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Original Research Article
Plant Cell Biotechnology and Molecular Biology 22(31&32):86-95; 2021 ISSN: 0972-2025
Amblystegium serpens AND Oxyrrhynchium speciosum
TWO NEW SPECIES FOR THE BRYOFLORA OF
MOROCCO
KHALID ACHOUAL
*
, IMANE FADEL, AOMAR DABGHI, KAMAL SAADI,
JAMILA DAHMANI AND NADIA BELAHBIB
Laboratory of Plants, Animals Productions and Agro-Industry, Faculty of Sciences, Ibn Tofail University,
BP 133, Kénitra 14000, Morocco [KA, IF, AD, KS, JD, NB].
[
*
For Correspondence: E-mail:
khalidachowal1987@gmail.com]
Article Information
Editor(s):
(1)
Dr. Nehal S. El-Mougy, National Research Centre, Egypt.
Reviewers:
(1)
Sawsan Reda Khader, Kuwait University, Kuwait.
(2)
Caren Lorensi, UNIVAP, Brazil.
Received: 08 February 2021
Accepted: 13 April 2021
Published: 23 April 2021
_______________________________________________________________________________________________
ABSTRACT
During an in-depth bryological survey of the Tafoughalet massif, a site of biological, ecological and
archaeological interest located in the north-east of Morocco, two new species were identified in the
bryophytic flora of Morocco. These species are pleurocarpous mosses: Amblystegium serpens
belonging to the Amblystegiaceae family, a mesophilic species collected on limestone rocks and on
the trunks of fig trees, then Oxyrrhynchium speciosum belonging to the Brachytheciaceae family, a
hygrophilic species colonising highly humid soils. These two taxa are qualified as very rare within
the study area.
Keywords: Amblystegium serpens; Oxyrrhynchium speciosum; Mosses; Tafoughalt Massif; Morocco.
INTRODUCTION
Within the framework of a study devoted to the
elaboration of a catalogue of the bryophyte flora
of the Tafoughalt massif in the North-East of
Morocco [1], a region which has never been
studied in depth by bryologists, we have collected
two new mosses for the bryophyte flora of
Morocco: Amblystegium serpens and
Oxyrrhynchium speciosum. In Morocco, the genus
Amblystegium comprises two taxa [2,3], while the
genus Oxyrrhynchium is represented by four
species [3,2].
This work presents the descriptive sheet for each
of these species based on the main morphological
and anatomical characteristics that have made it
possible to distinguish them from related species,
also specifying their distribution and ecology.
MATERIALS AND METHODS
Study Site
The study area is located in the area of the rural
commune of Tafoughalt, which lies at the heart of
the Beni Snassen range. It is a site of biological
Achoual et al.
87
and ecological interest (SIBE) located in the
north-east of Morocco [4]. The Beni-Snassen
range is a mountainous massif 130 km long and 25
km wide [5,6], which also includes the Tafoughalt
massif. This chain is bounded to the north by the
Triffa plain and the Kebdana chain, to the west by
the Beni Bou Yahi chain, to the south by the
neogenous Guercif-Angad basin and to the east by
the Traras chain, in Algeria [7]. The highest point
is Ras-Foural with an altitude of 1532 m [8]. The
dominant stage of vegetation in this region, the
whole Beni Snassen chain, is of thermo-
Mediterranean and Meso-Mediterranean type
according to the classification of Bagnuls and
Gaussen [9] with beautiful stands of Aleppo pine
(Pinus halepensis), and it corresponds to the
rainiest area in north-eastern Morocco [10]. The
bioclimate is of a semi-arid Mediterranean type
with alternating cold and wet and hot and dry
seasons. It is characterised by a spatio-temporal
irregularity of rainfall and by intense and violent
showers [11].
The village of Tafoughalt lies at an altitude of 850
m [10], 50 km west of Oujda and 18 km south-
west of the town of Berkane [4] in the Eastern Rif
region of Morocco (Fig. 1). It is surrounded by
2,876 ha of forest, representing 4.19% of the total
area of the provincial territory [12]. This "SIBE"
is also of great archaeological interest [4]. The
bioclimate of Tafoughalt is of the subhumid
Mediterranean type with a cool winter [9] with an
average annual precipitation of 334 mm and an
average annual temperature of 15.4°C [13,14].
The Tafoughalt site contains some seasonal wet
spots that favour the establishment of diverse
vegetation. The latter are essentially made up of
various herbaceous plants such as Lavandula
dentata, Drimia maritima, Rosmarinus officinalis,
Cistus villosus and also, we have noted the
Fig. 1. Geographical location of the sampling sites for the two species - Amblystegium serpens (red
star) and Oxyrrhynchium speciosum (white star)
Achoual et al.
88
presence of tree matorrals of various densities,
especially around humid areas; these matorrals are
formed by oleander, a few fig trees, cork oak and
holm oak roots. As mentioned earlier, this site is
marked by a remarkable landscape of Aleppo pine
stands that are well adapted to the climatic
conditions of the Tafoughalt site. This vegetation
creates a microclimate favourable to the
installation of bryophytes.
METHODOLOGY
In the framework of our scientific explorations
which consists in inventorying the bryological
flora of the Tafoughalt massif (north-east of
Morocco), extensive field research has been
conducted in different sites of the massif [1]. In
two sites close to the "Grotte des Pigeons",
prospected in spring (April 2017), we collected
two new species for the Moroccan bryoflora. The
samples were photographed in-situ and ex-situ in
humid and dry state, then transported to the
laboratory for description and identification.
Different macroscopic and microscopic
characteristics were described, such as: the general
shape of each plant, its colour, the presence or
absence of sporophyte, the size and shape of the
cells of the different parts of the plant, etc. This
allowed to determine the family, genus and
species from the determination keys [15,16,17,18].
The nomenclature used is that of the bryophytes of
North Africa and that of the Mediterranean basin
[19,2].
RESULTS AND DISCUSSION
Two small pleurocarpal mosses were reported
during our bryophytic expedition [1]. The first
belongs to the genus Amblystegium encountered
twice in the same place occupying the tree trunks
in the first sample and fixed directly to the rocks
in the second sample. The second species
belonging to the genus Oxyrrhynchium is
harvested directly from very wet soil.
Amblystegium serpens (Hedw.) Schimp. (1853)
It is a bryophyte which belongs to the
Amblystegiaceae family, the order of Hypnales
and the class of Bryopsida. It has been observed
on the road leading to the Pigeon Cave of
Tafoughalt, whose coordinates are N34°81′72″
and W2°41′04″, at an altitude of 755 m. It is a
pleurocarpal moss, small in size in the form of
loose clumps (Fig. 2, A and B), yellowish-green in
colour, shiny when wet, collected from limestone
rocks and the trunks of fig trees in more or less
damp and shady places very close to temporary
water sources. On this site, it is Saxicolous,
corticole and mesophilic.
The primary stem is creeping, not more than 20
mm long, it is strongly attached to the substrate by
very fine and short rhizoids. The stem is
irregularly branched into short twigs of variable
size not exceeding 0.5 cm. It lies flat when dry
and more or less erect when wet and its branches
are not pinnate. The leaves of the main stem are
inserted in several rows all around the stem (Fig. 2
C), they are spread when dry and erect-spread
when wet. They are similar to the leaves of twigs,
less than 1mm long, lanceolate to oval-lanceolate,
elongated (Fig. 2, D and E), slightly folded
lengthwise, tapering gradually at the acuminate to
acute apex (Fig. 2, F and G). The leaf margin is
whole or finely denticulate in the upper half and
slightly denticulate in the middle of the leaf (Fig.
2 H). The cells of the leaf blade are abruptly
uniform and isodiametric up to the base, they are
linear to elongated rhomboidal, narrow and
chlorophyllous (Fig. 2 I). The basal part of the leaf
blade consists of more or less short rectangular
cells (Fig. 2 K). The basal angles of the leaf blade
consist of small short cells (Fig. 2 J). The auricles
are poorly distinct. Marginal cells are narrow,
rectangular in the basal part of the leaf blade and
similar to the cells in the middle of the leaf blade
in the upper half of the leaf. The costa is strong,
not very broad at the base and slightly narrow at
the top, thick, simple, single and incomplete,
reaching 3/4 of the length of the leaf blade. The
capsule is cylindrical, inclined to subhorizontal
with a poorly developed neck and brownish in
color (Fig. 2, M and N). The operculum is obtuse
conical and the peristome is simple (Fig. 2 O). The
setae is straight, fine and reaches 6 mm in length.
The spores are isodiametric and spherical and
green in colour (Fig. 2 P).
Morphologically, the taxa of the Amblystegiaceae
show many similarities with those of the
Brachytheciaceae which are often found in the
same places. Generally, the species of the family
Amblystegiaceae have a small form compared to
Achoual et al.
89
those of the family Brachytheciaceae, which have
a silky appearance. The taxa of the genus
Amblystegium are very similar to each other, it is
very difficult to treat and determine these taxa;
their average size is small. Leaves are lanceolate
and the costate extends to the middle [15]. The
species of this genus can be distinguished
morphologically by their wet and dry shape, their
size, the cells (form and size) and length of
costate. Amblystegium serpens can be
distinguished from taxa of the same genus by its
small size and its leaves rarely exceeding 1 mm
with very weakly denticulate marginal cells,
especially in the middle of the leaf blade
[16,15,18]. Ecologically, it grows on different
types of substrates: widely found at the base of
trunks or on rocks [16], rarely on earth [15], in
humid areas and [17] and mesophilic corticoles
[20]. In our study site, it is harvested in humid and
wooded areas.
The family Amblystegiaceae comprises 20 to 30
genera worldwide and the number of species is
estimated at 100 to 150 [21]. It is widespread in
humid or wet temperate habitats and is mainly
present at higher altitudes and in tropical and
subtropical areas [22]. Amblystegium serpens is an
almost cosmopolitan but discreet species,
preferring wetlands [23,24]. It is widely reported
in the majority of Mediterranean countries except
in: Andorra, Corsica, Cyprus, Egypt, Libya,
Madeira and Malta [2,25,26]. In North Africa, it is
only reported in Algeria and Tunisia. [19].
Oxyrrhynchium speciosum (Brid.) Warnst.
It is a moss belonging to the family
Brachytheciaceae, subfamily Rhynchostegielloi-
deae [27], the order of Hypnales and the class of
Bryopsida; synonym: Eurhynchium speciosum
(Brid.) Jur., according to [18,17]. Oxyrrhynchium
speciosum is harvested in Tafoughalt, in a stream
under the waterfall of the Pigeon Cave, N
34°81'55″ and W-2°40'50'', at an altitude of 705
m. This species is found in association with
another moss Eurhynchium hians (Hedw.) Sande
Lac.
It is a pleurocarpal moss forming loose, more or
less dense, medium sized and dark green, often
shiny when dry (Fig. 3 A). It is harvested on
highly humid soils, near water sources in more or
less shaded places. It is a soil-hygrophilic species.
It is harvested in a vegetative state, but generally
speaking, it is often fructified [16]. The primary
stem is creeping and very long, from 6 to 8 cm
(Fig. 3 B), irregularly branched with fairly long
twigs (1 to 4 cm). These twigs, more or less
spaced and strongly arched, are pinnate once,
often bent when dry and relatively erect when wet
(Fig. 3 C). The stem is fixed to the substrates by
some fine brown rhizoids. The leaves of the main
stem are between 1 and 1.5 mm long, generally
not exceeding 2 mm; they are 2 to 3 times longer
than they are wide, inserted in several rows all
around the stem, widely spaced, more or less
nested. Sometimes the apex of the leaves are
slightly pointed in the same direction, especially
in the dry state. When moistened, the leaves
become erect to spread out. They are narrowly
decurrent, oval to oval-triangular, abruptly
narrowed with an acute apex, these leaves
narrowed at the base (Fig. 3 E et F). The leaf
margin is flat, irregularly toothed in the upper 3/4
to 2/3 of the leaf blade (Fig. 3 I), especially near
the apex on both sides (Fig. 3 G), more or less
whole in the lower part (1/4) of the sheet (Fig. 3
J). The leaves of the secondary shoots are small
compared to the leaves of the main stems; they are
1 mm long and triangular in shape (Fig. 3 D). The
cells of the limbus are unistratose. In the middle of
the leaf blade, the cells are rhomboidal and
elongated (Fig. 3 I) and become short rhomboidal
at the apex. The basal cells are short, rectangular
to square (Fig. 3 K). The cells in the basal corners
of the leaves are not differentiated into auricles
(Fig. 3 L). The costa is simple, single, thick,
slightly toothed at the top, wide near the base,
dark green and reaches 4/5 of the length of the leaf
(Fig. 3 H).
Based on phylogenetic and morphological
analyses, the family Brachytheciaceae includes
four subfamilies and 41 genera worldwide [27].
The number of species is estimated between 250
and 350 [27]. The study of the specific diversity
within this family is very difficult to specify
because the morphological and histological
characters between the taxa of the family are very
similar and also the latter are present in different
bioclimatic locations and stages. This creates
difficulties in determining the exact number of
genera and species of Brachytheciaceae.
Achoual et al.
90
Fig. 2. Macroscopic and microscopic views of Amblystegium serpens
A,B: Plant in wet state macroscopic view; C: End of a sterile branch (X40); D,E: Leaf (X100); F,G: Top of a leaf (X400); H:
Marginal cells at the top of the leaf limb (X 400); I: Cells in the middle of the leaf limb (X 400); J: Basal angle cells of the leaf limb
(X 400); K: The basal part of a leaf (X200); L: Cross section of the leaf (X400); M: Capsule without operculum (X20); N: Capsule
with operculum (X20); O: Peristome (X400); P: Spores (X400)
I
D
F
E
H
P
J
B
N
C
A
K
O
G
M
L
Achoual et al.
91
Fig. 3. Macroscopic and microscopic views of Oxyrrhynchium speciosum
A: Tuft of the plant in a dry state macroscopic view; B,C: Leafy stems of the fertile wet plant (X20 et X40); D: Branch leaf (X100);
E,F: Stem leaf (X100); G: Top of a leaf (X400); H: Costa (X 400); I: Marginal cells and cells of the leaf blade of the upper part of the
leaf (X400); J: Marginal cells and cells of the leaf blade of the basal part of the leaf (X400); K: The basal part of a leaf (X400);
L: Basal cells of the leaf blade (X400)
Achoual et al.
92
The genus Oxyrrhynchium contains 19 species
[28], it is a difficult genus to treat because it
contains mixed characteristics with other genera.
It has long been described as a sub-genre of
Eurhynchium [28]. These two genera are very
similar in terms of morphological, histological and
ecological observation. The genus Oxyrrhynchium
differs from Eurhynchium only in the flattening of
its twigs, the leaves never folded lengthwise and
the seta always rough [15] but smooth in
Eurhynchium [28]. The segregation of
Oxyrrhynchium with Eurhynchium was reported
by Warnstorf in 1905 cited by [29], but because of
the difficulties of morphological separation
between these two genera, it is often mentioned as
a subgenus or as a simple synonym of
Eurynynium. The genus Oxyrrhynchium belongs
to a different subfamily from Eurhynchium in the
strict sense (Rhynchostegioideae). Phylogenetic
analysis has shown that Oxyrrhynchium is
monophyletic while Eurhynchium is polyphyletic
[30].
From a morphological point of view, the most
similar species of Oxyrrhynchium speciosum is
Oxyrrhynchium hians which is characterised by its
generally greenish-yellow colour, the slightly
short, wider leaves and the twigs spaced further
apart. Cytological comparison between these two
taxa shows that the number of chromosomes is
n=15 in Oxyrrhynchium speciosum and n=7 in
Oxyrrhynchium hians [27].
Oxyrrhynchium speciosum has two forms: aquatic
and terrestrial. Ecologically, it is widely found in
highly humid environments, especially in
mountainous areas or near water sources. It is
known as an aquatic foam [31] in the island of
Cyprus and also terrestrial in the forest ecosystems
of Slovakia [32], mainly on soils, but it can also
be found on other substrates such as rocks, trunks
and roots [18,15]. The aquatic form is on average
12 times larger with more branched stems (4-30
times) and longer leaves compared to the
terrestrial form. But molecular studies show no
distinction between the two forms [33].
This moss is well adapted to the Mediterranean
climate and has a wide distribution in the
Mediterranean region [25,26], with the exception
of certain countries: Andorra, Albania, Jordan,
Lebanon, Libya and Tunisia [2]. Particularly in
North Africa, it is only mentioned in Algeria [19].
Oxyrrhynchium speciosum is relatively rare in
New Zealand [33] and Switzerland [23] and
abundant in Turkey, Bulgaria [34,35]. In
Morocco, its frequency in the study site is very
rare.
CONCLUSION
In this study, two species new to Moroccan
bryoflora were described: Amblystegium serpens
and Oxyrrhynchium speciosum. They respectively
occupy a mesophilic and hygrophilic habitat in our
study site. These two mosses are considered very
rare according to the index of ecological
significance (IES) [1] in the Tafoughalt massif,
but they are widely distributed in European
countries.
ACKNOWLEDGEMENTS
All authors read and approved the final
manuscript. I thank Mrs. Nadia Belahbib, my
supervisor, for the corrections made. I also thank
Mr. Dabghi O. and Mr. Saadi K. for their help
during the bryological expeditions.
COMPETING INTERESTS
Authors have declared that no competing interests
exist.
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