Remote, Inland Occurrence of the Oceanic Anogramma leptophylla (L.) Link (Pteridaceae: Taenitidoideae) in Hungary

2008·5.3 MB·English

by  Csaba Molnár

Remote, Inland Occurrence
Oceanic
of the
Anogramma
leptophylla Link
(Pteridaceae:
(L.)
Hungary
Taenitidoideae)
in
CSABA MOLNAR
52. Istvan utca, H-3036 Gyongyostarjan, Hungary
ZoltAn Baros
Department of Regional and Rural Development, Karoly Robert College, 36. Matrai H-32
fit,
Gyongyos, Hungary
IstvAn Pinter
Hungarian Academy Science—
of Eotvos Lorand University, Research Group
for Evolution;
Pazmany
Genetics 1/C, H-1117 Budapest, Hungary
P.
s.,
iLDIKd TlJRKE
}.
nstitute of Environmental and Landscape Management,
Szent
Istvan University, Pater Karol
1.
H-2100 Hungary
Godollo,
Abstract.—The presence Anogramma
of the fern species leptophylla was detected Zemplei
in the
ME
Hungary) in 1991. The species was known neither from the country nor from
th
whole
Carpatho-Pannonian Region known
(also as Carpathian Basin)
previously.
Its habital
on
situated a roadside bank, cut into an unstable rhyolite above
surface, the valley of the Cree
Kemence
near
— the village of Palhaza. The fern a cosmopolitan taxon humi
is restricted to
nd
is considered to be an oceanic-sub
Europe. Th
ien in
t
ungary is located more than 1000 km from the closest populations, on
thus, this
is
may
be due
a to the peculiar microclimatological conditions which
of the
habitat, are describe
e in order to give an exact explanation
for this outstanding occurrence. The chromosomes wei
counted some
a in individuals of the Hungarian population and were found be n = 26n
to
fc
In 1991, a population
of a small fern species was found Hungary
in the
for
time by Zolyomi
first Sz. in the Zempleni Mountains
(Hungarian
spelling:
Zempleni-hegyseg). The
was
species Anogramma
identified
as leptophylla
(L.)
Link
(Pteridaceae, Taenitidoideae; Tyron
et al, 1990; Sanchez-Baracaldo,
2004) using the "Collectio Pteridophytorum"
of the Hungarian
Natural History
Museum
(BP) and by counting
the chromosomes. The
species has not been
reported previously from
the Carpatho-Pannonian Region known
(also as
[email protected]
:
ANOGRAMMA
LEPTOPHYLLA
:
The newly found
Hungariai
Carpathian Thus, with
Basin).
sporophyte, A. leptophylla can
1
members
of the flora of the region.
The Anogramma
gene genus Tryon
center of the {sensu 1990)
et
al., is
and South America, where Anogramma
located in Central species except A.
all
ascensionis (Hook.) Diels can be found, including cosmopolitan A. lepto-
Anogramma
phylla. caespitosa Pic.Serm. reported from Tanzania was recently
synonymy
reduced under and
A. leptophylla (Nakazato Gastony,
to 2003).
Anogramma many humid
leptophylla can be found and mild
in regions
around the world. Besides the Americas, the range of the species includes the
and and
subtropical regions coastal regions of Africa, the subtropical regions of
Australia and Oceania. distribution range extends from Iran to Malaysia in
Its
Asia, and also inhabits the southern portion of the Himalayas, the Caucasus
it
and the Middle East. In Europe, occurs in the Mediterranean region, along
it
the Atlantic coast, the Mediterranean basin and in Crimea (Fig. Jalas and
1;
Suominen, Komarov,
1972; 1934; Pignatti, 1982; Castroviejo et 1986; Prelli,
al.,
1990; Stace, 1997).
The European distribution pattern of the species shows a conspicuous
preference for humid regions, and considered to be an oceanic-suboceanic
it is
element (Meusel et 1965; Dostal, 1984). In the western territories grows
al.,
it
and mountains under oceanic whereas
in coastal regions climatic influence,
suboceanic) occurrences
the vast majority of the eastern is restricted to the
(i.e.,
Some
shores of the Mediterranean Sea (Dostal, 1984). inland .occurrences have
under
also been detected, but these are either oceanic influence or influenced
by microclimatic conditions. This pioneer species prefers competition-
local
free habitats, where inhabits rock surfaces and crevices (Page, 1982; Jermy
it
and Camus,
1991).
AMERICAN FERN VOLUME NUMBER
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Anogramma
Location
of leptopbylla
Peak
Settlements
Area of Zempleni
the Mts.
Anogramma
Fig. 2. Location of leptophylla in Hungary.
In contrast to the above-mentioned, the Hungarian population
the
village
at
of Palhaza unique
being
(Fig. 2) is in situated in the inland region of the
km
European
continent 1000 from
ca. the nearest occurrences along the
southern
part of the Adriatic Sea and on the southern one
side of the Alps.
It is
of the northernmost populations
of this species in the world.
Although
presence Hungary
in unusual
its is at glance, the ecological
first
requirements
of this otherwise oceanic, and
subtropical, tropical species are
by
fulfilled the microclimate which
of this location, provides an
air
temperature above freezing and high humidity Some
throughout
the
year.
European
other inland
populations
of A. leptophylla, such as those on the
southern side of the Alps Switzerland and
(in northern the
Italy), live at
mouths
of caves (Chiodi, 1958; Hess
ef al, 1976) that provide a suitable
microclimate Our
the
for species. case although some
is different, similarity
may
between
exist the microclimate and
of caves the talus crevices Palhaza
at
may
in that they provide
similar climatic conditions. During the winter 2003
of
MOLNAR ANOGRAMMA HUNGARY
LEPTOPHYLLA
ET
AL.: IN 131
and 2004, our was entirely covered by snow with the exception of the
site ca.
m
2
10 area where adult sporophytes of the fern grow, surrounded by green
mats mosses on wet
of the rocks.
This paper presents the characteristics (geological, microclimatological, and
vegetational features] of the Hungarian population and the habitat preferences
of A. leptophylla in the Carpathian Basin.
Methods
and
For insight into the microclimatic conditions of the habitat to explore
how Anogramma
leptophylla can tolerate the unfavorable macroclimatic
measurements
conditions, seasonal expedition not repeated, but
field
(i.e.,
detailed observations of the climatic conditions that enable one to gain basic
were conducted
insight to the microclimatologic conditions of a site) at 12
designated points, which were intended to reflect the characteristic features
(i)
and them adequate permit
of the habitat represent in density;
itself
(ii)
comparison between the habitat and environment, and represent points
its (iii)
and
with various exposure, vegetation coverage.
units, soil
Measurements were carried out three times a day sunrise, around noon,
(at
and on two sporophyte
sunset) different dates in 2005: the height of
at at
rd
germination on February 19th, and at the height of spore ripening on April 3
.
The following parameters were determined: temperature and relative
air
humidity at a height of 2 m, soil temperature at depths of 2, 5, and 10 cm. and
wind speed and direction at a height of 2 m. In addition to the above, air
temperature and humidity were measured every 30 minutes, directly
relative
next to the sporophytes at the crevices of the substrate.
and humidity were measured using
Air temperature relative a digital
platinum resistance thermometer and capacitive hygrometer. Measurements of
temperature were carried out using a digital platinum resistance
soil
down wind
thermometer measurements 10 cm, whereas speed
suitable for to
wind and
was measured with cup anemometer. Additionally, directions
a
winds were determined.
microclimatological
was
The newly found population visited regularly to observe life-cycle.
its
known
Because no species of similar life-strategy has been in our region, the
was examined. Growth designed
of the fern also studies, to
life-cycle
humidity and water by watering and
investigate the impact of air availability
drying, were also carried out under laboratory conditions.
For chromosome counts, some adult sporophytes from the population were
and greenhouse. Chromosomes were counted from
collected transferred to a
methods (Manton, Photographs
spore mother following standard 1950).
cells,
DMRB.
model
were taken with a Leica microscope
—
The
and population
Lithology, climate vegetation of the habitat. in the
known
Mountains)
Zempleni Mountains the Eperjes-Tokaji located
(also as is
AMERICAN FERN VOLUME NUMBER
132 JOURNAL:
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3 (2008)
along a roadside bank near the Palhaza above Kemence
village the valley of
on
Creek, the hillside of Nagy-Gereben 48 25'N, 21 The
(Fig. 2) at 25'E.
m
elevation 365 above Exposure
is ca. sea level. to the east south-east
is
and
(Fig. the side of the most exposed The
5], is hill to solar radiation.
population found on
is talus slopes densely interlaced and developed on the
bedrock
rhyolite of the Palhaza-Telkibanya eruption center of the Lower
A
Sarmatian
volcanic cycle (Gyarmati, 1971). road was cut 1958 by
in forestry
(Nagy Imre
ex
retired forester, verb.) into this unstable, rubble rhyolitic surface
m m
forming a moderately steep scree slope, which provides 10 X
ca. of
1
The
habitat. population fluctuates between 400 and 800 sporophytes
ca.
The
annually. surrounding
vegetation can be described as a habitat-complex of
rocky grassland and embedded
debris slope forest all in acidophilic hornbeam-
A
woods
oak with beech
(Simon, voucher specimen
1977). deposited
the
is at
herbarium
"Collectio Pteridophytorum" in the Hungarian Natural History
Museum
(BP) under the number "689786"
collection
(collecting date:
11.04.2001, Csaba
collector: Molnar).
mm
The
annual
precipitation of the sample area 700-750 (Mersich
is ca. et
which
above
al., 2002), is the national average. Also, the highest values
of
humidity
relative in the Hungarian context found
are here
in seasons,
all
75%
approximately
during winter with
values
of actual evapotranspiration
mm
may
that exceed 100
in July (Mersich et 2002). Air temperature
al.,
conditions Zempleni
of the Mountains, with mean
a annual temperature
between and C
8.0 8.5 at the northern regions, are among the coolest compared
other mountains
to of the North Hungarian Middle As
Range. whole,
a the
climate of the region can be and humid
classified as cool moderately
(Justyak,
J
'
1998).
Microclimatic conditions habitat—
of the Results
of microclimatological
measurements
indicate a positive anomaly both
in air temperature ("heat
outflow"; and
Fig. 5 A) relative humidity at the center of the
locality, hereafter
referred to as the "cauldron",
the inner microclimatic space
i.e., of the
population The
location. rise in temperature quite conspicuous winter
is in
when,
seasons,
unlike the surrounding
area that usually covered by
is
snow
significant cover, the cauldron snow
and
is free of manifestly unfrozen.
Relative humidity remained 100%
constant between
and
at sunrise sunset
The
permanently
(Fig. moist
3). surface of the bedrock, and
the water droplets
on
the surrounding
moss-cushions
also
reflected saturated or over-saturated
m
present
air in the cauldron. At
a distance of above
2 the
surface, a significant
decrease
in the diurnal course
of humidity
relative can be observed
in front of
the cauldron on
the road
at the outer microclimatic
(i.e., space; Here,
Fig.
3).
ca^70%
the value
after sunrise decreases noon
until
44.6%), then reaches
(to
it
65%
The
ca. at sunset. values measured
here compare
well with
the values of
the remote measurement
points (51.0-58.1%
noon, 55.5-61.3%
at
at sunset).
Thus,
the outer microclimatic
space
reflects the
characteristic values
of the
In the period following
springtime
thaw,
a significant change was observed
in the diurnal rhythm
of relative humidity. The
cauldron and
close
its
MOLNAR ANOGRAMMA
ET LEPTOPHYLLA HUNGARY
AL.: IN
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80-
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cauldron"
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100-
"Cauldron"
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Time (UTM+1)
AMERICAN FERN VOLUME NUMBER
JOURNAL:
98
3
both maps.
i
environment switched showed
to a different course that conspicuous
congruency phenomenon may
This be due
(Fig. 4). to the cessation of the
continuous water supply
(snow-melt), that plays a crucial role in the existence
of the population by providing mild and humid
conditions during germination
dormant embryo
of sporophytes or gametophytic lobes from perennating
tubercules as illustrated by Nakazato and Gastony
(2003).
The
high humidity
relative values at the cauldron are coupled with high
air
temperature values that are 3-5 C higher than those in the nearby environment
(Fig. 5A). Temperature values measured here between sunrise and sunset were
not lower than 4 C, whereas, example, was
for 0.3-0.6°C observed
at the rest of
measurement
the
points sunset. Additionally, measurement
at at the point in
front of the cauldron (and on
close to the sections of the cart-road covered by
it,
snow)
the temperature
air increased
significantly, in contrast to the highly
where
forested parts, the most moderate
oscillation in temperature values
air
were
observed.
Soil temperature values measured at a depth of 2 cm in the cauldron were 7-
C
11 higher than in surroundings and were even
its higher at a depth of 5 cm.
The
increase in temperature was
soil after sunrise the most intense the depth
at
of cm, exceeding even
2 that of the temperature. However,
air the diurnal
course of these two parameters becomes
rather similar during the rest of the
day. This rise in temperature occurred somewhat cm
later at 5 depth and, in
was
spite of this short delay, characterized by
the highest values and also the
lowest daily fluctuation of temperature At depth cm
(Fig. 5B). the of 5 the
was between and
latter 9.1 11.9°C, whereas the between
difference the values
measured cm
depths and was
at of 2 5 only ±0.1 with
C, greater variation
at
MOLNAR ANOGRAMMA
ET LEPTOPHYLLA HUNGARY
AL.
IN
35
1
:
The
the shallower depth. measurements
results of carried out in the vicinity of
on mat moss
the fern, a of the most exposed
at side of the outcrop, indicated
and
higher
also non-fluctuating temperature
soil values.
Because flow can
air counteract the high humidity,
relative
characteristic
wind
conditions of the sample area should also be mentioned. The
results of
our measurements showed predominance
the meso- and
of macroclimatic
influences, channel
i.e., a so-called effect in accordance with the orientation of
the valley system. The mean values of wind speeds of 0.5 to nearly m/s
1
measured
during day
the indicate an intensive dehydrating
effect.
The
positive anomalies in temperature and humidity mentioned
relative
m
above diminished abruptly within even a few distance from the Anogramma
was
habitat patch. This most probably due the 0.5-1 m/s mean wind speed
to
may
values resulting from the channel This cause
effect. the suitable habitat
for
Anogramma
be highly
to restricted to the cauldron.
The
Life-cycle.— sporophyte generation of the studied population
starts to
develop from dormant embryos on
perennating tubercules cover
[see
illustration of Nakazato and Gastony (2003)] in December, and during the
crucial period of germination the above described
specific microclimatological
conditions provide favorable circumstances. Sporophytes reach maturity
the
at
beginning of April, and spores are ripe from April the beginning
to of June,
then the sporophytes wither. newly produced from
Prothallia, the spores,
photosynthesize during June and and then, since no growing
July, suitable
conditions exist in this period, they form a perennating organ, a dormant
tubercule, as found by Baroutsis (1976) and reported by Hagemann and
(1997)
new
Nakazato and Gastony The
sporophyte
(2003). generation from
arises
these prothallia or from their dormant tubercules as explained and
illustrated
by Nakasato and Gastony (2003). Our greenhouse observations have shown
that adequate humidity and mild temperature (above provided
if zero) are
under
controlled conditions, the sporophytes can arise at any time in the year,
and their life-cycle is completed within the same time as under natural
conditions. Regarding our desiccation tolerance observations, dormant
tubercules were found to be able to survive for 2.5 years in drought conditions.
5%
After watering, of the prothallial tubercules immediately developed
sporophytes.
—
Chromosome
counts. For the Hungarian population, the chromosome
n
=
number was
n 26 obtained spore mother However,
of (Fig. 6) in cells.
numbers
different are reported for A. leptophylla in the Brownlie
literature:
u n
= =
(1958) found n 29 in New-Zealand, Tutin (Fabri, 1963) found n 26 in the
n
=
Channel Verma and Khullar found n 29 Mickel
Islands, (1965) in India,
et
al.
= n
found n 27 and 29 11 Mexico, Baroutsis and Gastony found
(1966) in (1978)
=
n 29" in South Africa and n=29' in gametophytes from Mexico, Manton
et
al.
n
found n-26 11 Madeira, and Queiros found n=26
(1986) in et (1988) in
al.
Portugal. Baroutsis and Gastony (1978) discussed the tendency of mitotic and
chromosomes
meiotic of A. leptophylla (including A. guatemalensis (Domin)
C.Chr.) together and observed thin and knob-like short chromosomes,
to stick
Some
factors potentially contributing to the variation in reported numbers.
AMERICAN FERN VOLUME NUMBER
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6B
chromosomes
larger in the photograph of our specimen can be
(Fig.
6.)
observed. Rasbach and Reichstein and
[1990] Lovis re-examined
et (1993)
al.
n
chromosome numbers many =
the in populations, and they observed n 26
.
They many
concluded
that of the previously published counts presumably
are
incorrect, although, they urged further taxonomic research two
in cases.
Nakazato and Gastony
noted chromosome
(2003) that the reinterpretation of
if
number
in A. leptophylla correct, the number 26 an apomorphic reduction
is
is
common
in the ancestor of their A. leptophylla clade from the plesiomorphic
n
number =
base n 29
of in Taenitidoid Thus,
ferns. in spite of the cytological
Anogramma
variability within the genus (Gastony and
Baroutsis, 1975), the
world-wide Anogramma
distributed species leptophylla seems be uniform
to
in this respect according to the re-examinations of Rasbach and Reichstein
and
(1990) Lovis Our
et al. (1993). result in line with these re-examinations,
is
and chromosome
the count Hungarian
of the population supports the
karyological homogeneity
of the species.
Conclusions—The
surprisingly remote inland establishment extreme
of the
Anogramma
oceanic element Hungary may
leptophylla
in be
attributable to
certain favorable environmental
conditions. At
the habitat of the species,
special microclimatological
conditions (higher temperature and
air relative
humidity values compared
to its surroundings) can be observed the very
at
close surroundings
of the population, hence providing
suitable habitat the
for
colonization of the species. These conditions
are strictly restricted to the close
environment
of the habitat, and absence may
their not allow colonization
Anogramma
elsewhere by
in the surrounding The and
area. life-cycle the
chromosome
counts of the Hungarian
population accordance with
are in those
reported European
for other populations. The
relatively recent colonization of
the species probable since the was
is habitat created by 1958 (Nagy
forestry in
Imre, ex The
occurrence
verb.). of A. leptophylla
in this inland territory further
contributes our knowledge
to of the distribution of pteridophtyes, where
MOLNAR ANOGRAMMA
HUNGARY
ET LEPTOPHYLLA
AL.: IN
137
similar isolated occurrences have been witnessed before. also underlines the
It
and
success colonizing cosmopolitan
ability of this species.
who
improved
greatly
c
r
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