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Floral rewards and pollination in Cytiseae (Fabaceae)

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DOI: 10.1007/s00606-002-0270-5Floral

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Plant Syst. Evol. 238: 127–137 (2003)
DOI 10.1007/s00606-002-0270-5
Floral rewards and pollination in Cytiseae (Fabaceae)
M. Galloni and G. Cristofolini
Department of Biology, University of Bologna, Italy
Received August 12, 2002; accepted November 25, 2002
Published online: June 2, 2003
Ó Springer-Verlag 2003
Abstract. Cytiseae have been reported to be mostly
nectar-lacking, yet some taxa secrete nectar from
extrastaminal nectaries. We studied the pollination
biology of four shrubby species of Cytiseae (Cyti-
sophyllum sessilifolium (L.) Lang, Spartium junceum
L., Genista radiata (L.) Scop., Genista cilentina
Valsecchi) which differ for ecology, distribution
and population size. All species resulted obliged
xenogamous, insect visits being necessary for suc-
cessful pollination. Bumblebees and solitary bees
are the principal pollinators, but also many beetles,
some hover-flies, and few bugs visit flowers. Pol-
linator specificity is low, and this may be the reason
of the scarce seed set compared to the number
of ovules. Pollen is the main reward, but traces of
glucose were detected in all species, at the base of
vexillum or on the reproductive column. Nectar
production is irregular in time, and apparently
unpredictable. We suppose that nectar may play a
role in attracting pollinators determining their right
position for a successful pollination.
Key words: Brooms, Cytiseae, extrastaminal
nectaries, Fabaceae, nectar, pollen presentation,
pollination.
Introduction
The biological structure of plant species is
mainly determined by their reproductive strat-
egies.
Correlation between pollination ecology,
mating system and population size has been
demonstrated in several cases (Karron 1987,
Petanidou et al. 1995, Corbet 1996, López et al.
1998, Tepedino et al. 1999, Rodriguez-Riaño
et al. 1999, Utelli and Roy 2000, Bosch et al.
2002, Tandon and Shivanna 2001).
The mutualistic relationship between plant
and pollinators should be investigated espe-
cially in the case of predominantly outbreeding
animal-pollinated plant species, in order to
better understand which are the factors that
affect pollination fitness and consequently the
species’ reproductive output and dispersion.
This kind of studies could be very important
for conservation purposes.
The tribe Cytiseae Bercht. & J. Presl,
belonging to the family Leguminosae, is rep-
resented by 22 genera according to Bisby et al.
(2001). It includes several rare and threatened
species. In spite of the abundant taxonomic
and molecular information available, until
now few investigations on pollination ecology
have been done on representatives of this
group.
Molecular investigations on some species
of Leguminosae suggested that a relationship
between population size and the degree of self
compatibility should exist (Conte et al. 1998,
Conte and Cristofolini 2000).
128 M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae
Flowers of Cytiseae are characterised by
the typical papilionaceous structure. The sta-
mens’ filaments are united at the base, forming
a monadelphous androecium. By contrast, the
majority of Fabaceae present flowers with
diadelphous stamens, which produce nectar
as major attractant and reward for pollinators.
The functional connection between nectar
production and diadelphy was first discovered
by H. Müller (1873). In diadelphous flowers
nectar is usually secreted by an intrastaminal
nectary – a specialised glandular tissue of the
mesenchymatic type – which generally consti-
tutes a distinct perigynous disk (Vogel 1977).
Nectar accumulates between the carpel base
and the filaments, and then it is sought below
the vexillar petal at the base of the reproduc-
tive column, where there are two openings
(Vogel 1997).
Monadelphous flowers are supposed to be
nectar-lacking because their completely fused
staminal tube (without any opening at its base)
implies the absence of the perigynous disk.
Therefore, members of the tribe Cytiseae have
been considered nectar-lacking for a long time
(Delpino 1868/69 cited by Vogel 1997, Meeuse
1961).
Yet, in the tribe Cytiseae there are excep-
tions to this rule: in fact secretion of free floral
nectar has been detected in several taxa of
European and Mediterranean brooms and
gorses like Retama and Cytisus (Polhill 1976,
Bisby 1981, Vogel 1997, Westerkamp 1997,
Rodriguez-Riaño et al. 1999).
Past investigations proved that no glandu-
lar tissue is present on the surface of the flower
parts that could be responsible for the external
secretion of nectar (Vogel 1997). The nectary
of nectar-producing flowers with monadel-
phous staminal column has been described as
‘‘extrastaminal’’ by Bisby (1981), while Vogel
(1997) refers to ‘‘non-structural staminal nec-
tary’’: in fact, as no specialised tissue is
present, filament tube is itself the source of
nectar, which is then exposed to the base of
androecium, mainly inside the wings and
standard insertions. Nectar probably repre-
sents leaking phloematic exudate.
According to recent literature only few
species of Cytiseae produce nectar, while all
the remaining species are believed not to
produce any nectar. Evidence for nectar secre-
tion has been reported for the following species
from direct observations of the visitors’
behaviour and from tests by glucose reagent
strips: Cytisus hirsutus L., Cytisus purpureus
Scop., Cytisus ruthenicus Fischer ex Woloszak,
Cytisus proliferus L. fil., Cytisus proliferus var
palmensis Christ, Spartocytisus supranubius
Webb et Berth., Erinacea anthyllis Link, Pet-
teria ramentacea (Sieb.) Presl, Retama raetam
(Forsskål) Webb, Retama monosperma (L.)
Boiss. (Vogel 1997).
In other cases past and recent investiga-
tions led to different and contradictory results:
e. g. the flowers of Laburnum anagyroides have
been described as nectariferous by some
authors (Müller 1873, Daumann 1941), and
as nectar-lacking by others (Vogel 1997).
The present investigation was conducted
mainly in the field during three flowering
seasons (2000–2002). We studied the pollina-
tion biology of four shrubby species of Cyti-
seae (Cytisophyllum sessilifolium (L.) Lang,
Spartium junceum L., Genista radiata (L.)
Scop., Genista cilentina Valsecchi), which
present similar flower structure but differ for
ecology, distribution and population size.
Our aim was: (i) to investigate the breeding
system of the four species in natural condi-
tions, determining the dependence on insect
pollination to set seeds, (ii) to typify the floral
rewards offered to pollinators, (iii) to monitor
pollination activity and to define a complete
spectrum of pollinators and visitors, (iv) to
describe the pollination mechanisms.
Materials and methods
The species studied
Cytisophyllum sessilifolium (L.) Lang (= Cytisus
sessilifolius L.) is a south-west-european-mediter-
ranean species, growing in open broad-leaved
woods, thickets and wood-rims. Flowers produce
a delicate citron-like scent. Flowering season lasts
from mid-April to the end of May.
M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae
Spartium junceum L. is an eurimediterranean
species, widely distributed and quite ubiquitarian in
its geographic range, commonly present in post-
coltural vegetation. Flowers are strongly sweet-
scented. Blooming period lasts from May to end of
June.
Genista radiata (L.) Scop. (= Cytisanthus
radiatus Lang) is a south-european orophyte,
whose main distribution range is in the eastern
Alps; on the Apennines it is present in small and
isolated disjunct populations. It grows as a poly-
cormic cushion-like shrub up to 50 cm high,
forming extended compact populations in open
soils, but it behaves also as a coloniser of debris
and nude rocks. The flowers are sweet-scented. It
flowers from June to July.
Genista cilentina Valsecchi belongs to the
Genista ephedroides species-complex, which is dis-
tributed around the Thyrrenic sea. It is endemic to
the clay rocky cliffs and slopes near the sea between
Ascea Marina and Palinuro (Salerno – Southern
Italy), where it forms very small populations. The
flowers are sweet-scented. Anthesis occurs from
April to May.
Specimens of the four species have been
collected from each population studied: vouchers
are preserved in the Herbarium of the University of
Bologna (BOLO).
The study sites
Cytisophyllum sessilifolium and Spartium junceum
were monitored in an area within the natural
park ‘‘Parco dei Gessi bolognesi e dei Calanchi
dell’Abbadessa’’, about 10 km eastwards from
Bologna.
Genista radiata was studied at three different
sites: a pre-alpine population located in Pian delle
Fugazze (Trento), at 1160 m elevation, and two
populations on the Apennines, located respectively
within the ‘‘Corno alle Scale’’ natural park, at
1680 m elevation, and on Monte Beni, at 1090 m
elevation, at a Site of Community Importance,
included in the NATURA2000 European network
of protected areas.
Genista cilentina was studied on two popula-
tions located, respectively, on the ‘‘Punta della
Torre del Telegrafo’’ close to Ascea Marina, near
the sea coast, and on a south exposed cliff between
Pisciotta and Marina di Camerota, about 150 m
above the sea.
129
Flower morphology
The species studied present yellow flowers with
papilionaceous corolla: the immovable claw of
vexillum at the upper side, and the wing-keel
complex on the lower side. The two wings are
parallel to the keel. Anthers are dimorphic: linear-
oblong basifixed anthers are alternate with shorter
dorsifixed ones (with more or less triangular-
acuminate apex), corresponding respectively to
the lower and upper whorl of stamens.
The style is curved upwards in all species;
stigma is fibrous, introrse and pad-like in Spartium
junceum, papillate and extrorse in Cytisophyllum
sessilifolium, papillate and terminal in Genista
radiata, papillate and terminal-extrorse in Genista
cilentina.
Breeding system
The breeding system was studied in the field,
comparing fruit and seed set in a series of bagged
racemes (where only spontaneous self-pollination
was possible) and non-bagged control racemes
(open to natural pollination). Two racemes were
selected and marked at beginning of anthesis in 10
to 20 plants for each population. One raceme per
plant was bagged when flowers were still in
blossom; the bags were taken away at the end of
flowering season, and replaced before fruit matu-
ration to collect the seeds produced.
We determined ovule production per flower for
each species by direct observation of 40 flowers’
ovaries, from 20 plants per population.
Floral rewards
We checked the presence of nectar with a simple
device made up using Glukurtest strips (Roche),
especially designed to act as an insect’s ligula, so
that even very small amount of glucose could be
detected. This method is very useful for field
studies, especially when the study sites are located
far away from laboratories: it permits to detect the
presence of glucose on the surfaces of flower parts.
Past studies proved that ‘‘most fabaceous nectars
have a balanced content of the three main sugars’’
(Percival 1961); assuming that glucose is always
present in nectar composition of other species
belonging to the family Fabaceae, glucose-test
130 M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae

paper can be used to detect presence of nectar, Results

even if no information on sugar volumes and
percentage is given (Westerkamp 1997, Vogel
1997). For each species we repeated this test both
in flowers before triggering, which were stimulated
to trigger during the experiment, and in flowers
after triggering, at several times of the day
(possibly each hour from 9 a.m. to 18 p.m.),
during different days (at least six days) at different
flowering stages.
Insect activity
Direct observations of insect activity let us obtain
a spectrum of pollinators and visitors for each
species. Each flowering season we dedicated at
least two entire days of observation on insects’
activity for each population. We recorded the
behaviour of visitors on small groups of plants
over an area of about 4m2, and eventually
sampled them during periods of 15’, at intervals
of 30’. The total time spent in studying the
interactions between plants and pollinators was
21h for Cytisophyllum sessilifolium, 10h for Spar-
tium junceum, 8h for Genista radiata and 13h for
Genista cilentina. Sampled insects were identified,
and pollen on their bodies was collected, stained
with ‘‘Calberla solution’’ (Dafni 1992) and mount-
ed on microscope slides. To identify pollen grains
we analysed them under a light microscope and
compared with reference pollen collected from
known specimens.
Breeding system
Bagged flowers almost never produced seed
(Table 1); by contrast, control racemes did pro-
duce seeds; nevertheless, the seed set compared
to the number of flowers resulted quite low
(Table 2): considering the mean values between
years, Cytisophyllum sessilifolium produced 10
seeds per 100 flowers; Spartium junceum pro-
duced 72 seeds per 100 flowers; in Genista
radiata seed set was 25 seeds per 100 flowers; and
in Genista cilentina 16 seeds per 100 flowers.
Comparing mean values of each season, we
observed some fluctuations in seed production:
in some cases differences were particularly
marked (Genista radiata, Spartium junceum).
Seed set resulted even lower when compared
to the number of ovules (Table 3): in Cytiso-
phyllum sessilifolium a mean of 1.4 seeds reached
maturity for 100 ovules, while Genista radiata
and Genista cilentina yielded respectively a
mean of 5.3 and 2.8 mature seeds per 100
ovules. Considering the mean values between
years, Spartium junceum shows the highest
seeds/ovules ratio (4. 8 seeds per 100 ovules).
Floral rewards
Traces of glucose were detected in all
species studied (Table 4): in Cytisophyllum

Pollination mechanisms and stigma receptivity Table 1. Seed production in bagged flowers

We studied pollination mechanisms both by direct Species Year N° N° N°

field observation of insect activity and by manual Plants Flowers Seeds
stimulation of ‘‘closed’’ flowers, in order to observe
Cytisophyllum 2000 19 402 0
the mechanism of pollen release. We collected
sessilifolium 2001 18 855 1
flowers in different blooming stages to observe and
2002 21 736 0
record anthers’ stage and to check stigma receptiv-
ity. For this purpose we used special peroxidase Spartium junceum 2001 9 236 0
test-papers (Macherey-Nagel Peroxtesmo KO per- 2002 12 521 0
oxidase test paper) following the method described Genista radiata 2000 15 436 0
by Dafni and Maués (1998). (Corno alle 2001 15 1176 0
To detect the presence of pollenkitt, fresh Scale) 2002 15 843 7
pollen grains have been stained with Scarlet R (Monte Beni) 2001 20 1170 0
and observed at light microscope: pollenkitt ap-
Genista cilentina 2001 10 737 2
pears as red droplets over the surface of the exine
2002 13 1181 4
(Nepi and Franchi 2000).
M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae 131

Table 2. Seed production in control racemes (flowers free-pollinated): ratio seeds vs. flower
Species Year N° N° N° Seeds Seeds/Flower Seeds/Flower
Plants Flowers (mean ± s.e.) (Overall mean ± s.e.)
Cytisophyllum 2000 18 467 51 0.13 ± 0.05 0.10 ± 0.02
sessilifolium 2001 18 828 57 0.08 ± 0.02
2002 21 752 76 0.09 ± 0.03
Spartium junceum 2001 9 269 51 0.22 ± 0.09 0.72 ± 0.15
2002 12 616 565 1.09 ± 0.20
Genista radiata 2000 15 465 55 0.11 ± 0.02 0.25 ± 0.02
(Corno alle Scale) 2001 15 1247 394 0.32 ± 0.05
2002 15 1218 360 0.29 ± 0.05
(Monte Beni) 2001 20 1269 344 0.28 ± 0.04
Genista cilentina 2001 10 849 170 0.20 ± 0.03 0.16 ± 0.03
2002 13 1097 114 0.13 ± 0.04

Table 3. Seed production in control racemes (flowers free-pollinated): ratio seeds vs. ovules
Species Ovules/Flower Seeds/Flower Seeds/100 Ovules
(mean ± s.e.) (mean ± s.e.) (mean ± s.e.)
Cytisophyllum sessilifolium 7.1 ± 0.2 0.10 ± 0.02 1.4 ± 0.3
Spartium junceum 15.1 ± 0.3 0.72 ± 0.15 4.8 ± 1.0
Genista radiata 4.7 ± 0.1 0.25 ± 0.02 5.3 ± 0.4
Genista cilentina 5.8 ± 0.2 0.16 ± 0.03 2.8 ± 0.5

Table 4. Nectar production in closed flower (manually triggered during the test), and in naturally triggered
flowers. Positive reaction is expressed as ratio between number of flowers positive to glucose-test and total
number of flowers tested
Species Flowers before Flowers after
triggering triggering

N° flowers Positive reaction N° flowers Positive reaction

(mean ± s.e.) (mean ± s.e.)
Cytisophyllum sessilifolium 184 0.29 ± 0.05 85 0.16 ± 0.05
Spartium junceum 38 0.78 ± 0.08 12 0.70 ± 0.20
Genista radiata 27 0.91 ± 0.06 11 0.55 ± 0.16
Genista cilentina 47 0.86 ± 0.07 16 0

sessilifolium and Spartium junceum nectar is
present mainly at the base of vexillum, while in
Genista radiata and Genista cilentina it is
produced on the reproductive column. In both
Genista species secretion of nectar occurs only
in triggered flowers, immediately after the
release of reproductive column from the keel.
Nectar production is very irregular and un-
predictable. Apparently there is no correlation
between nectar secretion and the time of the
day.
In the case of Cytisophyllum sessilifolium
there was a remarkable difference between
different years: in 2001 nectar secretion was
observed very rarely, while in 2002 traces of
nectar were commonly detected at least
during the early and middle stages of
anthesis.
132 M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae

Visitor species and behaviour were congru-
ent with nectar analysis: during both years we
observed some nectar-feeding insects visiting
the flowers (such as long-tongued Antophora
and male solitary bees), and in 2002 we
recorded several Bombylius sp. clearly foraging
nectar on the flowers.
Pollination activity
Many different species of insects use to visit the
plants (Table 5): we recorded many beetles
(Order Coleoptera), some hover-flies
(Syrphidae, Order Diptera), few bugs (Order
Hemiptera); the most frequent visitors are
bumblebees and solitary bees (Order Hyme-
noptera), whose behaviour on the flowers
suggests their role as principal pollinators. In
fact, hymenopterans touch the floral repro-
ductive organs sternotribically: they usually
are female bees that actively gather pollen with
the aid of structural adaptations (corbiculae,
scopae, etc.) that help its collection and
transport back to the nest for larval nutrition.

Table 5. Insect taxa observed visiting flowers of the four Cytiseae. Cyt = Cytisophyllum sessilifolium,
Cil = Genista cilentina, Rad = Genista radiata, Spa = Spartium junceum
Cyt. Cil. Rad. Spa.
Hymenoptera Apoidea
Andrena sp. 4 4 4
Anthidium sp. 4 4 4
Anthophora plumipes 4 4
Bombus lapidarius 4 4
Bombus pascuorum 4
Bombus pratorum 4
Bombus terrestris 4 4 4
Ceratina cucurbitina 4
Eucera sp. 4
Halictus sp. 4
Megachile cfr.centuncularis 4
Megachile sp. 4
Osmia cornuta 4
Osmia rufa 4
Xylocopa violacea 4 4
Other Hymenoptera
Eumenidae 4 4
Coleoptera
Alleculidae (Omophlus sp.) 4
Attelabidae (Deporaus sp.) 4
Cantharidae (Rhagonicha cfr.fulva) 4 4
Chrisomelidae (Cryptocephalus sp.) 4
Dermestidae 4 4
Oedemeridae 4 4 4
Scarabaeidae (Tropinota hirta) 4
Staphylinidae 4 4
Diptera
Bombyliidae 4
Muscidae 4
Syrphidae 4 4 4 4
Hemiptera
Miridae 4 4
M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae 133

We observed 19 different species of visitors observed on the plants, but their behaviour
on Cytisophyllum sessilifolium. Among them, and function were not clear.
12 were Hymenopterans belonging to 5 differ-
ent families (Table 6): Halictidae (Halictus
Pollination mechanisms
sp.), Megachilidae (Osmia spp., Anthidium
sp.) Apidae (Bombus spp., Xylocopa violacea), Pollenkitt is present in all species.
Andrenidae (Andrena spp.) and Antophoridae Pollen release in the four species follows
(Antophora plumipes). This high variety indi- different ways, which cannot always be referred
cates low specificity in plant-pollinator rela- strictly to the four categories described for
tionship. Fabaceae (Delpino 1868/1869, Leppik 1966
The specificity is low also for Genista cited by López 1999).
cilentina: we sampled 15 visitor species, 9 of Cytisophyllum sessilifolium presents pollen
which were hymenopterans. Most visits were with a mixt piston-to-valvular mechanism: ini-
made by solitary bees and bumblebees belong- tially, when the insect move the keel down-
ing to the families Andrenidae, Apidae, Anto- wards, pollen is pushed out from the keel tip, the
phoridae and Megachilidae. petals remaining stiffened and the stigma con-
Genista radiata shows a higher specificity to cealed inside (see also Polhill 1976); after several
pollinators, being constantly visited by 2 Bom- visits, the keel opens along its total length
bus species and by a small bee belonging to the
genus Andrena in all the populations surveyed.
We suppose that the environmental conditions
(high altitude, low temperatures, strong wind)
act as selecting factors on insect diversity.
Surprisingly, the very common species
Spartium junceum shows the highest specificity
to pollinators, possibly because of the size of
its flowers, that need a strong force to be
triggered. The main pollinator is the big
solitary bee Xylocopa violacea, whose visits
always lead to explosive tripping. It is also
visited by smaller megachilids (Anthidium sp.),
but they often do not provoke flower trigger-
ing and consequently there is no pollen release.
Many beetles and hover-flies have also been

Table 6. Frequencies (%) of the different families

of Hymenoptera Apoidea observed visiting flowers
of the four Cytiseae. Cyt = Cytisophyllum sessili-
folium, Cil = Genista cilentina, Rad = Genista
radiata, Spa = Spartium junceum
Families Cyt. Cil. Rad. Spa.
Andrenidae 15.0 38.7 12.7 –
Antophoridae 12.0 17.6 – –
Fig. 1. Spartium junceum. Flowers before and after
Apidae 22.8 24.4 86.4 55.6
the explosive mechanism: after triggering the flower
Halictidae 26.9 – – –
remains open, with the reproductive column exposed.
Megachilidae 23.3 19.3 0.9 44.4
Scale bar ¼ 1 cm
134 M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae

releasing anthers and stigma, as long as the
pressure is maintained. Even after complete
triggering, flowers can still receive several fur-
ther visits.
Spartium junceum and Genista cilentina
present a typical explosive mechanism: when
the pollinator pulls the wing-keel complex
downwards, the central reproductive column
is forcefully pushed upwards, pollen is released
in a small cloud and flower parts remain
displaced (Fig. 1). In Genista cilentina there is
sometimes a partial return to the previous
arrangement, but at least the stigma remains
exerted, so that a single visit is sufficient for
successful pollination (Fig. 2).
Genista radiata also releases pollen with
explosive mechanism, but the flower’s parts
return to their initial position after triggering, so
that the reproductive column remains concealed
in the keel, and usually several visits occur.
All species are characterized by sternotribic
pollen presentation: during pollination, pollen

Fig. 2. Genista cilentina. Flowers before and after the explosive mechanism. a Reproductive column concealed
inside the keel in a closed flower. b After triggering, the reproductive column may return in its initial position,
but at least the stigma remains exposed. Scale bars ¼ 1 mm
M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae
Table 7. Stigma receptivity of the four Cytiseae.
Stage 1 = blossom, basifixed and dorsifixed anthers
indehiscent; Stage 2 = blossom, basifixed anthers
dehiscent, dorsifixed indehiscent; Stage 3 = flower
untriggered, basifixed and dorsifixed anthers dehis-
cent; Stage 4 = already triggered flower
Stage Cyt. Spa. Rad. Cil.
1 ) ) )/+ )
2 )/+ )/+ + +
3 + + + +
4 + + + + The species studied are obliged xenogamous.
grains adhere to the ventral surface of the
insect’s body.
The four species employ secondary pollen
presentation, as the mechanism of expulsion
involves contact of the pollen with other floral
parts than pollen-sacs (Yeo 1993). Basifixed
anthers dehisce first, when the flower is still
untriggered. In Cytisophyllum sessilifolium the
pollen released by basifixed anthers is pushed
to the tip of the keel by the extending
dorsifixed ones, and is collected by insects
during the first stage of anthesis with a piston-
like mechanism (Fig. 3).
In all cases both types of anthers are finally
arranged around the upper part of the style.
The keel can be opened (by valvular or
explosive mechanism) when the dorsifixed
anthers become dehiscent.
By Genista cilentina and Genista radiata
anthers dehisce before the standard opens: we
observed bumblebees and solitary bees visiting
135
Fig. 3. Cytisophyllum sessilifoli-
um. a Basifixed anthers. b Dorsi-
fixed anthers. c Reproductive
column with all anthers dehiscent.
Basifixed anthers dehisce first,
when the flower is still untrig-
gered. The pollen released by
basifixed anthers is pushed toward
the tip of the keel by the extending
dorsifixed ones. Scale bar ¼ 1 mm
the flowers at this early stage, indicating that
they are not attracted by the vexillar petal.
Stigma is receptive in all species from
the early flowering stages. Receptivity was
observed since basifixed anthers dehisce, and in
G. radiata even before anthers’ dehiscence, and
lasted until the opening of flowers (Table 7).
Discussion
Some incompatibility mechanisms must be
present, since anthers dehisce already in blos-
som, stigma is receptive at the same stage, and
yet no seed is produced. Insect visits are
necessary for successful pollination: the pro-
duction of seed is strictly linked to pollen
transport, which is performed by insects be-
longing mainly to the superfamily of Apoidea.
Principal pollinators are bumblebees and
solitary bees, but in some case also beetles
could play a role.
Insects are attracted mainly by the copious
pollen production: the pollenkitt, present in
the four species, produces odors which can
play a role as olfactory attractants (Dobson
and Bergström 2000), and keeps pollen grains
agglutinated helping the collection by pollina-
tors (Pacini and Franchi 1993).
Effective pollinators (bumblebees and sol-
itary bees) collect a great amount of pollen
grains to feed their brood, but the mechanisms
of pollen presentation and dispensing (differ-
ently specialized in each species) provoke
136 M. Galloni and G. Cristofolini: Floral rewards and pollination in Cytiseae
contamination of the ventral part of insects,
which then are apt to pollinate other flowers.
Pollinator specificity is very low both in
Cytisophyllum sessilifolium and Genista cilenti-
na; it is higher in Genista radiata and in
Spartium junceum.
This may be one of the causes of the low seed
set under natural conditions. These low values
were also partly due to beetles feeding on
broom, and probably to high within-pod abor-
tion of embryo (Herrera 1999). Predation by
beetles has been reported by previous authors
(Syrett and Emberson 1997), and was directly
observed by us on all species; predation was
especially severe in Cytisophyllum sessilifolium.
In contrast with past investigations, we
could show that nectar is produced, although
in small amount, in all the species studied. We
found traces of glucose at the base of vexillar
petal and on the reproductive column; nectar is
secreted, apparently as a phloematic exudate,
on the surface of monadelphous androecium,
at the base of the standard or on other basal
parts of the flower.
The behaviour of some visitors (Bombylius
sp.) that are strictly nectar feeding, provides
further, indirect evidence of the presence of
nectar. Anyhow, although the species studied
are not nectar-lacking and traces of nectar
represent an attraction at least for some
insects, pollen seems to be the main floral
reward. In fact, the majority of insects were
collecting pollen.
The inconstancy of nectar secretion is
probably due to many environmental factors,
which assume even greater importance when
the quantities are very small, like in our case.
We suppose that nectar, along with other
structures, such as tongue-guides present in the
lower side of the standard, may play a role in
attracting pollinators determining their right
position for a successful pollination.
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Address of the authors: Marta Galloni (e-mail:
martagalloni@libero.it), Giovanni Cristofolini
(e-mail: cristofo_g@biblio.cib.unibo.it), Diparti-
mento di Biologia Evoluzionistica Sperimentale,
Università di Bologna, Via Irnerio 42, I-40126
Bologna, Italy.