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Ecological studies on water plants of 14 sites around

Kangerlussuaq, southern West Greenland, with
special regard to Potamogeton

Margrit Vöge
Ecological studies on water plants of 14 sites around Kangerlussuaq,
southern West Greenland, with special regard to Potamogeton

Margrit Vöge (2002)

Abstract
Water plants were studied in 14 lakes and pools around Kangerlussuaq, southern West
Greenland. The study was carried out near the end of seasonal growth in 2000. The sites distinctly
vary in water colour, Secchi depth, conductivity, pH and number of plant species. A total of 18
species was observed, representing four life-forms. New to Kangerlussuaq surroundings, two
submersed species were found: Subularia aquatica and Potamogeton praelongus. Employing site
data, species and life-forms, plus the species´ demands, four lake groupings/ site types are proposed.
Plant zonation is sketched, plant communities, expressed in terms of synusiae and combinations,
exhibit eight zonation types. Shoot and leaf morphology of three Potamogeton species are related to
the environment: the internode length appears dependent on light availability or the depth of
settlement, whereas the high values of leaf length/ leaf width ratio and the large leaf surface area are
explained by the low water temperature. Due to short growing season and low water temperature the
Potamogeton shoot complexes appear rudimentary. Some growth-forms reflect particular site
conditions. Spike development is striking in Potamogeton species, but the number of mature seeds
at the end of seasonal growth is low. P. filiformis only makes up dense seed banks in lakes with
high conductivity. Positive correlations and compromises in plant development were recognized.
Though nearly all examined plant species developed flowers and/ or fruit, vegetative reproduction
and survival appear dominant.

Keywords: Greenland, water plants, Potamogeton, shoot morphology, shoot complexes,
leaf morphology, reproduction

Margrit Vöge, Pergamentweg 44b, D-22117 Hamburg, info@solo-tauchen.de

1. Introduction
A lot of work has been done on the flora of Greenland, including water plants (e.g. Böcher 1949,
1963). More recent investigations are reported by Fredskild (1992,1996) and Bay (1992). The area
around Kangerlussuaq is well known for the highest number of lakes in Greenland, with sites
distinctly varying in several aspects. This makes this region advantageous for ecological studies on
plants which additionally experience a short period of seasonal growth. Special attention has to be
paid to vegetative and sexual reproduction, being highly important phases of the life-cycle.
Phenotypic plasticity is frequent in Potamogeton. Plants grow by accumulating a varying number of
function-dependent modules. Responses to particular site conditions seem conceivable in a variation
of the basic module that is leaf morphology and internode length (1), and of this unity which is
specialized on reproduction that is spike stem and spike length plus the number of fruit (2). Barko et
al. (1982) report from laboratory studies leaf and shoot morphology of Potamogeton nodosus to be
dependent on water temperature and light. Law (1979) presumes the existence of positive
correlations between plant features only for such plants, growing in excellent conditions; otherwise
compromises are expected.

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The aim of the present field study are
1. to investigate plant species´ diversity of lakes/ ponds around Kangerlussuaq,
2. to examine plant communities and zonation along downward slope,
3. to measure module characters of Potamogeton plants,
4. to evaluate effects of temperature and light availability
5. to recognize correlations or compromises between plant features

2. Materials and methods
2.1. Investigation area and climate
The investigation was carried out in an area of some 50 km ² around Kangerlussuaq (N 67.02°,
W 50.69°), between August 19 th and September 17 th 2000. Lakes and ponds are frequent in this
landscape as a result of a periglacial and impeded drainage. The vegetation in the systems of valleys
that continue to fjord into the ice cap, is very stabilized. Due to the severe dessication, herb field
rich in species, is totally absent (Böcher 1948). Lowland plant communities are dominated by
Calamagrostis lapponica grassland; according to Birks & Penford (1990) variations in aspect and
drainage result in its replacement by shrub vegetation with Salix glauca, flourishing along stream
sides and on south facing slopes, whilst Betula nana dominates slopes with a more northerly aspect.
Due to the severe dessication shallow pools may dry out during summer. The climate is low-arctic
and continental. The average temperature is 4° C (minimum) and 13° C (maximum) in August, 0° C
(minimum) and 7° C (maximum) in September. The examined 14 lakes and pools are situated in
lowland between some 50 and 150 m a. s .l. The period of investigation was selected for
experiencing the end of seasonal growth.

2.2. Site characteristics
Water samples were taken at one point in the lakes near the water surface. Habitat conditions
were assessed by measuring the conductivity (LF 90, WTW), pH (digi 88, WTW) and water
transparency (modified Secchi disc). Ground sight is given instead of Secchi depth in some shallow
pools. Water temperature was measured regularly during the field studies in one deep lake (site 1)
and one shallow pond (site 6) for comparing the particular temperature courses. Hazen colour
number was determined by field method (Merck) in lake 1 and pool 6.

2.3. Species diversity and plant profiles
The submerged plants were recorded and living material for laboratory studies on plant growth
and fertility was gathered with access facilitated by the use of snorkelling equipment. As the lakes
in their entirety could not be surveyed, the species list does not always correspond to the species
inventory of the sites studied. Bryophytes, though present in most sites have not been taken into
account. The species were grouped according to the ecomorphological system of life-forms, which
has been established for aquatic macrophytes (Mäkirinta 1978a). The plants observed are classified
as a helophyte, isoëtid, elodëid or charid species. Helophytes (He) are adapted predominantly to
terrestrial life, e.g. they perform a high shoot/ root mass ratio; they inhabit the most shallow water.
Isoetids (Is) are characterized by the typical small thick and stiff leaves and by their high root/ shoot
mass ratio. Elodëids (El) are well adapted to aquatic life, with long shoots and submersed leaves of
simple anatomy. Charids (Ch) are thallophytes often inhabiting rather deep water. Further growth
forms, as Nymphaeids and Lemnids are not found in lakes of arctic Greenland (Fredskild 1992).
Helophyte species sometimes exhibit the elodëid life-form, growing completely submerged.
Mäkirinta prefers life-forms to growth-forms in his concept, since life-forms account for
morphplogical characters plus some ecomorphological adaptations to particular site conditions.

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The profiles were established employing depth meter und tape measure. In the studies of plant
sociology the concept developed by Mäkirinta (1978 b) was used.

2.4. Module characters of Potamogeton species
The shoot morphology of three Potamogeton species was characterized by measuring the length
of internodes, and the leaf morphology by establishing the length and width of leaves. Leaf length
was measured from the point of leaf intersection with the shoot to the leaf apex, leaf width was
measured at the widest point along the length of the leaf. Leaf surface area was established
according to the method of Steubing and Fangmeier (1992). Sexual reproduction includes the
development of spike stem and spike; length and width of spike stems and spikes plus the number
of fruit per spike were measured. Fully developed and rudimentary fruit at the same spike were
counted separatedly. Further surveys were carried out on the development of turions and winter
buds. Histograms were established; SPSS and Excel were used for statistics.

3. Results
3.1. Water conditions and species diversity
The water characteristics of the study lakes are listed in Table 1. The sites distinctly vary in
several aspects: size, conductivity (25° C), Secchi depth, and water colour. Most of the sites are
smaller than 1 ha, the remaining lakes are between 2 and 750 ha. Water is slightly acidic in site 5,
and highly alkaline in the sites 9 -14. It is approximately neutral in the remaining lakes 1 – 4 and 6 –
8. The conductivity ranges from rather electrolyte-poor (up to 179 µS cm-1 ) in half of the study
lakes through moderately rich (up to 633 µS cm -1) in four sites to electrolyte-rich (up to 3920 µS
cm -1) in the lakes 12 – 14. Secchi depth and water colour usually are correlated: water transparency
is high in clear water (lakes 1 and 9), and lowest in brown water (site 6). Twelve lakes/pools are
natural (e.g. site 11, Store Salt Sø, Fig.1) and the further two are man-made (e.g. quarry lake, site 9,
Fig.2).
The following plant species were observed in this study. The taxonomy follows Böcher et al.
(1978). According to Fredskild (1992) the species belong to three distribution types: Circum-
Greenlandic (1), low-arctic Greenland (2), plus South and/ or West Greenland (3).

(1) Hippuris vulgaris L.
Ranunculus confervoides Fr.
Ranunculus hyperboreus Rottb.
(2) Eleocharis acicularis (L.) R. et S.
Menyanthes trifoliata L.
Ranunculus reptans L.
Sparganium hyperboreum Laest.
Subularia aquatica L.
Potamogeton filiformis Pers.
(3) Potamogeton praelongus Wulfen
Myriophyllum spicatum L. ssp. exalbescens (Fern.) Hult.
Potamogeton alpinus Balb. ssp. tenuifolius (Raf.) Hult
Potamogeton gramineus L.
Potamogeton pusillus L. ssp. groenlandicus (Hagstr.) Böch.

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Charophytes, rare in Greenland to-day, occur in the continental low-arctic interior (Fredskild
1992).
Chara fragilis Desv.
Chara delicatula Ag.
Nitella flexilis (L.) Ag. em. R.D.W.
Tolypella sp.
The plants observed in the particular lakes are listed in Table 2, arranged according to their life-
forms (Mäkirinta 1978a). Many plant species (eight from eighteen) exhibit the elodëid life-form.
The other species are distributed to the same extent to the three remaining life-forms.
The species distinctly vary in the number of sites they inhabit: Potamogeton filiformis occurring
in nearly all sites and Subularia aquatica in only one lake. P. filiformis experiences the total range
of ion content, whereas S. aquatica inhabits electrolyte-poor lakes exclusively (Fig. 3). In spite of
the limited number of the study lakes, the conductivity ranges preferred by the particular species are
reasonably consistent with the data given by Fredskild (1992). Furthermore the species vary in their
nutrient demands and conductivity preferences (Fig. 3). Following Fredskild (1992), one
oligotrophic, five oligotrophic-mesotrophic, six mesotrophic and four indifferent species plus two
species preferring high conductivity are distinguished.

3.2. Site descriptions, depth profiles and plant communities
The sites substantially vary in bottom slope and light availability limiting the area of settlement;
furthermore, nutrient and ion content of lake water decide the number of plant species (Table 3).
Water plant symbols are demonstrated in Fig. 4 and applied in the depth profiles (Fig. 5a- j). Site
1, Tasersuatsiaq (Lake Ferguson) is approx. 4 km away from Kangerlussuaq Airport (Fig. 5a). All
helophyte species observed grow together with three isoëtid species in shallow water. Following the
depth gradient, six elodëid species occur with some charid species. Deepest water is inhabited by
Potamogeton praelongus and charid species.
The sites 2 (Fig. 5b) and 3 (Fig. 5c) are approx. 30 km apart, near the foot of Sugar Loaf. Pool 2
supports each two helophyte and isoëtid species plus some eloëid species in shallow water and
Potamogeton praelongus in deepest water. In pool 3 two helophyte and four eloëid species occur in
shallow water; deeper than 1.50 m Chara delicatula covers the lake bottom.
The shallow pond 4 (without sketch) is situated on the way to Qaarsorsuaq (Black Ridge) on a
plateau, 150 m a.s.l. Three helophytes plus P. filiformis and (more frequently) P. gramineus make
up dense stands, both showing spikes.
Site 5 (Fig. 5d) is a slightly dystrophic shallow pool 750 m (northwest direction) away from lake
1. Sparganium hyperboreum exhibits the elodëid life-form and is dotted about here and there,
accompanied by Potamogeton filiformis and P. gramineus, both with spikes.
Pool 6 (Fig. 5e), at half way to lake 1, supports vegetation down to only 50 cm, due to poorest
light availability from all sites studied, consisting of helophyte and elodëid species.
The ponds 7 (Fig. 5f) and 8 (Fig. 5g) are situated on the way to Sugar Loaf, approx. 400 m apart.
Vegetation of site 7 is made up by helophyte and elodëid species. Potamogeton alpinus occurs in
both sites. In very shallow water of pool 8 the species develops floating leaves exclusively. The
main vegetation belt consists of helophyte, isoëtid and elodëid species. Both sites 7 and 8 support
Myriophyllum spicatum in deepest water.
Lake 9 (Fig. 5h) is the only site with charid life-form dominating the vegetation, corresponding
with the steep bottom slope. The lake is 400 m apart from site 8.

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In sites 10 (Fig. 5i) and 11 (without sketch) Potamogeton filiformis is prevailing with
Ranunculus confervoides or R. hyperboreus in shallow water. The lakes are situated on the way to
Sugar Loaf, one 600 m away from the other.
In both sites 12 and 13 (Fig. 5j) Potamogeton filiformis grows near the water surface, followed
by charid vegetation with increasing depth. The lakes are nearby, approx. 10 m between them, on
the way to Umiarsualivik (harbour). P. filiformis plants experience optimal growth conditions in
water with high ion content; a dense seed bank is recognized at the shores (Fig. 6).
Site 14 (without sketch) harbours monospecific stands of Potamogeton filiformis exhibiting
luxurious growth.
Plant communities making up vegetation belts are best recognized in lake 1 due to an even slope
and transparent water. They are described in this study by means of synusiae and their
combinations. Synusiae consisting of plants which exhibit the same (or similar) life-forms, are
considered as ecologically uniform plant societies (Mäkirinta 1978b). They are denoted as “He”,
“Is”, “El” or “Ch”, a combination of the first two as “He-Is”.
The following six synusiae/ combinations were established in this investigation:
He-Is in lake 1,
He-Is-El in sites 2 and 3,
He-El in the pools 3, 4, 5, 6, 7,
El-Ch in lake 1,
El in the sites 1, 2, 10, 11, 12, 13, 14 ,
Ch in the lakes 3, 9, 12, 13.
The most frequent synusia is El, reflecting the high number of elodëids within the lakes.

3.3. Module parameters of Potamogeton plants
Potamogeton plants develop two kinds of modules, beginning with a basic module. It consists of
a leaf with an axillary bud and a shoot internode. Numerous modules are accumulated, until another
kind of module appears which is specialized on sexual reproduction. The vertical growth is finished
and a stem bearing a spike is developed. Only one such module per shoot is usually developed (Fig.
7). Modular growth may give rise to extreme shoot variation.
The module specialized on growth is characterized by the internode length, leaf width, leaf
length and leaf surface area. The data that were established on three Potamogeton species on each
25 shoots from site 1, are presented in Table 4. The species distinctly vary in mean leaf parameters,
but the mean internode length is similar.
Also for these species the internode length was determined in further sites 5, 6 and in deep water
of lake 1. Size classes for this parameter were defined and six histograms were established (Fig.
8–10). The histograms 8a, 9a, 10a are based on data obtained in lake 1 (more shallow water, Table
4), and the histograms 8b, 9b, 10b on data received in sites 5, 6 and lake 1 (deep water). Four
histograms show a peak in the 30-49 mm length class (P. gramineus, 8a, P. alpinus, 9b) or in the
50-69 mm length class (P. alpinus, 9a, P. praelongus, 10a). No peak is recognized in the histograms
8b and 10b, but with 5-6 similar size classes.
The module specialized on reproduction is characterized by length and width of the spike stem,
length of the spike and the number of fruit per spike. Length of spike stem proved to be correlated
with spike length for Potamogeton alpinus (Fig. 11). Furthermore, width of spike stem is related to
length of spike stem and length of spike. On P. praelongus a regression was established between
spike length and number of fruit per spike (Fig. 12). All four regressions are compiled in Fig. 13.
Hence, the wider and longer the spike stem is, the longer the spike and the more numerous are the
fruit.

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Not all fruit mature, a certain number remains rudimentary until autumn. Table 5 shows results
of fruit counting on spikes of Potamogeton species in different sites. The share of rudimentary
seeds of P. filiformis is lower in the sites 12 and 13 than in lake 1. Only half of the fruit of P.
gramineus mature in site 4. On spikes of P. alpinus (in the sites 6 and 8) and of P. praelongus (in
sites 2 and 3) all fruit mature. However, on P. alpinus plants developing more than one spike per
shoot, (as in very shallow water of site 8), the second and third spikes remained rudimentary. P.
praelongus plants produced numerous spikes per shoot in deep water of lake 1, setting of fruit was
scarce until decay of shoots in autumn.

3.4. Observations on survival and reproduction
With the exception of Potamogeton pusillus and Eleocharis acicularis, flowers / fruit were
observed on all plant species during the time of investigation. Antheridia and archegonia were
present in all charid species. The flowers of Subularia aquatica (Fig. 15) did not open below the
water surface, as was reported for flowers of Ranunculus confervoides (Böcher 1954). P. pusillus
plants develop unusual numerous turions in the leaf axils, appearing dominant in the phenotype. In
the plant depicted (Fig. 16) the turion, developed in autumn last year, is recognized, that gave rise to
this year shoot growth,and a new turion in its upper part. Several Potamogeton species (P.
filiformis, broad-leaved species) overwinter with rhizome buds (Fig. 16). Starch accumulation in
autumn is useful for new shoot growth in spring, after the period of dormancy. Myriophyllum
spicatum plants exhibit both inflorescence and many turions (Fig. 17). It is supposed that several
species overwinter in a green state.

3.5. The end of seasonal growth in the study lakes
The end of the growing season was in late August, in low-arctic, continental Greenland. Fig. 18
shows a population of Potamogeton alpinus, beginning to decay. Compared with the very low air
temperature during long winter time, water temperature is even, between 2 and 4° C under the ice.
Furthermore, during summer, water temperature does not follow the peaks of the air temperature.
During the investigation period water temperature fell from 12.6° to 8.8° C in the deep lake 1, with
ice covering the shallow bays. In shallow pond 6 the drop in temperature was essential, with 4° C
measured in the middle of September. Three temperature curves are presented in Fig. 19, two of
them representing the values established in the sites 1 and 6. The third gives the mean air
temperatures that were calculated from the daily minimum and maximum values at Kangerlussuaq
Airport. Usually seasonal growth begins in lakes when water temperature is at least 10° C, so plant
growth is assumed for approx. 3 months around Kangerlussuaq, from June to August. The aspect of
under water vegetation corresponded to late summer in the beginning of the investigation and to late
autumn in the end.

4. Discussion
4.1. Species diversity and environment
Except for two species, all the other are reported for Kangerlussuaq surroundings. Subularia
aquatica was observed south of 64.5° and north of 68° n. L (Fredskild 1992, 1996). This new
finding is in between them, at 67° n. l. For Potamogeton praelongus, the second species, a single
isolated finding from central East Greenland was first mentioned by Laegaard (1960); in spite of
intensive floristic work in this Greenland region during the eighties , the species had not been found
again (Fredskild et al. 1982). Bennicke & Anderson (1998) reported P. praelongus occurring in a
lake approx. 20 km east of Kangerlussuaq Airport and approx. 7 km west of the marging of the
Inland Ice. The three new findings of the present study are situated nearer to Kangerlussuaq Airport.
In contrast to the terrestrial vegetation, which is poor in species, Böcher (1954) stresses the real
vegetation of hydrophytes around Kangerlussuaq, due to the particular climate conditions. In the

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present study it is assumed that the high number of lakes and ponds in this area and their variation
in site characteristics are essential reasons. Water fowl enable plant dispersal from one lake to
another, where the plant species may settle to if the site conditions consist with the species´
demands. Chara fragilis typically makes up dense stands in the man-made site 9, representing a
pioneer vegetation, typical for man-made lakes (Krause 1981); according to Fredskild (1992) this is
the most frequent among the charid species. In shallow water one single Myriophyllum spicatum
plant has settled to this site, according to an optimal ion content, further dispersal of the species in
this lake appears likely.
Hansen (1967) stressed that from a limnological point of view the deeper Greenland lakes
display the greatest similarity with the sites on Moskenes island in the Lofoten group, northern
Norway. In Table 6 the species inventory of site 1 (= GR1), the deepest lake in the present study, is
compared with the species lists of some lakes in Norway, South Greenland and Iceland, resulting
from earlier studies (Vöge 1988, 1997a). Site N1 (lake Sottjun) is situated in the south boreal zone,
the sites N2 (lake Matthisvatnet) and N3 (lake Jansvatnet) are situated in the north boreal zone,
from which N3 is the most northern lake (70.5° n. l.) with some oceanic influence. Plants
experience a subarctic climate in the lakes GR0, IS1 and IS2. The ion content of all the lakes is
rather low and does not exceed 159 S cm, water is circumneutral with a pH between 6.8 and 7.5.
Sites N1 and GR1 are the richest in species.
Some trends are recognized when comparing the species lists (Table 6), from south boreal
through north boreal/ subarctic to low-arctic conditions: 1.) Nymphaëid species (though possessing
floating leaves, they are adapted preferably to terrestrial life) occur only in the southernmost lake
N1. 2) There is some evidence of species´ replacement with changing climate conditions within the
isoëtid species: Lobelia dortmanna and Littorella uniflora retire first, followed by Isoëtes lacustris
and then I. echinospora, whereas Subularia aquatica and Eleocharis acicularis tolerate a cold
climate. 3.) Regarding the remaining life-forms, similarities are displayed: species´ diversity is low
for the helophyte and charid life-forms with only four species each. The elodëid life-form is
represented by eight species. Myriophyllum alterniflorum is very rare and Ranunculus peltatus was
not found in the Greenland lakes, whereas Potamogeton filiformis was observed only in these lakes.
This species is known to be a salt tolerant fresh water species, dominating in saline lakes. P. alpinus
and P. gramineus are most frequent pond weeds within the selected lakes.

4.2. Groupings of the study lakes
Accounting for water characteristics, plant species and their life-forms, the following site
groupings are proposed as presented in Table 7:
1 ) Sites with approximately neutral water with low conductivity, (sites 1, 2, 3, 4, 5).
1a) Clear water lakes supporting isoëtids, Potamogeton gramineus, P. praelongus and Nitella; (sites
1, 2, 3).
1b) sites with yellowish water, inhabited by P. gramineus; (sites 4, 5).
2) Lakes with low conductivity and alkaline water, characterized by the elodëids P. alpinus and
Myriophyllum spicatum; (sites 6, 7, 8).
3 ) Clear water sites with medium ion content, dominated by a pioneer charid vegetation; (site 9).
4 ) Lakes with highly alkaline water; (sites 10, 11, 12, 13, 14).
4a) Clear water sites with medium ion content, characterized by P. filiformis, Ranunculus
confervoides or R. hyperboreus; (sites 10, 11).
4b) Brownish water lakes with high ion content, alkalinity and pH, inhabited by P. filiformis and
Chara fragilis; (sites 12, 13, 14).
Böcher (1954) reported of similar groupings, accounting for 16 lakes/ pools in southern West
Greenland, distinguishing a Potamogeton alpinus-Myriophyllum spicatum-type A and a

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Potamogeton filiformis-type B. Both types consist with the groupings 2 and 4 of the present study.
The additional groupings 1 and 3 are based on the findings of Subularia aquatica and Potamogeton
praelongus (group 1) and on the observed deep water charid vegetation (group 3). This group is
represented by only one lake. This site (9) is man-made with particular water conditions; such lakes
probably did not exist 50 years ago. It is supposed that man-made sites, e. g. quarry lakes, will be
more frequent in future.
The plant species surveyed essentially vary in their demands on nutrient/ ion content
corresponding to the variation of site characteristics. Accounting for the representation of
oligotrophic, oligo-mesotrophic, mesotrophic species, and species preferring high conductivity, plus
indifferent species in the particular sites (Fig. 3, Table 8), another site grouping appears reasonable,
distinguishing site type I, (consistent with former group 1a), site type II, (embracing the former
groups 1b and 2), site type III, (containing the former groups 3 and 4a), plus site type IV, (consistent
with former group 4b).
Each site type is uniform in light availability, as indicated by water colour and Secchi depth.
Mean conductivity was calculated for each site type. Regarding the species´ demands, the types are
characterized in the following way:
Site type I: oligotrophic-mesotrophic, high light availability, 114 µS cm-1, sites 1, 2, 3.
Site type II: (oligotrophic)-mesotrophic, reduced light availability, 139 µS cm-1 , sites 4, 5, 6, 7, 8.
Site type III: mesotrophic, high light availability, 620 µS cm-1 , sites 9, 10, 11.
Site type IV: saline, reduced light availability, 3500 µS cm -1 , sites 12, 13, 14.
Potamogeton species experience different light availability in site types I and II, P. filiformis,
growing in site types II and IV, experience different ion contents. These types are the most
informatives with regard to the environment influencing growth parameters.

4.3. Zonation of plant communities
Usually, different water plant communities are recognized to follow the bottom downward slope.
The zonation, characteristic for a particular lake, is described by a sequence of vegetation belts.
Plant communities are denoted by synusiae and combinations, in the present study. “HeIsVeg”
means a vegetation belt, representing a combination of helophyte and isoëtid synusia. The following
sequences of plant communities, so-called zonation types (ZT), were established in the study sites:
(ZT 1) HeIsVeg- ElVeg- ChVeg, in site 1
(ZT 2) HeIsElVeg- ElVeg, in site 2
(ZT 3) HeElVeg- ChVeg, in site 3
(ZT 4) HeElVeg, in sites 4, 5, 6. 7
(ZT 5) IsElVeg- ElVeg, in sites 2, 8
(ZT 6) ChVeg, in site 9
(ZT 7) ElVeg, in sites 10, 11, 14
(ZT 8) ElVeg- ChVeg, in sites 12, 13
The most frequent zonation type is (ZT 4) with one plant community, the most complete
zonation type is (ZT 1), in site type I/ lake 1), with three plant communities, comprising all life-
forms examined. This is possible only if all habitat conditions, required by the the life-forms are
fulfilled: oligo-mesotrophic conditions, high water transparency, and appropriate site morphology,
that is a deep lake with an even slope. Site 1 is the only one offering all these conditions. In the site
type II/ ponds 4, 5, 6, 7 supporting one plant community, low light availability causes a narrow
vegetation belt. Zonation types (ZT 7) and (ZT 8) characterize lakes with high ion content, site types
III and IV.

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4.4 .Leaf and shoot morphology of Potamogeton species
Shoots elongate due to repetitive growth, the length of modules agreeing with the internode
length. In type I/ site 1 leaf area and mean internode length increase from Potamogeton gramineus
through P. alpinus to P. praelongus, growing in shallow water (Tab. 4), corresponding with the
shoot length of the species. Usually, the internode length is rather constant for a particular species.
As was expected, this pattern is recognized in the size-class diagrams in Fig. 8a (P. gramineus), Fig.
9a (P. alpinus) and Fig. 10a. The diagram in Fig. 10a is compared with that in Fig. 20. Both
represent P. praelongus and they are much alike. Since data of lake Großensee in C Europe (Vöge,
unpubl.) were employed for Fig. 20, the comparison supports to the interpretation of the internode
length to be largely independent of climate conditions.
The diagrams in Fig. 8b (Potamogeton gramineus), Fig. 9b (P. alpinus) and Fig. 10b (P.
praelongus) exhibit more variation and no clear peak is recognized in the internode length of P.
gramineus or P. praelongus. The tend towards to more variable and longer internodes is explained
by particular site conditions. P. gramineus and P. alpinus inhabiting site type II/ ponds 5, 6
experience low light conditions, P. praelongus suffers from deep settlement in site type I/ lake 1.
Internode elongation due to a deficit in light makes sense, since shoot elongation results in
accelerating the shoot growth and approximating the shoot apex to the water surface with better
light conditions. The essential internode elongation in P. praelongus (Fig. 10a, compared with Fig.
10b) displays a within lake variation in clear water lake 1. Inflorescence usually is developed, when
the shoot apex is near the water surface. Ressource allocation assumably is insufficient for
producing a shoot of four meter length plus setting of fruit. There is some evidence indicating a
compromise: in spite of internode elongation the shoot apex remains far from the water surface ,
nevertheless inflorescence is developed but remain rudimentary until the end of seasonal growth.
Variation in the internode length and internode elongation are less distinct in Potamogeton
alpinus than in the other broad-leaved species. One explanation may be found in better tolerating of
low light conditions in P. alpinus, this view being supported by the preference of type II habitats in
the present study.
The product of leaf length and leaf width proved to be correlated with the leaf surface area, valid
for seven Potamogeton species, growing in different climate conditions (Vöge, in press). The
regression is depicted in Fig. 21. Data from the present study on three Potamogeton species (Table
4) are expressed by dots. The expression of this relationship is consistent between the earlier results
and the present study. It should be noted that the leaf surface area of P. alpinus is the same in site
type I/ lake 1 and site type II/ pond 6, independent of light availability.
The leaf surface area is larger on Potamogeton gramineus and P. praelongus in Greenland than
in a temperate climate. Furthermore, the leaf length/ leaf width ratio is higher in all three
Potamogeton species. Data established in lakes of C Europe (Vöge, in press) and from the present
study are listed in Table 9. It appears that the higher values are due to low-arctic climate.
The influences of light and water temperature on the growth of submersed macrophytes are
difficult to separate from one another during field studies. For a discussion of the variation studied
between the lakes around Kangerlussuaq water temperature influences are not to be considered.
However, compared with data established in a temperate climate, effects of cold water may be
evaluated. Barko & Smart (1981) emphasize that both light and water temperature may be of equal
importance with regard to macrophyte growth. According to laboratory studies, the leaf surface area
of Potamogeton nodosus Poiret and leaf length/ leaf width ratio are reported to increase with
decreasing water temperature (Barko & Smart, 1981), thus supporting the results from the three
Potamogeton species studied in Greenland (Table 9). The leaf-form was found unaffected by light
(Barko et al. 1982). This result is consistent with P. alpinus leaves exhibiting the same length and
width in site type I and II. Variations in shoot length effected by irradiance reflect internode
elongation which can be influenced by temperature as well; Berry & Bjorkman (1980) caution the

9
interactive relationship between light and temperature. Furthermore, the depth of settlement may
influence the internode length, as is recognized in the within-lake variation of P. praelongus in site
1 (Fig. 10).

4.5. Spike and fruit development of Potamogeton species
Phillip et al. (1990) emphasize sexual reproduction to be a common physiological reaction
toward extreme environment. On the other hand sexual reproduction is a rare event even though a
high number of seeds may be produced, up to 135 000 per square meter, and vegetative
reproduction is predominant in Potamogeton (Vöge 1997b, Wiegleb & Brux 1991). According to
Callaghan & Collins (1981) vegetative reproduction is preferred to sexual reproduction in arctic
areas, due to the limited ressource allocation and difficult establishment of seeds and shoots.
With regard to the number of seeds per shoot (Table 5) successful sexual reproduction appears
unlikely for four Potamogeton species. Only P. filiformis, exhibiting a remarkable shoot density and
higher seeds production in lakes with high ion content, effects a vigorous seed-bank. With the
exception of P. gramineus (Fig. 14) one spike per shoot is developed, however, two spikes are
frequently observed in P. filiformis growing in lakes with high conductivity (Fredskild 1992). Since
inflorescence is developed near the water surface, light conditions as indicated by Secchi depth and
water colour are presumed with low influence. The more numerous fruit per P. alpinus shoot in
extremely shallow water (site 8) compared with some deeper water (site 6) is supposed to be due to
varying ressource partitioning. Though sexual reproduction is unlikely, the importance of even
modest seed-banks is stressed as they may enable resettlement after disturbance (Haag 1983, Van de
Weyer 1988).
Spike stems and the (fruit-bearing) spikes characterize the reproduction-specialized modules.
The highly significant regression between length of spike stem and spike length of Potamogeton
alpinus (Fig. 11) established in sites 6 and 7, is not valid for such plants growing in the particular
conditions of site 8. The relationship between the spike length and the number of mature fruit per
spike, established on P. praelongus in sites 2 and 3 (Fig. 12), provides explanation for the
rudimentary spikes observed in deep water of lake 1: shorter than 2.5 cm, no fruit mature on such
spikes. In contrast to the spike length, the spike stems are as long as in sites 2, 3 and consistent with
data from lakes in C Europe (Tab.10), according to Casper & Krausch (1980). This contrast appears
due to limited ressource availability. Spike stems and spikes of P. gramineus are essentially shorter
in Greenland site 4 compared with data from C Europe (Table 10). It appears that this Potamogeton
species suffers most from the low-arctic climate. This trend is less pronounced in P. alpinus.
Summarizing, the mentioned authors support the observations of this study. Sexual reproduction
seems emphasized considering the conspicuous inflorescence development (e.g. Fig. 22), but the
number of mature fruit is low. So vegetative reproduction by means of specialized vegetative
propagules or fragmentation of clonal growing plants appear to prevail.

4.6. Diversity in shoot complex development
The life-cycle of Potamogeton species usually begins in late March, and is finished in October in
a temperate climate (Vöge 1997, Wiegleb & Todeskino 1985). After a period of dormancy of the
reproductive unit horizontal and vertical shoots are growing continually, sexual reproduction ends
in August/ September, fragmentation of the shoot complex and vegetative reproduction indicate the
end of seasonal growth, when shoot decomposition begins. The result of a growing season of six
months in displayed in Fig. 23: a shoot complex of P. perfoliatus in lake Hohendeicher See in C
Europe. From a horizontal shoot, two meters long, nineteen vertical shoots arise, with a total length
of more than ten meters. The branching of the horizontal shoot is recognized, giving rise to further
vertical shoots. These shoots vary in their length according to different age (Vöge 1997b). In
contrast, shoot complexes are somewhat rudimentary in low-arctic climate. The shoots within a
population are of approximately the same length, indicating that continued growth of shoots does

10
not belong to the set of options established in Potamogeton species growing in a temperate climate.
Further options, that is growth patterns which enable survival of a population (Chapleau et al 1988,
Van de Weyer 1997) are: clonal growth, including asexual reproduction, morphological plasticity
and sexual reproduction.
Particular development of shoot complex was observed in sites with very special growth
conditions. Fig. 24 shows a Potamogeton alpinus shoot complex from site 8 exhibiting the typical
feature as known from a temperate climate, but miniaturized due to low water level. Maximum
shoot length is only ten centimeters, reducing the energy that must be expended on vegetative
growth. Ressource partitioning favours sexual reproduction which is evident in the three spikes
developed on one shoot, however, only the fruit of one spike matured. P. alpinus in site type II/
pond 6 displays a divergent shoot complex, and is presumed an adaptation to low light availability
(Fig. 25). Shortcoming light is experienced by horizontal shoots near the bottom, whereas these
horizontal shoots arising from the upper vertical shoot are advantaged. Limited resources appear to
explain the absence of inflorescence. P. praelongus exhibits an unusual growth-form in deep water
of lake 1 (Fig. 26). One vertical shoot developed numerous side-shoots with one inflorescence each,
but all of them were rudimentary.
Though some positive correlations between plant parameters were established, compromises
appear frequent in Greenland climate conditions. This result supports the opinion of Law (1979),
accordingly positive correlations reflect optimal growth conditions, whereas compromisses are
expected in suboptimal environment.

5. Conclusions
Human influence on lakes and pools around Kangerlussuaq is low. For the last decades, man-
made lakes were added to the high number of natural sites in this region. Such young lakes/ pools
may support some pioneer vegetation in an early stage after sediment stabilization, until further
species settle if site characteristics are appropriate.
Water plants may experience stress and disturbance. Stress, following Grime (1979) affects the
increase of biomass, and results here from low water temperatures during the short growing season
and low light availability in dystrophic sites. Some further stress is experienced by Potamogeton
plants growing in deep water. Disturbance reduces (partly or totally) the biomass (Grime 1979). It
may be caused by the dry climate in this particular region. Due to lowering of water level the
submersed leaves of Potamogeton alpinus decayed, and a few floating leaves developed in site 8.
Vegetative reproduction by means of specialized vegetative propagules or fragmentation of
clonal growing plants appears to prevail. However, some modest seed banks may enable
resettlement of species after disturbance.

6. Acknowledgements
The author is greatly indebted to Danish Polar Center for the allowance to stay and work at
Kangerlussuaq International Science Support; to Christian Bay, Botanical Museum, University of
Copenhagen for valuable information; to Fleming Skou, Danish Meteorological Institute,
Copenhagen, for climate data; to Antje Eggers, Hamburg, who has corrected the English text; and to
my husband Harald Vöge for continued assistance.

7. References
Barko, J. W. & Smart, R. M., 1981: Comparitive influences of light and temperature on growth and
metabolism of selected submersed freshwater macrophytes. - Ecol. Monogr. 51: 219 - 235.
Barko, J. W. , Hardin, D. G. & Matthews, M. S., 1982: Growth and morphology of sumersed
freshwater macrophytes in relation to light and temperature. - Can J. Bot. 60: 877 - 887.

11
Bay, C., 1992: A phytogeographical study of the vascular plants of Northern Greenland- north of
74° northern latitude. - Meddr Grønland. Biosci. 36.
Bennicke, O. & Anderson, J.N., 1998: Potamogeton praelongus in West Greenland. - Nord. J.
Bot.18: 499 - 501.
Berry, J. B. & Bjorkman, O., 1980: Photosynthetic response and adaptation to temperature in higher
plants. - Annu. Rev. Plant Physiol. 31: 491 - 543.
Birks, J. D. S. & Penford, N., 1990: Observations on the ecology of arctic foxes Alopex lagopus in
Eqalummiut Nunaat, West Greenland. - Meddr Grønland Biosci. 32.
Böcher, T. W., 1949: Climate, soil and lakes in continental West Greenland in relation to the plant
life. - Meddr. Grønland 147.
Böcher, T. W., 1954: Oceanic continental vehetatuional complexes in southwest Greenland. -
Meddr. Grønland 148, Nr. 1.
Böcher, T. W., 1963: Physiography of middle West Greenland. - Meddr Grønland. 148, Nr. 3.
Böcher, T. W., Fredskild , B., Holmen, K. & Jakobsen, K., 1978: Grønlands Flora. - Copenhagen
Callaghan, T. V. & Collins, N. J., 1981: Life cycle, population dynamics and the growth of tundra
plants. - In: Bliss, L. C. Heal, O. W., Moore, J.J. (eds.): Tundra Ecosystems: a comparative
analysis. IBP 25: 257- 284, Cambridge.
Casper, S. j. & Krausch, H. -D., 1981: Süßwasserflora von Mitteleuropa. - Bd. 24: Jena.
Chapleau, F. Johanson, P.H. & Williamson, M., 1988: The distinction between pattern and process
in evolutionary biology: the use and abuse of the term "strategy". - Oikos 53; 136 - 138.
Fredskild, B., 1992: The green limnophytes- their present distribution and Holocene history. - Acta
Bot. Fennica 144: 93 - 113.
Fredskild, B., 1996: A phytogeographical study of the vascular plants of W Greenland (62°20´ -
74°00 N)´ - Meddr Greenlamd, Biosci. 45
Fredskild, B., Bay, C. & Holt, S., 1982: Botaniske undersøgelser på Jameson Land - Botanisk
Museum, København
Grime, J. P., 1979: Plant strategies and vegetation processes. - Wiley, Chichester, pp. 7-119.
Haag, W., 1983: Emergence of seedlings of aquatic macrophytes from lake seedlings. - Can. J. Bot.:
61: 148 - 156.
Hansen, K., 1967: The general limnology of arctic lakes as illustrated by examples from Greenland.
- Meddr. Grønland 178, Nr. 3.
Krause, W., 1981: Characeen als Bioindikatoren für den Gewässerzustand. - Limnologica. 13: 399-
418
Law, R., 1979: Ecological determinants in the evolution of life histories. - In: R. M. Anderson, B.
D. Turner & L. R. Taylor (eds), Population Dynamics, Oxford, 315 - 345.
Lægaard, S., 1960: Two species of Potamogeton new to Greenland. - Bot. Tidskr. 56: 247 – 251.
Mäkirinta, U., 1978a: Ein neues ökomorphologisches Lebensformensystem der aquatischen
Makrophyten. - Phytocoenologia 4: 446-470.
Mäkirinta, U., 1978b: Die pflanzensoziologische Gliederung der Wasservegetation im See Kukkia,
Südfinnland. - Acta Univ. Ouluensis, Ser. A, 75(5): 157 S., Oulu.

12
Philipp, M., Böcher, J., Mattsson, O. & Woodell, S. R. J., 1990: A quantitive approach to the sexual
reproductive biology and population structure in some arctic flowering plants: Dryas
integrifolia, Silene acaulis and Ranunculus nivalis. - Meddr. Grønland, Biosci. 34.
Steubing, L. & Fangmeier, 1992: Pflanzenökologisches Praktikum. - Stuttgart.
Vöge, M., 1988: Tauchuntersuchungen an der submersen Vegetation in skandinavischen Seen unter
besonderer Berücksichtigung der Isoetiden-Vegetation. - Limnologica 19/2: 89-107.
Vöge, M., 1997a: Plant size and fertility of Isoëtes lacustris L. in 20 lakes of Scandinavia: a field
study. - Arch. Hydrobiol. 139: 171-185.
Vöge, M., 1997b: Vegetationskundliche und populationsbiologische Untersuchungen im
Hohendeicher See in Hamburg. - Tuexenia 17: 109 – 123.
Vöge, M., 2001: Beschreibung von Wachstum und Entwicklung und Isoëtes mittels Korrelationen;
Ein Vergleich. - Tuexenia (in press).
Weyer, K. van de, 1988: Ein Wiederfund von Potamogeton coloratus VAHL am Niederrhein. -
Natur am Niederrhein N. F. 3: 46 - 48.
Weyer, K. van de, 1997: Untersuchungen zur Biologie und Ökologie von Potamogeton
polygonifolius Pourr. im Niederrheinischen Tiefland - Diss., Berlin. Stuttgart
Wiegleb, G. & Brux, H., 1991: Comparison of life history characters of broad-leaved species of the
genus Potamogeton L. General characterization of morphology and reproductive strategies. -
Aqua. Bot. 39: 131 - 146.
Wiegleb, G. & Todeskino, D., 1985: Der biologische Lebenszyklus vpn Potamogeton alpinus und
dessen Bedeutung für das Vorkommen der Art. - Verh. Ges. f. Ökologie XIII: 191-198.

13
Table 1: Characteristics of the study sites

Site Position Size pH Conductivity Secchi depth Water colour
no. (ha) (µS cm-1) (m)
1 N66.98342 W50.69003 Z750 7.5 75 10.0 clear
2 N67.03912 W50.56249 Z 2 7.7 130 1.2* clear
3 N67.03930 W50.56017 Z 10 7.5 137 1.5* clear
4 N67.00875 W50.64190 < 1 7.0 233 1.0* yellow
5 N66.98274 W50.70811 < 1 6.6 61 0.8* yellow
6 N66.99789 W50.67268 < 1 7.0 125 0.6 brown
7 N67.02941 W50.61394 < 1 7.7 179 2.0* slightly yellow
8 N67.02991 W50.60421 < 1 7.9 94 1.0* slightly yellow
9 N67.02989 W50.59450 < 1 8.2 419 >4.0 clear
10 N67.02271 W50.65397 < 1 8.4 607 1.0* slightly yellow
*
11 N67.02377 W50.66665 < 1 8.6 633 3.0 clear
12 N67.00062 W50.80527 < 1 8.7 3470 3.0* slightly brownish
13 N67.00010 W50.80560 < 1 8.7 3920 3.0* slightly brownish
14 N66.98829 W50.58403 Z 30 9.0 3110 2.0* slightly brownish

*)
Bottom sight

14
Table 2: Plant species observed in the study sites

Life Species Site no.
form 1 2 3 4 5 6 7 8 9 10 11 12 13 14
He, El Menyanthes trifoliata + + +
Hippuris vulgaris + + + + + + +
Sparganium hyperboreum + + + + + + + +
El Potamogeton alpinus + + + +
Potamogeton filiformis + + + + + + + + + + + +
Potamogeton gramineus + + + + +
Potamogeton praelongus + + +
Potamogeton pusillus + + +
Myriophyllum spicatum + + + +
Ranunculus confervoides + + + + + +
Ranunculus hyperboreus + +
Is Ranunculus reptans + + +
Subularia aquatica +
Eleocharis acicularis + +
Ch Chara fragilis + + +
Chara delicatula +
Nitella flexilis +
Tolypella sp. +

Table 3: Ranges of the site characteristics

Characteristics Range
Size (ha) <1 - 750
pH 6.6 - 9
Conductivity (µS cm -1) 61 - 3920
Secchi depth (m) 0.6 - 10
Water colour Clear - brown
Hazen colour number 5 - 300
No. of plants species per site 1 - 14
Depth limit of vegetation (m) 0.6 - 6

15
Table 4: Characteristics of vegetative growth for three Potamogeton species in lake 1

Species Mean internode Mean leaf Mean leaf Mean leaf
length length width area
(cm) (cm) (cm) (cm²)
Potamogeton gramineus 5.0 6.5 0.8 3.4
Potamogeton alpinus 5.2 11.9 0.9 7.5
Potamogeton praelongus 6.0 15.2 1.8 19.6

Table 5: Fruit production of four Potamogeton species in different sites

Species Site Mean number of seeds per spike Mean number of
seeds per shoot
G mature rudimentary
G mature
Potamogeton filiformis 12, 13 4 6 8-16
Potamogeton filiformis 1 2 7 6
Potamogeton gramineus 4 8 8 16
Potamogeton alpinus 8 109 - 109
Potamogeton alpinus 6 35 - 60 - 35 - 60
Potamogeton praelongus 2, 3 55 - 55

16
Table 6: Plant species in lakes of Norway, Greenland and Iceland

Life Species Lake no.
form N1 N3 GR0 IS1 N2 IS2 GR1
He, El Menyanthes trifoliata +
Hippuris vulgaris + + +
Sparganium angustifolium + + +
Sparganium hyperboreum + +
Ny Nymphaea candida +
Potamogeton natans +
El Myriophyllum alterniflorum + + + + +
Ranunculus peltatus +
Potamogeton alpinus + + + + +
Potamogeton filiformis + +
Potamogeton gramineus + + + +
Potamogeton praelongus + +
Potamogeton pusillus + +
Ranunculus confervoides +
Cer, El Utricularia ochroleuca +
Is Juncus bulbosus +
Lobelia dortmanna +
Littorella uniflora +
Isoëtes lacustris + +
Isoëtes echinospora + +
Ranunculus reptans + +
Subularia aquatica + + + +
Eleocharis acicularis +
Ch Chara delicatula +
Chara fragilis +
Nitella flexilis + + + +
Tolypella sp. +

17
Table 7: Site groupings according to site conditions and vegetation

Site Site no. Site conditions Vegetation
group
Water colour pH Ion content Dominant life- forms and species

Clear Yellow- Circum- alkaline Highly Low Medium High Isoëtids Elodëids Charids
brown neutral alkaline
Nitella Chara

a 1, 2, 3 + + + + P. gramineus, P. praelongus +
1
b 4, 5 + + + P. gramineus

2 6, 7, 8 + + + P. alpinus, M. spicatum

3 9 + + + +
a 10, 11 + + + P. filiformis, Ranunculus sp.
4
b 12, 13, 14 + + + P. filiformis +

Table 8: Number of species with different demands for nutrient / ion content in the sites studied

Species Site no.
1 2 3 4 5 6 7 8 9 10 11 12 13 14
Oligotrophic species 1
Oligotrophic-mesotrophic species 6 5 3 3 2 3 1 3 1
Mesotrophic species 3 1 1 1 1 1 1 1
High conductivity preferring species 1 1 2 1 1 1 2 1 1 1 1 1 1 1
Indifferent species 3 3 1 1 2 2 2 1

Table 9: Leaf area and leaf length / leaf width in a) low- arctic lake 1, b) temperate climate, for three
Potamogeton species.

Species Leaf area (cm²) Leaf length / leaf width
a b a b
Potamogeton praelongus 19.6 16.8 8.4 6.4
Potamogeton alpinus 7.5 11.0 13.3 8.2
Potamogeton gramineus 3.4 2.3 8.1 5.2
Table.10: Length of spike stems and spikes in Greenland sites compared with C European (CE) sites

Parameter Potamogeton Potamogeton
Potamogeton alpinus
praelongus gramineus
CE* 1** 2, 3 CE* 4 CE* 6 8***
Length of spike (cm) 2- 4 1.2 3.3 2-3 0.8 2- 4 1.6 2.3
Length of spike stem (cm) 5 - 30 24.5 26.0 2-6 1.8 5 - 15 4.3 3.3

*) According to Casper & Krausch (1980)
**) Rudimentary spikes in deep water
***) In extremely low water level

Fig. 1: Site 11

19
Fig. 2: Site 9, a quarry lake

20
12: Potamogeton filiformis 61 – 3920 d
8: Sparganium hyperboreum 61 - 233 b
7: Hippuris vulgaris 75 - 233 e
6: Ranunculus confervoides 61 - 633 e
5: Potamogeton gramineus 61 - 233 b
4: Potamogeton alpinus 75 - 179 c
4: Myriophyllum spicatum 94 - 419 d
3: Potamogeton praelongus 75 - 137 b
3: Potamogeton pusillus 75 - 137 b
3: Ranunculus reptans 75 - 130 b
3: Chara fragilis 419 – 3920 c
3: Menyanthes trifoliata 175 - 233 b
2: Eleocharis acicularis 75 - 130 e
2: Ranunculus hyperboreus 125 - 607 b
1: Chara delicatula 137 c
1: Nitella flexilis 75 c
1: Tolypella sp. 75 c
1: Subularia aquatica 75 a

Fig. 3: Number of findings, conductivity ranges (µS cm -1) and demands on nutrient/ ion content of
the species
a: Oligotrophic species
b: Oligo- mesotrophic species
c: Mesotrophic species
d: Species preferring high conductivity
e: Indifferent species

21
Menyanthes trifoliata Ranunculus confervoides

Hippuris vulgaris Ranunculu hyperboreus

Sparganium hyperboreum Ranunculus reptans

Potamogeton alpinus
(floating/submersed) Subularia aquatica

Potamogeton filiformis Eleocharis acicularis

Potamogeton gramineus Chara fragilis

Potamogeton praelongus Chara delicatula

Potamogeton pusillus Nittella flexilis

Myriophyllum spicatum Tolypella sp.

Fig. 4: Species symbols

22
2m

6m

Fig. 5a: Site 1

23
1.20m

Fig. 5b: Site 2

1.50m

Fig. 5c: Site 3

24
0.75m

Fig. 5d: Site 5

0.6 m

Fig. 5e: Site 6

25
1m
Fig. 5f: Site 7

1m

Fig. 5g: Site 8

26
3m

Fig. 5h: Site 9

0.5m

Fig. 5i: Site10

27
3m

Fig. 5j: Site 12, 13

Fig. 6: Seed bank of Potamogeton filiformis, site 12

28
Fig. 7: Spike of Potamogeton alpinus in site 6

29
52

36

22,2
20
15,5 14,4 15,5
12 10,2

2,2
0 0 0 0 0 0

-29 -49 -69 -89 -109 -129 -149 -169

Fig. 8: Size class diagrams: internode length for Potamogeton gramineus, site 1 (left), site 5 (right)

60

44,3

35

24,3

18,6

7,1
4 2,9 2,8
0 0
1 0 0 0 0

-29 -49 -69 -89 -109 -129 -149 -169
Fig. 9: Size class diagrams: internode length for Potamogeton alpinus, site 1 (left), site 6 (right)

30
37,5

27,5
23,6
21,8
20 18,3
15
12,7
8,8
6,3
3,6
2,5 1,2 1,2 0 0

-29 -49 -69 -89 -109 -129 -149 -169
Fig. 10: Size class diagrams: internode length for Potamogeton praelongus, site 1, shallow water
(left), site 1, deep water (right)

25 y = 0,3976x + 0,0758
2
R = 0,7339
20
Spike length (mm)

15

10

5

0
0 10 20 30 40 50 60
Length of spike stem (mm)

Fig. 11: Relationship between spike length and spike stem length for Potamogeton alpinus, site 6

31
100 y = 2,9007x - 39,85
2
90 R = 0,5424
Number of fruits per spike

80
70
60
50
40
30
20
10
0
0 5 10 15 20 25 30 35 40 45
Spike length (mm)

Fig. 12: Relationship between spike length and number of fruit per spike for Potamogeton
praelongus, sites 2,3

1 4
Mean length mean spike mean number of
of spike stem length seeds per spike

2 3 P < 0.001
P < 0.01
Mean width of
spike stem
Correlations between:

1: Length of spike stem and spike length
2: Length of spike stem and width of spike stem
3: Width of spike stem and spike length
4: Spike length and number of seeds per spike

1, 2, 3: Established on Potamogeton alpinus in site 6
4 : Established on Potamogeton praelongus in sites 2, 3

Fig. 13: Significant regressions between Potamogeton module parameters

32
Fig. 14: Spikes of Potamogeton gramineus in site 4 (left) and of Potamogeton filiformis in site 13
(right)

33
Fig. 15: Subularia aquatica in site 1

34
ø
2

ø
1

Fig. 16:(left) Potamogeton pusillus with last year turion (1) and this year turion (2).
(right) Rhizome bud of Potamogeton filiformis

35
Fig. 17: Myriophyllum spicatum with inflorescence and turions

36
Fig. 18: Potamogeton alpinus in site 1 in the end of seasonal growth

16

12 Water temperature
(surface)
Temperature (°C)

8 (Site 1)

(Site 6)
4 Air temperature
(means)

0
08/18/2000 08/25/2000 09/01/2000 09/08/2000 09/15/2000
Date

Fig. 19: Air and water temperature during the study time

37
47,4

28,9

10,5
7,9
5,3

0 0 0

Spalte 45

-29 -49 -69 -89 -109 -129 -149 -169
Fig. 20: Size class diagram: internode length for Potamogeton praelongus in temperate climate

50
y = 0,6994x + 0,3202
40 R2 = 0,9864
Leaf surface area (cm²)

30

20

10

0
0 10 20 30 40 50 60 70
Leaf width * leaf length (cm²)

Fig. 21: Correlation between leaf width * leaf length and leaf surface area

38
Fig. 22: Potamogeton alpinus shoot with 3 spikes in site 8

39
207 cm

203 cm

Fig. 23: Shoot complex of Potamogeton perfoliatus in temperate climate, in August

40
10 cm

Fig. 24: Shoot complex of Potamogeton alpinus in very low water level, site 8

3

2

1

Fig. 25: Shoot complex of Potamogeton alpinus with horizontal shoots 1,2,3, in site 6

41
12 mm

20 mm

8cm

43cm 10 mm

Internode length Spike length

11 mm

27cm

7 mm

28cm

38cm

Fig. 26: Potamogeton praelongus shoot, with side-shoot-bearing and apical inflorescence, in site 1,
4 m deep

42