PRELIMINARY NOTES ON THE POTENTILLA NIVEA COMPLEX IN NORTHWESTERN
EUROPE
Jorun Nyléhn |
References |
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The taxa
in the Potentilla nivea complex (section Niveae) are closely related
and morphologically similar, they are mainly separated on indumentum types and number of
leaflets: P. chamissonis Hulten has long, straight, verrucose hairs and ternate
leaves; P. nivea L. has short, floccose hairs and ternate leaves; and P.
insularis Soják has long, straight hairs less verrucose than P. chamissonis
and digitate leaves. Despite the close relationship, the taxa have been subdivided into a
series of subspecies, varieties, forms and postulated hybrids.
The subdivisions might reflect local races that have arisen through the
predominantly apomictic mode of reproduction in these plants. Many species of Potentilla
reproduce by pseudogamy, i.e. agamospermy with a demand for pollen to initiate seed set
(endosperm formation). Furthermore, the apomixis is often facultative, i.e. some seed
production might be sexual. The reproduction system may vary among and within taxa and
populations, see e.g. Eriksen (1996), Holm (1996), Asker (1977).
In this study taxa in the Potentilla nivea complex from
Svalbard and Fennoscandia have been analysed. Collections were made of Potentilla nivea
ssp subquinata (Lange) Hulten, P. chamissonis, and P. insularis from
Svalbard, P. chamissonis and P. nivea ssp nivea from Northern and
Southern Norway, and P. chamissonis from Northern Finland. The material is or will
be analysed by means of isozymes, RAPDs (Random Amplified Polymorphic DNAs) and
morphometry. Crossing experiments are carried out within and between the taxa from
Svalbard. Offspring from the crossing experiments will be analysed by means of isozymes
and morphometry.
The taxa have slightly different habitats. At Svalbard, P.
chamissonis occurs in bird manured scree slopes and flat ground close to the
shoreline; P. insularis occurs in south facing cliffs, often bird manured scree
slopes; and P. nivea occurs in south facing cliffs and on mountain plateaus. In
mainland Norway, P. chamissonis was found mainly in subalpine cliffs, and P.
nivea mainly on alpine mountain plateaus. However, in a few localities the two taxa
occurred together.
Soják (1986) proposed that P. insularis, a
Svalbard endemism, had arisen from P. chamissonis x P. lyngei Yurtsev &
Sojak (the latter in section Multifidae). Potentilla lyngei has been
reported once from Svalbard, but this collection has been questioned (Elven
& Elvebakk 1996). It probably represents a manured specimen of P. insularis
(cf. Eriksen & Nyléhn 1999). Elven & Elvebakk
(1996) proposed that rather P. chamissonis and P. pulchella R. Br. (the
latter in section Multifidae) might be the parent species of P. insularis.
Furthermore, Soják (1986) proposed that P. nivea ssp subquinata
is the hybrid between P. nivea ssp nivea and P. pulchella. These
hypothese of hybrid origin for the Svalbard taxa are not supported by molecular data.
Analyses of RAPDs (Hansen 1998), morphometry, and
isozymes (Hamre in prep.) have revealed that P. insularis, P. nivea ssp subquinata
and P. chamissonis on Svalbard are closely related, and distant from P.
pulchella. P. pulchella is neither a parent species of P. insularis nor
for P. nivea ssp. subquinata.
Potentilla pulchella has, however, been recorded with different
“morphotypes” closely connected to different habitats: the “beach
type” growing on flat ground mainly close to the fiords, and the “bird manured
scree slope type” (which have been denoted P. rubricaulis by Rønning,
1961) under bird mountains and in southfacing cliffs. Analyses by means of RAPDs (Hansen, 1998) and isozymes (Hamre, in prep.) showed almost no genetic
variation. The little genetic variation found did not correspond to the
“morphotypes”. Thus, P. pulchella should not be taxonomically subdivided
in Svalbard.
When material from Fennoscandia was included in the RAPD
analysis of taxa in the P. nivea complex, three different groups were established in the
PCO plot (Fig. 1). P. chamissonis from Finland and Southern Norway is grouped
together with P. chamissonis from Svalbard. P. nivea ssp. subquinata
from Svalbard and P. nivea ssp nivea from Southern Norway form separate
groups, as distant from each other as from P. chamissonis. The collections from
Northern Norway are not analysed at the moment. Thus, the so far analysed material (P.
chamissonis, P. nivea subsp. subquinata, P. nivea subsp. nivea
and P. insularis) are closely related, but should probably be regarded as separate
taxa. Whether the taxa will be given rank of species or subspecies will await further
research (especially analysis of the material from Northern Norway).
Although hybrids, hybrid swarms and complexes are proposed to be common
in these taxa, only few putative hybrids were found in our collections. No hybrids between
different sections were found. In two populations in Southern Norway where P. nivea
and P. chamissonis co-occured, no certain hybrids were found. The position of the
two lowermost South Norwegian P. chamissonis individuals, originated in a mixed
population, might indicate some introgression. Only a few plants in the Svalbard material
were probably of hybrid origin, possibly between P. nivea and P. insularis (Hansen, 1998).
The Svalbard taxa have less genetic variation than the mainland
populations, thus it is likely that the Svalbard populations are founded by one or few
events.
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Fig. 1.

PCO analysis of P. nivea and
P. chamissonis from Svalbard, Northern Finland and Southern Norway, based on RAPDs
analysis. Axes 1 and 2. |
References |
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Asker,
S. 1977. Pseudogamy, hybridization and evolution in Potentilla. - Hereditas, 87:
179-184.
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Elven, R. & Elvebakk,
A. 1996. A catalogue of Svalbard plants, fungi, algae, and cyanobacteria. Part 1. Vascular
plants. - In: Elvebakk, A. & Prestrud, P. (Eds) Norsk Polarinst. Skr. 198: 9-55.
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Eriksen, B. & Nyléhn,
J. 1999. Cases of phenotypic plasticity in leaves of Potentilla L. (Rosaceae). -
Det Norske Videnskaps-Akademi. I. Mat. Naturv. Klasse, Skrifter, Ny serie 38: 207-215.
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Eriksen, B. 1996. Mating
systems in two species of Potentilla from Alaska. - Folia Geobot. Phytotax. 31:
333-344.
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Hamre (in prep.).
Variation in Potentilla at Svalbard. Cand. Scient. Thesis, University of Oslo.
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Hansen, K. T. 1998.
Variation in the Potentilla nivea complex and P. pulchella in Svalbard,
analysed by morphometry and RAPD markers. Cand. Scient. Thesis, University of Oslo.
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Holm, S. 1996. Sexuality,
apomixis and polyploidy: Potentilla argentea as a critical example. PhD thesis,
Lund University.
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Rønning, O. I. 1961. Some
new contributions to the flora of Svalbard. - Norsk Polarinst. Skr. 124: 1-20.
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Soják, J. 1986. Notes on Potentilla.
I. Hybridogenous species derived from intersectional hybrids of sect. Niveae x
sect. Multifidae. - Bot. Jahrb. Syst., 106: 145-210.
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