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УДК 502.7:632.51
ЭВОЛЮЦИЯ ИНВАЗИВНОСТИ У ЭНОТЕР
Род Oenothera L. (subsect. Oenothera, Onagraceae) является очень удобным модельным объектом для изучения эволюции инвазивности заносных видов в природные местообитания. Это связано с генетическим механизмом размножения, который присущ энотерам (механизм перманентной гетерозиготной транслокации), который способствует тому, что в результате гибридизации двух видов может образовываться константный третий. В результате анализа оригинальных и литературных данных о распространении гибридных и паренталь-ных видов и их инвазивности в различных частях Европы были установлены некоторые особенности микроэволюции инвазивности у видов. Группа «очень инвазивных» гибридных видов чаще всего происходила от родителей из группы «очень инвазивных» парентальных видов. Изучение происхождения парен-тальных видов свидетельствует о том, что наиболее инвазивные гибриды образовались в результате гибридизации между североамериканскими и европейскими видами. Частота встречаемости парентальных видов также как и генетические особенности близких морфологически видов также являются значимыми характеристиками, влияющими на степень инвазивности видов.
Ключевые слова: Oenothera, эволюция, инвазивность, гибриды.
Invasive species are of great interest to evolutionary biologists and ecologists because they represent historical examples of dramatic evolutionary and ecological change [1]. The genus Oenothera L. (subsect. Oenothera, Onagraceae) is one of the most widespread genera of American origin in Europe. Oenothera species are successful invaders of disturbed riverbanks, ruderal places, parks, roads, sandy dunes, natural reserves and sea coasts in Europe [2, 3]. In result of their intercontinental spread many species and forms have been originated in Europe that has resulted in a number of taxonomic problems in the genus (Table 1). The species possess a special breeding system in Evening Primroses (permanent heterozygous translocation (PTH) mechanism) which promotes the hybridization (also introgressive one) among several species resulting in the formation of a stable (constant) third taxon [4, 5]. 14 chromosomes of oenotheras from the subsection can merge by different way and form rings during meiosis (so called Renner’s rings). And many combinations of these rings can be formed in hybrid plants. Therefore, this genus is a good object for research of microevolutionary changes and adaptation of plants and evolution of their invasiveness.
Material and methods
Hybridization among species may serve as a stimulus for the evolution of invasiveness and leads to critical evolutionary changes that create an opportunity for increased invasiveness [1]. The analysis of hybrid species distribution in Europe shows existence of groups with different degree of invasiveness (Table 2). The hybrid Evening primroses were analyzed by the following criteria which can related to evolution of their invasiveness: 1) time of origin of hybrid,
2) degree of invasiveness of the parental species, 3) geographical origin of parental species, 4) frequency of occurrence of parental species, 5) their invasiveness, 6) genetic factors (hetero-sis; possibility of hybridization between: cultivated and natural species, ring formed and bivalent species), 7) environmental factors (climatic and ecological), 8) biological peculiarities (life history traits and morphology) of newly created species, 9) presence of accompanying organisms on hybrids (EICA hypothesis). Some of the mentioned traits are analyzed in the Table 2.
Results and discussion
The time of origin is not determined factors in plant invasiveness of Evening primroses. It is perhaps connected with the fact that majority of species have been originated practically
В.К. Тохтарь1, Р. Виттиг2
1 Белгородский
государственный
университет
Россия, 308015, г. Белгород,
ул. Победы, 85
e-mail: [email protected]
2Университет
им. И.В. Гете, Институт
ботаники
Германия, Д-60323,
г. Франфуркт на Майне,
п/о 1119 32,
Сиесмайерштрассе, 70
e-mail:
simultaneously in time. Therefore, considering this criterion there are no sufficient differences on in the plant group.
The degree of invasiveness of the parental species is influential but not good understood trait impacting to invasiveness of hybrids. Our analysis shows that in the group of “very invasive” hybrid species there are also “very invasive” parental species (table 2). One of the basic species for such hybrids often is Oe. biennis. And there is only one exception from the rule with Oe. punctulata which are not invasive at the moment and their parental species belong to “very invasive” species group. However it also can be connected with the lack of reliable data on distribution of the species in Europe.
The analysis of the parental species origins testifies that the most invasive hybrids are created from crossing among North American and European species. In result of hybridization the heterosis effect can arise among isolated lines promoting viability of newly created species.
The frequency of the parental species is of important value for the invasiveness of hybrid species because the hybridization probability between parental species is increased with frequency of their presence at the same habitat.
Sometimes the resulting hybrids are morphologically similar to the parental species and, at the same time, are very different by genetic nature. The invasiveness of all these species is also distinguished from one another. In this case evolution of invasiveness depends on genetic nature of the species most of all.
Table 1
Oenothera species in Europe based on literary data
Oenothera species Origin Remarks
1 2 3
acutifolia Europe, Oe. silesiaca x rubricaulis Crossing between European species
affnis South America
albipercurva Europe, Oe. biennis x ammophila Crossing between European and North American species
ammophila N America
angustissima N. America = Oe. rubricuspis, introduced before 1900, species escaped from gardens
biennis s.str. Europe (Rostanski), N.America (Raven et al.)
braunii hybrid origin crossing between European and N. American species
britannica Oe. glazioviana x cambrica (R., 2003) crossing between N. American species
cambrica N. America = Oe. nova-scotiae
canovirens North America = Oe. renneri, introduced in 20th century, after II World War
chicaginensis = Oe. pycnocarpa
coronifera Perhaps Oe. parviflora x glazioviana (R., 2003) crossing between N. American species
cruciata N. America = Oe. atrovirens, introduced before 1900, species escaped from gardens
deflexa North America, Europe = Oe. lipsiensis, introduced in 20th century, after II World War
depressa North America = Oe. salicifolia, introduced before 1900, species escaped from gardens
drawertii Oe. depressa x suaveolens crossing between European and N. American species
ersteinensis N. America = Oe. perangusta
erythrosepala N. America =Oe. glazioviana
fallax glazioviana x biennis crossing between European and N. American species
flaemingina Europe related with Oe. rubricaulis
flava ? ?
glazioviana North America = Oe. erythrosepala, introduced before 1900, species escaped from gardens
Тохтарь В.К., Виттиг Р. Эволюция инвазионности у энотер
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The end of the table і
і 2 3
hoelscheri rubricaulis x depressa crossing between European and N. American species
indecora South America
issleri biennis x oakesiana crossing between European and N. American species
italica p ?
jamesii N. America introduced before 1900, species escaped from gardens
jueterbogensis Europe (Germany) hybrid origin, recently noted in Poland
laciniata N. America
longiflora p ?
missouriensis N. America
moravica Europe, fallax x victorini
nuda N. America = Oe. nutans, introduced in 20th century, after II World War
oakesiana North America = Oe. syrticola, introduced before 1900, species escaped from gardens
oehlkersii Europe, glazioviana x suaveolens crossing between European and N. American species
paradoxa Perhaps Oe. depressa x subterminalis (R., 2003) crossing between N. American species
parviflora North America introduced before 1900, species escaped from gardens
perangusta N. America (Rost., 2003) = Oe. ersteinensis
perennis N. America 7
Oe.polgari Oe. suaveolens x depressa (R., 2003) crossing between N. American species
punctulata Oe. biennis x pycnocarpa crossing between European and N. American species
purpurans Oe. glazioviana x depressa (R., 2003) =Oe. hungarica, crossing between European and N. American species
crossing between European species
pycnocarpa North America =Oe. chicaginensis, introduced in 20th century, after II World War
rigirubata
rosea South America pink flowers, different sect.
royfraseri N. America = Oe. turoviensis, introduced in 20th century, after II World War
rubricalix N. America
rubricaulis East European species (Rostanski, 2003) Recently originated
salicifolia = Oe. depressa
silesiaca = Oe. subterminalis
stricta South America
stuchii Perhaps Oe. jamesii x suaveolens (R., 2003) crossing between N. American species
suaveolens South Europe (Rostanski, 2003)
subterminalis North America = Oe. silesiaca
syrticola N. America = Oe. chicaginensis
tacikii Oe. suaveolens x rubricaulis Crossing between European species
tetragona
tetraptera
turoviensis = Oe. royfraseri, introduced in 20th century, after II World War
victorini North America =Oe. nissensis, = Oe. rostanskii Jehlik, introduced in 20th century, after II World War
wienii Oe. rubricaulis x depressa crossing between European and N. American species
Table 2
Analysis of European Oenothera species on different invasiveness groups
Invasive- ness Name of hybrid Parental species Frequency of occurrence of hybrids related to parental species Frequency of occurrence of hybrids Origin of parental species Invasiveness of parental species and their distribution Predominant distribution of hybrid
Very invasive x Oe. fallax x Oe. issleri x Oe. hoelscheri Oe. biennis s.str. x Oe. glazioviana Oe. biennis s.str. x Oe. oakesiana Oe. rubricaulis x Oe. depressa > < > Often Often Often Europe ? Europe Europe NA Very invasive (E) Sometimes invasive (WE, CE) Very invasive (E) Non-invasive or rare-invasive (WE, CE) Invasive (CE, EE) Invasive (CE, EE) WE, CE Mostly WE and less in CE CE, EE
Middle- invasive x Oe. oehlkersii x Oe. acutifolia x Oe. paradoxa Oe. suaveolens x Oe. glazioviana Oe. rubricaulis x Oe. silesiaca ? < < < ? Not often Not often Not often Europe NA Europe Europe ? Middle invasive (WE, CE, EE) Middle invasive (WE, CE) Invasive (CE, EE) Invasive (WE, CE) ? WE, CE, EE CE CE
Non- invasive, Locally distrib- uted x Oe. drawertii x Oe. punctulata x Oe. wienii x Oe. coronifera x Oe. moravica x Oe. jeter-bogensis Oe.depressa x Oe. suaveolens Oe. biennis s.str. x Oe. pycnocarpa Oe. rubricaulis x Oe. depressa Oe. glazioviana x Oe. parviflora Oe. fallax x Oe. victorini ? < < < < < < < < < ? ? Rare Not often Rare Rare Not often Very rare NA Europe Europe NA Europe NA NA NA Europe ? ? Non-invasive (WE, CE) Middle-invasive (WE, CE, EE) Very invasive (E) Very invasive (WE, CE) Invasive (CE, EE) Invasive (CE, EE) Middle invasive (WE, CE) Non-invasive (WE, CE, EE) Very invasive (WE, CE) Non-invasive (CE, FE) ? WE, CE WE, CE CE, EE CE CE CE
WE -West Europe, CE - Central Europe, EE - East Europe, FE - Far East, E - everywhere, NA - North America.
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Toxrapb B.K., Burrur P. Эво^wцмa MHBa3MOHHOCTM y энотер
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Oe. glazioviana also has presumable hybrid nature. And there are a lot of hybrid species with undetermined status and unclear range of distribution and putative parental species. These species are very difficult to recognize them reliably without experience. Many of them are recently originated and locally distributed, mostly from places where they were described.
Beside those hybrid species, which were noted in the table 2 K. Rostanski reported also 33 hybrid species locally distributed in Europe. Sometimes such species are known only from one locality: Oe. braunii Doell, Oe. brevispicata Hudziok, Oe. canovertex Hudziok, Oe. clavifera Hudziok, Oe. coloratissima Hudziok, Oe. compacta Hudziok, Oe. conferta Renner, Oe. editicau-lis Hudziok, Oe. flaemingina Hudziok, Oe. inconspecta Hudziok, Oe. indivisa Hudziok, Oe. macrosperma Hudziok, Oe. mediomarchica Hudziok, Oe. obscurifolia Hudziok, Oe. octolineata Hudziok, Oe. pseudocernua Hudziok, Oe. rigirubata Renner ex Gutte et Rostanski (all are distributed in Germany), Oe. adriatica Soldano, Oe. fallacoides Soldano et Rostanski, Oe. marinella Soldano, Oe. pedemontana Soldano, Oe. pellegrinii Soldano, Oe. sesitensis Soldano, Oe. stuchii Soldano (all are distributed in Italy), Oe. pseudochicaginensis Rostanski, Oe. tacikii Rostanski, Oe. wratislaveinsis Rostanski (all are distributed in Poland), Oe. polgari Rostanski, Oe. purpurans Borbas (Hungary), Oe. carinthiaca Rosanski, Oe. heiniana Teyber (Austria), Oe. slovaca Jehlik et Rostanski (Slovakia), Oe. britannica Rostanski (Great Britain). That is why there are many perspectives to provide new insights studying invasiveness in this, very suitable for these goals, plant group in future.
Conclusions
The analysis of hybrid species distribution in Europe shows existence of groups with different degree of invasiveness. The degree of invasiveness of the parental species is influential but not good understood trait impacting to invasiveness of hybrids. Our analysis shows that in the group of “very invasive” hybrid species there are also “very invasive” parental species. The analysis of the parental species origins testifies that the most invasive hybrids are created from crossing among North American and European species. In result of hybridization the heterosis effect can arise among isolated lines promoting viability of newly created species. The frequency of the parental species is of important value for the invasiveness of hybrid species because the hybridization probability between parental species is increased with frequency of their presence at the same habitat. Sometimes the resulting hybrids are morphologically similar to the parental species and, at the same time, are very different by genetic nature. The invasiveness of all these species is also distinguished from one another. In this case evolution of invasiveness depends on genetic nature of the species most of all.
Acknowledgements
First author of this research work was partly supported by the Russian Foundation for Basic Research (RFBR), grant № 08-04-00239, “Evolution of alien plants invasiveness: mechanisms of microevolution and methodical aspects of alien species distribution prognosis”.
Literature
1. Ellstrand N.C. & Schierenbeck K.A. Hybridization as a stimulus for the evolution of invasiveness in plants? / / PNAS - 2000. - Vol. 97, № 13 - P. 7043-7050.
2. Rostanski K., Tokarska-Guzik B. Distribution of the American epecophytes of Oenothera in Poland. / / Phytocoenosis. - 1998. - Vol. 10. - P. 117-130.
3. Wittig R., Lenker K.-H., Tokhtar V.K. Zur Sociologie von Arten der Gattung Oenothera L. im Rheintal von Arnheim (NL) bis MUlhouse (F) // Tuexenia. - 1999. - Bd. 19. - S. 447 - 467.
4. Renner O. Versuche uber die gametische Konstitution der Oenothera // Z. Abst. Vererb. - 1917. - Vol. 18. - P. 121-294.
5. Cleland R.E. The evolutionary history of the North American evening primroses of the “biennis group” / / Proc. Amer. Phil. Soc. - 1964. - Vol. 108. - P. 88 - 98.
V.K. Tokhtar'1,
R. Wittig2
1 Belgorod State University Pobedy Str., 85,308015, Belgorod, Russia
e-mail: [email protected]
2 Frankfurt-on-Main University, Institute of Botany Siesmayerstrasse 70,
Postfach 1119 32, D-60323, Frankfurt-on-Main, Germany
e-mail:
EVOLUTION OF INVASIVENESS IN OENOTHERA
The genus Oenothera L. (subsect. Oenothera, Onagraceae) is very convenient modelling object for evolution studying of inadventive invasiveness species in natural habitats. It is connected with the genetic mechanism of reproduction, which is inherent oenotheras (the mechanism of a permanent heterozygotic translocation) which promotes that as a result of hybridization of two kinds the third can be formed constant. As a result of the analysis of original and literary data about distribution of hybrid and parental species and their invasiveness in various parts of Europe some features of microevolution invasiveness at species have been established. The group «very invasiveness » hybrid species more often occurred from parents from group «very invasiveness» parental species. Origin studying parental species testifies that most invasiveness hybrids were formed as a result of hybridization between North American and European species. Frequency of occurrence of parental species as well as genetic features of species close by morphology is also the significant characteristics influencing on degree of species invasiveness.
Key words: Oenothera, evolution, invasiveness, hybrids.
