
Quatre espèces polyvalentes du genre
Vitex (Lamiaceae) de la République
démocratique du Congo réagissent
diéremment aux techniques de
multiplication pour la production en
pépinière
Les plantes médicinales ligneuses
constituent l’un des principaux produits
écosystémiques fournis par la forêt claire
sèche du HautKatanga (République démo-
cratique du Congo). La déforestation rapide,
et la demande croissante en médicaments
traditionnels mettent en péril la sécurité de
l’approvisionnement. Dans ce contexte, la
domestication et la production en culture
peuvent représenter une solution durable.
Les espèces du genre Vitex (Lamiaceae)
sont bien connues pour leur utilisation en
médecine traditionnelle. Cette étude vise à
optimiser les techniques de propagation de
quatre espèces du genre Vitex (V.doniana,
V.madiensis subsp. milanjiensis, V.fischeri,
V.mombassae) qui coexistent dans la forêt
claire sèche du HautKatanga. Quatre tech-
niques, à savoir le semis, le bouturage de
racines, le bouturage de tiges et le mar-
cottage aérien, ont été testées. Les quatre
espèces montrent des réponses remarqua-
blement contrastées. Vitexmadiensis n’est
pas apte à la multiplication générative (0%
de taux de germination), mais les graines
mûres scarifiées ont enregistré des taux de
levée plus élevés chez V.doniana (90 %),
tout comme les graines scarifiées récoltées
à l’état vert chez V.mombassae (75%). Le
bouturage de racines a été plus perfor-
mant chez V. doniana (87%), V. madien-
sis (80 %) et V. mombassae (77 %), mais
pas pour V. fischeri (0%). Le marcottage
aérien convient bien à V.fischeri (83%) et
V.madiensis (77%), mais pas pour V.mom-
bassae (0%) dans les conditions de notre
étude. Les boutures de tiges n’ont été e-
caces pour aucune des quatre espèces. Des
protocoles spécifiques ecaces pour la
propagation représentent un premier pas
vers la domestication de ces espèces en
République démocratique du Congo.
Mots-clés: marcottage, bouturage, semis,
multiplication végétative, domestication,
Vitex, plante médicinale, miombo,
République démocratique du Congo.
ABSTRACT
Four multipurpose species of
the genus Vitex (Lamiaceae) in
the Democratic Republic of the
Congo show dierent responses to
propagation techniques for nursery
production
Woody medicinal plants are one of the
main ecosystem products found in the dry
woodlands of the province of HautKatanga
(Democratic Republic of the Congo), where
rapid deforestation coupled with increasing
demand for traditional medicine are jeo-
pardising supplies. In this context, domesti-
cation and cultivation may be a sustainable
solution. The species of the genus Vitex
(Lamiaceae) are well known for their uses
in traditional medicine. This study aims to
optimise propagation techniques for four
species of the genus Vitex (V. doniana,
V.madiensis subsp. milanjiensis, V.fischeri,
V.mombassae) that coexist in the dry woo-
dlands of Haut-Katanga. Four techniques,
i.e., sowing, root cuttings, stem cuttings and
air layering, were tested experimentally.
The four species show strikingly contras-
ting responses. Sowing was ill-suited to
V. madiensis (0% germination rate), but
scarified ripe seeds produced high germi-
nation rates in V.doniana (90%) as did sca-
rified seeds harvested from unripe (green)
seeds of V.mombassae (75%). Root cuttings
were most suitable for V. doniana (87%),
V. madiensis (80%) and V. mombassae
(77%), but not for V.fischeri (0%). Air laye-
ring was well suited for V.fischeri (83%) and
V.madiensis (77%), but not for V.mombas-
sae (0%) in the conditions of our study. Stem
cuttings were not ecient for any of the
four species in the conditions of our study.
Eective and specific propagation protocols
would be a first step towards domestication
of these species in the Democratic Republic
of the Congo.
Keywords: sowing, root cuttings,
stem cuttings, air layering, vegetative
propagation, domestication, Vitex,
medicinal plant, miombo, Democratic
Republic of the Congo.
RESUMEN
Cuatro especies polivalentes del género
Vitex (Lamiaceae) de la República
Democrática del Congo reaccionan
de forma diferente a las técnicas de
multiplicación para la producción en
vivero
Las plantas medicinales leñosas consti-
tuyen uno de los principales productos eco-
sistémicos proporcionados por el bosque
abierto seco del Alto Katanga (República
Democrática del Congo). La deforestación
rápida, y la creciente demanda de medica-
mentos tradicionales ponen en peligro la
seguridad del aprovisionamiento. En este
contexto, la domesticación y la producción
en cultivo pueden representar una solu-
ción sostenible. Las especies del género
Vitex (Lamiaceae) son bien conocidas por
su uso en medicina tradicional. Este estu-
dio pretende optimizar las técnicas de pro-
pagación de cuatro especies del género
Vitex (V. doniana, V. madiensis subsp.
milanjiensis, V.fischeri, V.mombassae), que
coexisten en el bosque abierto seco del
Alto Katanga. Se ensayaron cuatro técnicas,
es decir, el sembrado, los esquejes de raíz,
los esquejes de tallos y el acodo aéreo. Las
cuatro especies muestran respuestas clara-
mente contrastadas. Vitex Madiensis no es
apta para la multiplicación generativa (0%
de tasa de germinación), pero las semillas
maduras escarificadas registraron tasas de
crecimiento más elevadas en V. doniana
(90 %), al igual que las semillas escarifi-
cadas recogidas siendo verdes en V.mom-
bassae (75%). El esqueje de raíces fue más
efectivo en V. doniana (87%), V.madien-
sis (80%) y V.mombassae (77%), aunque
no para V.fischeri (0 %). La reproducción
por esquejes aéreos es conveniente para
V.fischeri (83%) y V.madiensis (77%), pero
no para V.mombassae (0%) en las condi-
ciones de nuestro estudio. Los esquejes de
tallos no fueron eficaces para ninguna de
las cuatro especies. Los protocolos especí-
ficos eficaces para la propagación repre-
sentan un primer paso hacia la domesti-
cación de estas especies en la República
Democrática del Congo.
Palabras clave: acodo, reproducción
por esqueje, siembra, multiplicación
vegetativa, domesticación, Vitex, planta
medicinal, miombo, República Democrática
del Congo.
D. N M, P. M, B. M L,
M. N S

TECHNIQUES DE MULTIPLICATION DE VITEX SP. / RECHERCHE
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Volume 357 – 3e trimestre – octobre 2023 – p. 43-56 45
Introduction
Medicinal plants are used in the treatment of many
diseases in traditional African medicine. Although tradi-
tional medicine cannot be reduced to herbal medicine
alone, the vast majority of traditional recipes and treat-
ments are plant-based. In the Democratic Republic of the
Congo (D.R. Congo), more than 80% of the population uses
traditional medicine to treat themselves (Okombe et al.,
2014; Bakariet al., 2017) with, in particular, a massive use
of woody medicinal plants (Lejolyetal., 1992; Vwakyana-
kazi and Petit, 2004; Muyaetal., 2014; Kondaet al., 2012;
Mbayoetal., 2016; Mongekeetal., 2018; Rusaatietal., 2021).
The dry tropical woodlands in the south of the country,
also known as miombo woodlands, are being rapidly
degraded by the unsustainable harvesting of wood for char-
coal making, inappropriate wildfire practices and cleared
for slash-and-burn agriculture, mining activities, and urba-
nization (Potapovetal., 2012; Usenietal., 2017). This repre-
sents a serious threat to woody medicinal plants. At the
same time, the demand for medicinal plants is increasing
due to population growth. Supply challenges could lead to
substitution with other plants, which may be inactive or
even toxic, thus threatening the health and livelihoods of
the people depending on them (Yamanietal., 2015).
In this context, strategies for conservation and sus-
tainable use of medicinal plants need to be developed
(Maghembeetal., 1998; Akinnifesietal., 2007; Meunieretal.,
2008; Leakey and Van Damme, 2014; Mapongmetsem and
Diksia, 2014). Cultivation of overexploited plants and exclu-
sively harvested in the wild would be a sustainable alterna-
tive as it would not only reduce pressure on wild popula-
tions, but also secure supply and generate more income for
households (Simons and Leakey, 2004; Amujoyegbeetal.,
2012). Cultivation also has the advantage of combating
adulteration by providing reliable botanical identification.
However, lack of knowledge on the most appropriate pro-
pagation techniques is one of the main constraints to culti-
vation (Gbenatoetal., 2014).
Research is currently being developed in dierent
regions of tropical Africa to assess the domestication
potential of overexploited or endangered medicinal plants.
This is the case of, among others, Irvingia gabonensis
(Atangana et al., 2001), Prunus africana and Pausinysta-
lia johimbe (Tchoundjeu et al., 2002, 2004), Spathodea
campanulata (Meunier et al., 2006), Warburgia salutaris
(Hannweg et al., 2016), Ximenia americana (Fawa et al.,
2015), etc.
In the Katanga region, several species of the genus
Vitex (V.madiensis, V.mombassae, V.doniana and V.fischeri)
of the Lamiaceae family are well known by local communi-
ties for their medicinal and food (fruit) value. Ethnobotani-
cal studies in this and neighboring regions have shown that
these species are widely used in traditional medicine to
treat various diseases including diabetes, diarrhea, asthma,
gastrointestinal parasitosis, anemia, etc. (Augustino, 2011;
Bruschietal., 2014; Bakarietal., 2017; Mpasiwakomu, 2021;
Rusaatietal., 2021).
Vitex doniana, the species whose propagation has
been most extensively studied in Central and West Africa,
has been shown to have a stronger ability to propagate by
root cuttings compared to stem cuttings (Sanoussietal.,
2012; Mapongmetsem et al., 2016a). Germination is
enhanced by scarification (Ahotonet al., 2011; Neyaetal.,
2017b), because seeds are surrounded by a hard endocarp
(physical dormancy). Available data on the propagation of
V.madiensis show that the species is unsuitable for seeding
(Mapongmetsem, 2006). The size (diameter at chest height)
of the mother plant (parent) could influence the perfor-
mance of the propagation material (Mapongmetsemetal.,
2016b). Further investigations on the propagation in ex-situ
conditions of other species of the genus Vitex occurring in
Katanga would make available to users’ protocols for culti-
vation, a necessary step for domestication.
In this paper, we compare propagation protocols for
seeds, root and stem cuttings, and aerial layering for these
four Vitex species (V. doniana, V. fischeri, V. madiensis,
V.mombassae). We test i) if the protocols developed for
V.doniana in Central and West Africa can be applied to other
species of the same genus, or whether, specific protocols
should be used for each of these four species; ii) whether,
at the intraspecific level, the success of the propagation
methods is influenced by phenotypic characteristics of the
material, especially the vigour of the cuttings (stem or root).
Material and methods
Study environment
The study area, called “Plaine de Lubumbashi” (Schmitz,
1971), is located in the south of the Haut-Katanga province, in
D.R. Congo. The experiments were conducted in-situ and in
the experimental field of the Faculty of Agronomic Sciences
of the University of Lubumbashi (UNILU) (E27°48’61’’,
S11°61’55,3’’; 1,257m a.s.l.). The climate is characterized by
a 5-month rainy season (November-March) and a 5-month
dry season (May-September) separated by two transition
months (October and April). The average annual rainfall is
ca. 1,300mm. The average annual temperature is 20°C with
the lowest temperatures in June-July (temperatures < 10°C
can be recorded at night) and the highest temperatures
in October (maximum temperatures of 39°C have been
recorded in recent years). Ferralsols are the major soil type
of the region. The most common vegetation formation, is
miombo woodland, i.e. dry tropical woodland dominated by
three legume genera of the Caesalpinioideae subfamily, i.e.
Brachystegia, Julbernardia, and Isoberlinia (Campbelletal.,
2007). Industrial and artisanal mining and slash-and-burn
agriculture are the main activities of the population of the
area.

Plant material
The genus Vitex comprises ca. 270 species of trees and
shrubs, formerly included in the Verbenaceae family, but
which phylogenetic classification now places in the Lamia-
ceae family (Harleyetal., 2004). They grow mostly in tropi-
cal regions and are easily recognized by their opposite, pal-
mate compound leaves. The fruit is a drupe with a 4-celled
pit, each cell containing a seed (Mabberley, 2017). Vitex
doniana is widely distributed in tropical Africa; V.fischeri
and V. mombassae are distributed in east and southern
Africa, while V.madiensis subsp. milanjiensis is restricted
to the Zambezian region. For more details on the morpholo-
gical traits of the four-study species, see Meerts (2018) and
Paton and Meerts (2020). Within the miombo woodland, the
four species occupy slightly dierent niches, with V.fischeri
occurring almost exclusively on termite mounds, on soil
with high base cation contents (Nkuluetal., 2022).
Germination trial
The seeds were collected from five sites: Mikembo
(V. fischeri, V. madiensis and V. mombassae), Lwiswi-
shi (V.fischeri, V. madiensis and V. mombassae), Kasapa
(V.doniana, V.fischeri, V.madiensis and V. mombassae),
Village Katuba (V.doniana) and Kipopo (V.doniana). The
sites of origin of propagation materials are presented in
figure1 and appendixI. These sites were chosen based on
the availability of the material. In each site and for each
species, an average of five vigorous individuals (parent), ca.
200m from each other and in good health were selected.
As V.madiensis and V.mombassae showed variation in the
leaflet number in all sites, seeds were collected from indivi-
duals with five leaflets, individuals with 3leaflets and indi-
viduals with 3&5leaflets (in the same tree). Twelve fruits
were harvested from each individual. Seeds were sown in
a manner that allowed identification of the parent plant in
the nursery. Parent plant characteristics are presented in
appendixI.
A total of 160fruits per species were harvested and
sown in March2019 (end of the rainy season). Fruits were
manually pulped with a knife. The pits were divided into four
dormancy-breaking pre-treatment batches (manual scarifi-
cation, soaking in boiling water until cooled, soaking in tap
water at room temperature for 24h, untreated control) and
two maturity levels (ripe fruit and green fruit). Ripe fruits
Figure 1.
Location of the sites of origin of propagation materials (Mujike, 2022).
27°22’30’’E 27°27’0’’E 27°31’30’’E 27°36’0’’E 27°40’30’’E
27°22’30’’E 27°27’0’’E 27°31’30’’E 27°36’0’’E 27°40’30’’E
11°46’30’’S 11°42’0’’S 11°37’30’’S 11°33’0’’S 11°28’30’’S
11°46’30’’S 11°42’0’’S 11°37’30’’S 11°33’0’’S 11°28’30’’S
INERA Kipopo
Campus Kasapa
Experimental garden
Sample plot
Lubumbashi town border
Ferme Jacaranda
02,5 5 10 km
Village Mimbulu
Ecole Chemaf
Légende
Village Katuba
Lwiswishi Mikembo
Congo (Democratic Republic of the)
Kinshasa
Lubumbashi
Angola
Zambia
Malawi
Mozambique
Tanzania
Rwanda
Burundi
Kenya
Uganda
Keny
a
Sudan
Ethiopia
Central African Republic
Chad
Nigeria
GabonCongo
RESEARCH / PROPAGATION TECHNIQUES FOR VITEX SP.
Bois et Forêts des Tropiques – ISSN: L-0006-579X
Volume 357 – 3rd quarter – October 2023 – p. 43-56
46

TECHNIQUES DE MULTIPLICATION DE VITEX SP. / RECHERCHE
Bois et Forêts des Tropiques – ISSN : L-0006-579X
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were identified by the blackish (V. doniana, V. fischeri,
V.madiensis) or brownish (V.mombassae) color of the peri-
carp. The ripe fruits were collected on the ground, while the
green fruits were collected from the tree. Scarification was
done by breaking the pit with a hammer. The use of green
fruits was chosen because Vitex seeds are known to be phy-
sically dormant (Belhadjetal., 1998), and we thought that
the endocarp of fruits harvested in the ripe state would be
strongly lignified and could therefore prevent the contact
of water and embryo necessary for any germination. The
green fruits would have a less lignified (porous) endocarp.
The black polyethylene bags of 12cmx20cm containing the
soil of the experimental garden (of the ferralsol type) were
installed under a shade in a rectilinear way (in rows spaced
of 10cm, with a spacing of 15cm).
The experiment was conducted in a randomized block
design of eight treatments repeated twenty times with three
factors: species, dormancy breaking pre-treatment and fruit
maturity level. One pit per bag was sown at a depth of about
2cm. Germination rate, lag time (period between sowing
and first germination), number of leaves and plant height
were observed. Approximately 22cl of water was supplied
daily to every bag for six months.
Root cuttings
This experiment was conducted in February 2019,
near the end of the rainy season. Propagation materials
were collected in four sites: Chemaf (V.doniana), Lwiswi-
shi (V.fischeri, V. madiensis and V. mombassae), Kasapa
(V.doniana, V.fischeri, V.madiensis and V.mombassae) and
Kipopo (V.doniana). In average ten trees per species and
per site were selected for collection of one root (diameter
1.1–2.5cm) within a radius of 1.5m around the parent, fol-
lowing the protocol of Sanoussietal. (2012) and Mapong-
metsemetal. (2016a). The mother plants have on average
a minimum diameter of 5cm and a maximum of 15 cm at
breast height. The leaf trait characteristics were the same
as those of the seedlings.
A refrigerated box was used to transport the roots to
the nursery. The roots were cut into 15cm long fragments
and placed horizontally in the propagator at a depth of
about 2-3cm. The experimental design was a randomized
block repeated nine times with two factors: (i) the species
and (ii) the rooting substrate (decomposed sawdust and
potting soil). Five root cuttings were planted for each treat-
ment (species x substrate). The propagators are installed in
a shadehouse.
A total of 360root cuttings, i.e. 90per species, were
grown. The propagator is a wooden box of 3m length by
1m width and 1m height divided into three equal compart-
ments, following the model of Leakeyetal. (1990). Recovery
rate, lag time, number of stems, leaves, roots, root length,
rooting rate (proportion of cuttings with at least one root)
and plant height were observed. A cutting was considered
as rooted when it had at least one root of 1cm in length
(Mapongmetsem et al., 2016b). Photos 1 illustrate some
images of the results obtained in the propagator (a), aerial
layering (b) and plants in black polyethylene bags (c).
Stem cuttings
This experiment was conducted in April 2019. The
collection sites were the same as for the root cuttings
and mother plants with the same characteristics. A total
of 240stem cuttings (60 per species) of about 30 cm in
length and 2.5-4 cm in diameter (Sanoussi et al., 2012;
Gbenatoet al., 2014; Mapongmetsem et al., 2016ab) were
harvested and cultured in black polyethylene bags of
16cmx30cm in a factorial design with three blocks. Soil
rich in organic matter constituted the substrate of culture.
Prior to plantation, cuttings were soaken for a few seconds
in a solution of auxin (Rootone). The bags were arranged in
a straight line and placed under a shade (with a shade net
that covers the entire nursery). A quantity of 15cl of water
was given to the cutting in the morning and evening for six
months. Budding rate, rooting rate, number of new stems,
number of leaves and stem height were monitored for six
months.
Aerial layering
Aerial layering was carried out between December
2019 and January 2020 (in the middle of the rainy sea-
son; trees are in fruiting state), in seven sites: Lwiswishi
(V.fischeri, V.madiensis and V.mombassae), INERA Kipopo
(V. fischeri, V. madiensis and V. mombassae), Chemaf
(V.doniana), Katuba (V.doniana and V.mombassae), Mim-
bulu 2 (V.madiensis) and Jacaranda farm (V.fischeri and
V. madiensis). Marcottes were prepared from orthotropic
branches of 3-7cm in diameter, with a maximum of 2mar-
cottes per individual. The mother plants had a range dbh of
5-12cm (V.doniana) and 4-10cm (V.fischeri, V.madiensis
and V.mombassae). A total of 360marcottes were prepared,
i.e. 30marcottes per site for each species. The experimental
design was a main factor design with species and sites as
factors. It was not possible to evaluate the species x site
interaction because the four species where not present in
the same sites. Thus, the number of sites varied among the
species.
Each marcotte was wrapped in translucent plastic film.
The plastic sleeve was filled with decomposed sawdust
as a rooting substrate. To avoid contamination, the knife
was regularly washed and disinfected. The substrate was
moistened every month with a syringe. Rooting rate and lag
time were our observed parameters.
Influence of mother plants
The morphotype (i.e. 3 or 5 leaflets) and diameter
at breast height (DBH) of all individuals or mother trees
used for seed collection, stem and root cuttings and aerial
layering were recorded in all sites in order to assess their
influence on the seedling’s performance (Momoetal., 2017).
Statistical analysis
For the germination trial, ANOVA was not applied to
test the eect of treatment on the germination rate as there

was only one seed per pot for each treatment. The number
of germinations for each treatment was calculated based on
the twenty pots in the experimental device.
Two-way ANOVA without replication was applied to
the data from lag time, number of leaves and seedling
height. For the lag time, species and blocks were the main
factors. For V.fischeri, two-way ANOVA was applied to test
the eect of dormancy breaking (scarification and soaking
in cold water) and blocks. Fruit maturity and blocks were
the main factors for V.mombassae. The eect of interac-
tions (scarification x block, dormancy breaking x block
and fruit maturity x block), were not calculated.
As for root cuttings, three-way ANOVA without repli-
cation was used to test the species eect, the rooting
substrate eect, the block eect as well as the species x
rooting substrate interactions.
For aerial layering, two one-way ANOVA was perfor-
med successively. First, one-way ANOVA was applied to
test the influence of site for each species separately. As
there was no significant eect of site, all the sites were
pooled and a one-way ANOVA was applied to test the
eect of species (all sites pooled).
A one-factor ANOVA was also per-
formed to test the eect of the mor-
photype (i.e. number of leaflets) of the
mother plant on the performance of
the propagation material. To test the
eect of mother plant diameter on the
performance of the propagation mate-
rial, multivariate analyses of variance
(MANOVA) were performed between DBH
and the dierent variables (emergence
rate, recovery rate, number of leaves per
plant, plant height, etc.).
In order to respect the normality
and homogeneity of the variance of the
residuals, the data of lag time, number
of leaves, roots and plant height were
transformed into Log 10 while those
expressed in percentage were transfor-
med into square root. The method used
to compare mean values was the least
significant dierence at the 5% proba-
bility level (LSD) Tukey HSD test. All the
statistical analyses were performed
using R sotware 64.4.0.5.
Figure 2.
Germination rate of four species of Vitex. 160 seeds per species. MF = mature fruit,
GF = green fruit, SC = scarified fruit, SW = soaked in water at room temperature,
TC = soaked in boiling water, T = control.
90
65
0
0
60
65
0 00 0
75
00 0 0 0
0000000000000000
0
10
20
30
40
50
60
70
80
90
100
Vitex donianaVitex fischeriVitex mombassaeVitex madiensis
Germination rate (%)
MF+SCMF+SWGF+SC
GF+SWMF+TCGF+TC
MF+T GF+T
Figure 3.
Lag time of germination of mature, scarified fruits in three
species of Vitex. 160 seeds per species. A, b, c: values sharing
the same letter are not significantly dierent at the 5% pro-
bability threshold (Tukey’s test).
c
a
b
0
10
20
30
40
50
60
70
Vitex donianaVitex fischeriVitex
mombassae
Lag time (days)
Figure 4.
Number of leaves per plant as well as plant height of Vitex in the nursery. Comparison of MF+SC treatment by species.
160 seeds per species. a, b, c: values sharing the same letter are not significantly dierent at the 5% probability threshold
(Tukey test). Legend: MF = mature fruit, SC = scarified fruit.
a
c
b
0
5
10
15
20
25
30
35
40
45
Vitexdoniana Vitexfischeri Vitex
mombassae
Height
(
cm
)
Treatment MF + SC
a
a
b
0
2
4
6
8
10
12
14
16
18
Vitexdoniana Vitexfischeri Vitex
mombassae
Number of leaves
Treatment MF + SC
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48

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Results
Germination
Ater six months, no V. madiensis seeds germi-
nated regardless of fruit maturity and pretreatment to
break dormancy. Similarly, there was no germination with
the untreated controls, with the combination of Green
FruitxBoiling Water Dip and Green FruitxRoom Tempera-
ture Water Dip regardless of species. The other three spe-
cies showed dierent germination patterns (figure2). For
V.doniana, only seeds from ripe and manually scarified fruit
germinated with a rate of 90% (18/20). For V.fischeri, germi-
nation was recorded only for seeds from ripe fruits scarified
manually (13/20, 65%) or soaked in tap water (12/20, 60%).
For V.mombassae, germination occurred only for manually
scarified seeds from green fruits (15/20, 75%) or from ripe
fruits (13/20, 65%).
ANOVA (appendicesII) shows that the lag time, the
number of leaves as well as plant height are significantly
influenced by the species (p = 0.000). The shortest lag
time was obtained with scarified ripe seeds of V.doniana
(18±4days), followed by scarified ripe seeds of V.fischeri
(33 ± 3 days) while scarified ripe fruits of V. mombassae
recorded the longest lag time (45±2days) (figure3). The
eect of block was not significant for all observed parame-
ters.
In V.fischeri, lag time, number of leaves per plant and
plant height were not influenced by the dormancy breaking
technique (manually scarified ripe fruits and cold water
soaked ripe fruits) (figure 4). In V.mombassae, none of the
parameters were influenced by the physiological maturity of
the seed. ANOVA results are presented in the appendicesII.
Root cuttings
Ater 6 months, the recovery rate ranged from 0%
(V. fischeri) to 79 ± 7% for V. mombassae, 80 ± 4% for
V.madiensis and 87±3% for V.doniana. The lag time was
about 75days for V.doniana and V.madiensis and about 85
days for V.mombassae.
No parameter was significantly influenced by the
block. The rooting substrate significantly influenced the
recovery rate, root emission rate and plant height (p<0.05).
The species eect is very highly significant on all observed
parameters (p<0.05). The interaction of species with roo-
ting substrate was significant on recovery rate, lag time,
number of leaves and root emission rate (p<0.05). Figure 5
shows the results obtained for budding rate, recovery rate,
lag time and number of leaves per plant. The other parame-
ters were not significant. The table summarizing the results
of the ANOVAs is presented in appendixIII.
a
c
c
d
bcc
d
0
10
20
30
40
50
60
70
80
90
100
Vitex donianaVitex fischeriVitex
mombassaeVitex madiensi s
Root emission rate (%)
SawdustBlack soil
a
d
c
b
e
c
0
10
20
30
40
50
60
70
80
90
100
Vitex donianaVitex mombassaeVitex madiensis
Recovery rate (%)
SawdustBlack soil
bbb
b
a
b
0
20
40
60
80
100
120
Vitex donianaVitex mombassae Vitex madiensis
Lag time (days)
SawdustBlack soil
a
bc c
ab
b
c
0
5
10
15
20
25
Vitex donianaVitex mombassae Vitex madiensis
Number of leaves
SawdustBlack soil
Figure 5.
Root cuttings of species of the genus Vitex. Budding rate, recovery rate, lag time and number of leaves per plant ater six months.
Mean ± Standard deviation. 360 root cuttings, 90 per species, 30 per treatment (5 x 9 replicates). a, b, c: values sharing the same letter
are not significantly dierent at the 5% probability threshold (Tukey’s test).

Stem cuttings
The recovery rate of stem cuttings was 25 ± 9%
(V.madiensis), 30±10% (V.fischeri), 43±10 (V.doniana)
and 63±15% (V.mombassae). The lag time was 22±4days
(V. mombassae), 24± 4 days (V. madiensis), 29 ± 3 days
(V.doniana) and 34±3days (V.fischeri). However, no roots
were produced, regardless of species. A mortality rate of
100% was recorded for all cuttings four months ater plan-
ting in pots. The figure illustrating the evolution of the sur-
vival rate of stem cuttings in the nursery is presented in
appendixIV.
Aerial layering
Data were pooled ater non-significant ANOVA test
for site eect. The results obtained show that no rooting
of marcottes could be obtained for V.mombassae. For the
other three species, the rooting rate was high, ranging
from 67±5% (V.doniana), 77±5% (V.madiensis) to 83±4%
(V.fischeri). The lag time was 99 ±20days (V.madiensis),
99±14days (V.fischeri) and 108±16 days (V.doniana).
The influence of species was very highly significant on
rooting rate (F(3,8)= 5903.94; p=0.000) as well as on lag
time (F(2,201)=6.55, p=0.002). Figure 6 shows schemati-
cally the results for root emission rate and the lag time. The
results of the ANOVAs for the site eect are presented in
appendixV.
Influence of the mother plant
The MANOVA results showed a positive significant
influence of mother plant diameter on the recovery of
root cuttings of V. doniana (P= 0.000) and V. mombas-
sae (p = 0.006). The diameter of mother plants also had
a positive significant influence on the number of roots of
V.doniana (p=0.002). The recovery and the number of roots
increased with the diameter of the mother plants. For all
the other propagation techniques and parameters there
was no significant influence of mother plant diameter.
Discussion
Dierent responses of the four Vitex species tested
We have compared the suitability of dierent germi-
nation techniques in four congeneric species of Vitex, a
genus of woody species occurring in tropical Africa. Apart
from the negative results obtained for stem cuttings, and
the lack of germination of unscarred seeds for all the spe-
cies, there were striking dierences in the responses of
the four species to all the other techniques. The failure of
stem cuttings under the conditions of our study is surpri-
sing because previous works used this technique success-
fully for V.doniana and V.madiensis (Mapongmetsemetal.,
2016a; Gbenato et al., 2014; Sanoussi et al., 2012). Future
work should use cuttings from 1-or 2-yold shoots and/or
test other growth conditions.
Concerning the germination of seeds, our results show
that all the species have physical dormancy (lignified endo-
carp), and require dormancy breaking. This confirms pre-
vious results for V.doniana and V.madiensis (Belhadjetal.,
1998; Mapongmetsem, 2006; Gbenatoet al., 2014; N’Dani-
kouetal., 2014; Neyaetal., 2017ab). Interestingly, the spe-
cies show contrasting response to the dierent dormancy
breaking techniques, with V. fischeri seeds being able to
germinate ater soaking in water at room temperature while
only mechanical scarification with a hammer was ecient
for the other species. This is likely due to the smaller pit
size and thinner endocarp of V.fischeri. V.madiensis subsp.
milanjiensis was found to be unsuitable for propagation by
seed. In Western Africa, dicult germination of the other
subspecies of V.madiensis (subsp. madiensis) was reported
by Mapongmetsem (2006). Seed viability should be assessed
in V.madiensis. Nkulu (2022) found very high rates of selfing
Figure 6.
Aerial potting of species of the genus Vitex. Root emission rate and lag time ater 150 days of observations. Total number of marcottes was
360 or, 90 per species. a, b, c, d: Values sharing the same letter are not significantly dierent at the 5% probability threshold (Tukey’s test).
cb
a
d
0
10
20
30
40
50
60
70
80
90
100
Vitex donianaVitex fischeriVitex
mombassae Vitex
madiensis
Emission of roots (%)
a
bb
0
20
40
60
80
100
120
140
Vitex donianaVitex fischeriVitex madiensis
Lag time (days)
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in V.madiensis, therefore inbreeding depression could pos-
sibly aect seed viability.
For all the other techniques, the contrasting responses
of the four species are surprising, because all our species
co-occur in sympatry in the dry woodlands of Haut-Katanga.
Interestingly, the four species occupy somewhat dierent
niches. In particular, V.fischeri occurs only on high Macro-
termes termite mounds, which have quite dierent soil
conditions compared to the surrounding woodland (Muji-
nyaetal., 2010; Cuma Mushagalusaetal., 2020). V.madiensis
and V.mombassae are never found on termite mounds, and
tend to occur in dierent plant communities (Nkuluet al.,
2022). It is possible that the regeneration niches of the four
species are somewhat dierent.
Previous studies (Sanoussi et al., 2012; Mapongmet-
sem, 2006; Mapongmetsem and Diksia, 2014; Mapong-
metsem et al., 2016a,b) suggested that V. doniana and
V.madiensis were more suitable to multiplication by root
cuttings than stem cuttings. This is confirmed by the results
of this study. Interestingly, the high recovery rate of root
cuttings of V.doniana can be related to the frequent obser-
vation of suckers in this shallow-rooted species in nature
(Mujike, 2022). The complete failure of aerial layering in
V.mombassae, in contrast to the other species, is intriguing
and deserves further investigation.
Conclusion : Practical implications
for seedling production and pro-
pagation of the four Vitex species
The objective of this study was to identify the most
suitable propagation techniques for the four species of the
genus Vitex from miombo in Haut-Katanga through a com-
parison of several propagation methods in a domestication
perspective. This has made it possible to provide users with
the first data on the propagation of V.fischeri and V.mom-
bassae. The results show that the same protocol does not
apply to all four species. Each of them presents one or two
most appropriate modes of propagation. Stem cuttings were
identified as the least appropriate mode of propagation for
the four species under the conditions of this study. These
results lead to the following practical recommendations for
the propagation of Vitex species in Haut-Katanga (tableI).
All the results, especially those from the root cuttings,
showed a more significant eect of species compared to
substrate (sawdust and potting soil). This suggests that the
use of sawdust or potting soil in the production of propaga-
ting material of the four Vitex will depend on their availa-
bility.
Acknowledgments
We sincerely thank the authorities of ARES for having
financed this study in the framework of the doctoral
research through the PHYTOKAT project.
Funding
This research was made possible by funding from the Bel-
gium Kingdom through ARES as part of the PHYTOKAT pro-
ject.
Data access
The data collected in the field and used for the statistical
analysis and writing of this article are freely available on the
ZENODO platform. https://zenodo.org/deposit/8355009#
Table I.
Preferred propagation techniques for four Vitex species occurring in Haut-Katanga dry tropical woodlands.
Propagation technique Vitex doniana Vitex fischeriVitex madiensis
subsp. milanjiensis Vitex mombassae
Stem cuttings
----
Root cuttings ++ -+
++
+
Aerial layering ++++
+-
Seed germination
(soaking in water) -+
--
Seed germination
(mechanical scarifi cation) ++ +-++
- = poor, + = good, ++ = very good.

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52
Table II.
Parent plant characteristics. Characteristics of seedlings (mother
plants) of 4 species of the genus Vitex used in the experiments.
Species Vegetative propagation Generative propagation
Sample (n) DBH (cm) Sample (n) Fruit diameter (cm)
Vitex doniana 15 5-13 30 1.8±0.1
Vitex fi scheri 15 6-12 30 0.7±0.06
Vitex madiensis 15 5-10 30 1.6±0.1
Vitex mombassae 15 5-10 30 2.2±0.2
DBH = diameter at breast height.
Table III.
E ect of species for the scarifi ed wall fruit treatment. 2-factor ANOVA
without replication: ***: p < 0.01, **: p < 0.01, *: p < 0.05, NS: p > 0.05.
Observed parameters Source of variation F value P value
Lag time Block (11,22) = 1.30 NS
Species (2,22) = 34.8 ***
Number of leaves Block (11,22) = 1.243 NS
Species (2,22) = 13.2 ***
Plant height Block (11,22) = 1.24 NS
Species (2,22) = 65.
4*
**
Table IV.
E ect of scarifi cation vs. cold water soaking for V. fi scheri, 2-factor ANOVA
without replication: ***: p<0.01, **: p<0.01, *: p<0.05, NS: p>0.05.
Observed parameters Source of variationF value P value
Lag time Block (11,11) = 2.17 NS
Treatment (1,11) = 2.92 NS
Number of leaves Block (11,11) = 0.3
6N
S
Treatment (1,11) = 12.69 **
Plant height Block (11,11) = 0.82
3N
S
Treatment (1,11) = 6.4
1*
*
Appendix I.
Appendices II: ANOVA germination.

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Table V.
E ect of fruit maturity vs. scarifi ed seeds of V. mombassae. Two-factor ANOVA
without replication: ***: p<0.01, **: p<0.01, *: p<0.05, NS: p>0.05.
Observed parameters Source of variation F value P value
Lag time Block (12,12) = 0.558 NS
Treatment (1,12) = 2.63
0N
S
Number of leaves Block (12,12) = 1.28
1N
S
Treatment (1,12) = 0.357 NS
Plant height Block (12,12) = 2.91
0N
S
Treatment (1,12) = 0.178 NS
Table VI.
E ect of species, substrate and interaction (sub x sp) on performance of root
cuttings. 3-factor ANOVA without replication: ***: p<0.01, **: p<0.01, *: p<0.05,
NS: p>0.05.
Observed parameters Source of variation F value P value
Budding rate
Block (8,40)=1.07
5N
S
Substrate(1,40)=5.16
5*
Species (2,40)=41.636 ***
Sub. x sp. (2,40)=6.701 **
Lag time
Block (8,40)=1.35
1N
S
Substrate(1,40)=1.53
9N
S
Species (2,40)=14.043 ***
Sub. x sp. (2,40)=7.268 **
Number of leaves
Block (8,40)=1.11
0N
S
Substrate(1,40)=0.069 NS
Species (2,40)=24.626***
Sub. x sp. (2,40)=8.512***
Recovery rate
Block (8,40)=2.98 NS
Substrate(1,40)=41.98 ***
Species (2,40)=4,095.97 ***
Sub. x sp. (2,40)=27.60 ***
Number of roots
Block (8,40)=1.817 NS
Substrate(1,40)=0.77
2N
S
Species (2,40)=25.714 ***
Sub. x sp. (2,40)=1.09
7N
S
Plant height
Block (8,40)=0.719 NS
Substrate(1,40)=19.006 ***
Species (2,40)=19.960 ***
Sub. x sp. (2,40)=1.231 NS
Appendix III: ANOVA root cuttings.

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54
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Table VII.
One-factor ANOVA to test the layering site e ect: ***: p<0.01, **: p<0.01,
*: p<0.05, NS: p>0.05.
Species Observed
parametersF value P value
Vitex doniana Lag time (2,57)=0.0
7N
S
Recovery rate (2,8)=0.057 NS
Vitex fi scheri Lag time (2,72)=3.30 NS
Recovery rate (2,8)=0.092 NS
Vitex mombassae Lag time (2,66)=6.9
4N
S
Recovery rate (2,8)=5.49 NS
Appendix IV: Stem cuttings.
Appendix V: ANOVA aerial layering.
Figure 7.
Stem cuttings. Evolution of the survival rate of stem cuttings in the nursery. Do = V. doniana; Fi = V. fischeri;
Ma = V. madiensis; Mo = V. mombassae.
0
10
20
30
40
50
60
70
80
90
100
12345678910111213141
51
6
Survival rate (%)
Weeks
Do Fi Ma Mo

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Numbi Mujikeetal. – Contribution des auteurs
Contributor role
Conceptualization
Formal Analysis
Funding Acquisition
Investigation
Methodology
Resources
Validation
Visualization
Writing – Review & Editing
Contributor names
D. Numbi Mujike, M. Ngoy Shutcha, P. Meerts
D. Numbi Mujike, M. Ngoy Shutcha, P. Meerts
P. Meerts
D. Numbi Mujike, B. Mbinga Lokoto
D. Numbi Mujike, M. Ngoy Shutcha, P. Meerts
B. Mbinga Lokoto
M. Ngoy Shutcha, P. Meerts
M. Ngoy Shutcha, P. Meerts
D. Numbi Mujike, M. Ngoy Shutcha, P. Meerts