Journal of Research in Ecology ISSN No: Print: 2319 –1546; Online: 2319– 1554

An International Scientific Research Journal

Original Research

Morphological and molecular diagnosis of Cuscuta sp parasitizing

Solanaceae plants in the middle of Iraqi provinces
Authors: ABSTRACT:
Journal of Research in Ecology

Bashar Al-Gburi K1, The genus Cuscuta is represented with 14 species distributed in all the
Al-Sahaf FH1, regions of Iraq. Most species belonging to this genus cause a major problem in the
Al-Fadhal FA1 and fields of many agricultural crops including Solanaceae members which are severely
Del Monte JP2 affected by these parasitic plants. Geographical distribution, host-range and
phenotypic diagnosis supported by molecular diagnosis as well as host-parasitism
Institution: (histopathology) studies were conducted to detect the parasitic species of Cuscuta on
1. Department of Plant
the Solanaceae, and the most widespread. We aimed to create a database facility for
Protection, Faculty of
Agriculture, Kufa providing an Integrated Weed Management (IWM) program in order to increase plant
University, Iraq. health in Solanaceae cultivation fields in Iraq. The average infection ratio of Cuscuta
sp on this family members was as high as 70.39 % in the provinces of Najaf, Karbala
2. Department of and Babylon. 10 Cuscuta races were registered as new races nationally and then
Agricultural Production, submitted to NCBI database. C. pentagona and C. australis were recorded for the first
E.T.S.I.A.A.B., Polytechnic time in Iraq in the presented study. However, C. campestris was found to be most
University of Madrid, Spain.
predominant over other species of Solanaceae. In this study we showed for the first
time the ability of C. campestris to parasitize on eggplant fruits, which have been
Corresponding author:
proved anatomically in addition to some other traits.
Bashar Al-Gburi K
Keywords:
Haustoria, PCR, C. pentagona, C. australis, C. campestris, Eggplant.

Article Citation:
Bashar Al-Gburi K, Al-Sahaf FH, Al-Fadhal FA and Del Monte JP
Morphological and molecular diagnosis of Cuscuta sp parasitizing Solanaceae plants in
middle of Iraqi provinces
Journal of Research in Ecology (2018) 6(2): 2315-2333

Web Address:
http://ecologyresearch.info/
documents/EC0611.pdf
Dates:
Received: 01 Aug 2018 Accepted: 27 Aug 2018 Published: 30 Sep 2018
This article is governed by the Creative Commons Attribution License (http://creativecommons.org/
licenses/by/4.0), which gives permission for unrestricted use, non-commercial, distribution and
reproduction in all medium, provided the original work is properly cited.

Journal of Research 2315-2333| JRE | 2018 | Vol 6 | No 2

in Ecology
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INTRODUCTION
Cuscuta species is considered to be a plant para-
sitic, pest and pathogen. At the same time, its effect on
the host plant causes physiological, chemical and bio-
logical damage, which leads to the reduction of produc-
tion or sometimes lead to the death of host plants (Smith
et al., 2013). Also, Cuscuta causes general weakness in
the infected plants, which leads to exposure to insects
and pathogens, thus increasing the damage to the host-
plant (Lanini et al., 2002). Cuscuta sp (Convolvulaceae)
is one of the most important parasitic weed, and the
name Cuscuta originated from the Arabic word kushkut,
which is translated as a tangled wisp of hair (Austin,
1980). It covers the host plant by tangling filaments of
yellow to orange colour with different diameters, which
are rootless and leafless. Cuscuta sp has different names
according to the country and sometimes region, but the
most common name is „Dodder‟, that probably derived
from the old German word “dotter” which means
„yolk‟ (Dawson et al., 1994). Identification of Cuscuta
to species level is difficult due to the few morphological
characters that do not exceed the form of fruits and
flowers with many differences of opinion among re-
searchers in the identification of species ranging from
about 100-200 species (Soliman et al., 2005). Further,
McNeal et al. (2007) stated that Cuscuta sp is difficult
to diagnose through flower and it is almost impossible
to identify from vegetative parts. Through molecular
studies, Cuscuta was identified to contain 194 species
spread across the world (Costea et al., 2015).
In Iraq, Cuscuta sp (dodder) is known by three
various common names, such as dodder, cancer and
troublesome weed. Agronomically, Mamluk and
Weltzien (1978) reported that 26% of fields cultivated
with eggplant (Solanum melongena), onions (Allium
cepa), cucumbers (Cucumis sativus), pepper
(Capsicum annuum), potatoes (Solanum tuberosum),
chili (Capsicum annuum), as well as alfalfa (Medicago
sativa) and tomatoes (Solanum lycopersicum) were in-
2316
Al-Gburi et al., 2018
fected by Cuscuta sp. Therefore, the weed study in Iraq
placed Cuscuta as the primary noxious weeds (Al-
Jubouri et al., 1985). Moreover, taxonomic studies have
shown that there are 14 species of Cuscuta which wide-
spread in the country. Nine species (C. campestris,
C. chinensis, C. monogyna, C. lehmanniana, C. pedicel-
lata, C. babylonica, C. palaestina, C. brevistyla, and
C. planiflora) were diagnosed by Rechinger (1964) and
then Chakravarty (1976) detected four other species
(C. kurdica, C. reflexa, C. arabica, and C. kotschyana),
and only one species (C. viticis) of Cuscuta was identi-
fied by Yonis (2016). Most researchers focused their
studies only on diseases transmitted by Cuscuta or on
Cuscuta control (Al-Tamimi, 2015); that lead to lack of
information in covering the diagnosis of Cuscuta on
specific host plant. These studies were not up to the
level of ambition and does not give a better understand-
ing of the Cuscuta sp in the Flora of Iraq yet. A survey
of Cuscuta sp as parasite on Solanaceae in the city of
Najaf, Karbala and Babylon was conducted in the cur-
rent study to cover the missing part of previous studies.
Studies were undertaken to understand the parasitic spe-
cies of Cuscuta on the Solanaceae. The most wide-
spread species (C. campestris) was identified by the
morphological and anatomical characteristics and study
of convergence and genetic differences between these
species were also done. Presented study will also pro-
vide a foundation for future studies on the biology and
ecology of Cuscuta species. Also, the present study fa-
cilitates to provide an Integrated Weed Management
(IWM) program in order to increase plant health in Sol-
anaceae cultivation fields in Iraq.
MATERIALS AND METHODS
Specimens collection
Cuscuta species were from three provinces viz:
Najaf, Karbala and Babylon of Iraq from the Solanaceae
crops cultivated during the period from 2016 - 2017.
The areas on the map were identified by GPS. The sam-
Journal of Research in Ecology (2018) 6(2): 2315-2333
Al-Gburi et al., 2018

C. pentagona

C. campestris C. australis

ples were collected from different plant hosts including:
tomato, eggplant, potatoes, chili and pepper. Cuscuta
species samples (filament and flowers) were placed in
50 mL sterile cans and preserved at refrigerator at the
temperature of 4 - 10°C, while the seeds were preserved
at the room temperature. After diagnosis and dissection,
samples were kept in deep freezer at -17°C for molecu-
lar diagnosis.
Cuscuta species with host
According to the method proposed by Naik and

Table 1. Cuscuta species infection rate (%) on Solanaceous plants in the field survey of three provinces of Iraq
S. No Host Plant Region Infection ratio
1 Chili Jabla/ Babylon 62.3%
2 Eggplant Haidariyah / Najaf 76.2%
3 Tomato Haidariyah / Najaf 74.1%
4 Eggplant Al-abbasiya / Najaf 64.8%
5 Eggplant Jabla / Babylon 88.7%
6 Eggplant Tuwerj / Karbala 50.5%
7 Tomato Jabla/ Babylon 85.6%
8 Tomato Tuwerj / Karbala 73.8%
9 Potato Musayyib / Babylon 62.1%
10 Pepper Jabla / Babylon 69.5%
11 Average 70.39%
Journal of Research in Ecology (2018) 6(2): 2315-2333 2317

Lakkund (1997) and Singh et al. (2004), infection ratio
was estimated in the areas covered by the survey of
Cuscuta species.
Infection ratio% = Number of infected plants
× 100
Number of total plants
Morphological diagnosis of Cuscuta species
Cuscuta species samples (flowers and seeds)
were the phenotypically by anatomical microscope ac-
cording to the classification key suggested by Flora of
lowland Iraq (1964), Plant wealth of Iraq (1976), and
others (Rhui-Cheng et al., 1995; Costea et al., 2011;
Spaulding, 2013).
Molecular diagnosis of Cuscuta species
DNA extraction and PCR conditions
DNA was extracted using the adjusted proce-
dure of Cenis (1992) from ten different isolates of
Cuscuta sp which were isolated from different hosts,
following the manufacturer‟s instructions
(FAVORGENR Biotech Corp, Plant DNA Extraction
Mini kit). ITS was amplified using a forward primer
(ITS1) and a reverse primer (ITS4) according to Alsaadi
et al. (2016) (Intron, Korea). PCR products were sub-
jected to agarose gel electrophoresis at 50 V for 120 min
on a 1% (w/v) agarose gel containing 0.1% GelRed™
Biotium Inc. (United States) and then visualized under
UV light. Aliquots of PCR products of 650 bp (30 μl of
each) were sent to Macrogen Inc. (Korea) for sequenc-
ing.
DNA sequence alignment and analysis were
conducted using Geneious v. 9.1.7 (Biomatters Ltd). All
sequences were aligned using Geneious alignment with
cost matrix of 70% similarity (IUB) (5.0/-4.5), gap open
penalty: 13, gap extension penalty: 3 and global align-
ment with free end gaps as the alignment type. A neigh-
bour-joining tree was constructed using the function
„Geneious Tree Builder‟ and adopting the Tamura-Nei
genetic distance model and the bootstrap resampling
method (random seed 900,000, number of replications
2318
Al-Gburi et al., 2018
1,500 and support threshold higher than 70%). The
sequence of ribosomal DNA (rDNA) gene of Solanum
lycopersicum KC213739 from GenBank was used as the
out-group.
Anatomical study of C. campestris
Anatomy of C. campestris was studied by using
a compound microscope (Altay type) under the magnifi-
cation of 10-40X and the dissection microscope (Krüss
type) under the magnification of 1-3X, with AmScope
MD35 USB digital camera microscope.
The haustoria with host
Free hand sectioning was done as indicated by
Alnomani (2012).
Study of other characteristics
This included anatomy of C. campestris for
studying some possibilities of parasitism on the host,
mechanism of non-sexual reproduction and seed germi-
nation by gibberellic acid (250 ppm) in the incubator set
at 25°C (Mustapha et al., 2015).
RESULTS
The persistence and duration of Cuscuta species
plants depend on the survival and persistence of the host
plant. Longevity of the host plant always determines the
growing period and longevity of the Cuscuta species.
Dodder (Cuscuta species) parasitizing annual plants will
be annual and the same case will be with biennial and
perennial plants. The field survey of ten fields cultivated
with solanaceous plants distributed in three provinces
showed that Cuscuta species weeds were widely distrib-
uted and infectious in nature (Figure 1).
Infection rates (%)
Ranged from 50.5 to 88.7%, the highest infec-
tion rate was recorded on eggplants (88.7%) followed
by tomato (85.6%) in the district of Jbala (Babylon pre-
view) (Table 1).
Morphological diagnosis of Cuscuta species
The morphological diagnosis of Cuscuta species
resulted in the identification of three species,
Journal of Research in Ecology (2018) 6(2): 2315-2333
 Table 2. Typical morphological characteristics of the Cuscuta species flower listed by species (C. campestris (C), C. pentagona (P), C. australis (A)).

Ovary length (mm) Ovary width (mm) Translucence of ovary

Species Thickenings in the distal part of ovary
Minimum Maximum Average Minimum Maximum Average wall
Al-Gburi et al., 2018

C 0.72 0.90 0.81 0.84 1.30 0.90 Interstylar aperture margins thickened Transparent

P 0.61 0.75 0.67 0.62 0.90 0.67 Tip of ovary forms a broad depression Transparent
from which styles emerge

A 0.88 1.07 0.98 1.03 1.42 1.21 Interstylar aperture margins thickened Opaque
Style length (mm) Style width (mm)
Species Number of styles Unequal styles
Minimum Maximum Average Minimum Maximum Average
C 0.25 0.44 0.33 0.10 0.15 0.13 One bifid style Present
P 0.28 0.48 0.37 0.06 0.10 0.08 One bifid style Present

Journal of Research in Ecology (2018) 6(2): 2315-2333

A 0.35 0.57 0.49 0.12 0.22 0.17 One bifid style Present
Stigma length (mm) Stigma width (mm) Stigma
Species Stigma surface area (mm2)
Minimum Maximum Average Minimum Maximum Average surface
C 0.11 0.16 0.13 0.17 0.23 0.20 0.11 Smooth
P 0.09 0.16 0.12 0.11 0.19 0.15 0.09 Smooth
Irregular lobed or wrin-
A 0.13 0.19 0.16 0.22 0.37 0.29 0.20
kled
Flower length (mm)
Species Stigma type Calyx
Minimum Maximum Average
C Depressed-glopbose 2 3 2.5 Lobes not strongly overlapping at the base of mature flowers.
P Glopbose, depressed-glopbose 1.5 2 1.8 Lobes strongly overlapping at base.
A Depressed-glopbose (flattened) 1.9 2.03 2 Lobes not strongly overlapping
Capsule (mm)
Species Seeds per capsule Seeds (mm) Seeds form Seeds number of 1 gram Seed weight (g)
Minimum Maximum
C 2 3 3-4 1-(1.3)-1.5 Renal-inverted oval 1612 0.0006-0.0007
P 1 2 3-4 1-(1.08)-1.2 Pear-shaped 1626 0.0006- 0.0007
A 1.9 2.05 3-4 1-(1.4)-1.6 Spherical-inverted oval 1604 0.0006-0.0007

2319
 Al-Gburi et al., 2018

Figure 2. Diagram to facilitate discrimination and consequently species identification in Cuscuta sp ( A: flow-
er; B: opened corolla; C: capsule; D: matured capsule; E: opened calyx and F: seed)

C. campestris, C. pentagona and C. australis. Species
differed for each area of the samples collected. These
species were characterized by several distinct morpho-
logical characteristics especially of the flower. These
species were taxonomically far from other species in the
plant encyclopedia but they are closely related to each
other Table 2. Since the three Cuscuta species
(C. australis, C. pentagona and C. campestris) are very
similar to each other, some of morphological differences
among these three species studied were diagramed as
shown in Figure 2 to facilitate discrimination and conse-
quently species identification as the style followed in
plants encyclopedia. The flowers, are compact heads 8-
12 mm across, corolla lobes lance-acuminate, calyx is
imbricate, forming 4 to 5 strong angles at the sinuses of
mature flowers and sometimes obscured in fruit; corolla
lobes in C. pentagona; But in C. campestris, corolla
lobes deltoid-ovate, calyx lobes not strongly overlap-
ping at the base of mature flowers, therefore not dis-
tinctly 5-angled. While in C. australis, corolla lobes
obtuse, sometimes reddish glandular on capsule, distinct
crater between styles and infrastaminal scales bifid
(Figure 3).
Molecular diagnosis of Cuscuta species
Ten races were recorded in this study as new
races nationally; seven of them were C. campestris, two
races belong to C. pentagona and only one race under
C. australis of the genus Cuscuta were recorded in the
world. Moreover, the two species namely C. pentagona
and C. australis were recorded for the first time in Iraq.

2320 Journal of Research in Ecology (2018) 6(2): 2315-2333

Al-Gburi et al., 2018

1 mm
1 mm
C.campestris C.campestris

1 mm 1 mm
C. pentagona C. pentagona

1 mm 1 mm
C. australis C. australis
Figure 3. Flowers and seeds of Cuscuta sp

Journal of Research in Ecology (2018) 6(2): 2315-2333 2321

 Al-Gburi et al., 2018

Figure 4. PCR product (band size 650bp), visualized under UV light. M: Marker’ (1): C. australis;
(2-8): C. campestris; (9,10); C. pentagona

All of these races have been registered at the National
Center for Biotechnology Information (NCBI). These
results were beneficial to facilitate and solve the diffi-
culties in the differentiation between species of this
genus depending only on morphological bases. Results
showed that C. campestris is the most prevalent in the
areas where the survey was conducted. This is mostly
due to its wide range of parasitism. It was found on po-
tato and tomato as well as widely on eggplant compared
with C. pentagona which was moderately parasitizing
eggplant and chili. However, C. australis had the lowest
rate of parasitism and was only found on pepper (Table
3 and Figure 4).
DNA sequence alignment and analysis
The phylogenetic tree of Cuscuta genotypes
shows that 100% of the genotype was identical to the
Genbank, while 84.13% were identical to the second
group, 74.8% of corresponding percentage were identi-
cal to the third group, and the fourth group had 75.2%
identity to that of the gene bank. Phylogenetically, there
was close relationship among races within group but
also there were genetic differences between groups 1, 2,

Table 3. Cuscuta species with races recorded in Iraq and their prevalence by region
S. No Accession number Cuscuta sp Parasitism on Region
1 MG669311 Cuscuta pentagona strain Bashar_5 Chili Jabla
2 MG669312 Cuscuta pentagona strain Bashar_4 Eggplant Haidariyah
3 MG669313 Cuscuta campestris strain Bashar_3 Tomato Haidariyah
4 MG669314 Cuscuta campestris strain Bashar_2 Eggplant Al abassiya
5 MG669315 Cuscuta campestris strain Bashar_1 Eggplant Jabla
6 MG669316 Cuscuta campestris strain Bashar_H Eggplant Tuwerj
7 MG669317 Cuscuta campestris strain Bashar_S Tomato Jabla
8 MG669318 Cuscuta campestris strain Bashar_K Tomato Tuwerj
9 MG669319 Cuscuta campestris strain Bashar_Z Potato Musayyib
10 MG669320 Cuscuta australis strain Bashar_A pepper Jabla

2322 Journal of Research in Ecology (2018) 6(2): 2315-2333

Al-Gburi et al., 2018
Figure 5. DNA sequence alignment and analysis of Cuscuta species
3 and 4. In addition, it was noted that the C. campestris
species recorded in different regions showed genetic
differences between these races within the species.
While, in the case of C. pentagona, species were genet-
ically convergent and closely related despite the differ-
ence in area of collection and the crop type (Figure 5).
The haustoria with host
As shown in Figure 6 the beginning of pre-
haustoria of C. campestris before occurred by intruding
and complete parasitism on the host was done with an
enzymatic helper to facilitate the adhesion and haustoria
attachment and then parasitism. After the successful of
penetration, it was also observed that haustoria enlarge-
Journal of Research in Ecology (2018) 6(2): 2315-2333
ment due to the degradation of the host tissue by the
degradation enzymes, which leads to the development
of haustoria to pseudo-haustorium to increase compati-
bility between C. campestris tissue and the host tissue
and thus increasing the solute absorption area.
In Figure 7, C. campestris parasitizing on fruit
peduncle of eggplant was shown. The haustoria parts
development inside the fruit peduncle, was observed.
The final stage of haustorium development was where it
overcomes the host defenses and increased solute ab-
sorption. The effect of C. campestris can also be identi-
fied from the shape of hyphae end, which lead to cell
2323
 Al-Gburi et al., 2018

h hs

p-h

p-h

hs
p-h
h ps-h

p-h hs

ps-h

Figure 6. C. campestris haustoria in host (p-h: Pre-haustoria; h: haustoria; ps-h: pseudo-haustorium and
hs: with host)

death due to withdrawal of all the host cells content
(amino acids and elements).
Study of other characteristics
The self-parasitism is a common phenomenon in
C. campestris where the stem raps on itself or on anoth-
er stem of the same dodder body. The parasitic stems
were dissected to observe differences in color on differ-
ent stems (filaments) due to nutritional content (Figure
8).
In this study Figure 9, the C. campestris at seed
germination stage was observed and does not to have
any rootlet or root-like organs, but the occurrence of an

2324 Journal of Research in Ecology (2018) 6(2): 2315-2333

Al-Gburi et al., 2018

C. campestris

h-h
h

pl

h-h

p-h

p-h

p-h h-h cd

Figure 7 . C. campestris parasitizing on fruit peduncle of eggplant (p-h: pre-haustoria; h: haustoria; h-h: haus-
torial hyphaes; pl: peduncle of eggplant and cd: cell death ).

Journal of Research in Ecology (2018) 6(2): 2315-2333 2325


df
df
h
Figure 8. Self-parasitism in the stems of C. campestris (df: dodder filament and h: haustoria)
incision (a small tiny slit) of the seed carrying out a very
thin thread-like flap at the end of which a viscous mate-
rial similar to the probes or sensors to guide the thread
path towards the host was recorded.
This study recorded the asexual reproduction of
C. campestris by forming many filaments from one ac-
tive stem (Figure 10). The field survey also showed the
high ability of C. campestris to parasitize on seeds of
weeds (Figure 11). The current study recorded the pres-
ence of mites near the C. campestris haustoria parasitiz-
ing site on the host plant (Figure 12). For the first time,
in this study C. campestris was observed to have high
ability to parasitize eggplant fruits. This can be seen on
the fruit peel where the color of which changed to red-
dish-brown indicating the penetration and development
of dodder haustoria within eggplant fruit tissue (Figure
13).
df
Figure 9. C. campestris seeds germination without host (df: dodder filament; p: probe and s: seed)
2326
Al-Gburi et al., 2018
df
h
df
DISCUSSION
Results of the field survey of current study on
areas cultivated with solanceous plants showed that the
adoption of traditional cultural practices and the ineffi-
cient mechanical or hand removing of Cuscuta species
filaments increased spreading and infection in fields of
eggplant, tomato, pepper, chili and potatoes (Figure 1).
This wide spread was mostly due to the high capability
of Cuscuta species to adhere firmly and attaching of one
host to another to start over a new growth confirming
former studies (Hassawy, 1974; Dawson, 1984; Kelly
et al., 2001 and Al-Jubouri, 2002). Vaughn (2002) also
pointed out that the enzymes involved in the connection
of haustoria to the host increased the efficiency of para-
sitism on solanaceous plants. The production of the
large quantities of seeds that remain dormant in the soil
had the most substantial role in increasing the spread of
p
Journal of Research in Ecology (2018) 6(2): 2315-2333
Al-Gburi et al., 2018
yf
as
Figure 10. The asexual reproduction of C. campestris (as: active stem and yf: young filament)
Cuscuta species for several years (Costea et al., 2004;
Mishra, 2009).
Parker and Riches (1993) showed that the ap-
propriate host and temperate humidity and temperature,
as well as the movement of hormones responsible for
flowering and fruit formation in the host, have crucial
role in timing of flowering and increasing its survival on
the host plant, likewise, Roney et al. (2007) reported
similarly. Due to these factors, the rate of infection was
70.39 % on the solanaceous plants grown in the prov-
inces of Najaf, Karbala and Babylon Table 1. This may
seriously threaten the agricultural production and cause
crop losses (Jayasinghe et al., 2004; Nickrent and
Musselman, 2004; Fathoulla and Duhoky, 2008).
df
Seed
h
Figure 11. Ability of C. campestris to parasitize on seeds (df: dodder filament and h: haustoria)
Journal of Research in Ecology (2018) 6(2): 2315-2333
The morphological diagnosis for species identi-
fication in this survey agreed with previous studies
(Flora of lowland Iraq, 1964; Plant wealth of Iraq, 1976;
Rhui-Cheng et al., 1995; Costea et al., 2011; Spaulding,
2013). This study recorded two new species of Cuscuta
(C. pentagona and C. australis) to be added to the list of
Cuscuta species so far identified in Iraq. There is no
evidence that these two species were recorded in Iraq
according to previous survey studies (Rechinger, 1964;
Chakravarty, 1974; Yonis, 2016; Taher, 2017) (Table
2).
Despite the use of classified taxonomic guides
such as chemical, environmental, and cellular, which led
to the clarification of many complex taxonomic prob-
lems, the phenotypic characteristics remained the basis
df
2327
 Al-Gburi et al., 2018

df
h

h
mi h

mite
Figure 12. Mites near the C. campestris haustoria (df: dodder filament and h: haustoria)

in the diagnosis of plants. For more reliability, this basic
and classic diagnostic tool has to have supportive and
confirmative methods. Thus molecular diagnosis came
to be an integration method supporting morphological
diagnosis in identification of plant species and their
races. There is a great morphological similarity in the
flowers between C. pentagona, C. australis and
C. campestris confirming the results of several taxo-
nomic studies (McNeal et al., 2007; Stefanovoic et al.,
2007; Costea et al., 2008 a and b) (Figure 2 and 3).
The current study recorded ten species of
Cuscuta to be registered in the NCBI in addition to the
registration of C. pentagona and C. australis for the first
time in Iraq Table 3. The presence of different species

df

df

Ef

ps-h

Figure 13. Ability of C. campestris to parasitize on eggplant fruits (df: dodder filament; h: haustoria; ps-
h: pseudo-haustorium and Ef: Eggplant fruit)

2328 Journal of Research in Ecology (2018) 6(2): 2315-2333

Al-Gburi et al., 2018
of Cuscuta sp in different parts of Iraq may be attributed
to their small size and weight of seeds, which can be
spread easily from a place to another, and for quite long
distances, and this is in consistent with previous studies
(Holm et al., 1997). We believe that the emergence of
the two new species in Iraq is due to the dissemination
of seeds by wind, river water, floods, and or crop seeds.
Organic fertilizers were imported for cultivation purpos-
es, especially from untrusted sources usually contain
Cuscuta seeds.
Phylogenetically Figure 5, Cuscuta sp belongs
to the plant family Convolvulaceae and the subfamily
Cuscuteae which was classified by (Garcia, 2004; Gar-
cia et al., 2014). The abundance and persistence of C.
campestris seeds for many years makes it a active viable
seed and thus spawning in many crops (Hutchinson and
Ashton, 1980). In addition to the ability of C. cam-
pestris to choose the type and age of the appropriate
host, it has the capability to coil around the host stem
within 5 h of adhesion (Dawson et al., 1994). These
factors make C. campestris dominant in parasitism and
proliferation compared to other species, and this is in
consistent with Dawson et al. (1994); Orr et al., (1996)
and Koch et al. (2004). C. campestris weed is one of the
most herbicide tolerant including glyphosate and sul-
fometuron-methyl. This is considered to be an additive
mechanism for increasing its spread and persistence in
different fields (Hassar and Rubin, 2003). The ratio of
genetic close relation and divergence among different
species is mostly due to the type of host and its contents
of the minerals and organic acids, as well as its toler-
ance to herbicides (Kim et al., 2004). It was also be-
lieved that environ-mental conditions of the region and
their altitude (location) have a role in the genetic make
up differences among races within a species.
C. campestris penetrates the host tissue with the
aid of enzymes (methylesterases, pectinases and cellu-
lases) (Vaughn, 2002). Then the haustorium cells begin
to elongate and grow toward the phloem and xylem tis-
Journal of Research in Ecology (2018) 6(2): 2315-2333
sues of the host. Developed pseudo-haustorium will be
released searching hyphae, which continues to grow
through the mesophyll without distinguishable hyphae
for phloem or xylem, which was confirmed previously
(Vaughn, 2003; Hong et al., 2011) (Figure 6). The com-
patible host of C. camestris will support haustorial hy-
phaes development to increase absorption and establish
successful connections with the host vascular tissues,
and thus lead to cell death due to withdrawing of their
vital contents (Farah et al., 2014) (Figure 7). The cur-
rent study does correspond to Taher (2017) who stated
that Cuscuta self-parasitism occurs between C. cam-
pestris stems or filaments. But, it can be considered that
this phenomena to be more like mutualism as nutrition
and amino acids exchange between the mature and the
young filament to increase parasitism efficiency, which
can be explained by differentiation of the two filaments.
In addition, the absence of any defense reaction and
only occurrence of the entry of haustoria and fusion
with the tissue of the other stems Figure 8 was noted.
C. campestris germination did not require the
presence of the host or its secretions (Benvenuti et al.,
2005). Possession of no regular roots is mostly because
of the low level of plasmodesmata in the cells, in addi-
tion to the removal of root like structures, if found, did
not affect the growth of C. campestris seedling
(Jianhong and YangHan, 1993; Sherman and Vaughn,
2003) (Figure 9). If believed that the probe was located
in the end of filament to facilitate the exit of filament in
the soil to the external periphery. C. campestris is prop-
agated by stem fragments and seeds (Lanini and Kogan
2005). However, many studies have not mentioned that
the young or old stem of C. campestris can produce a
gelatinous substance from which many young stems
will be released. This is confirmed by the current study
(Figure 10).
Results of the current study also showed that for
the first time globally that C. campestris has the ability
to parasite on seeds of weeds (Figure 11), in addition to
2329

the presence of mites on the pre haustoria feeding on
available nutrients without direct parasitism on the host
tissues (Figure 12). This may confirming that dodder
presence increases plant exposure to insects and patho-
gens (Agrios, 2005). Vaughn, (2002 and 2003) con-
firmed that C. campestris parasitizes on the plant leaves,
petioles and the stem. However, in the current study for
the first time the ability of C. campestris to parasitize on
the fruits of eggplant was noted. This level of parasitism
(Figure 13) aggressiveness may be related to mutations
in this parasite due to the environmental and nutritional
changes, through parasitism on different species of
plants.
CONCLUSION
Current study revealed the impact of Cuscuta
species and its spread in solanaceous fields. However,
C. campestris was the most dominant in the survey
regions. Morphological, genetical, and molecular diag-
nosis showed the presence of two other species C. pen-
tagona and C. australis in addition to different races
belonging to C. campestris. Anatomical study exhibited
the stage of C. campestris parasitism where haustoria
first penetrate the cortex of stem with the help of degra-
dative enzymes and grow toward phloem and xylem
tissues. C. campestris (dodder) was found to parasitize
eggplant‟s fruit peduncle and peel and this is regarded
as a new findings. The results of this study expand our
understanding of Cuscuta species and may help in
designing a reliable IWM program in Iraq.
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