ONOSMA (BORAGINACEAE) POLLEN INTERACTIONS WITH BOMBUS TERRESTRIS (BUFF-TAILED BUMBLEBEE) SPECIES IN JORDAN

2015·14.2 MB·

by  S Al-Qura’N

irnaldoa 21 343 - 358, 2014 ISSN: 1815-8242
(2):
Onosma
(Boraginaceae) pollen
interactions
Bombus
with
terrestris (buff-tailed
bumblebee)
Jordán
species
in
Onosma
Interacciones polen de (Boraginaceae)
del
Bombus
común)
con
especie en
(abejorro
terrestris
la
Jordania
S.Al-Qura’n
Mu’tah University, Department of Biology, Faculty of Science, Karak, Jordán.
ARNALDOA
343
Diciembre
21 Julio -
(2):
I
Bombus
\I-Qura’n: ¿7/705/773 pollen interactio terrestris species in
1
Abstrae!
The between morphology Onosma and
relationships pollen of three species their interactions
with their insect visitors Bombus terrestris during springs of 2012 and 2013 were studied in Jordán.
morphology and
Pollen characteristics (type size of pollen grains, pollen shape, symmetrical
structures between pollen grains and the pollen polar and equatorial measurements, P/E
visitors,
way
ratios, structures of endo- and ecto- apertures) were investgated to formúlate any possible of
nter ct on Certain kind of bees Bombus terrestris (buff-tailed bumblebee) was observed as the most
many
abundant visitor especially on the flowers of Onosma in April although other honey bees were
observed like Apis mellifera, Bombus impatiens, Apis dorsata and A. florea but in very low frequencies.
Onosma species in Jordán as boraginacean species produces a large amount of whitish polliniferous
all
Bombus
dust, thus considered as a potential source for pollen visitor species of bees, especially
all
known
terrestris as buff-tailed bumblebee. Three Onosma species were investigated palynologically
LM
by and SEM. The pollen class is either 3 zonocolporate {Onosma aleppica Boiss and Onosma
frutescens Lam.) or 4-zonocolporate {Onosma cinérea Schreber). Polar measurements ranges are (37-
pm pm pm
Onosma Onosma and Onosma
43) for aleppica Boiss, (42-45) for cinérea Screber (41-44) for
pm pm
Lam. measurement Onosma
frutescens Equatorial ranges are (26-34) for aleppica Boiss, (28-34)
pm
for Onosma cinérea Screber and (28-33) for Onosma frutescens Lam. P/E ratio ranges between
Onosma Onosma and
Schreber
(1.41-1.39) for aleppica Boiss, (1.49-1.33) for cinérea (1.45-1.28) for
Onosma frutescens Lam. Pollen outline has been recognized, the equatorial view for all Onosma species
was erect for species examined. The most important diagonistic factor in recognition of pollen
all
morphology of Onosma species in the endoaperture and ectoaperture which showed some kind
is
on The morphology
of variation reflected their pollen visitor association. of ectoaperture structures
and
(colpi, fastigium margins) for the three species exhibited short colpi, absence of fastigium
with indistinct margins for Onosma aleppica Boiss, long colpi, presence of fastigium and indistinct
margin for Onosma cinérea Schreber and short colpi, presence of fastigium and distinct margin of
Onosma frutescens Lam. The morphology of endoaperture structures (colpi, costae and margins) has
Onosma showed
reflected also distinguished variation; aleppica Boiss short colpi, presence of costae
and indistinct margin, Onosma cinérea Schreber showed short to long colpi in general, absence of
Keywords: Onosma, Boraginaceae, pollen interactions, Bombus Jordán
terrestris,
Se estudiaron las relaciones entre la morfología del polen de tres especies de Onosma sus
y
interacciones con los insectos visitantes Bombus terrestris durante las primaveras de 2012 y 2013 en
Jordania. Las características morfológicas del polen (tipo tamaño de los granos de polen, forma
y
del polen, estructuras simétricas entre los granos de polen los visitantes del polen, medidas
y
polares ecuatoriales, relaciones P/E, estructuras de las endo ecto aperturas) fueron investigadas
y y
modo
para establecer cualquier posible de interacción. Cierto grupo de abejas Bombus
terrestris
común) fueron observadas como más abundantes en de Onosma
(abejorro las visitantes las flores
especialmente en abril, aunque fueron observadas muchas otras abejas de miel, tales como Apis
muy
Bombus impatiens. Apis dorsata A. pero con poca frecuencia. Las especies de
mellifera, y florea,
Onosma en Jordania, al igual que todas las especies de boragináceas, producen una gran cantidad
como
de polvillo polinífero blanquecino, por lo tanto son consideradas atractivo potencial de todas
como
las especies de abejas visitantes del polen, especialmente Bombus terrestris conocida abejorro
común. Tres especies de Onosma fueron estudiadas palinológicamente con microscopio óptico
y
microscopio electrónico de barrido. El tipo de polen es 3-zonocolporado {Onosma aleppica Boiss
y
Onosma Lam.) 4-zonocolporado {Onosma Los rangos de medidas
frutescens cinérea Schreber). las
y
gm gm
polares son (37-43) para Onosma aleppica Boiss, (42-45) para Onosma cinérea Schreber y (41-44)
ARNALDOA
344 2014
21 Julio - Diciembre,
(2):
I
Bombus
^I-Qura’n: 0/705/773 pollen innteractions with terrestris species in Jordán
|xm para Onosma frutescens Lam. Los rangos de las medidas ecuatoriales son (26-34) \im para Onosma
gm gm
Onosma Onosma Lam.
para Schreber para Las
aleppica Boiss, (28-34) cinérea y (28-33) frutescens
P/E Onosma Onosma
relaciones varían entre (1.41-1.39) para aleppica Boiss, (1.49-1.33) para cinérea
Schreber (1.45-1.28) para Onosma frutescens Lam. Los perfiles del polen han sido reconocidos. La
y
vista ecuatorial para todas las especies de Onosma examinadas fue rectangular-elíptica; la vista
P/E
polar fue triangular para todas las especies. La vista de la relación fue levantada para todas las
más
especies examinadas. El importante factor de diagnóstico en el reconocimiento de la morfología
del polen de las especies de Onosma, está en la endoapertura la ectoapertura, las cuales mostraron
y
cierto tipo de variación reflejada en su asociación con los visitantes del polen. La morfología de las
estructuras de ectoapertura (colpos, fastigio márgenes) exhibió colpos cortos, ausencia de fastigio
y
con márgenes indistintos para Onosma aleppica Boiss; colpos largos, presencia de fastigio margen
y
indistinto para Onosma cinérea Schreber; colpos cortos, presencia de fastigio margen distinto
y y
para Onosma frutescens Lam. La morfología de las estructuras de endoapertura (colpos, costillas
márgenes) ha reflejado también variaciones distinguibles; Onosma aleppica Boiss mostró colpos
y
cortos, presencia de costillas margen indistinto; Onosma cinérea Schreber mostró colpos cortos
y y
largos en general, ausencia de costillas márgeni
y
Bombus
aes del polen, Jordania.
terrestris,
complex from
nectar-pollen secreted a
Introduction
mixed with white
certain secretory
cells
Onosma genus from Boraginaceae
as
pollineferous material attracting different
known
well and widely distributed in
is
hymenopteran The
kinds
of species.
which unique
Jordán, located in position
is
morphology
investigation of pollen of
middle between
in the heart of the east
Onosma considered very important
a
is
longitudes 53° 40' and 39° and between
E,
morphological evidence
to investígate
29° 30' and 34° N,
latitudes this location of
some
aspects related to pollen class,
Jordán gives country extraordinary
this
endoaperture, ectoaperture structures, polar
main
opportunity comprise four
to at least
and measurements, P/E
equatorial ratio
phytogeographical elements
different
and pollen outline shape and structure, to
from north south and from west
to to east
whether some
see palynological variations
(mediteranean, irano-teranean, saharo-
among
examined and
the species will
exist,
and
arabian nubo-sudanian), surely
this
on
be such
will relied for characterizations
which
leaded to geobotanical variation
among
and Onosma
differentiation the
enriches the plant biodiversity in jordanian
having such
species the role for attraction
territories, and reflected over the high
between and
the pollen grains their visitors
spectrum Onosma widely
of plant species.
is
from Bombus
especially species
terrestris
producing different kinds of pollineferous
&
(Zohary Feibrun-Dothan, 1962-1986;
dusts especially during spring pollination
&
Adam,
Khayyat
Mursi,
2001; 1981; Peter,
which
times, interconnected large variation
Punt
Prance,
1994; 2001; et al, 1994).
bomblebees
of insects especially buff-tailed
The Onosma
as pollen visitors between the wild and floral parts of species
gametophytic male
cultivated species (Bender, 1974; Al-Quran, are consisting of
by androecium and
structures represented
1986; 1988; 2010; 2011).
gametophytic female structures represented
Onosma genus has unique
potential
by gynoecium
located in the central part
in production of a vast quantities of
surrounded by
of the flowers externally
ARNALDOA
345
Diciembre
21 Julio -
(2):
I
Bombus
Al-Qura’n: ¿7/705/773 pollen interactions with terrestrís in Jordán
perianth parts represented by calyx and addition to that, these species are producing
corolla, this kind of floral arrangement certain kinds of pollen grains visited by
is
typically found as general morphological a different types of insects representing
shape of this family enabling the flowers different orders and classes. Absolutely
Onosma Onosma
of to attract certain types of buff- the floral structures of species
tailed bomblebees (Bombus during with their pollen grains are considered
terrestris)
Also have form
the pollination times. the inflorescence to the potentiality to this kinf
of Onosma similar to other boraginacean, of specialized intimate relationship with
is
many
composed
of simple clusters of several pollen visitors to reinforce this association
flowers. The presence of colored lobulated with pollen and néctar as rewards for the
and was
external purple glandular trichomes doubtless there a correlations
visitors.
form which was
hairs in of papillae near the base of representing similarities lead to
floral parts especially corolla, considered some kind of superimposing between the
is
the source of néctar production release anther structures as the source of pollen
high quantities of nectar-pollen substrato grains, corolla bases as the source of néctar
and
to facilitate this kind of pollen visitors' the pollen visitors attracted, so these
&
Anderson and
attraction (Zohary, 1973; pollen morphological characteristics
Gensel 1976; Boulos,1979; Al-Quran, 2005). the pollen-nectar as rewards to pollen
orming what blooming
visitors similar to
f is
and
All the palynological terms
works on
time. All previous are stressing
definitions used in general in such studies
by
the seasonality followed the different
LM
are related to the structures seen by and
pollen visitors of insects as the ultímate
ultrastructural parts of the pollen grains
factor of initiation reflecting different
SEM
by
seen are focusing in the ectoaperture
observed
patterns of visitor distribution
and
endoaperture of pollen grains, these
among
examines Onosma
the species of for
complex
structures are considered as a
the collection of pollen-nectar as potential
which with
structures especially deais colpi,
rewards introduced by these plant flowers
vary
costae, fastigia; these structures are
Adailkan
(Thorp,
1979; Stickel et al, 2000;
from These
species to species. structures
& &
Guathaman,
Boulos Al-Eisawi,
2001;
which and
are similar to echinae micro-
&
Lehham,
Boulos
1977; 1978).
form
echinae in of spine-like projections
giving the pollen exine an ornamental The formulation of pollen-insect
may
view be investigated to see whether association between the plant and the
they can depend upon as an important pollen morphology of Onosma species
complimentary with
distinguishing characters in interactions their insect visitors
among
same Bombus bomble
delimitation the species of the (buff-tailed bees)
terestris is
genus or the subspecies of the same species surely reflecting some kind of specialized
morphology
(Rodriguez et al, 1998; Al-Quran, 2010; structural adaptations. Pollen
2011; Sharma, 1974; Samways, 1990). traits regarding ectoaperture, endoaperture
morphology
of pollen grains, fastigia, colpi
obvious from the previous studies the
It is
margins and
of pollen grains, of polar
any image
lack of pollen insect associations
measurements,
equatorial pollen type, size
between Onosma and
the pollen grains of
its
of pollen grains, pollen shape, symmetrical
pollen visitors of buff-tailed bumblebees,
between and
structures pollen grains the
which was most
the intrinsic factor to
and P/E
pollen visitors, therefore ratio
and
kind
initiate this of investigation, in
ARNALDOA
346 2014
21 Julio - Diciembre,
(2):
I
Bombus
^I-Qura’n: 0/705/773 pollen innteractions with terrestris species in Jordán
are playing a major rules to formúlate any taken by using Nikon HFX-11 microscope
way
possible of interactions. Buff-tailed using ocular micrometer scale by glycerin
bomble bees (Bombus was the method.
terrestris) Jelly
most observed and abundant on
especially
SEM
For studies, the treated pollen
Onosma
the flowers of in April.
grains were coated by carbón layer
first,
Four main objectives of this study, they then by gold layer to increase the conduction
whether Onosma and micrographs were
are: to speculate plant electrón yield rates,
1)
flower having some kind of particular taken by SEM.
and
morphological palynological
were
All observations taking place
groups
characteristics to attract certain of
in the area of study of Ajlun in northern
pollen visitors over others. to speculate
2)
heights of Jordán. These observations were
whether the understanding of the flower
from April 2011 to April 2012 during the
rewards form complex
in of nectar-pollen
booming syndrome The methodology used
considered as the motivating factor that
is
Onosma
for recording the species of as plant
may
play the major role of such pollen
by bomble
species visited buff- tañed bees as
whether
visitor attractions. to speculate
3)
was and
pollen according
visitors to visitors
between
the attractions the pollen visitors
and
pollen counts, photographic records
with the pollen grains of flowering Onosma
specimens with entomological
collection
have
particularly in the field area a clear
investigated image concerning this kind of
This study of plant-insect interaction
is
occurred in the plant communities mainly
dominated by certain types of boraginacean
Onosma and
plants especially species
The
collected fresh polliniferous
most them commonly known
of are as
from
material different sites representing
entomophilous plant species. In each
the different jordanian territories, belonging
Onosma and
species studied visited regularly
Onosma (Onosma
to three species aleppica
by pollen visitors, showed some kind of
Onosma and Onosma It
Schreber
Boiss, cinérea
between phenology
correlation flowering
Lam.) by using
frutescens field collecting
and censuses with bomble
insect buff-tailed
and
tools (field viales small brushes).
bees (Bombus the pollen grains
was terrestris),
This material acetolysed typically
LM
were and photographed by
collected
Erdtman
according standard
to
(1960)
and SEM.
method. So 25 randomly chosen acetolysed
from were
pollen grains 5 different slides
and
Results discussion
prepared representing the pollen grains
LM
SEM
Onosma have been prepared All and micrographs taken for
of species to
Onosma
the pollen grains of species studied
be studied to collect the data required in
form measurements by LM. were exposed with their full captions (Fig.
of pollen All 1,
and
2 All palynological data concerning
measurements
palynological related to the 3).
and measurements
the observations of
pollen morphological structures with their
Onosma
were the pollen grains of the studied
calculations estimated or the
statistical f
were and
morphometric species constructed (table 1
data. All the terminologies
The two main
adopted were based on Punt data are collected in
All 2).
et (1994).
al.
LM
measurements concerning
categories
micrographs and measurements were
(1)
ARNALDOA
347
Diciembre
21 Julio -
(2):
I
Bombus
Al-Qura’n: ¿7/705/773 pollen interactions with terrestrís in Jordán
the pollen morphology of the examined important diagonistic factor in recognition
Onosma morphology Onosma
pollen grains All observations of pollen of species
(2)
collected from the field related to pollen- in the endoaperture and ectoaperture
is
insect interactions concerning the pollen which showed some kind of variation
Onosma and bomble on
of buff-tailed bees as reflected their pollen visitor association.
The morphology
pollen visitors. Related to the first category; of ectoaperture structures
examined Onosma and
the pollen class of the (colpi, fastigium margins) for the
species either 3- zonocolporate {Onosma three species exhibited short colpi, absence
is
aleppica Boiss and Onosma frutescens of fastigium with indistinct margins for
Lam.) or 4-zonocolporate {Onosma cinérea Onosma aleppica Boiss, long colpi, presence
measurements and margin
Schreber). Polar ranges of fastigium indistinct for
were Onosma and
are identified, they as follows: (37- cinérea Schreber short colpi,
pm
Onosma and margin
for aleppica Boiss, (42-45) presence of fastigium distinct of
43)
pm
Onosma and Onosma Lam. The morphology
for cinérea Screber (41-44) frutescens of
pm
for Onosma frutescens Lam. Equatorial endoaperture structures (colpi, costae and
measurement
ranges are also identified margins) has reflected also distinguished
pm
Onosma Onosma showed
as follows: (26-34) for aleppica variation; aleppica Boiss
pm
Onosma and
Boiss, (28-34) for cinérea Screber short colpi, presence of costae indistinct
pm
and Onosma Lam. Onosma showed
margin, Schreber
(28-33) for frutescens cinérea
P/E
ratio are found to be ranged between short to long colpi in general, absence of
Onosma and
(1.41-1.39) for aleppica Boiss, (1.49- costae distinct margins.
Onosma and
Schreber
for cinérea
1.33)
With second
respect the category;
to
Onosma Lam. The
(1.45-1.28) for frutescens
the insect-plant initiation of interactions
is
morphology
of ectoaperture structures
morphology
refered to the pollen of studied
and
(colpi, fastigium margins) for the
species (long or short colpi, distinct or
three species exhibited short absence
colpi,
and
endoapertures
indistinct ectoapertures,
of fastigium with indistinct margins for
and
presence or absence of fastigia costae).
Onosma long presence
aleppica Boiss, colpi,
Onosma
Also species in Jordán as well as
and margin
of fastigium indistinct for
for boraginacean pollen grains produces
all
Onosma and
Schreber short
cinérea colpi,
amount
a large of whitish polliniferous
and margin
presence of fastigium distinct of
and
dust constitutes a potential source of
Onosma Lam. The morphology
frutescens of
pollen for different species of buff-tailed
and
endoaperture
structures costae
(colpi,
bomble providing an
bees, interesting field
margins) has reflected also distinguished
such worthy
for attraction of insects.
It is
Onosma showed
variation; aleppica Boiss
depending on
to say that the previous
and
short colpi, presence of costae indistinct
and measurements
palynological results
Onosma showed
margin, Schreber
cinérea
from Onosma
obtained pollen grains of
short to long colpi in general, absence of
studied, obviously cannot be relied on in
is
and
costae distinct margins. Pollen outline
Onosma
delimitations of species species
at
has been recognized, the equatorial view
on
level, but surely can be relied to explain
Onosma examined was
for species
all
between
the intímate relationship the
view was
rectangular-elliptic; the polar
morphology of pollen grains with pollen
P/E
view
triangular for species. ratio
all
Bomhus
visitors of to explain this
terrestris,
was examined. The most
erect for species
all
point very important reconize
clearly, to
it is
ARNALDOA
348 2014
21 Julio - Diciembre,
(2):
I
Bombus
^I-Qura’n: 0/705/773 pollen innteractions with terrestris species in Jordán
the presence of such differences between deposits blocking the anther anthesis to
the pollen grains related to polar (P) and expel the pollen grains is rising. So in this
equatorial measurements of pollen grains way, worthy to assure the presence of
it is
by an between and
exhibited ecto-and endo-apertures cióse relationship the type size
morphology which morphology and
are playing the role of pollen the pollination
syndrome by "buzz
of variations in pollen-visitor attraction pollination" reinforcing
among
and from
the different species of Onosma, the observations collected the
fields.
this was seen clearly in most of the pollen- Easily also, is obvious that small sized
it
showed
insect communities studied the pollen grains with ornamentation will
little
and more from
presence of specialized intímate insect- be expelled easily the poricidal
flower associations, this refers in some anthers during the vibration of the honey
Onosma Edmond, Adam,
aspects to the complicated flower bees (Erickson,1975; 1984;
morphologies which hinders or facilitates 2001; Al-Quran, 2004a, 2004b; Buchman,
kind depending on Thorp,
this of visitation the 1986; 1979).
species visited by wide range of other insect
The and
observations data collected
groups. So clear that Bombus
it is terrestris
showed
the presence of other types of
colonias are active approximately in over
all
honey
insects as bees (Apis
mellifera),
from
the year seasons extending winter to
Bombus and
Apis A.
impatiens, dorsata
florea
summer autumn
to late to collect as large
and wasps from
(Pterygophorus
insignis)
possible quantities of néctar as they can
low
species but in frequencies. All the
so these foragers appeared in the study
, pollen visitors observed used the "buzz
on show
area early in the season plants that
procedure during
pollination" their
high densities and high néctar productions
The
foraging behaviour. previous published
on Onosma
especially flowers.
"buzz
studies report that pollination
more syndrome" hymenopteran
In the investigated pollen requires species
details,
grains of Onosma were indicating the with a specific behavior for pollen removal
some which
presence or absence of morphological typically applied to the bees
is
from and Bombus
characters especially in the pollen Apis
mellifera terrestris
ornamentations, the size of the pollen species. important to demónstrate
It is
grains and the occurrence of the the importance of this kind of association
little
complex some between improve
variated of pollen ecto-and the partners to the
endo-apertures of some species of Onosma apiculture management in honey. So the
highly related to the shedding process of previous studies were trying to provide
is
these grains inside of the poricidal anthers evidences of such pollen collection strategy
during anthesis process. The shedding from pollen grains of Onosma species by
Adam,
process playing the most important role such kind of bees (Buchman, 1986;
is
was
in playing the execution through pollen 2001). There a lot of previous workings
visitor vibrations or "buzz pollination" by investigating similar aspects concerning
bomble by working with
visiting buff-tailed bees. Exactly the interrelationship Vespa
on same on Anchusa
relying the principie of expression, orientalis L. the pollen grains of
wherever the larger the pollen grains of some species. These results obtained from the
studied Onosma species with high complex previous workings approved these results
and showed
pollen endoaperture ectoaperture obtained since they that the
ornamentations, wherever the case to form collection of pollen grains by the pollen
ARNALDOA
349
Diciembre
21 Julio -
(2):
I
Bombus
Al-Qura’n: ¿7/705/773 pollen interactions with terrestrís in Jordán
visitors due to such pollen characteristics the bees the major aspect. Of course the
is
amount
relatad to the ornamentations of pollen bees are exerting a larga of energy
move
endo-and and muscles and complete
ecto-apertures, these results to their flight to
Buchman
consequently are clerifying the importance the vibration process.
(1986)
of the palynological data to understand confirmad the presenceto a certain degree
this association depending on pollen the presence of waste of pollen grains in
morphology of pollen grains and the this process, and only pollen with a high
visitors (Pyle et al, 1991; Rates, 2001; Peter, protein level could justify this plant-insect
&
Anderson
1994; Ricklefs, 2004; Gensel, relationship.
Friedman
Joud
1976; et al, 1986; et al, 2001;
Conclusión
Eddouks
etal, 2002).
The building of such specialized and
Edmonds
from
obtained similar
(1984)
between
intimate associations the pollen
on some
studies Solanaceae species
morphology Onosma and
of the pollen visitor
showing
(Solanum) species that the
{Bombus due need
clearly to the
terrestris) is
morphological variations in pollen grains
of such bees to collect the néctar and pollen
genus Solanum
of the (Solanaceae) reflectad
by preelude
default, this certainly a certain
no any morphological
significant variations,
morphological adaptational
similarities
because theyn refered only exine
to
between he
partners in the plant-pollinator
ornamentation
level related to the certain
community
interactions of the to collect a
structures of ecto-and endo-apertures
high quantities of néctar forage as soon as
morphology, what happening
similar to
is
possible in short time and straight way.
Anchusa
in species.
may
These an
results give indication that
The ornamentation
variation in character
flowers with similar reward composition
morphology
of ecto-and endo-apertures of
tended groups
to attract similar of visitors.
pollen grains considerad the crucial factor
is
In this regard, the pollen visitor responsos
determining the phylogenetic relationships
reward and
to flower néctar as floral
among
genomic
the species in addition to
main
considerad as motivating factor rather
and
combinations,
isolation speciation,
than responsing to the flower morphology.
confirmad by Al-Quran
this result
(2004)
The
using of the palynological study as
through his study of pollen grains of 11
taxonomy and
palynological evidence in
Hyoscyamus from
species of collected
Onosma
delimitation of the three species
is
different parts of Jordán.
mainly not worthful, because the depending
Other (Buchmann,
researches 1986;
on and measurements
the pollen structures
Erickson, 1975; Thorp, 1979) were going LM SEM
seen by and be depend
can't
further by demonstration the presence of
upon
in such kind of delimitation because
certain electrostatic forces, facilitating the
of the high degree of similarities between
body
attachment of the pollen to the of
examinad which showed
the species
moment
by
the insect at the of pollination
no
clear differences in the diagonistic
through blooming syndrome,
vibration the
features exhibited, consequently they have
which
transference
facilitates finally to
its same
with
exhibited the inter-relationship
the stigma. So the presence of a cost-benefit
hymenopteran By same
species. the reason,
an important involved
relationships fact in
worthy depend on morphological
the
to
it is
syndrome by
the pollination vibrations of why
variations of pollen grains to explain
ARNALDOA
350 2014
21 Julio - Diciembre,
(2):
I
.
Bombus
^I-Qura’n: 0/705/773 pollen innteractions with terrestris species in Jordán
do some
pollen visitors are attracted to noptera, Apoidea) flowers of Solanum palina-
in
more Dunal (Solanaceae). Revista Brasileira de
certain plant pollens than the others C3/7//7Í//77
35-44.
Zoociéncias,
3
(1):
The by
previous results obtained the
&
Adailkan, R G. K. Gauthaman. 2001 History of her-
.
showed
researcher have the presence
bal medicines with an insight on the pharmacologi-
must
of four different aspects be taken cal properties of Tribulus terrestris. The Aging Male,
4:163-169.
in consideration to reach a better
understanding the linkage between pollen
Al-Eisawi, D. 1982. List of Jordán Vascular plañís. The
Ammán,
morphology of Onosma and the pollen Jordán University Press, pp 22-39.
{Bombus phenology
visitor
terrestris):
(1) Al-Qura’n, S. 2004a. Pollen Morphology of Plantagina-
between 1594-1602.
construction the the partners cae in Jordán. Pak. J. Biol. Sci, 7 (9):
morphology
associated of the pollen
(2) Al-Qura’n, S. 2004b. Pollen Morphology of Solanaceae
morphology,
visitors, pollen nectar- in Jordán. Pak. J. Biol. Sci, 7(9): 1586-1593.
(2) (3)
pollen rewards given by the flower the
to
Al-Qura’n, S. 2005. Ethnobotanical Survey of Folk To-
the pollen visitor. In other words, the better xic Plañís Southern Parí of Jordán. Toxicon, 46:
In
119-129.
understanding importance between
of
morphology, rewards and
pollen
floral
Al-Quran, S. 2010. Pollen characteristics of three Pa-
flowering time of each plant species very species and the observation of Polyphylla as
is
why thew main pollen collector. Pak. Enntomol. 32
important do (2):
to to interpreting certain
116-124
insects to be attracted to a certain flowers
Al-Quran, S. 201 Pollen characteristics of three An-
not 1
definitely to others. .
chusa species and the observation of bees as the
3-
This kind of studies with other pollen collector Jordán. Pak. Etomol. 33
in (2): 1 1
117.
complementary should be
studies intensified
further to have a better understanding Anderson, G. & R G. Gensel. 1976. Pollen morpho-
J.
between logy and the systematics of Solanum Section. Ba-
of the existing relationships the
sarthrum. Pollen et Spores, 18: 533-552.
and
pollinators the attracted species to justify
the occurrence of pollination syndrome Bender, F. 1974. Geology of Jordán. Berlín, Stuttgart.
by vibration found in the observed bees in &
Boulos, L. D. El-Eisawi. 1977. On the flora of Ras en
form of "buzz pollination", so this kind Naqab. Candollea 32: 81-98.
of association notonly foraging strategy,
is &
Lehham. On
Boulos, L. J. 977. the desert flora north
1
but in rather is intímate reciprocal kind of east of Aqaba. Candollea, 32: 99-1 1 0.
exchange between
benefits the partners.
On
Boulos, L. 1977. the flora of El-Jafer- Bayir desert.
32:99-110.
Candollea
main
Finally the conclusión can be
Buchmann, Solanum
1986.
relied on wherever the pollen grain has a S. L. Vibratile pollination in
is,
and Lycopersicorr. a look at pollen chemistry, W.
reduced ornamentation
the aspects in
size,
G. D’Arcy (ed.). ///Solanaceae: biology and syste-
and
ecto-and endo-apertures psilated exine
New
Colombia
matics. University Press, York.
pp.
together sharing to give the outline of 218-252.
all
moprphology
pollen to appropriate certain
L
Eddouks, M.; M. Maghrani; A. Lemhadri; M. Quahi-
behavioral form, allowing pollen grain &
H. Joud. 2002. Ethnopharm-acological sur-
di.
access by small sized bees as observed with vey of medicinal plants used for the treatement of
Diabetes mellitus, hypertension and cardiac disea-
bomble
buff-tailed bees species.
morocco
ses in the south-eastern región of (Tafila-
Ethnopharmacology, 82: 97-103.
let). J.
Literature cited
Edmonds, M. 1984. morphology Solanum
J. Pollen of
Adam, M. S. 2001 Foraging behavior of bees (Flyme-
.
L. section Solanum. Bot. J. Linn. Soc. 88: 237-251
ARNALDOA
Diciembre 351
21 Julio -
(2):
I
Bombus
Al-Qura’n: ¿7/705/773 pollen interactions with terrestrís in Jordán
Erdtman, G. 1960. The acetolysis method in a revised forests in Venezuela? Biol. Conserv., 83: 69-76.
description. Svensk Botanisk Tidskrift, Lund, 54
Samways, M. 1990. Insect conservaron ethics. En-
J.
561-564.
(4):
7-8.
viron. Conserv., 17:
Erickson, E. H. 1975. Surface electric potentials on
Sharma, B. D. 1974. Contributions to the palynota-
worker honeybees and
leaving entering the hive.
J.
xonomy genus Solanum
of the Linn. J. Paly., 10:
141-147.
Apic. Res., 14:
51-68.
&
Friedman, J.; Z. Yaniv; A. Dafni D. Palevitch. 1986.
&
Stickel, R; G. Egerer H. K. Seitz. 2000. Hepatotoxici-
A
preliminary classification of the healing potential
113-124.
ty of botanicals. Pub. Heal. Nutr., 3:
of medicinal plants, based on a rational analysis of
among
an ethnopharmacological fieid survey Be- Thorp, R. W. 1979. Structural, behavioral, and physio-
douins in the Negev desert, Israel. J. Ethnopharma- logical adaptations of bees (Apoidea) for collecting
275-278.
col., 16: pollen. Ann. Miss. Bot. Gard., 66: 788-812.
&
Joud, H.; M. Haloui; H. Rhiouani; J. Ehilaly M. Zohary, M. 1973. Geobotanicalfoundationsofthe midd-
Eddouks. 2001 Ethnobotanical survey of medici- Gustav.FisherVerlag 30-55.
east. Stuttgart,
le pp
.
nal plants used for the treatment of diabetes, car-
&
Zohary, M. N. Feinbrun-Dothan. 1962-1686: Flora
Morocco
diac diseases the north center región of
in
Palaestina, Flebron University Press, Jerusaiem,
pp
175-
(Fez-Boulemane). Ethnopharmacol. 77:
J.
77-90.
182.
&
Karim, S. Al-Qura’n. 1988: Wild Flowers of Jordán.
F.
Yarmouk
University Press, Jordán.
Irbid,
&
Karim, S. Al-Qura’n. 1986. Medicinal plants of
F.
Yarmouk
Jordán. University Press, Jordán.
Irbid,
Pp 11-30.
&
Khayyat, A. A. M. Mursi. 1981. Pharmacology and
veterinary toxicology in Iraq, Ist ed., Ministry of
higher Education Press, Baghdad. pp14-33.
Peter, K. E. 1994. Special differentiations associated
and
with pollinator attraction. Diversity Evolutio-
In:
nary Biology of Tropical Flowers. Cambridge Tropi-
148-189.
cal Biology Series, pp.
A
Prance, G. 2001 dictionary of natural products:
T.
.
Terms phamacognosy
in the filed of relating to
medicinal and pharmaceutical materials and the
whom
plants, animáis, and minarais from are de-
301-302.
rivad. Biodivers. Conserv. 10:
Puní, W.; S. Blackmore & A. Le Thomas. 1994. Glos-
sary of Pollen and Spores Terminology. Utrecht,
LPP
Foundation.
&
Pyle, R. M.; Bentzien R Opler. 1991. Insect conser-
233-258.
varon. Ann. Rev. Entomol., 26:
Rales, S. M. K. 2001 Plants as source of drugs. Toxi-
.
con,39: 603-613.
A
2004. comprehensivo framework
Rickiefs, E. A. for
1-15.
global pattern in biodiversity. Ecol. Lett, 7:
&
Rodríguez, R; D. Pearson R. R. Barrera. 1998.
J. L.
A
test for the adequacy of bioindicator taxa: Are
ti-
ger beetles (Coleóptera: Cicindelidae) appropriate
indicators for monitoring the degradation of tropical
ARNALDOA
352 2014
21 Julio - Diciembre,
(2):
I