International Journal of Agricultural Sciences and Veterinary Medicine Vol.

7 (4), November (2019)

Seed germination and seedling morphology as a tool for

taxonomic study on Vachellia nilotica (L.) P.J.H.Hurter
and Mabb. (Synonym: Acacia nilotica (L.) Willd. ex
Delile) in the Sudan
Haytham Hashim Gibreel1, Reem Rabie Mohammed Salih2* and Yousif Elnour Yagoub3
1. Department of Silviculture, Faculty of Forestry, University of Khartoum, SUDAN
2. Department of Clinical Medicine, Faculty of Veterinary Medicine, University of Khartoum, SUDAN
3. Department of Forest Conservation and Protection, Faculty of Forestry, University of Khartoum, SUDAN
*reemat7@yahoo.com

Abstract Acacia nilotica population in the Sudan.

Variation in seed germination behavior and early
growth morphology of seedlings was studied durng Keywords: Seed germination, seedling morphology, Acacia
2018-2019 among the subspecies nilotica, tomentosa, nilotica, taxonomy, Sudan.
adstringens and the new group of Vachellianilotica (L.)
P.J.H.Hurter and Mabb. (Synonym: Acacia nilotica (L.) Introduction
The study of seedling morphology has become a current
Willd. ex Delile) in the Sudan. Materials (pods and
trend in taxonomic research57. A seedling characteristic is
seeds) were collected from 13 variable sites. The one of the disciplines which are generally considered by
experiment was laid in completely randomized design taxonomists to offer comprehensive taxonomic and
under nursery condition. In total, 1500 pure seeds were phylogenetic trends in a taxon44, selection and classification
used for each sub-taxonomy and subjected to three of a species30,57 and has emerged as an essential discipline
treatments (soaking in H2SO4 60 min, soaking in for angiosperms taxonomy50.
boiling water and no treatment as control) 500 seeds
each. The treated seeds were sown in rate of 10 seeds Furthermore, seedling morphology has also been utilized in
in 50 polythene tube filled with clay and sand (2:1) clay plant classification at different taxonomic levels as
(3 treatments x 4 subtaxa x50 replicates x 10 seeds per families57, genera26,57 and species18,55 as well as in the
replicate) and kept under controlled nursey condition. preparation of seedling flora54 and construction of keys for
the identification of seedlings of some prominent forestry
The greminated seeds were recorded on the day 7th,
woody species15.
14th, 21th and 28th after sowing. Morphological
measurements of seedlings were made in early Das et al14 stated that seedling morphological characteristics
germination, after two months up to one year old. are very relevant for determining taxonomy of a species.
Seedling features such as collet, cotyledons shape, epicotyls,
Data subjected to analysis of variance (ANOVA) and phyllotaxy provide distinguishing information about
means were separated using Duncan’s multiple range particular plant species seedlings47. Throughout arid and
test. Significant variation (P < 0.0001) was recorded semiarid area of the world, Acacia play important role in life
among subtaxa in germination behavior (germination of local community as it provides food, fibre, fodder, timber,
rate, germination rate index and mean germination gum or resin, medicine, apiculture, energy fuel, katha,
tannins and also gives contribution in soil erosion control,
time) as affected by treatments and high germination
nitrogen fixation, soil improvement, intercropping,
characteristics were obtained with sulphuric acid boundary or barrier or support47.
followed by boiled water and less in the control.
Variation in seedling morphology among the studied Acacia nilotica (L.) Willd. ex Del. is a multipurpose
subtaxa was significant (P < 0.0001) in early leguminous species39,45,53 in the family Fabaceae, subfamily
germination and late (two-one year old) growth Mimosoideae and tribus Acacieae (Angiosperm Phylogeny
development. The new group showed among the other Group, 2009 and 2016). It has been accepted in recent
studied subtaxa of Acacia nilotica the best germination nomenclature as Vachellia nilotica (L.) P.J.H. Hurter and
behavior in all treatments as well as seedling growth Mabb.36,37,41 The species has a wide range of distribution
development. The study concluded that seedling through the tropical and subtropical area of the world and
characters are as important and reliable as that of occupies wide range of natural habitats7,16,43,53.
floral one in the delimitation of Acacia nilotica
The species provides goods as timbers and fuel wood27,
subspecies and the new group. Further studies in tannin8,19,33 and also has medicinal attributes8,22,23,35,45,52 and
seedling anatomy, cytology and molecular are needed role in Agroforestry9 and environmental conservation56. Its
to confirm the taxonomic status of the new group within taxonomy to subspecies was based on variation in pods and

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International Journal of Agricultural Sciences and Veterinary Medicine Vol. 7 (4), November (2019)

branchlets morphological characteristics at mature stage, N, 32° 30' E) and Alsunt riverine forest reserve (15°35' N, 32°
thus nine subspecies were recognized worldwide29,43, among 30' E) in semi-desert: Acacia tortilis – Maerua crassifolia
these the subspecies indica, cupressiformis and Desert Scrub, where mean annual rainfall is 75 - 300 mm/year;
hemispherica are native from India29 and subspecies El Ain natural forest reserve (12° 59' N, 30° 20' E) in semi-
nilotica, tomentosa, leiocarpa, subalata, kraussiana and desert: Grassland on sandy soils, annual rainfall 150-450 mm;
adstringens are from Africa12. Aldenaigila riverine forest reserve (13° 53' N, 32° 37' E),
Alsnait riverine forest reserve (14 ° 44 'N, 33° 22' E) in semi-
In the Sudan the species is commonly known by11 Sunt" or desert: Grassland on clay soils, with dominance of Acacia
"Grad" and includes the subspecies nilotica and tomentosa mellifera, annual rainfall 400 mm; Allmbwa riverine forest
in the flooded sites along the River Nile bank and its reserve (13° 00' N, 33° 59' E), Abu Geili riverine forest reserve
tributaries; subspecies adstringen in the dry areas on gravely (13° 34' N, 32° 35' E.) and Um Zibil riverine forest reserve
or alluvial soils in Kordofan and Darfur mainly along (13° 23' N, 34° 38' E) in Low rainfall woodland savanna on
seasonal rivers and valleys; subspecies subulata in savanna clay soils and Klikis natural forest reserve (12° 58' N, 32° 53'
grassland forming thicket with Commiphora africana in E) on sandy soils, annual rainfall varies from isohyets 400-600
northern and southeast Equatoria13,17,19. Furthermore, Sahni mm; Aldeisa riverine forest reserve (12° 03' N, 34° 18' E.) and
(1968) stated the presence of subspecies indica as exotic Sawlail riverine forest reserve (12° 19' N, 34° 21' E) in low
species similar to subspecies tomentosa in pods appearance rainfall woodland savanna on clay soils, mean annual rainfalls
and characteristics, this occurrence needs more vary from isohyets 600-800 mm; Nabak natural forest reserve
confirmations. (12° 31' N, 29° 55' E) Kadogli natural areas out of forests
reserve (11° 01' N, 29° 42' E and 11° 02' N, 30° 04' E) in low
Furthermore, Gibreel29 reported for the presence of a new rainfall savanna on sandy soils where mean annual rainfalls
group within riverine population of Acacia nilotica range between 300-500 mm and 600-800 mm southwards
population in the Sudan with intermediate in floral and respectively (Figure 3.1).
vegetative morphology between the subspecies tomentosa
and nilotica. Generally, absence of reproductive organs at Mature pods (50 per single tree) were harvested from the
early stage of seedling development makes identification of crown (top, bottom and lower) of 100 trees (dbh 25-35)
Acacia species so difficult43,47. selected for the subspecies nilotica, tomentosa, adstringens17
and the new group29 of Acacia nilotica and surveyed according
However, it is unfortunate that in spite of considerable to Abdelkheir et al2 and kept separately in clean polythene bags
taxonomic potentiality and prospects of seedling for further seeds extraction.
morphology, very little attention has been paid to this field
in Acacia taxonomy.18 However, Miller and Miller48 stated Seeds extraction and purification process: In the
that problem related to the identification and classification laboratory of the Department of Silviculture, Faculty of
of Acacia seedling at juvenile stage can be resolved by Forestry, University of Khartoum, the seeds were extracted
studying seedlings morphology and also determining their from pods and tested for viability by floating method3 to
actual position in taxonomic classes. discard damaged and insect infected seeds and eliminate the
empty one. The pure seeds were kept separately for each
Moreover, discrimination of plant species by seedling single tree (each group/each site) in clean polythene bags
blastogenic characters plays very important role in stored at laboratory conditions at 2-5°C and 80% of
identification, classification, conservation and delimitation humidity10 for further use.
of taxa at juvenile stage of plants47. With the fact that, some
distinct morphological changes occur between the seedling Effects of pre-sowing treatments on seeds germination:
and adult stages, it is very difficult to identify Acacia nilotica The experiments were carried out under controlled nursery
subspecies in their early growth based on seedling's condition11 at the nursery of Faculty of Forestry, University
morphological and/or anatomical characteristics29. In this of Khartoum (15° 39' N, 32° 30' E). 1500 seeds were used
study and since seedling morphology is very important to for the subspecies nilotica, tomentosa, adstringens and the
conserve plants permanently; makes an easy identification new group (15 seeds x 100 tree) and subjected to three pre-
and description at the juvenile stage and gives good data for germination treatments for 500 seeds in each treatment. We
classification of these plant species. We designed this study applied soaking in concentrated sulphuric acid (98% H2SO4)
aiming to assess the variation on seed germination and early for one hour followed by washing and soaking overnight in
growth characteristics of seedlings among Acacia nilotica distilled water28; we also used soaking in boiled water for
subtaxa occur in the Sudan. one hour, left to cool and soaked overnight10 and finally
soaking in tap water overnight as control. For each subtaxon
Material and Methods 150 polythene bags (20x25 cm) were prepared by mixture of
Seed sources, tree sampling procedure and pods collection: clay and sand (2:1) as sowing unit (50 polythene bags x 3
The study was conducted during 2018-2019. Seeds sources treatments).
were 13 sites across the agro-ecological zones of the Sudan34
including Shambat natural areas out of forests reserve (15° 39' The seeds per treatment (500 seeds) were sown in rate of 10

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International Journal of Agricultural Sciences and Veterinary Medicine Vol. 7 (4), November (2019)

seeds per polythene bag (50 polythene bags per treatment) diameter (RotC. D), leaf length (LF. L), petiole length (Pt.
and kept at the nursery condition under controlled watering. L), pinnae length (Pinn. L) and width (Pinn. W), leaflet
Germination was marked according to Kheloufi et al40. The length (LFt. L) and width (LFt. W) and stipular spine length
counting of germinated seeds (the emergence of cotyledon (Stip.spin. L); also, number of compound leaves per seedling
and 2 mm radicle through the seed coat) was made on the (LF. No), number of pinnae pairs per leaf (Pinn. No), number
7th, 14th and 21th and 28th day after sowing and was expressed of leaflets pairs per pinnae (LFt. No), number of stipular
in percentage. spines pairs per seedling (Stip.spin. No) were counted.

Other calculations were made as adopted by Kheloufi et al40 The qualitative evaluation of seedlings included stem bark
and included the germination rate index (GRI) which reflects colour (Stm. Clr) and texture (Stm. Tex), leaflets shape (LFt.
the percentage of germination on each day of the Shp) and color (LFt. Clr) and spine colour (Spin. Clr).
germination period (Equation 1) and mean germination time Digital photographs were taken at different stages of
(MGT) which represents the mean time a seed requires lot to seedlings development using digital camera (SAMSUNG
initiate and end germination (Equation 2). 18x, WB200F) to complete seedling description. A
dicotomus key constructs using the quantitative and
Equation 1: GRI (%) = ΣNo: of Germinated Seeds qualitative seedlings vegetative characteristics to facilitate
No: of Days identification of Acacia nilotica to subspecies, hybrid and/or
new group in the early stage of seedling growth and
Equation 2: MGT (days) = Σn.D development.
Σn
Statistical analysis: Experimental layouts were in a
where n: number of seeds newly germinated at time D; D: completely randomized design with 50 replications of 10
days from the beginning of the germination test and Σn: final seeds. The morphological data of seedlings (stem, leaves,
germination. stipular spines and roots) arranged in Excel sheets and mean
were subjected to analysis of variance (one-way, two-way
Morphological characterization of seedlings ANOVA) by using GLM procedure (Generalized Linear
Seedlings morphological variation in early germination: Model: GLM) calculated using SAS Version 9.0 (Statistical
Erely seedling morphological characters were described Analysis System) (2002) software. The mean separations
after germination up to one week following the terminology were carried out using Duncan’s multiple range tests and
proposed and adopted by several authors14,44 for pattern of significance was determined at p < 0.05.
seedling development either phanerocotylar when
cotyledons emerge from the test) or cryptocotylar when they Results
have not; cotyledon manifestation either food-storage In this study the environmentally induced variation was
(fleshy) or photosynthetic (paracotyledons); the position of eliminated during seedlings preparation by germinating the
the cotyledons (Seed-leaves) against the soil level as seed under the same environmental conditions. Thus, the
“epigeal” when they raised above the soil surface or as observed variation in seeds germination (Table 1, 2 and
“hypogeal” when they remain under the soil. Figure 2) and morphological growth development of
seedlings (Table 3, 4, 5, 6) among the three subspecies
Cotyledons were measured for its length (cm), width (cm), (nilotica, tomentosa, adstringens) and the new group, are
thickness (cm) and described for shape, phyllotaxy, texture most likely to have a taxonomic significance.
and colour; seedling hypocotyls and apicotyls described for
colour, form and texture and measured for length (in cm) and Variation in germination and early growth morphology
its relative growth was expressed as "slightly elongating", of seedlings: The treatments (H2SO4, Boiling water,
"elongating" and "strongly elongating" for length that were Control) effect was highly significant (P < 0.0001) on the
<2 cm, 5 cm and >5 cm, respectively.14 germination (%), germination rate index and the mean time
of germination (Table 1, 2 and Figure 2) within and among
Seedlings morphological variation after two months up the studied subtaxa of Acacia nilotica (subsp. nilotica,
to one year: The germinated seedlings were singled out at subsp. tomentosa, subsp. adstringens and the new group).
one month old to 3-4 healthy seedlings per each polythene The seeds of the new group showed varied behavior towards
bag (200 seedlings each subtaxon) and kept at the nursery treatments than other by having high germination percentage
with irrigation regime of three-day intervals for one year. (52.5±5.00% in 7 days, 80±8.16% in 14 days, 90±0.00% in
Morphological characterization of adult seedlings was made 21 days, 97.5±5.00% in 28 days) associated with high total
in two destructive harvests after two months and one year germination rate index (52.46±2.23) and low mean time of
after germination 100 seedlings each harvest (2 seedlings per germination (11.98±1.37) was in the sulphuric acid followed
single tree). Digital Vernier caliper (Model: GT-DC-01, by treatment with boiling water and less values of
Globaltronics GmbH and Co. KG) and ruler were used in germination percentage (0.00±0.00% in 7 days, 7.5±9.57%
measuring some quantitative variable of seedlings in “cm” in 14 days, 12.5±5.00% in 21 days, 15±5.77% in 28 days)
as shoot height (Sht. H), root length (Rot. L), root collar associated with high total germination rate index

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International Journal of Agricultural Sciences and Veterinary Medicine Vol. 7 (4), November (2019)

(4.17±2.49) and low mean time of germination (20.13±4.40) average of 0.15±0.03, while the subspecies nilotica and
were in control. tomentosa had leaflets with similar width 0.14±0.04 and
0.14±0.05 respectively.
The subspecies nilotica, tomentosa, adstringens and the new
group showed similar pattern of seedlings morphology. The Variation in seedling morphology after two months up to
seedlings were epigeal (phanerocotylar) with cotyledons that one year: The results of analysis variance indicated high
were food-storage (reserve or fleshy), coriaceous (leathery significant differences between the subspecies niloica,
form), semi-sessile, opposite, glossy and with moucronate tomentosa, adstringens and the new group in seedlings
apex and lobed base (Figure 3). The variation among the morphology after two months after nursery sowing (Table 4,
three subspecies and the new group was significant (P < 6 and Figure 4 B and C) up to one year (Table 5, 6 and Figure
0.0001) in measured traits of cotyledons, cotyledonary 4 D and E). Two months old seedlings of Acacia nilotica
leaves and hypocotyle (Table 3). subspecies nilotica had greenish shoot with average height,
root collar diameter and number of leaves of 12.64±1.68 cm,
Cotyledons were sub-orbicular-ovate, dark green for the 0.43±0.07 cm and 7±1.10 pairs respectively.
subspecies nilotica and pale green for the subspecies
adstringens with an average of 0.78±0.06, 0.71±0.09 and Leaves were bi-imparipinnate, pale green-greenish in
0.37±0.07 cm and 0.98±0.05, 0.77±0.07 and 0.44±0.05 cm colour, with average length of 3.34±0.60 cm and average
for the length, width and the thickness for the mentioned petiole length of 0.86±0.09 cm; average number of pinnae
subspecies respectively while they were oblong, pale green (pairs) was 1.95±0.43 with average dimension (length x
in both of the subspecies tomentosa and hybrid group with width) of 1.51±0.21 x 1.16±0.22 cm (oval shape); leaflets
an average length, width and the thickness of 0.81±0.06, were oblong-elongate in shape, sessile, opposite, with an
0.62±0.06 and 0.38±0.07 and 0.84±0.07, 0.58±0.05 and average number in pairs of 7.38±0.67 per pinnae and
0.39±0.06 respectively. 0.48±0.11 x 0.16±0.03 cm in dimension (length x width);
stipules spinest in leaves axils, with greenish base, brownish
Furthermore, the new group and the subspecies nilotica had tip, average number in pairs of 7.47±1.05 and average
no significant difference the longest hypocotyle with an length of 0.43±0.10 cm.
average length of 4.67±0.11 and 4.65±0.12 respectively
followed by the subspecies tomentosa 4.15±0.10 and Roots were tap, elongating, creamy white, cylinder with few
shortest hypocotyle was in the subspecies adstringens side root and average of 10.60±1.64 cm in length. The new
3.94±0.13. The subspecies nilotica, the new group and the group of Acacia nilotica showed after two months old
subspecies tomentosa had respectively and with no seedlings of greenish shoot with average height, root collar
significant difference the shortest cotyledonary leaves length diameter and number of leaves of 13.08±1.14 cm, 0.43±0.07
with an average of 2.92±0.13, 2.95±0.24 and 2.65±0.19 cm cm and 12.09±1.72 pairs respectively; leaves were bi-
than the longest one with an average of 3.95±0.11 cm in the imparipinnate, pale green-greenish in colour, average length
subspecies adstringens. of 4.33±1.01 cm, average petiole length of 0.82±0.11 cm;
average number of pinnae (pairs) was 2.32±0.58 with
The results indicated that the width of the cotyledonary average dimension (length x width) of 2.48±0.28 x
leaves depends on the leaflets length, thus the new group had 1.50±0.18 cm (oval shape); leaflets were sessile, opposite,
the widest cotyledonary leaves 1.55±0.03 and the longest with an average number in pairs of 8.68±0.65 per pinnae and
leaflets 0.78±0.04, followed by the subspecies tomentosa 0.68±0.08 x 0.17±0.14 cm in dimension (length x width)
1.44±0.05 and 0.72±0.06 and the subspecies nilotica giving an oblong-elongate shape; stipules in leaves axils
1.42±0.04 and 0.71±0.03, while the subspecies adstringens were spinest, greenish in base and brownish in tip with an
had the narrowest cotyledonary leaves length 1.35±0.04 as average number in pairs of 12.09±1.71 and average length
well as the shortest leaflets 0.68±0.05. Also, significant of 0.67±0.08 cm.
variation (Pr<0.0001) was also observed among the studied
sub-taxa of Acacia nilotica in number of leaflets (in pairs) Roots were tap, elongating, creamy white, cylinder with few
per cotyledonary leaves and it was considered to be side root and average of 10.12±1.31 cm in length.
diagnostic character in early identification of the subspecies Furthermore, the seedling’s shoot was also greenish in
and the new group of the species. colour at two months old for the subspecies tomentosa with
average height, root collar diameter and number of leaves of
However, the subspecies nilotica, tomentosa and the new 12.96±1.01 cm, 0.52±0.21 cm and 7.92±1.16 pairs
group had respectively and with no significant difference the respectively; leaves were bi-imparipinnate, pale green-
highest number of leaflets (pairs) per cotyledonary leaves greenish in colour, with average length and petiole length of
with an average of 9.54±0.06, 9.65±0.02 and 9.75±0.03 3.58±0.80 cm and 0.79±0.09 cm respectively; average
while the lowest number was in the subspecies adstringens number of pinnae (pairs) was 2.17±0.44 with average
with an average of 8.5±0.04. Furthermore, the subspecies dimension (length x width) of 2.00±0.33 x 1.25±0.27 cm
adstringens was distinct by its widest leaflets with an (oval shape); leaflets were oblong-elongate in shape, sessile,
average of 0.21±0.02 cm followed by the new group with an opposite, with an average number in pairs of 8.10±0.70 per

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International Journal of Agricultural Sciences and Veterinary Medicine Vol. 7 (4), November (2019)

pinnae and 0.51±0.05 x 0.16±0.04 cm in dimension (length 0.60±0.08 and 0.15±0.14; stipules spinset in leaves axils,
x width); stipules in leaves axils were spinest, greenish in white-greenish in base and dark brown in tip with average
base and brownish in tip with an average number in pairs of number in pairs and length of 36.40±4.08 and 3.44±0.23 cm
7.76±1.13 and average length of 0.50±0.19 cm. Roots were respectively.
tap, elongating, creamy white, cylinder with few side root
and average of 9.72±0.93 cm in length. One-year old seedlings of the subspecies tomentosa had also
woody stem, reddish-pale brown with longitudinal ridges in
The two month old seedlings of subspecies adstringens lower portion with an average height, root collar diameter
showed shoot colour very similar to other sub-taxa with and number of leaves of 65.42±5.08 cm, 1.03±0.09 cm and
average height, root collar diameter and number of leaves of 35.12±7.67 pairs respectively (Table 4.9 and 4.10). Leaves
12.19c (±1.16) cm, 0.54±0.16 cm and 7.16±0.68 pairs were developed and reached average 4.51±0.70 cm in length
respectively; leaves were bi-imparipinnate, pale green- and 1.18±0.11 cm in petiole length; average number of
greenish in colour, average length of 3.04±0.49 cm, petiolate pinnae (pairs), pinnae length (cm) and pinnae width (cm)
with average petiole length of 0.79±0.08 cm; average were developed to 2.81±0.54, 2.67±0.24 and 1.22±0.18
number of pinnae (pairs) was 2.05±0.38 with average respectively (oblong shape); leaflets were oblong, sessile,
dimension (length x width) of 2.22±0.33 x 1.11±0.13 cm opposite with an average number (pairs), length and width
(oval shape); leaflets were sessile, opposite, with an average (cm) of 11.35±4.34, 0.57±0.09 and 0.13±0.02; stipules
number in pairs of 8.24±0.79 per pinnae and 0.55±0.07 x spinset in leaves axils, white-greenish in base and dark
0.25±0.03 cm in dimension (length x width) giving an brown in tip with average number in pairs and length of
oblong-elongate shape; stipules in leaves axils were spinest, 34.16±7.08 and 4.40±0.52 cm respectively.
greenish in base and brownish in tip with an average number
in pairs of 7.16±0.68 and average length of 0.53±0.06 cm. The subspecies adstringens seedlings had woody stem after
a year old, reddish-pale brown with longitudinal ridges in
Roots were tap, elongating, creamy white, cylinder with few lower portion with an average shoot height, root collar
side root and average of 9.72±0.93 cm in length. The diameter and number of leaves of 61.52±5.87 cm, 0.98±0.09
seedlings vegetative traits (quantitative and qualitative) cm and 32.90±4.04 pairs respectively.
among the studied sub-taxa of Acacia nilotica showed
considerable development and variation after one year old Leaf average length was 4.05±0.47 and average petiole
(Table 5, 6). length was 1.29±0.13 cm; average number of pinnae (pairs),
pinnae length (cm) and pinnae width (cm) were 2.15±0.36,
The subspecies nilotica seedlings showed reddish-pale 3.24±0.36 and 1.02±0.18 respectively (oblong shape);
brown had woody stem, with longitudinal ridges in lower leaflets were oblong, sessile, opposite with an average
portion with an average shoot height, root collar diameter number (pairs), length (cm) and width (cm) of 9.43±0.85,
and number of leaves of 63.75±8.49 cm, 0.99±0.18 cm and 0.51±0.09 and 0.26±0.05; stipules spinset, with white-
26.65±3.28 pairs respectively; leaves type and colour were greenish base and dark brown tip, with average number in
the same as after one month with great development in pairs and length of 32.42±4.27 and 3.95±0.18 cm
average length that was 4.12±0.78 cm and average petiole respectively.
length that was 1.19±0.11cm; average number of pinnae
(pairs), pinnae length (cm) and pinnae width (cm) were Discussion
developed to 2.73±0.57, 2.73±0.57 and 1.37±0.27 In general, genetic make-up and/or environmental influence
respectively (oblong shape); leaflets were oblong, sessile, play a vital role in controlling the natural variation that
opposite with an average number (pairs), length and width occurs among organisms21. Elmund et al20 stated that the
of 10.13±0.94, 0.57±0.08 and 0.14±0.03 (reduced); stipules phenotype of members of the same species may vary
spinset, white-greenish in base and dark brown in tip with continuously while their genotype is relatively stable
average number in pairs and length of 26.64±3.28 and throughout the life of the organisms. Therefore, variation
4.03±0.29 cm respectively. based on the genetic make-up of the organisms is of more
taxonomic reliability than environmentally induced one5.
The seedlings of the new group at one year old showed According to Akinyele and Osekita4, the environment may
woody stem, reddish-pale brown with longitudinal ridges in affect the genetic constitution of a population by the pressure
lower portion with an average shoot height, root collar of selection it exercises on the population which in the longer
diameter and number of leaves of 66.01±5.75 cm, 1.02±0.13 term, may lead to evolutionary changes.
cm and 39.31±4.45 pairs respectively. Leaves were same in
colour as after one month, average leaf length was 3.62±0.39 Acacia seeds have a coat dormancy that acts primarily as
and average petiole length was 1.24±0.15; average number mechanical barrier limiting water and oxygen entrance10.
of pinnae, pinnae length and pinnae width were developed Luna et al42 stated fire, extreme temperatures and digestive
to 3.1±0.46, 3.07±1.69 and 1.28±0.16 respectively (oblong acids in animals’ stomachs or the abrasion of blowing sand
shape); leaflets were oblong, sessile, opposite with an as factors that break hard seed coats in natural conditions.
average number (pairs), length and width of 11.87±0.82,

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 International Journal of Agricultural Sciences and Veterinary Medicine Vol. 7 (4), November (2019)

Figure 1: Map of the Sudan showing the seeds sources. Where: Alde: Aldeisa forest; Saw: Sawlail forest; Alm:
Allmbwa forest; AbG: Abu Geili forest; Alden: Aldenaigila forest; Alst: Alsnait forest; Alsu: Alsunt forest; Sha:
Shambat area; UmZ: Um Zibil forest; Kli: Klikis natural forest; Ela: El Ain natural forest; Nab: Nabak natural
forest; Kadg: Kadogli natural mountainous area

Table 1
Variation in germination (%) among the four Acacia nilotica sub-taxa in four seeds treatments after 7th, 14th, 21st and
28th days after initial seeds sowing.
Subtaxa H2SO4 (60 min) Boiling Water Control
7 14 21 28 7 14 21 28 7 14 21 28
(days) (days) (days) (days) (days) (days) (days) (days) (days) (days) (days) (days)
Subsp. nilotica 40b 70b 80c 92.5b 20b 60a 77.5c 90b 0.00a 0.00c 7.5b 15a
(±8.16) (±18.16) (±8.16) (±5.00) (±8.16) (±8.16) (±5.00) (±0.00) (±0.00) (±0.00) (±9.57) (±5.77)
Subsp. tomentosa 47.5ab 75b 87.5b 95ab 30ab 67.5a 82.5b 95ab 0.00a 2.5b 10ab 15a
(±5.00) (±10.00) (±5.00) (±5.77) (±8.16) (±5.00) (±9.57) (±5.77) (±0.00) (±5.00) (±8.16) (±5.77)
NewG 52.5a 80a 90a 97.5a 37.5a 70a 90a 97.5a 0.00a 7.5a 12.5a 15a
(±5.00) (±8.16) (±0.00) (±5.00) (±9.57) (±8.16) (±5.83) (±5.00) (±0.00) (±9.57) (±5.00) (±5.77)
Subsp. 35b 67.5b 80.5c 90c 17.5b 60a 77.5c 87.5c 0.00a 0.00c 0.00c 7.5b
adstringens (±12.91) (±5.00) (±8.16) (±8.16) (±5.00) (±8.16) (±5.00) (±5.00) (±0.00) (±0.00) (±0.00) (±5.00)
F Value 3.41 1.9 2.68 1.11 5.47 1.85 2.68 4 0.00 1.71 2.55 1.80
Pr > F 0.0530 0.1829 0.0943 0.3829 0.0133 0.1853 0.0943 0.0346 0.00 0.2170 0.1051 0.2008
Means sharing the same letter in the same columns do not differ significantly at P= 0.05 according to Duncan's Multiple Range Test;
Means ± Standard deviation.

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Figure 2: Total germination rate index (%) and mean germination time (days) among four Acacia nilotica sub-taxa in
three seeds treatments after 7th, 14th, 21th and 28th days after initial seeds sowing. Where: NI: subsp. nilotica; to:
subsp. tomentosa; New G: the new group; ads: subsp. adstringens.

Table 2
Germination Rat Index % (GRI) among the four Acacia nilotica sub-taxa in four seeds treatments after 7th, 14th, 21st
and 28th days after initial seeds sowing.
Subtaxa H2SO4 (60 min) Boiling Water Control
7 14 21 28 7 14 21 28 7 14 21 28
(days) (days) (days) (days) (days) (days) (days) (days) (days) (days) (days) (days)
Subsp. 57.14b 50b 38.10b 33.04b 28.57b 42.86a 36.91b 32.14b 0.00a 0.00c 3.57b 5.36a
nilotica (±11.66) (±5.83) (±3.89) (±1.79) (±11.66) (±5.83) (±2.38) (±0.00) (±0.00) (±0.00) (±4.56) (±2.06)
Subsp. 67.86ab 53.57ab 41.67ab 33.93ab 42.86ab 48.22a 39.29ab 33.93ab 0.00a 1.79b 4.76ab 5.36a
tomentosa (±7.14) (±7.14) (±2.38) (±2.06) (±11.66) (±3.57) (±4.56) (±2.06) (±0.00) (±0.00) (±3.89) (±2.06)
New G 75a 57.14a 42.86a 34.82a 53.57a 50a 42.86a 34.82a 0.00a 5.36a 5.95a 5.36a
(±7.14) (±5.83) (±0.00) (±1.79) (±13.68) (±5.83) (±3.89) (±1.79) (±0.00) (±6.84) (±2.38) ±2.06)
Subsp. 50b 48.21b 38.10b 32.14b 25b 42.86a 36.91b 31.25c 0.00a 0.00c 0.00c 2.68c
adstringens (±18.44) (±3.57) (±3.89) (±2.92) (±7.14) (±5.83) (±2.38) (±1.79) (±0.00) (±0.00) (±0.00) (±0.00)
F Value 3.41 1.9 2.68 1.11 5.47 1.85 2.68 4.00 0.00 1.71 2.55 1.80
Pr > F 0.0530 0.1829 0.0943 0.3829 0.0133 0.1853 0.0941 0.0346 0.00 0.2170 0.1051 0.2008
Means sharing the same letter in the same columns do not differ significantly at P= 0.05 according to Duncan's Multiple Range
Test; Means ± Standard deviation

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Figure 3: Early morphology of germinated seedlings of the subspecies and the new group of Acacia nilotica where C:
the cotyledons; L: the cotyledon leaves

Table 3
Morphological traits variation of seedlings at early germination, among the subspecies and the new group of Acacia
nilotica in the Sudan.
Subtaxa
Characters Subsp. nilotica New group Subsp. tomentosa Subsp.
adstringens
Cotyledons length (cm) 0.78d (±0.06) 0.84b (±0.07) 0.81c (±0.06) 0.98a (±0.05)
Cotyledon width (cm) 0.71b (±0.09) 0.58c (±0.05) 0.62c (±0.06) 0.77a (±0.07)
Cotyledons thickness (cm) 0.37c (±0.07) 0.39b (±0.06) 0.38b (±0.07) 0.44a (±0.05)
Hybocotyle length (cm) 4.65a (±0.12) 4.67a (±0.11) 4.15b (±0.10) 3.94c (±0.13)
Cotyledon leaf length (cm) 2.92b (±0.13) 2.95b (±0.24) 2.65b (±0.19) 3.95a (±0.11)
Cotyledon leaf width (cm) 1.42b (±0.04) 1.55a (±0.03) 1.44b (±0.05) 1.35c (±0.04)
Number of leaflets (pairs) per Cotyledon leaf 9.54a (±0.06) 9.65a (±0.02) 9.75a (±0.03) 8.5b (±0.04)
Cotyledon’s leaflets length (cm) 0.71b (±0.03) 0.78a (±0.04) 0.72b (±0.06) 0.68c (±0.05)
Cotyledon’s leaflets width (cm) 0.14c (±0.04) 0.15b (±0.03) 0.14c (±0.05) 0.21a (±0.02)
Cotyledons colour Dark green, shinny Pale green- grayish Pale green Pale green
Cotyledons shape Oval Oblong Oblong Oval and Oblong
Means sharing the same letter in the same row do not differ significantly at P= 0.05 according to Duncan's Multiple Range Test;
Means ± Standard deviation

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Figure 4: Seedlings morphology of the subspecies nilotica, tomentosa, adstringens and the new group of Acacia nilotica
in the early germination (A), after two months (B and C) and one year (D and E).

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Table 4
Morphological variation of two months old seedlings among the subspecies and the new group of Acacia nilotica.
Subtaxa
Characters Subsp. nilotica New group Subsp. tomentosa Subsp.
adstringens
Shoot height (cm) 12.64b (±1.68) 13.08a (±1.14) 12.96a (±1.01) 12.19c (±1.16)
Root length (cm) 10.60a (±1.64) 10.12b (±1.31) 10.62a (±1.16) 9.72c (±0.93)
Root colar diameter (cm) 0.43b (±0.07) 0.43b (±0.07) 0.52a (±0.21) 0.54a (±0.16)
Number of leaves per stem 7.52c (±1.10) 12.09a (±1.72) 7.92b (±1.16) 7.16d (±0.68)
Leaf length (cm) 3.34c (±0.60) 4.33a (±1.01) 3.58b (±0.80) 3.04d (±0.49)
Petiole length (cm) 0.86a (±0.09) 0.82b (±0.11) 0.79c (±0.09) 0.79c (±0.08)
Number of pinnae (pairs) per leaf 1.95d (±0.43) 2.32a (±0.58) 2.17b (±0.44) 2.05c (±0.38)
Pinnae length (cm) 1.51d (±0.21) 2.48a (±0.28) 2.00c (±0.33) 2.22b (±0.33)
Pinnae width (cm) 1.16c (±0.22) 1.50a (±0.18) 1.25b (±0.27) 1.11d (±0.13)
Number of leaflets (pairs) per pinnae 7.38d (±0.67) 8.68a (±0.65) 8.10c (±0.70) 8.24b (±0.79)
Leaflet length (cm) 0.48d (±0.11) 0.68a (±0.08) 0.51c (±0.05) 0.55b (±0.07)
Leaflet width (cm) 0.16bc (±0.03) 0.17b (±0.14) 0.16c (±0.04) 0.25a (±0.03)
Number of stipule spines (pairs) per stem 7.47c (±1.05) 12.09a (±1.71) 7.76b (±1.13) 7.16d (±0.68)
Stipule spine length (cm) 0.43d (±0.10) 0.67a (±0.08) 0.50c (±0.19) 0.53b (±0.06)
Means sharing the same letter in the same row do not differ significantly at P= 0.05 according to Duncan's Multiple Range Test;
Means ± Standard deviation
Table 5
Morphological variation of one-year old seedlings among the subspecies and the new group of Acacia nilotica.
Subtaxa
Characters Subsp. nilotica New group Subsp. tomentosa Subsp.
adstringens
Shoot height (cm) 63.75b (±8.49) 66.01a (±5.75) 65.42a (±5.08) 61.52c (±5.87)
Root length (cm) 32.39a (±5.03) 30.94b (±4.00) 32.46a (±3.54) 29.70c (±2.83)
Root colar diameter (cm) 0.99b (±0.18) 1.02a (±0.13) 1.03a (±0.09) 0.98b (±0.09)
Number of leaves per stem 26.65d (±3.28) 39.31a (±4.45) 35.12b (±7.67) 32.90c (±4.04)
Leaf length (cm) 4.12b (±0.78) 3.62c (±0.39) 4.51a (±0.70) 4.05b (±0.47)
Petiole length (cm) 1.19c (±0.11) 1.24b (±0.15) 1.18c (±0.11) 1.29a (±0.13)
Number of pinnae (pairs) per leaf 2.73b (±0.57) 3.1a (±0.46) 2.81b (±0.54) 2.15c (±0.36)
Pinnae length (cm) 2.49d (±0.26) 3.07b (±1.69) 2.67c (±0.24) 3.24a (±0.36)
Pinnae width (cm) 1.37a (±0.27) 1.28b (±0.16) 1.22c (±0.18) 1.02d (±0.18)
Number of leaflets (pairs) per pinnae 10.13b (±0.94) 11.87a (±0.82) 11.35a (±4.34) 9.43c (±0.85)
Leaflet length (cm) 0.57b (±0.08) 0.60a (±0.08) 0.57b (±0.09) 0.51c (±0.09)
Leaflet width (cm) 0.14c (±0.03) 0.15b (±0.14) 0.13c (±0.02) 0.26a (±0.05)
Number of stipule spines (pairs) per stem 26.64d (±3.28) 36.40a (±4.08) 34.16b (±7.08) 32.42c (±4.27)
Stipule spine length (cm) 1.21d (±0.29) 1.87a (±0.23) 1.40c (±0.52) 1.49b (±0.18)
Means sharing the same letter in the same row do not differ significantly at P= 0.05 according to Duncan's Multiple Range Test;
Means ± Standard deviation

In the same way breaking seed coat dormancy of Acacia treatments showed potentiality to break down the coat
nilotica seeds under simulated natural habitat condition was inhibition1,32,38,51. In our study, we applied three treatments
studied in Sudan57. namely soaking in sulphuric acid, soaking in boiled water
and soaking in normal water treatments to overcome seed
On the other hand, several artificial methods are used in dormancy in Acacia nilotica subtaxa (nilotica, tomentosa,
laboratory and nursery conditions to break seed dormany adstringens and the new group). The results indicated that
exhibited by hard seed coat. The application of manual the subtaxa showed variation in seeds germination behavior
scarification, sulphuric acid, hot water and dry heat as affected by applied treatments.

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Table 6
Shoot and root traits development of seedlings after two months (8 weeks) up to one year (48 weeks) after sowing
within the subspecies and the new group of Acacia nilotica.
Subtaxa
Characters Subsp. nilotica New group Subsp. tomentosa Subsp. adstringens
Tow One year Tow One year Tow One year Tow One year
month month month month
Shoot height (cm) 12.64b 63.75a 13.08b 66.01a 12.96b 65.42a 12.19b 61.52a
(±1.68) (±8.49) (±1.14) (±5.75) (±1.01) (±5.08) (±1.16) (±5.87)
Root length (cm) 10.60b 32.39a 10.12b 30.94a 10.62b 32.46a 9.72b 29.70a
(±1.64) (±5.03) (±1.31) (±4.00) (±1.16) (±3.54) (±0.93) (±2.83)
Root collar diameter (cm) 0.43b 0.99a 0.43b 1.02a 0.52b 1.03a 0.54b 0.98a
(±0.07) (±0.18) (±0.07) (±0.13) (±0.21) (±0.09) (±0.16) (±0.09)
Number of leaves per stem 7.52b 26.65a 12.09b 39.31a 7.92b 35.12a 7.16b 32.90a
(±1.10) (±3.28) (±1.72) (±4.45) (±1.16) (±7.67) (±0.68) (±4.04)
Leaf length (cm) 3.34b 4.12a 4.33a 3.62b 3.58b 4.51a 3.04b 4.05a
(±0.60) (±0.78) (±1.01) (±0.39) (±0.80) (±0.70) (±0.49) (±0.47)
Petiole length (cm) 0.86b 1.19a 0.82b 1.24a 0.79b 1.18a 0.79b 1.29a
(±0.09) (±0.11) (±0.11) (±0.15) (±0.09) (±0.11) (±0.08) (±0.13)
Number of pinnae (pairs) per leaf 1.95b 2.73a 2.32b 3.1a 2.17b 2.81a 2.05b 2.15a
(±0.43) (±0.57) (±0.58) (±0.46) (±0.44) (±0.54) (±0.38) (±0.36)
Pinnae length (cm) 1.51b 2.49a 2.48b 3.07a 2.00b 2.67a 2.22b 3.24a
(±0.21) (±0.26) (±0.28) (±1.69) (±0.33) (±0.24) (±0.33) (±0.36)
Pinnae width (cm) 1.16b 1.37a 1.50a 1.28b 1.25a 1.22b 1.11a 1.02b
(±0.22) (±0.27) (±0.18) (±0.16) (±0.27) (±0.18) (±0.13) (±0.18)
Number of leaflets (pairs) per pinnae 7.38b 10.13a 8.68b 11.87a 8.10b 11.35a 8.24b 9.43a
(±0.67) (±0.94) (±0.65) (±0.82) (±0.70) (±4.34) (±0.79) (±0.85)
Leaflet length (cm) 0.48b 0.57a 0.68a 0.60b 0.51b 0.57a 0.55a 0.51b
(±0.11) (±0.08) (±0.08) (±0.08) (±0.05) (±0.09) (±0.07) (±0.09)
Leaflet width (cm) 0.16a 0.14b 0.17a 0.15a 0.16a 0.13b 0.25a 0.26a
(±0.03) (±0.03) (±0.14) (±0.14) (±0.04) (±0.02) (±0.03) (±0.05)
Number of stipule spines (pairs) per 7.47b 26.64a 12.09b 36.40a 7.76b 34.16a 7.16b 32.42a
stem (±1.05) (±3.28) (±1.71) (±4.08) (±1.13) (±7.08) (±0.68) (±4.27)
Stipule spine length (cm) 0.43b 1.21a 0.67b 1.87a 0.50b 1.40a 0.53b 1.49a
(±0.10) (±0.29) (±0.08) (±0.23) (±0.19) (±0.52) (±0.06) (±0.18)
Means sharing the same letter in the same row do not differ significantly at P= 0.05 according to Duncan's Multiple Range Test;
Means ± Standard deviation

However, sulphuric acid treatments (98% for 60 min) as According to Mohamed Ahmed49, the seeds of Acacia
compared by control had the best effect on Acacia nilotica nilotica subspecies tomentosa and subspecies adstingens
subtaxa seeds where the average germination percentage need fewer days for breaking dormancy with the increase in
reached 80% (New group), 76.25% (subsp. tomentosa), emersion length in sulfuric acid. Kheloufi et al40 reported
70.63% (subsp. nilotica) and 68.13% (subsp. adstringens) that the increase of the soaking time in the sulphuric acid
followed by boil water treatment where average germination from 60 to 120 min is favorable for the seeds of A.
rate of 73.75%, 68.75%, 61.88% and 60.63% was recorded cyanophylla and those of Acacia farnesiana improving the
for the new group, subsp. tomentosa, subsp. nilotica and increase of the germination rate index (GRI) from 29.1% to
subsp. adstringens, respectively. 73.61% and from 29.86% to 71.87% respectively.

Similar to our findings, Ben Zetta et al10 stated that manual Goda31reported that seeds of Acacia nilotica when treated
scarification and sulphuric acid treatments are the best pre- with H2SO4 for 90 minutes produced vigorous seedlings and
germination treatments for A. ehrenbergiana Hayne while higher per cent germination. In contrast of our findings, the
sulphuric acid treatment enhanced the germination rate of work of Aref et al6 showed that boiling water had positive
Acacia seyal Del., as compared to the boiling water which effect on Acacia seyalDel. germination percentage (34.25%)
presented low germination rates for both species. while it has less effect on Acacia ehrenbergiana Hayne
germination with a germination percentage of 9.35%. The
Furthermore, application of sulphuric acid and boiling water subspecies nilotica, tomentosa, adstringens and the new
as pre-sowing treatments for Acacia ehrenbergiana Hayne group showed significant variation in morphological
seeds revealed germination percentages of 94% and 69 % characteristics of seedlings.
compared to 96 % obtained by manual scarification6.

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International Journal of Agricultural Sciences and Veterinary Medicine Vol. 7 (4), November (2019)

In this study, epigeal (phanerocotylar) pattern of seedlings presence of the new group within Acacia nilotica population
growth with food-storage (reserve or fleshy) cotyledons that in the Sudan.
were coriaceous (leathery form), semi-sessile, opposite,
glossy and with moucronate apex and lobed base was Acknowledgement
observed in all seedlings of the subspecies nilotica, The authors would like to extend their sincere appreciation
tomentosa, adstringens and the new group. This result to all the authority of Forest National Corporation (FNC) in
agreed with Meena and Datta47 who reported epigeal, the different Sudan’s States (Khartoum, Geziera, Sennar,
phanerocotylar seedling type in Acacia nilotica subsp. Gedarif, Blue Nile, North and South Kordofan) for their help
indica (Benth.) Brenan. In the early growth coltyledons in providing official permission and transportation to access
shape, size, texture as well as colour showed variability a the forest for material collection.
high taxonomic value to distinguish among the three
subspecies and the new group than other the vegetative traits References
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