Vol. 37, no. 1
Journal of Vector Ecology
1
Tree-hole breeding mosquitoes in Israel
Günter C. Müller1, Vasiliy D. Kravchenko2, Amy Junnila1, and Yosef Schlein1
1
Department of Microbiology and Molecular Genetics , IMRIC, Kuvin Centre for the Study of Infectious and Tropical Diseases,
Faculty of Medicine, Hebrew University, Jerusalem, Israel 91120,
guntercmuller@hotmail.com
2
George S. Wise Faculty of Life Sciences, Department of Zoology, Tel Aviv University, Ramat Aviv, Tel Aviv, Israel 69978
Received 20 August 2011; Accepted 22 December 2011
ABSTRACT: A survey was conducted to evaluate the number of tree-hole breeding mosquito species and their distribution
in the six principal woodland types in Israel. Out of approximately 3,000 mature trees examined, only 38 contained holes
that retained water for extended periods of time, and breeding mosquitoes were observed in 27 of them. Two specialized
tree-hole breeders, Aedes pulchritarsis Rondani and Aedes geniculatus Oliver, were found breeding at several sites in northern
Israel, always at locations 500 m above sea level (a.s.l) and with high annual precipitation. Aedes albopictus Skuse which, in
Israel, is known as an opportunistic container breeder, was found in this study to have adapted remarkably well to breeding
in tree holes and was found in most forest types investigated and in most tree species which had adequate tree holes. Two
other species, Culiseta annulata Schrank and Culex pipiens Linnaeus instars, were found in one of the tree holes, but did not
survive to reach maturity. Journal of Vector Ecology 37 (1): xxx-xxx. 2012.
Keyword Index: Mosquito fauna, tree-hole breeders, Aedes pulchritarsis, Aedes geniculatus, Aedes albopictus, Israel.
INTRODUCTION
Water-illed tree holes provide a unique habitat to which
only a few highly specialized invertebrate communities are
adapted. his type of microhabitat is typically restricted
to mature, oten deciduous trees (Jenkins and Carpenter
1946). Due to a long history of human habitation, most
of the woodlands in the Mediterranean are synanthropic;
mature forests are rare, and degraded shrub lands are oten
the rule. As such, only six stenobiontic tree-hole breeding
mosquito species, and three eurybiontic species, which
only occasionally breed in tree holes (Table 1), have been
identiied in the Eastern Mediterranean (Ramsdale et al.
2000, Samanidou-Voyadjoglou and Darsie 1993, Shannon
and Hadjinicolaou 1937). hough some studies on the
ecology of these species were conducted in western Europe,
(Bradshaw and Holzapfel 1986, 1992, Marshall 1938, Service
1971, Yates 1979), northern Europe (Wesenber-Lund 1920,
[AU:
Natvig 1948), and central Europe (Rohnert 1950), little is
Rohnert
known from the Levant, which has been documented as the
not in
natural southern distribution border of tree-hole breeders
References
in the eastern Mediterranean (Knio et al. 2005, Margalit
Cited]
and Tahori 1970). his survey was conducted to evaluate
the distribution and abundance of tree-hole breeding
mosquitoes in the principal woodland areas of Israel.
MATERIALS AND METHODS
Climatology of Israel
Israel can be roughly divided into three longitudinal
landscape units: the Coastal Plain, the Central Mountain
Ridge, and the Rit Valley (Orni and Efrat 1980). his
division, along with north-south and east-west temperature
and precipitation gradients, creates a diverse range of
microhabitats over a relatively small area (Figure 1). he
northern part of Israel includes Mt. Hermon (2,200 m a.s.l.),
which receives annual snow and contains typical tragacanth
vegetation, whereas the Dead Sea area is about 400 m below
sea level (b.s.l.) and contains pockets rich in Afrotropical
fauna and lora (Zohary and Orshansky 1949, Bytinski-Salz
1961). he north and center of the country is temperate
(Mediterranean), while the southern and eastern parts are
semi-arid (Irano-Turanian grassland) and arid (desert)
(Danin 1992). In terms of precipitation, the short winter
accounts for 70% of the annual rainfall that occurs between
November and February. Rain from May to September is
negligible; the dry season lasts from June to August.
he principal forest types in Israel are: OroMediterranean (Montane) forest, Maquis forest, Oak
Park forest, Carob Park forest, Savannoid Mediterranean
woodlands, and Synanthropic woodlands. OroMediterranean forests are located on Mt. Hermon (above
1,300 m). he dominant trees are deciduous oaks (Quercus
boissieri Reut., Q. libani Olivier Fagaceae) and maples (Acer
microphyllum Hort. Aceraceae). hese forests are scattered
on slopes facing south, but in canyons they are contracted
and occasionally closed. Evergreen sclerophyllous Maquis
covers most of the northern central mountain range and
in areas with high precipitation the area is dominated
by dense thickets of evergreen oak (Quercus calliprinos
Webb), and its companions. In the Judean Mountains, the
number of mesophytic components decreases gradually,
allowing for woodlands dominated by Pinus halepensis
Mill Pinaceae and Arbutus andrachne L. Ericaceae. In
drier areas, only small isolated woods and hedges can be
found, occasionally accompanied by Q. calliprinos, Pistacia
2
Journal of Vector Ecology
June 2012
Figure 1. Larval tree hole sites of Ae. geniculatus, Ae. pulchritarsis, and Ae. albopictus in conjunction with the
major forest types and annual precipitation.
palaestina (Boiss.), and P. atlantica (Desf. Anacardiaceae).
North-facing slopes and narrow canyons, especially in
the northern Galilee where humidity is relatively high
throughout the year, are usually inhabited by pockets of
winter deciduous woodlands dominated by Q. boissieri.
Some of the streamlets in the northern Mediterranean zone,
like Nahal Skziv, are interspersed with small patches of
riparian forests dominated by sycamore (Platanus orientalis
L. Platanaceae), poplar trees (Populus euphratica Oliv.), and
willows (Salix L. species) Salicaceae are oten present with a
rich undergrowth of herbaceous plants.
here are two types of park forest in Israel: Oak Park
forest (Quercus ithaburensis Decne.), and Carob Park forest
(Ceratonia siliqua L. Park Forest). he Oak Park forests
are xerotherm, yet occur exclusively in the northern parts
of the country. Depending on the stand, Q. ithaburensis
is accompanied by a large variety of other trees. Scattered
forests usually have rich undergrowth composed of shrubs,
semi-shrubs, grasses, and many herbaceous species.
he Carob Park forests are found in hot and dry areas
from 0 to 300 m a.s.l. hey are found on all the limestone
hills at the foot of the Central Mountain Range of the
Mediterranean zone in Judea, Samaria, Carmel, Gilboa, and
the Galilee. Depending on the stand, these communities are
oten accompanied by olive trees (Olea europaea L. Oleaceae)
and Q. calliprinos trees and bushes. he undergrowth of this
scattered woodland is not as rich as in Q. ithaburensis Park
forests.
he Savannoid Mediterranean shrub and woodland is
restricted to the lower regions of the southeastern Galilee,
especially by the Sea of Galilee and the adjacent southern
Jordan Valley. his dry, forested grassland is dominated
by Ziziphus lotus (L.) Lam., Z. spina-christi (L.) Desf.
Rhamnaceae, and occasionally Faidherbia (Acacia) albida
(Delile) A.Chev. Mimosaceae.
Synanthropic woodland, intensively managed or
planted by humans, replaced many natural forests in Israel.
Typical elements on the Coastal Plain are citrus, mangos
(Mangifera indica L. Anacaridiaceae), and to a smaller
extent hickory orchards (Carya illinoinensis (Wangenh.)
Koch Juglandaceae). On the central mountain range, olive
and almond trees (Amygdalus communis L. Rosaceae) are
most abundant, whereas in the Northern Galilee, apple
and cherry plantations are more common. Pine forests,
mainly P. halepensis, and to a smaller extent Pinus pinea L.
and Cupressus sempervirens L. Cupressaceae, were planted
throughout Israel as far south as the northern and western
Negev Desert. Eucalyptus groves, (several species, but
especially Eucalyptus camaldulensis Dehn. Myrtaceae), are
abundant along the coastal plain, the larger inland valleys,
along streamlets, springs, around settlements, and along
roads. Numerous exotic species of trees can be found in
parklands and gardens in most towns in Israel. (Danin
1992, Eig 1926, Zohary 1973).
Sampling larvae
All principal woodland types found in Israel were
surveyed during mid-January to mid-February, 2006.
Vol. 37, no. 1
3
Journal of Vector Ecology
In each forest type, about 500 mature trees were checked
for holes. Potential areas were investigated by direct sight
and also using a round mirror (15 cm diameter) mounted
on a telescopic stick which could stretch up to 3 m long.
For preliminary veriication of water presence and depth,
holes were examined using a lexible wire probe with paper
wrapped around the distal end (50 cm). he following
parameters were recorded for each hole: forest type, sun
exposure (north or south-facing slope), tree species, height
above ground, opening size, and whether it was a rot or a
pan hole. All tree holes with water were then visited every
ten days from February 20, 2006 until May 10, 2006 and
then again before the irst rains: from the irst of November
until the end of April, 2007. Later, during the dry summer
months, the holes were visited only once a month. On each
visit, if water was present, the entire water content of the
holes was siphoned, its volume measured, the larvae deined
and counted, and they were returned with all the water back
into the tree hole (Bradshaw and Holzapfel 1986).
Sampling eggs during the dry season
Tree-holes in which water was observed at any time
during the survey (65) were sampled during August, 2007,
ater all the holes had dried up. Samples of 50 g of detritus
were scratched from the bottom and the side walls of the
holes and incubated in the laboratory, under standard
insectary conditions, in beakers with one liter of distilled
water for up to one month. Hatching was observed during
the following three months.
RESULTS
Suitable tree holes for breeding
During the survey, almost 3,000 mature trees were
checked, from which about 10% had some kind of holes,
with most either permanently dry or illed with detritus. In
about 2% of the inspected trees, water was found at some
point in the survey. Most of the holes with water were found
in broad-leafed trees, including oaks, eucalyptus, hickory,
olive, sycamore, pistachio, mango, poplar, and willow
(Table 1). Very few were found in conifers, including C.
sempervirens, P. pinea, and P. halepensis. Most of the holes
with water were rot holes, but a few were pan holes found
in the partially looded forest of Tel Dan Nature Reserve,
in the bottom of the Nahal Sksiv canyon and on the coastal
plain east of Tel Aviv. Water capacity of pan-holes never
exceeded 500 ml, which was typically maintained for less
then two weeks ater rainfall, and larvae or pupae were
never observed in them. he height of rot holes ranged from
10 to 235 cm above the ground, the size of their apertures
from 30 to 4,200 cm², and the maximum encountered water
capacity during the study period in these holes was from
100 to 8,500 ml. Only about two-thirds of these rot holes
(38) held water longer than one month, and larvae/ pupae
of mosquitoes were observed in only 27 of them. hese
tree holes retained water from the beginning of the rainy
season in early November, 2007, and were completely dry
four weeks ater the last strong rains, in mid-April. With the
exception of a single tree hole in an oak tree, all the active
holes were found in mesophile forests located in shady
areas, such as north-facing slopes, river canyons, and in one
speciic case, the shade of a church.
Tree-hole mosquitoes associated with the principal
forest types
Evidence of larvae and/or pupae was discovered in 27
out of 38 suitable tree holes containing water for more than
one month. hese 27 tree holes were distributed among nine
tree species found in the six principal forest types of Israel
(Table 2). Five mosquito species were breeding in their tree
holes. hey were Aedes geniculatus, Aedes pulchritarsis, Ae.
albopictus, Cs. annulata, and Cx. pipiens.
Four larval sites of Ae. geniculatus were found, all in
the northern-most part of Israel. hree were located in
Table 1. Variety and number of tree holes suitable for larvae; 65 out of a possible 267 tree holes contained water at some point
during the survey.
Tree variety
Oak
Total tree holes with water
Rot holes with water
Pan holes with water
17
16/17
1/17
Eucalyptus
7
4/7
3/7
Hickory
10
8/10
2/10
Olive
9
9/9
-
Sycamore
4
2/4
2/4
Pistachio
6
6/6
-
Mango
5
5/5
-
Poplar
2
1/2
1/2
Willow
1
-
1/1
C. semperviens
2
2/2
-
P. pinea
1
1/1
-
P. halepensis
1
1/1
-
4
June 2012
Journal of Vector Ecology
deciduous Maquis and one in the nearby Oro-Mediterranean
forests on Mt. Hermon. he irst site was in a narrow, shady
canyon in the upper part of Nahal Sksiv, about 650 m a.s.l.
Ae. geniculatus breeding was observed in the large rot hole
of a partially burnt stump of a P. orientalis. he second site
was on the north-facing slope of Mt. Meron, about 950 m
a.s.l. in a dense shady oak stand. Larvae of Ae. genculatus
were observed in a deep hole with a narrow opening in a
mature specimen of Q. boissieri. he third site was found on
the north-facing slope of Mas’ada, a broad-leaved deciduous
forest in the Golan Heights (1,000 m a.s.l). In the montane
forest of Mt. Hermon, one Ae. geniculatus larval site was
found in a severely damaged Q. boissieri at the bottom of a
canyon (1,650 m a.s.l).
First and second instar larvae of Ae. geniculatus were
collected throughout the entire period in which water was
present. Later instars were only observed from March until
the holes dried up and again in small amounts several
weeks ater the irst rainfall from mid-November to early
December. Although water contents of Q. calliprinos tree
holes were consistently less than those in the P. orientalis
tree holes, the former were preferred, containing several
times more larvae then the latter.
Breeding of Ae. pulchritarsis was mostly conined to
northern Israel; ive sites were in the Galilee, one was on
Mt. Hermon, one was in the Golan Heights, and one was
on the Carmel Mountain Ridge; all of these are located in
mountainous areas. Half of the breeding sites were in Maquis,
about a third in Q. ithaburensis Park Forests and only one in
Synanthropic Parkland (Table 2). Most breeding sites were
in shady locations like north-facing slopes and canyons.
he exception is one site in a scattered Q. ithaburensis park
forest on a south-facing slope. In the montane forests of Mt.
Hermon, breeding of Ae. pulchritarsis was observed in the
same tree hole utilized by Ae. geniculatus, described above.
he only true Synanthropic breeding site was in urban Beit
Shalla, east of Jerusalem, in a mature eucalyptus tree. First
and second instar larvae were observed in the tree holes
throughout the entire period in which water was present,
although more were observed during the spring. Older
instars were seen from March until the holes dried up.
An exceptionally proliic Ae. pulchritarsis site was
found close by in Mas’ada village. Hundreds of larvae of all
instars were observed in two ornamental wooden buckets
that were situated in a shady area of a garden with several
mature olive and oak trees. he water content of the buckets
luctuated between 1.5 to 6.0 liters between mid-July to
the end of August, which was the period the instars were
observed.
First and second instar larvae, a single egg rat of Cx.
pipiens, and a few second instar larvae of Cs. annulata,
together with Ae. pulchritarsis, were discovered once in
February in the biggest rot hole with the largest water
capacity in a Q. ithaburensis tree. he site was located in
a Q. ithaburensis Park Forest, on a south-facing slope. On
consecutive visits, only early stages of Ae. pulchritarsis were
found; Cx. pipiens and Cs. annulata were absent.
During the survey, no active Ae. albopictus breeding
habitats were observed above 500 m, at least in natural
tree holes. Ae. albopictus was by far the most common
and versatile species. It was found mainly in Synanthropic
woodland like mature mango and hickory orchards, and in
Synanthropic parkland (Table 2). In natural habitats, it was
found in Maquis, Quercus ithaburensis, and C. siliqua/P.
lentiscus park forests (Table 2).
All instars could be found in the tree holes one month
ater the hole illed with water and they remained until
the holes dried up. he productivity of the holes was not
connected to their size, water capacity, or tree species.
Actually, Ae. albopictus was able to utilize the smallest holes,
oten with very small water capacities (Table 2). Wherever
Ae. albopictus was found, no larvae of any other species
were ever observed. During the survey, this species was
frequently observed in natural forests breeding in various
types of disposed containers such as plastic bags, beverage
cans, buckets, and tires. In synanthropic woodlands and
gardens, it was found breeding in ornamental and disposed
containers, as well as tiny irrigation puddles.
Detritus samples were collected from all 65 tree holes
in which water was observed and brought to the laboratory
for incubation. No hatching of mosquito larvae was
observed from the ten pan hole samples. From 26 out of 27
active rot holes, viable eggs of all three Aedes species were
recovered and successfully hatched in the laboratory. he
bulk of Ae. geniculatus larvae hatched quickly; 87% hatched
ater ive days, 96% ater ten days, and the remainder during
the following ive days. On the other hand, Ae. pulchritarsis
eggs hatched much later; 45% hatched ater ive days, 63%
ater ten days, 86% ater 20 days, and 96% ater 30 days. Ae.
albopictus eggs hatched in a time frame between the other
Aedes species; 61% hatched ater ive days, 75% ater 10
days, 94% ater 20 days, and the rest in the next ten days.
In one sample taken from an Ae. geniculatus site, no
larva were observed following incubation. Generally, the
newly hatched larvae were from the same species that
were observed occupying the site; the exception was an
Ae. geniculatus and a Ae. pulchritarsis site where only Ae.
albopictus were hatching. From another Ae. pulchritarsis
site (500 m a.s.l.), about 80% of the hatching larvae were
Ae. albopictus and the rest were Ae. pulchritarsis. From the
11 tree holes in which water was present for more than
one month, but without any record, two samples with Ae.
albopictus were recovered. From the 17 tree holes with
water retained less than one month and no breeding record,
three samples with hatching Ae. albopictus were obtained.
hese ive sites were on the coastal plain/foothills along
the Central and Carmel Mountain Ridges. he number of
hatching eggs varied among species: out of 50 g of detritus,
an average of 35 Ae. geniculatus larvae hatched, while from
similar samples an average of 59 Ae. pulchritarsis larvae and
253 Ae. albopictus larvae were counted.
DISCUSSION
To date, about 30 mosquito species have been
documented in Israel (Orshan et al. 2008, Margalit and
Vol. 37, no. 1
5
Journal of Vector Ecology
Table 2. he trees harboring larval developmental sites of mosquitoes and the physical properties of the holes.
Tree species
Max
water vol.
(ml)
Diameter
opening/depth
(cm)
Period water found
3400
10/60
mid Jan. - early May
late Nov. - mid Mar.
Ae. pulcritarsis
Ae. pulcritarsis
2700
12/80
early Feb. - late Apr.
late Nov. – late Mar.
Ae. pulcritarsis
Ae. pulcritarsis
1500
8/50
mid Jan. - mid Apr.
late Jun. - late Mar.
Ae. pulcritarsis
Ae. geniculatus
Ae. pulcritarsis
Ae. geniculatus
900
15/50
mid Jan. - mid Mar.
late Nov.- late Mar.
Ae. geniculatus
Ae. geniculatus
8500
60/70
mid Jan. - early Jun.
late Nov. - late Jun.
Ae. pulcritarsis
Cx. pipiens
Cs. annulata
---
2400
22/90
early Feb. - mid Mar.
mid Nov. - early Mar.
Ae. albopictus
Ae. albopictus
2600
10/70
mid Feb. - late Jun.
early Nov. - mid Mar.
Ae. pulcritarsis
Ae. pulcritarsis
3600
20/100
early Feb - mid Jun
mid Dec. - late Mar.
Ae. pulcritarsis
Ae. pulcritarsis
---
4700
15/120
late Jan. - early Jun.
late Nov. - mid Mar.
Ae. geniculatus
--Ae. albopictus
2400
17/90
early Feb. - mid Jun.
late Oct. - mid Mar.
Ae. pulcritarsis
---
500
21/25
early Feb. - mid Mar.
early Nov. -early Mar.
Ae. albopictus
Ae. albopictus
2000
15/65
early Feb. - late Mar.
early Nov. - mid Mar.
900
7/20
mid Jan. - late Mar.
late Nov. - late Feb.
Ae. albopictus
Ae. albopictus
400
10/30
mid Feb. - early Mar.
late Nov. – late Feb.
Ae. albopictus
Ae. albopictus
200
5/20
mid Feb. - mid Jun.
mid Nov. - late Feb.
Ae. albopictus
Ae. albopictus
100
8/20
early Feb. - mid Mar.
mid Nov. - late Mar.
Ae. albopictus
Ae. albopictus
Q. calliprinos
Q. boissieri
Q. ithaburensis
P. palestinensis
P. orientalis
Eucalyptus sp.
M. indica
C. illinoinensisa
Mosquito species Viable eggs recov.
breeding
in August
Ae. pulcritarsis
Ae. pulcritarsis
6
Journal of Vector Ecology
Tahori 1970). Of these, only three species Ae. geniculatus,
Ae. pulchritarsis and Ae. albopictus were observed regularly
in tree holes during our survey. Ae. albopictus, an invasive
species, has been in Israel since 2002 and since then, it has
continuously spread along the coastal plain (Pener et al.
2003). his species is a known to opportunistically occupy
containers, rather than obligately (Tandon and Ray 2000).
A fourth native container/tree-hole breeder, Ae.
aegypti, was widespread in the lowlands of the temperate
zone in northern and central Israel until the 1930s. Despite
a brief reoccurrence in the mid 1970s along the coastal
plain, it most likely became extinct shortly thereater
(Pener-Salomon and Vardi 1975).
In previous surveys which utilized either CDC
light traps (incandescent light) or CO2-baited traps, Ae.
pulcritarsalis was rarely collected and Ae. geniculatus was
recorded only once (Margalit and Tahori 1970). However,
a recent faunistic survey of Lepidoptera and Coleoptera
with miniature UV traps yielded fairly good catches of both
species (Müller et al. 2005, 2006). In Israel, Ae. pulchritarsis
feeds readily on people, while Ae. geniculatus was never
observed to do so, although this is usual behavior in its
northern distribution area (Yates 1979).
he distribution of tree-hole breeding mosquitoes
depends on the distribution of mature deciduous trees, the
precipitation frequency, and the evaporation rate due to
solar radiation. Accordingly, the distribution center of the
two palearctic species Ae. geniculatus and Ae. pulchritarsis is
in the forest belt of temperate Europe and Asia. Many animal
and plant species found in Israel reach their most marginal
point of geographic distribution, and this is especially true
for woodland species (Bodenheimer 1935, Furth 1975).
Generally, both plant and animal species gradually become
infrequent at the periphery of their geographic distribution
before they inally vanish (Hengefeld and Haeck 1982,
Brown 1984). his also seems to be the case with the two
Aedes species.
hough Ae. geniculatus and Ae. pulcritarsalis are
generally woodland species, they are speciic in their
habitat preferences. Both species were only found in
forested mountainous areas (≥500 m a.s.l.) with high annual
precipitation (≥700 mm). Both species were most common
in the north, and Ae. geniculatus was not observed south
of the Nehring Line (Northern Galilee – Golan Heights).
Both Ae. pulchritarsis and Ae. geniculatus were found in
mesophilic woodlands with mature trees, the latter species
exclusively. Although Ae. pulchritarsis penetrated into
xerotherm forests, breeding was still restricted to shady
areas with high precipitation. In areas with low precipitation
and along the coastal plain, both species were absent despite
the abundance of suitable tree holes which retained water
for ample durations. Ae. geniculatus has thus far only been
found in natural habitats, whereas Ae. pulchritarsis has
been collected in both natural woodlands and synanthropic
woodlands. At some sites, Ae. pulchritarsis were found in
wooden buckets, although suitable tree holes were present
in the immediate vicinity. his occurred throughout the
year, including the dry season (unpublished data). he same
June 2012
species was observed in Greek villages without any mature
trees and far from forests (Shannon and Madjinicolaou
1937). It is not clear how important these secondary habitats
are for Ae. pulchritarsis, but they might partly explain the
numerous records of adults from urban Jerusalem and its
surrounding villages (unpublished data).
During the dry season, viable eggs were recovered
from both of the local Aedes species and from almost all
of the sites with evidence of previous breeding. In a small
riverine forest (500 m a.s.l.), close to the coastal plain,
the two species were possibly replaced by the invasive Ae.
albopictus. It is important to point out that no hatching
larvae were observed from any other samples taken
from the water-illed holes. It seems that both species are
k-strategists with a low rate of dispersal from their breeding
habitats (McNeely 1994). hough some adult females of Ae.
pulchritarsis and Ae. geniculatus were regularly caught in UV
CDC traps and were observed resting during the daytime
in tree holes, it remains unclear if these specimens were
old aestivating specimens, or if there was some continuous
breeding activity in unidentiied tree holes or even in other
secondary habitats. Nevertheless, in Israel the bulk of the
population of Ae. pulchritarsis and Ae. geniculatus seem
to pass the dry season in the oval stage. he diferential
hatching time of the two species might also explain their
diferent distribution patterns. Delayed hatching of Ae.
pulchritarsis might be the reason why this species is more
widespread in Israel then Ae. geniculatus, especially towards
the drier south. Half of the eggs hatch within a very short
period following introduction of water and develop rapidly,
while the other half hatch gradually. his mechanism might
ensure survival of viable eggs for the next rainfall, if water
availability is sporadic due to low precipitation or high
evaporation rates. he few eggs of Ae. geniculatus hatch
faster, which might explain its restriction to places with
high and reliable precipitation; dispersion would be more of
a risk than an advantage.
Ae. albopictus is mainly an opportunistic container
breeder (Comiskey et al. 1999, Knudsen 1995). In our survey,
this is the irst time this species has been found breeding in
natural tree-hole habitats in Israel. Moreover, it was the most
versatile tree-hole breeder found in all types of woodlands,
from sea level up to 500 m a.s.l., excluding the Montane
forests on Mt. Hermon. From the nine tree species that were
occupied by tree-hole breeders (Table 2), Ae. albopictus was
found in eight. However, in Israel its main breeding site is in
garbage and ornamental containers as well as in irrigation
puddles. Garbage in Israel is oten deposited in and around
forests and might have been a stepping stone to tree holes.
Most locations where Ae. albopictus was found would
probably not have been suitable for the other Aedes species
to breed; Ae. albopictus is invading the natural habitats of
the two local species. Along the coastal plain, Ae. albopictus
might replace the two species in the long run. It appears that
Ae. albopictus is also invading habitats of stenobiontic treehole breeding mosquitoes in the Americas (Barrera 1996,
Juliano et al. 2004, Sota et al. 1992).
Tree holes are unique breeding habitats, and only a
Vol. 37, no. 1
Journal of Vector Ecology
few mosquito species are fully adapted to this specialized
ecological niche (Bradshaw and Holzapfel 1986, Woodward
et al. 1988). It is therefore not surprising that Cx. pipiens
and Cs. annulata were observed for the irst time in this
habitat in Israel. hey apparently did not manage to breed
there successfully. he large opening of this particular hole,
the exposure to sunlight, or the relative scarcity of breeding
sites in general might have encouraged them to oviposit
there.
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