Modern Phytomorphology, Vol. 1, 2012

MODERNPHYTOMORPHOLOGY

Volume 1

Volum

e 12

012

2012

ISSN 2226-3063

M O D E R N P H Y T O M O R P H O L O G Y 12 0 1 2Editor-in-ChiefEditorial AssistantExecutive Editor

Tasenkevich L.O.Kondratyuk S.Ya.

Novikoff A.V.

Berko Yo.N.Budzhak V.V.Bukhtiyarova L.N.Danyluk K.N.Deroin T.Eberwein R.Kalinovych N.O.Klymyshyn A.S.Korzhenevsky V.V.Korzeniak J.Lobachevska .V.Lyakh A.M.Mamchur Z.I.Mitka J.Odintsova A.V.Ostash B.O.Peruzzi L.Savinykh N.P.Terek O.I.Tiezzi A.Fedorenko V.A.Tsaryk Yo.V.Chernobay Yu.M.Chornej I.I.Shipunov A.Shevchenko S.V.Szczepanek K.

S.Z. Gzhytskyj Lviv National University of Veterinary Medicine and Biotechnologies, Lviv, UkraineYuriy Fedkovich Chernivtsi National University, Chernivtsi, UkraineM.G. Kholodny Institute of Botany NASU, Kyiv, UkraineState Natural History Museum NASU, Lviv, UkraineNational Museum of Natural History, Paris, FranceCarinthian Botanic Center, Klagenfurt am Woerthersee, AustriaIvan Franko National University of Lviv, Lviv, UkraineState Natural History Museum NASU, Lviv, UkraineNikitsky Botanical Gardens National Scientific Centre,Yalta, UkraineInstitute for Nature Conservation PAS, Cracow, PolandInstitute of Ecology of the Carpathians of NAS of Ukraine, Lviv, UkraineA.O. Kovalevsky Institute of Biology of the Southern Seas NASU, Sevastopol, UkraineIvan Franko National University of Lviv, Lviv, UkraineInstitute of Botany PAS, Cracow, PolandIvan Franko National University of Lviv, Lviv, UkraineIvan Franko National University of Lviv, Lviv, UkraineUniversity of Pisa, Pisa, ItalyVyatka State University, Kirov, RussiaIvan Franko National University of Lviv, Lviv, UkraineTuscia University, Viterbo, ItalyIvan Franko National University of Lviv, Lviv, UkraineIvan Franko National University of Lviv, Lviv, UkraineState Natural History Museum NASU, Lviv, UkraineYuriy Fedkovich Chernivtsi National University, Chernivtsi, UkraineMinot State University, Minot, USANikitsky Botanical Gardens National Scientific Centre,Yalta, UkraineInstitute of Botany PAS, Cracow, Poland

Editorial Board

Modern Phytomorphology: Proceedings of the 1st International Scientific Conference on Plant Morphology (24-26 April 2012, Lviv, Ukraine). Lviv, 2012. Vol. 1. 208 p.

www.phytomorphology.org

Modern Phytomorphology

Technical EditorLayoutDesignProofreaders

Cover photo

Novikoff A.V.Novikoff A.V.Novikoff-Supp M.R., Novikoff A.V.Pochynok T.V., Gavrylov A.I., Klymovska D.T.

Sunset in Marmarosh Mts., Eastern Carpathians; Novikoff A.V.

Ivan Franko National University of Lviv, Lviv, UkraineM.G. Kholodny Institute of Botany NASU, Kyiv, UkraineState Natural History Museum NASU, Lviv, Ukraine

Supported by

Zhukovsky P.Yu.

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Contents

Mitka J. Aconitum in Central Europe: from Linnaean taxonomy to molecular markers .............................. 7

Sotys-Lelek A., Barabasz-Krasny B., Turis P., Turisov I. Taxonomic revision of roses (Rosa L.) of selected areas in buffer zone of the Low Tatras National Park (Slovakia) ....................... 11

Chernetskyy M.A. The role of morpho-anatomical traits of the leaves in the taxonomy of KalanchoideaeBerg. subfamily (Crassulaceae DC.) ................................................................................ 15

Dbrowska A. Morpho-anatomical structure of the leaves of Festuca trachyphylla (Hack.) Krajina in the ecological aspect ................................................................................................................................. 19

dos Santos P., Brockington S., Glover B. & Ronse de Craene L.P. Micromorphological evidence for androecium origin of Claytonia (Montiaceae) petaloids .................................................. 23

Novikoff A.V. & Kazemirska M.A. Vascular anatomy and morphology of the flowerin Fritillaria montana Hoppe (Liliaceae) ......................................................................................................................... 27

.., .., .., .., .. .................................................. 37

Savinykh N.P., Degtereva O.P., Zhuravleva I.A., Chuprakova E.I., Shabalkina S.V. Structural multivariate of plants from the position of modular organization ..................................... 37

.. - ........................................ 43

Kondratyuk T.O. Morphological and physiological characters of microscopic fungi damaging objects and materials under biocide influence .......................................................................................... 43

.., .-., .-., .. Cladosporium sphaerospermum ............................. 47

Kondratyuk T.O., Jeong M.-H., Hur J.-S., Kondratyuk S.Y. identification of Cladosporium sphaerospermum species complex after morphology and molecular phylogeny .................................. 47

.., .-., ., .., .-., . Teloschistaceae (Ascomycota): .............. 53

Kondratyuk S.Y., Jeong M.-H. , Krnefelt I., Elix J.A., Hur J.-S., Thell A. Phylogeny and taxonomy of the Teloschistaceae (Ascomycota): importance of monophyletic groups ..................................................... 53

- .. - Atlas flory Polskiej i ziem ociennych .......................................................................................................................................... 59

Novikoff-Supp M.R. An artistic analysis of morpho-anatomical illustrations from Atlas flory Polskiej i ziem ociennych ......................................................................................................................... 59

.. ........................................ 71Odintsova A.V. Toward the principles of syncarpous gynoecia classification .............................................. 71

.. Trapa natans L. . 77Nedukha .M. Ultrastructur of epidermal surface in floating and submerged leaves of Trapa natans L. . 77

.. Trapa natans L. .......................................................................................................... 81

Nedukha .M. Ultrastructural characteristic of cells and pigment analysis in floating and submerged leaves of Trapa natans L. .............................................................................................................................. 81

.. Bacillariophyta . i. Gomphonema Ehrenb. .............................................................. 85

Bukhtiyarova L.N. Morphology of the new for Ukraine Bacillariophyta from the hydrotopes of the Rightbank forest-steepe. i. Species of Gomphonema Ehrenb. .......................................................... 85

.. 3D-Microalga ............................................................................................ 89

Lyakh A.M. 3D-Microalgae software used for the estimation of microalgae biovolumes and surface area ....89

.. - Rorippa anceps (Wahlenb.) Reichenb. (Cruciferae) ................................................................................................................................ 93

Shabalkina S.V. Ecological and biomorphological features of Rorippa anceps (Wahlenb.) Reichenb. (Cruciferae) ................................................................................................................................. 93

.. .. ........................................................................................... 99

Kharchenko V.E. & Cherskiaya N.A. An using of biometric methods for the delineation of floral units on the plant .......................................................................................................................................... 99

.. .. Rapunculus L. (Fourr.) Boiss. Campanula L. ............................................................................................ 103

Dremliuga N.G. & Futorna O.A. The leaf surface strucure of the species from the section Rapunculus L. (Fourr.) Boiss. of the genus Campanula L. in the flora of Ukraine ............................. 103

.. Bromopsis ramosa (Huds.) Holub B. benekenii (Lange) Holub (Poaceae) ...................................................................................... 107

Krasniak O.I. The characters of epidermal ultrastructure for the taxonomy of Bromopsis ramosa (Huds.) Holub and B. benekenii (Lange) Holub (Poaceae).................................................................... 107

.. .. Stichococcus Ngeli (Trebouxiophyceae, Chlorophyta) ............................................................................................... 111

Karbovska V.M. & Kostikov I.Yu. New morphological features of the members of genus Stichococcus Ngeli (Trebouxiophyceae, Chlorophyta).......................................................................... 111

.. . - ....................................................................... 115

Ezhkin A.K. Morphological changes of the lichens in anthropogenic modified habitats in Yuzhno-Sakhalinsk City suburbs .............................................................................................................. 115

.. ....................................... 119Chrynova T.R. A morphometry of overground shoot of spring wheat plants ............................................ 119

.. Pinus sylvestris L. - ... 125Skliar V.G. Tthe morphology of Pinus sylvestris L. young undergrowth in the forests of

Novgorod-Siversk Polissia ........................................................................................................................ 125

.. Tofieldia calyculata (L.) Wahlenb. (Melanthiaceae) ......................................................................................................................... 129

Kuzyarin A.T. The biomorphological keys of age stages of Tofieldia calyculata (L.) Wahlenb. (Melanthiaceae) ........................................................................................................................................... 129

.. .. Fragaria L. ............................................................................................................................. 133

Biryulyova E.G. & Nikolenko V.V. Structural organization of the foliage of ornamental varieties of genus Fragaria L. .................................................................................................................................... 133

.. ................................................................................................................................ 137

Gorelov A.M. The role of phytogenous field in the formation of woody plants space structure ............. 137

.. .. .................................................................................................................. 143

Antyufeyev V.V. & Antyufeyeva L.I. Reconstruction of the insolation conditions of the past by the growth rings of the trees ...................................................................................................................... 143

.. Gentiana L. .................................................... 149Prokopiv .. Structural organization of shoot system of Gentiana L. ........................................................ 149

.. Festuca valesiaca agg. (Poaceae) .................................................. 153

Bednarska I.O. The method of analysis of the leaf blade anatomical structure of narrow-leaves fescues on example of populations of Festuca valesiaca agg. (Poaceae)............................................... 153

.. (Luzula taurica (V.i. Krecz.) Novikov, Juncaceae) .................................................................................. 157

Olshanskyi I.G. Morphology and surface ultrastructure of seed in Luzula taurica (V.i. Krecz.) Novikov (Juncaceae) ................................................................................................................................... 157

.. Celastraceae R. Br. Cassinoideae Loes. ....................................................... 161

Savinov I.A. The levels of fruit organization in Celastraceae and structural diversity of pirenariums in Cassinoideae ............................................................................................................................................ 161

.., .., .. Suaeda acuminata (C.A. Mey.) Moq., Suaeda prostrata Pall. Tamarix ramosissima Ledeb. ......................... 167

Tsymbalyuk Z.M., Bezusko L.G., Tsymbalyuk T.I. Palynomorphological features of Suaeda acuminata (C.A. Mey.) Moq., Suaeda prostrata Pall. and Tamarix ramosissima Ledeb. ................... 167

.. - Ruppia cirrhosa (Petagna) Grande ......... 173

Kireeva E.V. An usage of morpho-anatomical features of Ruppia cirrhosa (Petagna) vegetative organs for determination of the optimal depth of growing ................................................................... 173

.. .. ................................................................. 179

Voloshina N.Yu. & Belyavskaya N.A. Dependence of leaf structural indices in two forest maple species from within-crown irradiance ..................................................................................................... 179

.. .. ...................................................................................................................... 185

Zhuk I.V. & Musiyenko M.M. The influence of nitric oxide and mercury chloride on leaf mesophyll structure under natural drought conditions ............................................................................................ 185

.., .., .. (Rubus arcticus L.) .......................... 189

Konstantinov A.V., Khimchenko E.N., Kulagin D.V. Rejuvenation of the aseptic culture and obtaining of planting stock of arctic bramble (Rubus arcticus L.) ........................................................ 189

.., .., .. Sansevieria thyrsiflora Thunb. S. grandis Hook. f. (Dracenaceae) ........................................................................ 193

Shkrum I.V., Gajdarji .., Badanina V.. Anatomical features of the leaves of Sansevieria thyrsiflora Thunb. and S. grandis Hook. f. (Dracenaceae) ..................................................................... 193

.., .., .. Polystichum Roth ......................................................................................................................................... 197

Polyshchuk I.O., Vasheka O.V., Tyshchenko O.V. Anatomical structure of Polystichum Roth ferns rachises ............................................................................................................................................... 197

.. ................................................................................................................... 201

Klymyshyn A.S. Adaptive changes in biomorphs of grass creeping perennials in anthropogenic succession ..................................................................................................................................................... 201

7 Modern Phytomorphology 1: 79, 2012

ACONITUM IN CENTRAL EUROPE:FROM LINNAEAN TAXONOMY TO MOLECULAR MARKERS

Jzef Mitka

Abstract. A role of the Linnaean taxonomy in the arising of historical-biogeographical hypotheses is envisioned. The first example concerns the presumed hybrid origin of an Eastern-Sudetic endemic Aconium plicatum subsp. sudeticum. it was described on the basis of a unique character set including glandular hairiness of the indumentum. A PCR-RAPD+iSSR fingerpriting confirmed the supposition based on the morphological analysis. The second example is dealt with the marginal populations. They are of special interest because of the ecological and population genetic phenomena, including genetic drift and subsequent schizoendemism (endemovicarism), a form of the peripatric speciation. Aconitum bucovinense occurs in two marginal, isolated populations in the Western Bieszczady Mts. (E Carpathians). They form a unique morphotype recognized by a taxonomic revision. A PCR-iSSR protocol was used to check a hypothesis on the genetic distinctness of the small, isolated populations. in the effect the lowering by 13% of genetic diversity in the marginal populations, in comparison to the core population, was noted. However, the genetic depauperation was accompanied by the existence of unique bands leading to the distinct genetic stocks in the marginal populations.

Key words: Carpathians, marginal population, peripatric speciation, reticulate evolution, schizoendemism, Sudetes

Jagiellonian University, Institute of Botany, Botanical Garden, Kopernika 27, 31-501 Krakw, Poland; [email protected]

Introduction

The genus Aconitum in Central Europe possesses some 10% of a total of 300-400 species (Liangqian L. & Kadota 2001; Mitka 2003), mostly known from central and eastern Asia (Kadota 1987). in Europe its taxonomic division, in opposite to Asia, is clear. The subgenus Lycoctonum is confined to a few species, however morphologically variable (Warncke 1964). in the Carpathians it is represented by three species and their hybrids (Mitka 2008). The most abundant subgen. Aconitum consists of two section: the diploid sect. Cammarum and the tetraploid sect. Aconitum (Joachimiak et al. 1999, Mitka 2003, ilnicki & Mitka 2009). A. anthora should be included into a separate subgen. Anthora, together with some other Asian species.

The aim of the present paper is to show, how the Linnaean taxonomy may aide to understand the evolution of the genus in a regional scale. The molecular DNA analyses may yield fruitful results while considered in the framework of the proper classification system, which forms a working hypothesis on the evolutionary relationships within a taxon (Mitka 2004).

Evolutionary links between the Sudetes and Carpathians

The two neighbor mountain systems belong to two

different orogenic systems. The Eastern Carpathians and Western Carpathians were formed at the Oligocene/Miocene boundary (2622 myr BP), that the whole of the Carpathians were united some 14 myr BP (in the Middle Miocene), and that this event was accompanied by regression of the sea from the Alpine-Carpathian Foredeep. On the other hand, the Sudetes, a part of the Hercynides, were uplifted much earlier at the Cretaceous/Paleocene boundary during the Laramian tectonic phase some 6560 myr BP. From that time to the present they have formed a stable land mass (see Mitka et al. 2007).

Taking into consideration the geological-historical context of the biotas evolution in the Central-European mountain ranges one may hypothesize that the Carpathian flora has its roots in the old, Sudetic flora (Syabryay 1995). A taxonomic revision of the genus Aconitum in Poland and adjacent countries enabled an endemic species to the Eastern Sudetes, i.e. A. plicatum subsp. sudeticum Mitka (2003), to be described. it has pilose indumetum that relates it to the Western Carpathian A. firmum subsp. maninense (Skalick) Starmhl. and A. firmum subsp. moravicum Skalick. Thus, a hypothesis was that A. sudeticum is a hybrid between the Sudetic A. plicatum and one of the Carpathian species from the A. firmum-group. To check the hypothesis the molecular and cytogenetic studies on the Western-Carpathian and Eastern-Sudetic Aconitum were carried out (Mitka et al. 2007). A PCR-iSRR+RAPD protocol and C-Giemza

J. Mitka, 2012

8 Modern Phytomorphology 1 (2012)

heterochromatine staining showed the distinctness of the two species at the genome level and some similarities at the population genetic level. First of all, A. plicatum is an autotetraploid and A. firmum is an allopolyploid. The pattern of the heterochromatine bands show that A. firmum could be a hybrid between A. plicatum and a diploid species related to A. variegatum. The latter species could be an ancestral to the European Aconitum sect. Cammarum since it has the simplest in structure, homozygous the NOR-chromosome pairs (Joachimiak et al. 1999). The phenetic UPGMA classification revealed the close relationships between A. maninense and A.sudeticum at the bootstrap value 58% (Mitka et al. 2007). Here, we used the same data to perform a population-genetic analysis with the use of a reticulate evolution algorithm. The result displayed in Fig. 1 confirms the shared genetic genomes of the Sudetic A. sudeticum and Carpathian A. maninense, which are nested within A. plicatum with a high bootstrap 82%. Both species are allopatric, narrow endemics. A. sudeticum occurs in the Hrub Jesenk Mts. and Mt. Snenik (locus classicus), and A. maninense in the Straovske vrchy Mts. (locus classicus) and the Tatra Mts. (Mitka 2003). Their geographical ranges could meet in the forelands during one of the Quaternary pleniglacials. At those periods the high-mountain species extended their areas. A secondary contact, probably occurred somewhere in the Moravian Gate, might result in the gene exchange between the Sudetic and Carpathian genetic stocks. in interglacial period the geographical ranges of the parental species and their hybrid were again restricted to the ancestral areas. However, the process was asymmetric, because the putative hybrid A. sudeticum occurs today only in the one, Sudetic, area. Another hypothesis is

that both species: A. maninense and A. sudeticum are putative hybrids and represent opposite ends of the hybrids morphospace. if so, the parental species could be A. plicatum and one of the forms of A. firmum, for example A. f. subsp. moravicum. To test the hypothesis more molecular DNA and cytogenetic studies on Aconitum in the Eastern Sudetes and Western Carpathians are needed.

Populations at the range margin

Aconitum bucovinense is a high-mountain species, Southern/Eastern Carpathian endemic. Previous phenetic studies on a whole tetraploid group of Aconitum in the Eastern Carpathian showed the distinctiveness of the speciess populations at the range margin in the Western Bieszczady Mts. (Mitka 2002). The morphological traits specific to the region include: the type of indumentum hairiness (see Fig. 1 in Mitka 2000) and the shape of the spur. Recently, we checked a hypothesis on the peripatric speciation of Aconitum bucovinense in the Western Bieszczady Mts. (Boro et al. 2011). Peripatric speciation is a form of a schizoenedmism or endemovicarism. The latter terms denote continuous diversification of an ancestral taxon into derived taxa of identical chromosome number in various parts of the range. The process is dealt with founder effect and subsequent accidental elimination of genes (genetic bottleneck). in the effect the speciation process is accelerated by the isolation of relatively small, marginal populations. We checked the hypothesis

Fig. 1. Consensus Network (SplitsTree4, Huson & Bryant 2006) based on Reynoldss distances from 255 iSSR+RAPD bands. Bootstrap values (1000 permutations) are given.

Population Caryska Halicz Core

n 23 45 21

PPL95% 50.0 70.20 67.84

h 0.0713 0.0745 0.0791

S.D. h 0.1139 0.1027 0.0833

I 0.1270 0.1395 0.1521

S.D. I 0.1737 0.1567 0.1384

DW 2.5739 2.4753 4.0193

Rsign small small large

Table 1. Genetic diversity of the two marginal and core populations of Aconitum bucovinense in the Carpathians. PPL95% percentage of polymorphic loci, h Neis gene diversity, S.D. standard deviation, I Shannons index of diversity (POPGENE ver. 1.32, Yeh 1999); DW rarity index (Schnswetter & Tribsch 2005), RSign statistical significance (p 0.05) based on 1000 permutations (AFLPdat, Ehrich 2006). The analysis based on 258 iSSR bands (Boro et al. 2011).

9

Source of variation Sum of squares Variance components Percent variation P

Mariginal vs. Core population

Among populations 79.743 1.95674 10.34866 < 0.001

Within populations 1474.774 16.95142 89.65134

Total 1554.517 18.90816

Between Marginal populations

Among populations 127.683 3.70486 19.91069 < 0.001

Within populations 983.567 14.90253 80.08931

Total 1111.250 18.60739

Table 2. Analysis of molecular variance (AMOVA) of populations of Aconitum bucovinense in the Carpathians (Arlequin software ver. 3.1, Excofier et al. 2006), based on 258 iSSR bands (Boro et al. 2011).

Mitka J. Aconitum in Central Europe: from Linnaean taxonomy to molecular markers

that the marginal populations are geneticallyuniform and deprived of the genetic diversity in comparison to the core area (Boro et al. 2011).

The indices of population genetic diversity: percentage of polymorphic PLP95% and Neis genetic diversity h have not differed among the core and marginal population, in spite of the Shannons diversity index I that tends to be lower in the in the marginal populations, and rarity index DW that is statistically significantly greater in the core area, and statistically significantly lower in the marginal populations (Tab. 1). At the same time the among-population component of the total molecular variance is nearly twice as great in between the marginal population as between the marginal and core populations (Tab. 2).

in sum, the marginal populations differ each of other by the possessing of unique bands. However, Shannons index of diversity is on average 13% lower in the marginal populations in comparison to the core area. However, the genetic depletion in the marginal populations is accompanied by rare bands occurrence. These population genetic phenomena are linked with special morphological characters, unique to the populations at the range margin. They need a special conservation program.

References

Boro P., Zalewska-Gaosz J., Sutkowska A., Zemanek B, Mitka J. 2011. iSSR analysis points to relict character of Aconitum bucovinense Zapa. (Ranunculaceae) at the range margin. Acta Soc. Bot. Pol. (submitted).

Ehrich D. 2006. AFLPdat: a collection of r functions for convenient handling of AFLP data. Mol. Ecol. Notes 6: 603604.

Excoffier L., Guillaume L., Schneider S. 2006. Arlequin ver. 3.01: an integrated software package for population genetics data analysis. Computational and Molecular Population Lab, Univ. of Berne.

Huson D.H. & Bryant D. 2006. Application of phylogenetic networks in evolutionary studies. Mol. Biol. Evol. 23(2): 254267.

Ilnicki T. & Mitka J. 2009. Chromosome numbers on Aconitum sect Aconitum (Ranunculaceae) from the Carpathians. Caryologia 62(3):198203.

Joachimiak A., Ilnicki T., Mitka J. 1999. Karyological studies on Aconitum lasiocarpum (Rchb.) Gyer (Ranunculaceae). Acta Biol. Cracoviensia, ser. Bot. 41: 205211.

Kadota Y. 1987. A revision of Aconitum Subgenus Aconitum (Ranunculaceae) of East Asia. Sanwa Shoyaku Company, Ltd., Utsunomiya.

Liangqian L. & Kadota Y. 2001. Aconitum L. in: Zhengyi W., Raven P.H., Deyuan H. (eds). Flora of China. T. 6. Bejing: Science Press, St. Louis: Missouri Botanical Garden: 149222.

Mitka J. 2003. The genus Aconitum (Ranunculaceae) in Poland and adjacent countries. institute of Botany of the Jagiellonian University, Krakw.

Mitka J. 2004. Taksonomia linneuszowska w dobie biologii molekularnej. Fragm. Flor. Geobot. Polonica Suppl. 6: 931.

Mitka J. 2008. Aconitum moldavicum Hacq. (Ranunculaceae) and its hybrids in the Carpathians and adjacent regions. Roczn. Bieszczadzkie 16: 233252.

Mitka J., Sutkowska A., Ilnicki T., Joachimiak A.J. 2007. Reticulate evolution of high-alpine Aconitum (Ranunculaceae) in the Eastern Sudetes and Western Carpathians (Central Europe). Acta Biol. Cracoviensia, ser. Bot. 49(2): 1526.

Schnswetter P. & Tribsch A. 2005. Vicariance and dispersal in the Alpine perennial Bupleurum stellatum L. (Apiaceae). Taxon 54: 725732.

Syabryay S.V. 1995. The formation of the Neogene Carpathian flora. Ukrainian Bot. J. 52: 174180.

Yeh F., Yang R., Boyle T. 1999. POPGENE version 1.32. Microsoft-based freeware for population genetic analysis. Canada: Molecular Biology and Biotechnology Center, University of Alberta.

Warncke K. 1964. Die europischen Sippen der Aconitum lycoctonum-gruppe. Mnchen.

Zieliski R. 1982a. An electrophoretic and cytological study of hybridisation between Aconitum napellus ssp. skerisorae (2n=32) and A. variegatum (2n=16). i. Electrophoretic evidence. Acta Soc. Bot. Pol. 51: 453464.

Zieliski R. 1982b. An electrophoretic and cytological study of hybridisation between Aconitum napellus ssp. skerisorae (2n=32) and A. variegatum (2n=16). ii. Acta Soc. Bot. Pol. 51: 465471.


TAXONOMIC REVISION OF ROSES (ROSA L.) OF SELECTED AREASIN BUFFER ZONE OF THE LOW TATRAS NATIONAL PARK (SLOVAKIA)

Anna Sotys-Lelek1, Beata Barabasz-Krasny2, Peter Turis3, ingrid Turisov4

Abstract. The paper presents results of preliminary research carried out in 2011 in the area of two Protected Sites CHA Jakub, CHA Kopec and Nature Reserve PR Mackov bok in the buffer zone of the Low Tatras National Park. Six native species and one native hybrid of roses were found there (1 from the Pimpinellifoliae section and 6 from the Caninae section) and their 13 varieties. The most numerous among them are Rosa canina var. dumalis Baker and R. subcanina (H. Christ) R. Keller. Such great diversity of species and varieties in such a small area (only 20.2 ha) proves considerable floristic richness, and occurrence of habitats preferred by roses.

Key words: Rosa, Rosaceae, taxonomic revision, Low Tatras National Park, Carpathians, Slovakia

1 Ojcw National Park, 32-047 Ojcw 9, Poland; [email protected] Institute of Biology, Departament of Botany, Pedagogical University, Podbrzezie 3, 31-054 Krakw, Poland; [email protected] Low Tatras National Park, Lazovn 10, 974 01 Bansk Bystrica, Slovakia; [email protected] Faculty of Natural Sciences, Department of the Environment Sciences, Matej Bel University, Tajovskho 40, 974 01 Bansk Bystrica, Slovakia; [email protected]

Introduction

The genus Rosa L. is one of the most complicated ones as far as the systematics is concerned, hence, for over 200 years it has been the subject of numerous taxonomic investigations. Hybridisation, especially the introgressive one, is a crucial problem in identification of rose species. Rose species are formed by mixed ones not only closely related, but also species originating from different sections. Hence, it is sometimes difficult to determine species boundaries between them (Zieliski 1985).

Rodological literature applies morphological traits of flower shoots to distinguish the species e.g.: the shape of a disc, the width of an orifice, the shape of prickles, intensity of hairiness and the most important glandularity of leaves and position and shape of sepals. At the beginning of the 19th century the above mentioned features were discussed separately which resulted in multiplication of species names and many forms described today as intraspecies had been described as species (Zieliski 1985). According to the latest rodological systematics the morphological features have to be considered comprehensively and in various combinations. it gives a chance to apply simple selection of forms which might be treated as species and can be easy to characterise (Popek 1996, 2002, 2007; Zieliski 1985, 1987).

Occurrence of wild species of roses in the Low Tatras National Park and its buffer zone has not been thoroughly investigated so far. The data from

the area are scattered among several publications. Occurrence of 14 taxa of roses were determined after literature. Most of them are listed in the works of V. Vtvika (1992) and B. Benaov & K. Ujhzy (1998 ) and F. Prochzka & F. Krahulec (1982), where 11 species and one hybrid form were presented. Additionally J. Tmk (1886) and E.Martincova (1989) mentioned two more species in their publications. Partial data on occurrence of the species from the genus Rosa as well as the change of concepts concerning their systematics gave an impulse to undertake investigations which aim was to create a complete list of species of roses and their varieties wildly growing in the area. The results presented in the paper cover selected protected areas and they are an introduction to a further more detailed investigation.

Material and methods

Field investigations were carried out in the buffer zone of the Low Tatras National Park in the area of the Protected Sites CHA Jakub (12,71 ha), CHA Kopec (3,76 ha) and Nature Reserve PR Mackov bok (3,75 ha) (Fig. 1) in the vegetation season of 2011.

Fruiting short shoots were collected in the field and their following features were recorded: the shape of the prickles (straight, hooked or falcate), the shape of a disc (flat, conical), hypanthium opening (diameter bigger or smaller than 1/3 of the disc), position of sepals and their durability and also

A. Sotys-Lelek, B. Barabasz-Krasny, P. Turis, ingrid Turisov, 2012


intensity of hairiness and glandularity of leaves. in case of the leaves, middle and top parts were taken into consideration, while in case of the prickles the top parts of one-year or two-year-old long shoots, which had already completed the process of growth and their prickles were not changing, were studied.

Systematic approach and the nomenclature of the species were adopted after the works of H. Henker (2000), R. Popek (1996, 2002, 2007) and J. Zieliski (1985, 1987). Herbarium material was deposited in the herbarium of the Ojcw National Park.

Symbols used in article: CHA Protected Site, leg. legit, OPN Herbarium of the Ojcw National Park, PR Nature Reserve.

Results

Six native species and one native hybrid of wild roses (one of them from section Pimpinellifoliae and 6 from section Caninae) and their 13 varieties were recorded in the study area.

I. Sect. Pimpinellifoliae DC.1. Rosa spinosissima L. (syn.: R. pimpinellifolia L.)R. spinosissima appears in variety spinosissima

which is characterized by simple serrate leaflets and glandular pedicels; 13 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN).

II. Sect. Caninae DC. emend. H. Christ.2. Rosa dumalis Bechst. (syn.: R. vosagiaca Desp.)The species was found in the study area in

three varieties, distinctly differing in hairiness and serration of leaves.

- var. afzeliana (Fr.) Boulenger (syn.: R. afzeliana Fries.) is characterized by glabrous, simple or doubly serrate leaflets, normally glandless margin. Pedicels usually glabrous; 6 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN); CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN); PR Mackov bok, leg. Sotys-Lelek, 24.09.2011 (OPN).

- var. dumalis is characterized by glabrous, complex serrate leaflets, glandless or glandular on the underside. Pedicels usually glabrous; 3 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN); PR Mackov bok, leg. Sotys-Lelek, 24.09.2011 (OPN).

- var. coriifolia (Fr.) Boulenger (syn.: R. coriifolia Fries.) simple or doubly serrate leaflets, bilaterally haired or only on the underside; 2 records: CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN); PR Mackov bok, leg. Sotys-Lelek, 24.09.2011 (OPN).

3. Rosa tomentosa Sm. var. tomentosaR. tomentosa was found in the studied area in

one variety, which is characterized by complex serrate leaflets glandular on the underside and glandular pedicels as well; 2 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN).

4. Rosa inodora Fr. var. inodora (syn.: R. elliptica Tausch.)

R. inodora occurs in one variety with glabrous, not glandular pedicels; 3 records: CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN); PR Mackov bok, leg. Sotys-Lelek, 24.09.2011 (OPN).

5. Rosa agrestis SaviFound in the study area in four varieties

differing in hairiness of the leaflets and the degree of glandularity of pedicels.

- var. agrestis leaflets glabrous or haired. Pedicels glabrous; 7 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN); CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN).

- var. schulzei R. Keller leaflets glabrous or haired. Pedicels glandular; 5 records: CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN).

- var. albiflora (Opiz.) Degen bilaterally haired leaflets. Not glandular pedicels; 2 records: CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN).

- var. gizellae (Borbs) R. Keller bilaterally haired leaflets. Pedicels glandular; 5 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN); CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN).

6. Rosa canina L.

Fig. 1. Distribution of study areas: 1 Protected Site CHA Jakub, 2 Protected Site CHA Kopec, 3 Nature Reserve PR Mackov bok. Abbreviations: NAPANT Low Tatras National Park, Lipt. Hrdok Liptovsk Hrdok, Lipt. Mikul Liptovsk Mikul.

13 Sotys-Lelek A. et al. Taxonomic revision of roses (Rosa L.) of the Low Tatras NP

R. canina occurs in the study area in four varieties, distinctly differing in hairiness and serration of leaves and degree of glandularity of pedicels.

- var. canina glabrous, mostly simple serrate leaflets; 1 record: CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN).

- var. andegavensis (Bastard.) Desp. simple or complex-glandular serrate leaflets, glabrous and pedicels glandular with stalked glands; 2 records: CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN).

- var. dumalis Baker glabrous, doubly or complex serrate leaflets; 61 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN); CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN); PR Mackov bok, leg. Sotys-Lelek, 24.09.2011 (OPN).

- var. corymbifera (Borkh.) Boulenger leaflets bilaterally haired or only on the underside. Pedicels not glandular; 13 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN); CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN).

7. Rosa subcanina (H. Christ) R. KellerR. subcanina is a hybrid form characterized by

intermediate features between R. dumalis Bechst. and R. canina L. This hybrid is separated by some researches (Henker 2000) as independent species; 36 records: CHA Jakub, leg. Sotys-Lelek, 20.09.2011 (OPN); CHA Kopec, leg. Sotys-Lelek, 21.09.2011 (OPN); PR Mackov bok, leg. Sotys-Lelek, 24.09.2011 (OPN).

Discussion and summary

The investigated area is characterised by abundance of species from the genus Rosa L. More than a half of the species that were recorded in the whole region of the Low Tatras National Park occurred in the area which covered only about 20 ha. Roses with exceptionally varied morphological features, especially R. canina and R. agrestis (4 varieties were distinguished among each of them) occurred there. in case of R. agestis all varieties described by Popek (1996) for Central Europe were in the site. The varieties of R. agrestis found in the investigated region are presented in Fig. 2.

R. canina features the highest variety among all of the recorded roses. Several hundreds of species currently included into the complex of R.canina were described already at the beginning of the 20th century. Many of them have already been synonymised at the level of variety e.g.: R. corymbifera = R. canina

var. corymbifera, R. dumalis Baker = R.canina var. dumalis (Popek 1996). The varieties of R. canina which were found in the investigated region are not characterised by a single feature but a group of them (Fig. 3). Similar groups of features are applied to distinguish varieties within other species of roses. Within critical genus like Rosa, morphological and geographical diversity of particular species and their varieties can be described only by such comprehensive approach to the features.

Fig. 2. Morphological differentiation within the distinguished varieties of Rosa agrestis Savi; A part of long shoot; B, C, D: fruit; E: part of leaf (upperside); F, G, H: part of leaf (underside).Complexes of characteristics: R. agrestis var. agrestis (A, B, E, F), R. agrestis var. schulzei (A, C, D, E, F), R. agrestis var. albiflora (A, B, G, H), R. agrestis var. gizellae (A, C, D, G, H).

Fig. 3. Morphological differentiation within the distinguished varieties of Rosa canina L.; A part of long shoot; B, H: fruit, C - conical disc and styles of a spray type, D sepal; E, F, G: part of leaf (underside). Complexes of characteristics: R. canina var. dumalis (A, B, C, D, E), R. canina var. canina (A, B, C, D, G), R. canina var. corymbifera (A, B, C, D, F), R. canina var. andegavensis (A, C, G, H, E).


References

Benaov B. & Ujhzy K. 1998. Floristick kurz Zvolen 1997 (Zbornk vsledkov Floristickho kurzu konanho vo Zvolene 6.11.7.1997). Technick univerzita vo Zvolene.

Henker H. 2000. Rosa. in: Illustrierte Flora von Mitteleuropa. Band 4. Parey Buchverlag, Berlin.

Martincov E. 1989. Spis fondov Stredoslovenskho mzea. Botanika. Vyie rastliny. Bansk Bystrica.

Popek R. 1996. Biosystematyczne studia nad rodzajem Rosa L. w Polsce i krajach ociennych. Prace monograficzne 218. Wyd. Nauk. WSP, Krakw.

Popek R. 2002. Re dziko rosnce Polski. Klucz-Atlas. Plantpress, Krakw.

Popek R. 2007. Dziko rosnce re Europy. Officina Botanica, Krakw.

Prochzka F., Krahulec F. 1982. Kvtena okol Motenice v Nzkch Tatrch. Preslia, Praha, 54: 167184.

Tmk J. 1886. Adatok Zlyom megye szaknyugati rsznek flrjhoz. Besztercebnyai kath. Gymn. rt. 18851886: 1326.

Vtvika V. 1992. Rosa L. in: Flra Slovenska IV/3: 4289. Veda, Bratislava.

Zieliski J. 1985. Studia nad rodzajem Rosa L. systematyka sekcji Caninae DC. em Christ. Arbor. Krnickie 30: 3109.

Zieliski J. 1987. Rodzaj Rosa L. in: Flora Polski. T. 5: 74. PWN, Warszawa8.


THE ROLE OF MORPHO-ANATOMICAL TRAITS OF THE LEAVESIN THE TAXONOMY OF KALANCHOIDEAE BERG. SUBFAMILY (CRASSULACEAE DC.)

Myhajlo A. Chernetskyy

Abstract. The paper presents the results of the analysis of the morphological and anatomical structure of leaves of 35 species of the genus Kalancho Adans. (Crassulaceae DC.) in the taxonomic aspect of the subfamily Kalanchoideae Berg. Based on own studies and literature analyses of the flower morphology, embryology, karyology, vascular anatomy of stem and molecular genetics, the author has found that the most appropriate taxonomic system of the subfamily Kalanchoideae assumes existence of one genus Kalancho divided into three sections: Bryophyllum (Salisb.) Boit. & Mann., Eukalancho Boit. & Mann. and Kitchingia (Bak.) Boit. & Mann. Distinguishing three separate genera Bryophyllum Salisb., Kalancho Adans. and Kitchingia Bak., as has been the case throughout the history of the subfamily Kalanchoideae, is hardly possible due to existence of intermediate species.

Key words: morphology, anatomy, leaf, species, Kalancho, Kalanchoideae, Crassulaceae, taxonomy, nomenclature

Botanical Garden, Maria Curie-Skodowska University in Lublin, Sawinkowska str. 3, Lublin, 20-810, Poland;[email protected]

Introduction

The taxonomy and nomenclature of the subfamily Kalanchoideae Berg. (Crassulaceae DC.) have not been unquestionably established (Descoings 2006; Chernetskyy 2011). Two contrasting views have been prevailing throughout the history of research on the representatives of the subfamily. Some researchers distinguished separate genera Bryophyllum Salisb., Kalancho Adans. and Kitchingia Bak. (Salisbury 1805; Endlicher 1839; Bentham & Hooker 1865; Baker 1881; Baillon 1885; Schnland 1891; Stapf 1908; Berger 1930; Tillson 1940; Hutchinson & Dalziel 1954; Airy-Shaw 1966; Lauzak-Marchal 1974; Zepkova 1980; Wickens 1982; Forster 1985; Tlken 1985; t Hart 1995; Byalt 2000, 2008 and oth.), whereas others combined all species of the subfamily Kalanchoideae into one genus (Dalzell 1852; Hance 1873; Hamet 1907, 1908, 1963; Perrier de la Bthie 1923, 1928; Mauritzon 1933; Baldwin 1938; Decary 1962; Hamet & Marnier-Lapostolle 1964; Jensen 1968; Friedmann 1971, 1975; Raadts 1977; Rauh 1995; Gehrig et al. 2001; Mort et al. 2001 and oth.) divided into subgenera (Bryophyllum (Salisb.) Koorders, Calophygia Desc., Kalancho) (Descoings 2006) or sections (Bryophyllum (Salisb.) Boit. & Mann., Eukalancho (Kalancho) Boit. & Mann., Kitchingia (Bak.) Boit. & Mann. or Bryophyllum, Kalancho) (Boiteau & Mannoni 1948-1949; Friedmann 1975; Jacobsen 1981; Boiteau & Allorge-Boiteau 1995; Descoings 2003; Chernetskyy 2011).

The genus Kalancho includes approximately 150 species (Descoings 2006) distributed mainly in the arid areas of East and South-West Africa and some of its adjacent islands, and in South-East Asia. Almost half of all the Kalancho species described occur in the Madagascan flora as endemic species growing in the diverse climatic conditions of the island (Boiteau & Allorge-Boiteau 1995; Rauh 1995; Descoings 2003).

Numerous papers presenting the structure of organs of Kalancho genus plants focus primarily on the macro- and microstructure of flowers, stems or roots. Available reports on the structure of Kalancho leaves present mainly their macromorphology, including presence of adventitious buds (vivipary). in contrast, there is little information about the micromorphology and anatomy of Kalancho leaves. The available data involve only a few and more common species of the genus. Results of investigations on the leaf structure may provide knowledge of essential taxonomic traits that would be helpful in solving the current problem of the taxonomy and nomenclature of the subfamily Kalanchoideae.


The study involved 35 species representing various life forms, which, according to the systematic division (Jacobsen 1981), belong to three sections: 1) Bryophyllum (K. beauverdii Hamet, K. daigremontiana Hamet & Perr., K. fedtschenkoi Hamet & Perr., K. gastonis-bonnieri Hamet & Perr.,

M.A. Chernetskyy, 2012


K. laetivirens Desc., K. laxiflora Bak., K. manginii Hamet & Perr., K. marnieriana Jacobs., K. pinnata (Lam.) Pers., K. porphyrocalyx (Bak.) Baill., K.prolifera (Bowie ex Hook.) Hamet, K. rosei Hamet & Perr., K. tubiflora (Harv.) Hamet, K. uniflora (Stapf) Hamet); 2) Eukalancho (K. beharensis Drake, K. bipartita Chiov., K. blossfeldiana v. Poelln., K. crenata (Andrews) Haw., K. eriophylla Hilst. & Bojer ex Tulanse, K. hildebrandtii Baill., K. longiflora Schltr. ex Wood, K. marmorata Bak., K. millotii Hamet & Perr., K. nyikae Engl., K. orgyalis Bak., K.prittwitzii Engl., K. pumila Bak., K. rhombopilosa Mann. & Boit., K. schimperiana A. Rich., K.thyrsiflora Harv., K. tomentosa Bak., K. velutina Welw. ex Britten); 3) Kitchingia (K. campanulata (Bak.) Baill., K.gracilipes (Bak.) Baill., K. peltata (Bak.) Baill.).

The study material was obtained from the collection of greenhouse plants at the Botanical Garden of the Maria Curie-Skodowska University in Lublin. investigations of the micromorphology and anatomy of typical leaves of selected Kalancho species were conducted using stereoscopic microscopy (SM), light microscopy (LM) and scanning electron microscopy (SEM).

Results and discussion

The leaves of individual species of the genus Kalancho differ distinctly in terms of their shape, colour and parameters. They exhibit permanent species-specific macromorphological traits. The polymorphism observed in the leaves (simple or compound, sessile or petiolate) is characteristic for some species and closely related to changes in external factors in both artificial growing conditions and their homeland. A common feature for all taxa is the succulent leaf structure, which is unequally manifested due to the diverse habitat conditions. Leaf vivipary is a typical trait of most representatives of the section Bryophyllum, while infloral vivipary is a general feature of the genus Kalancho.

in terms of their morpho-anatomical organization, leaves of the genus Kalancho belong mostly to the bifacial structure type. However, the anatomical structure in many species reveals the unifacial petiole structure; hence, the leaf may be classified as the mixed (unifacial-bifacial) type. The representatives of the particular sections of the genus Kalancho share the leaf anatomy (Tab. 1), but the species differ distinctly in terms of appearance (the type of trichomes, the contours of the anticlinal walls

of epidermal cells, etc.), parameters or numbers, for instance, the length of the stomata (23-39 m) and non-glandular (42-1258 m) or glandular trichomes (28-155 m); the thickness of the epidermis (11-48 m); the number of epidermal cells (159-1075), stomata (17-97) and non-glandular trichomes (19-52) per 1 mm2 of the leaf surface. The epidermis in Kalancho leaves produces papillary structures composed of highly convex epidermal cells and one or more stomata. The structure of some Kalancho taxa is characterized by presence of surface calcium carbonate deposits (K. gastonis-bonnieri, K.pumila, K. thyrsiflora), papillary marginal cells (K.prolifera), non-glandular trichomes (K.beharensis, K.orgyalis, K. eriophylla, K. hildebrandtii, K. rhombopilosa, K. millotii, K. tomentosa and oth.), glandular trichomes (K. beharensis, K. gastonis-bonnieri, K. eriophylla, K. manginii, K. orgyalis and oth.), angular collenchyma (K. campanulata, K. daigremontiana, K. laxiflora, K. marnieriana, K. pinnata, K. prolifera and oth.), tangential collenchyma (K. beharensis, K. blossfeldiana and oth.) and stomata in the elongated aggregations on the petiole (K. pinnata, K. prolifera). The chlorenchymatous tissue in Kalancho leaves is not differentiated into palisade and spongy mesophyll, but is divided into small-celled subepidermal mesophyll and large-celled mesophyll (water transporting, CAM type). The leaf assimilation tissue contains pleomorphic chloroplasts. Green plastids have been detected also in the cytoplasm of cells: epiderma, stomata, collenchyma, tannin-bearing and epithem hydathodes.

The details of the microstructure of Kalancho leaves may constitute important taxonomic traits of the species of the subfamily Kalanchoideae. The leaf srtructure did not exhibit distinct differences between the species of the three sections (Bryophyllum, Eukalancho, Kitchingia) of the genus Kalancho. Based on own studies (Chernetskyy 2007) and other authors investigations concerning the flower morphology (Hamet 1907, 1908; Berger 1930; Boiteau & Mannoni 1948-1949 and oth.), embryology (Mauritzon 1933), karyology (Baldwin 1938; Uhl, 1948; Friedmann 1971; Zepkova 1976; Raadts 1983, 1985, 1989), vascular anatomy of stem (Jensen 1968) and molecular genetics (Ham & t Hart 1998; Gehrig et al. 2001; Mort et al. 2001), it is difficult to distinguish three genera in the subfamily Kalanchoideae due to existence of intermediate species. Descoings (2006) suggests a division of the genus Kalancho

17

Traits of leaf structureSections

Bryophyllum Eukalancho Kitchingia

polymorphism + + +

unifacial petiole + + +

peltate lamina + + +

viviparia +

non-glandular trichomes +

glandular trichomes + +

calcium carbonate deposits on the surface + +

epithemal hydathodes + + + +

papillary structures of the epidermis + + + + + +

amphistomatic + + + + + +

lower epidermis producing more stomata + + + + +

anisocytic stomata + + + + + +

well-developed outer cuticular ledges of stomatal cells + + + + + +

epicuticular wax + + + + + +

thick-layered cuticle + + + + + +

striated cuticle + + +

smooth cuticle + + +

single-layer epidermis + + + + + +

papillary epidermal cells +

slightly or distinctly convex outer epidermal cell walls + + + +

thickened outer walls of epidermal cells + + + + + +

the contours of the anticlinal walls in epidermal cells:

undulating + + +

straight + + +

the cells of the lower epidermis are smaller than the cells of the upper epidermis + + + + +

anthocyanin pigments in epidermal cells + + + + + +

small-celled, subepidermal mesophyll + + + + + +

water transporting mesophyll + + + + + +

tannins in some parenchymal cells + + + + + +

calcium oxalate crystals in parenchymal cells + + + + + +

lack of the trait in the species of the section, + + characteristic trait for the species of the section,+ characteristic trait for most species of the section, + characteristic trait for some species of the section.

Table 1. Morpho-anatomical traits of the leaf structure in the genus Kalancho.

into three subgenera: Bryophyllum, Kalancho and Calophygia. The author classifies intermediate species between Kalancho and Bryophyllum into the subgenus Calophygia, excluding species that have common traits of Bryophyllum and Kitchingia. He introduces a new taxon the subgenus Calophygia to the subfamily Kalanchoideae, and disregards the genus Bryokalancho Res. proposed previously by F. Resende in 1956 (see Boiteau & Allorge-Boiteau 1995). Generally, this gives no grounds for distinguishing three separate genera Bryophyllum, Kalancho and Kitchingia in the subfamily

Kalanchoideae, as has been done throughout the history of this systematic group.

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Rauh W. 1995. Succulent and xerophytic plants of Madagascar. Vol. 1. Strawberry Press, Mill Valley (US: CA).

Salisbury R.A. 1805. The paradisus Londinensis. Vol. 1, T. 3. D.N. Shury & W.W.Hooker, London.

Schnland S. 1891. Crassulaceae. in: Engler A. & Prantl K. (eds). Die Natrlichen Pflanzenfamilien. 1st ed. 3 (2a): 23-38. Verlag von Wilhelm Engelmann, Leipzig.

Stapf O. 1908. Kitchingia uniflora Stapf. Bull. Misc. Inform. Kew: 258259.

t Hart H. 1995. infrafamilial and generic classification of the Crassulaceae. in: t Hart H. & Eggi U. (eds). Evolution and Systematics of the Crassulaceae: 159172. Backhuys Publishers, Leiden.

Tillson A.H. 1940. The floral anatomy of the Kalanchoideae. Amer. J. Bot. 27: 596600.

Tlken H.R. 1985. Crassulaceae. in: Leistner O.A. (ed.). Flora of Southern Africa. Vol. 14:1244. Botanical Research institute, Department of Agriculture & Water Supply, Pretoria (RSA).

Uhl C.H. 1948. Cytotaxonomic studies in the subfamilies Crassuloideae, Kalanchoideae and Cotyledonoideae of the Crassulaceae. Amer. J. Bot. 35: 695706.

Wickens G. 1982. Miscellaneous notes on Crassula, Bryophyllum and Kalancho. Kew Bull. 36: 665674.

Zepkova N.L. 1976. Division main numbers of chromosomes and polyploidy of rows in the family Crassulaceae. Voprosy Sist. Pokrytosem. Rast.: 7781. (in Russian).

Zepkova N.L. 1980. Comparison of proposing a system the family Crassulaceae of system A. Bergers. Voprosy Bot.: 138-145. (in Russian).

19

Introduction

Festuca trachyphylla (Hack.) Krajina belongs to the group F. ovina agg. (Hackel 1882; Markgraf-Dannenberg 1980; Pawlus 1983-1985; Wilkinson & Stace 1991; Dengler 1996, 1998). its geographical range is limited to the temperate climate zone in the northern hemisphere and cool subtropical regions. it occurs in diverse communities: insolated hill slopes, steep slopes of calcareous or loess valleys, dry meadows and mid-field sand dunes, as well as scrub and well-lighted forest (Fijakowski & Warmiska 1972; Medwecka-Korna & Korna 1977; Falkowski 1982; uszczyska 2001). it is a co-dominant or dominant component of xerothermic grasslands from the associations Seslerio-Festucion duriusculae and Cirsio-Brachypodion pinnati (Matuszkiewicz 2007). it often occurs in anthropogenic habitats. F. trachyphylla is a morphologically variable species of the group F. ovina agg. and it thus poses problems to systematicians. its variability is manifested mainly in the morphological and anatomical structure of the leaf blade of sterile shoots (Pawlus 1983-1985; Conert 1996; Szczniak 2005).

The aim of the paper was to identify the variability of morphological and anatomical traits in F. trachyphylla leaves in the southeastern part of Poland.


The plant material included samples of F. trachyphylla (Hack.) Krajina from 24 sub-populations (432 individuals) collected from different habitats in the southeastern part of Poland (Tab. 1). Thirteen of the sub-populations originated from calcium carbonate-rich xerothermic grassland habitats, while the others grew on sand dunes. The studies involved thirteen traits: 9 quantitative and 4 qualitative (Tab. 2 and 3), and consisted mainly of biometric measurements of the traits that were variable in this species. Some observations of traits, for example the length of leaf and cauline leaf or leaf colour, were carried out directly on living plant specimens in their natural habitats. The other characters that demanded precise observation of leaf elements were counted or measured with a ruler or an Opta-Tech stereoscopic zoom microscope. The anatomical analysis was performed with the use of a Nikon light microscope. Differences among the means of the characters were tested using a one-way ANOVA version 7.1 of the STATiSTiCA programme (StatSoft inc. 2007).

Results and discussion

Quantitative charactersThe results of the ANOVA analysis showed a

significant variability (p


Habitat Origion AbbreviationGeographical factors

latitude (N) longitude (E)

xerothermic grassland Opoka Dua T-2 5052 42 2256 62

Lublin T-24 5116 52 2238 93

Rudnik T-25 5117 24 2236 13

Ciechanki T-27 5117 36 2252 22

iowiec T-30 5050 05 232448

Kazimierz Dolny T-41 5119 53 2155 50

Bychawa T-43 5101 21 2231 37

Naczw T-49 5117 26 2214 53

Czumw T-50 5047 49 2358 18

Niedzieliska T-51 5042 56 2304 33

Tarnogra T-65 5053 47 2307 21

Staw T-96 5112 21 2324 50

Parchatka T-131 5122 56 2149 96

sand dunes Chotyw T-16 5200 46 2323 49

Zwierzyniec T-23 5036 19 2259 56

Aleksandrw T-26 5154 28 2229 09

Konstantynw T-28 5112 43 2305 43

Puawy T-29 5128 42 2155 58

Orchwek T-31 5132 16 2336 38

Maaszewicze T-32 5101 56 2332 34

Sobibr T-44 5128 46 2339 46

Krnica Jara T-48 5109 49 2228 25

Potok Wielki T-61 5047 20 2213 72

Stoczek T-141 5138 45 2242 53

Table 1. Characteristics of the Festuca trachyphylla habitats investigated.

Table 2. The range for the nine quantitative characters and the results of ANOVA. (Differences traits were considered significant at the level of p

21

The highest variability was found for the length of the leaf, width of the cauline leaf, number of ribs in the leaf, length of cauline leaf, and length of hairs in the leaf. The other quantitative characters were permanent in this species and did not exhibit variability between the different types of habitats. The results revealed that the plants from xerothermic grasslands differed from the plants from sand dunes in terms of the following combination of characters: shorter leaf, narrower and shorter cauline leaf, and the leaf, which tends to have fewer ribs and longer hair on the upper surface of the leaf blade (Tab. 1).

Qualitative charactersAll individuals of the F. trachyphylla species were

studied for leaf colour, hairiness of the upper part leaf, cross-section of the leaf and distribution of the sclerenchymatous tissue in the leaf blade (Tab. 2). Plants growing in shallow and dry soil, full sun conditions, had filiform, blue-green, leaf blades with dense hairs on the lower part. in cross-section, the leaf blades were ovoid-shaped. Plants growing in sand dunes were characterized by green leaves, which had delicate pubescence on the lower part, or were hairless. in cross-section, the leaf blades were

wedge-shaped with convex sides and a rounded tip. Additionally, the plants growing on the sand habitats exhibited two or three rows of sclerenchymatous cell layer, clustered in three corner rows; in the plants from the calcareous soil in the xerothermic habitats, the single-row layer formed a separate ring. The cross-section diversity is presented in Fig. 1.

There are some morphological differences between plants of different habitats within F. trachyphylla; these differences, however, are too inconsistent and imprecise to allow recognition of two intraspecific taxa within F. trachyphylla. it could be expected that under such circumstances, a significant degree of morphological diversification would have occurred, making it possible to visually distinguish plants of different habitats. The results obtained in the present study are partly consistent with investigation data reported by other authors (Markgraf-Dannenberd 1980; Falkowski 1982; Szafer et al. 1986; Conert 1996) who found that the morphological and anatomical traits of the leaf blade in this species were considerably unstable and modified by the environmental effect.

Dbrowska A. Morpho-anatomical structure of the Festuca trachyphylla leaves

Descriptions of character Abbreviation of character[attributes]

Frequency of F. trachyphylla(N=432)

leaf colour [green/blue-green] 185/247

leaf: hairiness, lower part [glabrous/hairy] 28/404

cross-section of leaf [wedge-shape/ovoid-shape] 210/222

sclerenchyma [1-layer /2-3-layer] 193/239

Table 3. Descriptions, abbreviations and attributes of qualitative characters and their frequencies in Festuca trachyphylla.

A B C DFig. 1. Cross-sections of Festuca trachyphylla from different habitats:A-B xerothermic grasslands, C-D sand dunes. Scale bar=1 mm.


References

Conert H.J. 1996. Festuca. in: Hegi G. (ed.) Illustrierte Flora von Mitteleuropa 1(3): 561633. Paul Parey, Berlin-Hamburg.

Dengler J. 1996. Anmerkungen zur Taxonomie und Bestimmung von Schaf-Schwingeln i. w. S. (Festuca ovina agg.) in Deutschland mit besonderer Bercksichtigung Schleswig-Holsteins. Kieler Notizen zur Pflanzenkunde in Schleswig-Holstein und Hamburg 24: 129.

Dengler J. 1998. Neues von den schmalblttrigen Schwingel-Sippen (Festuca ovina agg. und F. rubra agg.) in Deutschland mit besonderer Bercksichtigung von Schleswig-Holstein und Hamburg. Kieler Notizen zur Pflanzenkunde in Schleswig-Holstein und Hamburg 25/26: 632.

Falkowski M. 1982. Trawy polskie. PWRiL, Warszawa.Fijakowski D. & Warmiska B. 1972. Zmienno kostrzew

kpowych (Festuca sp.) wojewdztwa lubelskiego. Ann. Univ. Mariae Curie-Skodowska, sec. C. 27(18): 189198.

Hackel E. 1882. Monographia Festucarum Europaearum. Kassel, Berlin.

uszczyska B. 2001. Distribution of Festuca ovina group (Poaceae) species in the xerothermic communities of the Piczw Hump and adjacent areas (southern Poland). in: Frey L. (ed.) Studies on grasses in Poland: 201209. W. Szafer institute of Botany, Polish Academy of Sciences, Krakw.

Markgraf-Dannenberg I. 1980. Festuca L. in: Tutin T.G., Heywood V.H., Burges N.A., Moore D.M., Valentine D.H., Walters S.M. & Webb D.A. (eds). Flora Europaea 5: 125153. Cambridge University Press, Cambridge.

Matuszkiewicz W. 2007. Przewodnik do oznaczania zbiorowisk rolinnych Polski. Wydawnictwo Naukowe PWN, Warszawa.

Medwecka-Korna A. & Korna J. 1977. Zespoy stepw i suchych muraw. in: Szafer W. & Zarzycki K. (eds). Szata rolinna Polski: 352366. PWN, Warszawa.

Pawlus M. 1983 (1985). Systematyka i rozmieszczenie gatunkw grupy Festuca ovina L. w Polsce. Taxonomy and distribution of the Festuca ovina group in Poland. Fragm. Flor. Geobot. 29(2): 219295.

Szafer W., Kulczyski S. & Pawowski B. 1986. Roliny polskie. Opisy i klucze do oznaczania wszystkich gatunkw rolin naczyniowych rosncych w Polsce bd dziko, bd te zdziczaych lub czciej hodowlanych. PWN, Warszawa.

Szczniak E. 2005. Species of Festuca ovina group (Poaceae) on the serpentine rocks in the Sudety Foreland. Acta Bot. Siles. 2: 121129.

Wilkinson M.J. & Stace C.A. 1991. A new taxonomic treatment of Festuca ovina L. aggregate (Poaceae) in the British isles. Bot. J. Linn. Soc. 106: 347397.


MICROMORPHOLOGICAL EVIDENCE FOR ANDROECIUM ORIGINOF Claytonia (MONTIACEAE) PETALOIDS

Patrcia dos Santos 1*,2, Sam Brockington 2*, Beverley Glover 2** & Louis P. Ronse de Craene 1**

Abstract. Caryophyllales is an order distinguished for having flowers with only one perianth whorl the perigone. The perigone is a calyx derived structure that can have either petaloid or sepaloid appearance. Members of the Portulacinae suborder have tendency to have a false bipartite perianth, forming a petaloid perigone and an epicalyx with the subtending bracts of the flower. Although Claytonia belongs to the Portulacinae suborder, previous studies have suggested a different origin for its petaloid organs other than the sepals. in this study we investigated the floral development of Claytonia sibirica and Claytonia perfoliata using Scanning Electron Microscopy (SEM) to understand the origin of the petaloid organs in the genus. Our results show that petaloid organs in Claytonia are of androecium origin and can be interpreted as the expression of the typical Caryophyllales perigone growing in androecium tissue.

Key words: Claytonia, Caryophyllales, Portulacinae, Floral development, perigone, perianth

1 Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland, UK;1* [email protected]; 1** [email protected] Department of Plant Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EA, UK;2* [email protected]; 2** [email protected]

Introduction

Caryophyllales have a fascinating evolutionary history, which is deeply related with the evolution of the perianth of their flowers. The perianth is usually formed by two series or whorls, a protective whorl formed by sepals (the calyx) and a pollinator-attractive whorl formed by petals (the corolla). in Caryophyllales the perianth is formed by one single whorl (the perigone) due to the loss of the corolla, which is thought to be related with a wind-pollinated ancestor (Brockington et al., 2009). Even though the perigone might have sepaloid or petaloid appearance it is of calyx origin, having a typical 2/5 initiation pattern and imbricate aestivation (Ronse de Craene 2008). Another important feature found in Caryophyllales that might be associated with the loss and reinvention of petaloid organs is the centrifugal stamen initiation (Brockington et al. 2009; Ronse de Craene 2008).

The typical flower of the Portulacinae suborder has a pentamerous petaloid calyx (Nyffeler & Eggli 2010). As the sepals have gained petal function, the protection role was taken by the two bracteoles acting as an epicalyx. Even though this has been largely assumed, previous studies on Claytonia L. (Montiaceae) have suggested that petaloids might have a different origin from the traditionally assumed for Portulacinae. The anatomical studies of Milby (1980) suggest that petaloids and stamens are

originated from the same primordium, suggesting an androecium origin for the petaloids in Claytonia. As the typical Portulacinae, flowers of Claytonia also have five petal-like organs and two bracteoles (Nyffeler & Eggli 2010), however the petaloids in Claytonia are fused at the base with the stamens, and both organs appear to share the same vascular bundle (Milby 1980).

in this study a micromorphological analysis was conducted using Scanning Electron Microscopy (SEM) to investigate the origin, development and evolution of the petaloids of Claytonia. This study is part of an MSc dissertation.


Several floral buds in different stages of development of Claytonia sibirica L. and Claytonia perfoliata Donn ex Willd. were collected and fixed in FAA (90% ethanol; 5% formaldehyde; 5% acetic acid). Samples were previously stored in 70% ethanol and analyzed under a ZEiSS Stemi SV6 Dissecting Microscope. The material was critical point dried, mounted on SEM pin stubs and coated with platinum. The specimens were observed and photographed using a LEO Supra 55VP Scanning Electron Microscope. SEM images were processed using Adobe Photoshop CS2.

P. dos Santos, S. Brockington, B. Glover, L.P. Ronse de Craene, 2012


Results

The inflorescence type in Claytonia is a monochasium each axillary bud originates lateral branches in alternation that give rise to other younger flower buds (Fig 1 A-B). From each undifferentiated flower bud, two opposite bracts are formed; the first one arises at the apex and the second one at the base of the flower (Fig 1 A-C). An invariable number of 5 stamens initiate more or less simultaneously (Fig 1 C-E). Only after this stage, the initiation of the petaloids is visible, arising from the base of each stamen (Fig 1 E-G). Petaloids grow simultaneously and centrifugally from the base of each stamen apparently in an imbricate aestivation (Fig 1 H-i). in the mature flowers it is possible to see the 5 stamens opposite to the 5 petaloids, all fused at the base forming a ring surrounding the gynaecium (Fig 1 i). The analysis of the developmental stages suggests a congenital fusion of stamen petaloid and a postgenital fusion of the androecium, originating the ring that surrounds the gynoecium. The gynoecium is usually 3-loculate (rarely 2 or 4; Fig 1 i) and differentiate more or less simultaneously with the androecium (Fig 1 C-E).

Discussion

in this study we suggest two hypotheses for the origin of petaloids in Claytonia. There is evidence for an androecium origin of the petaloids, although the differentiation of these organs can also be interpreted as sepal-derived petaloids.

The timing and pattern of initiation of the perianth and the petaloids position relatively to the stamens are evidence for petaloids with androecium origin. The petaloid initiation is much delayed, always arising after the stamens initiation and development. The results from the SEM analysis suggest that petaloids arise as part of the androecium from the base of the filaments, going in accordance with Milbys (1980) results. The position of the petaloids relatively to the stamens also suggest that the perianth is not a second whorl of staminodes as they are not alternating with the stamens, arising opposite to the stamens. This is also evidence for the theory that petaloids in Claytonia are outgrowths of the stamen filaments, and thus part of the androecium. This point of view assumes that the perianth is totally lost in this group and new organs are formed previously from the filaments.

The theory of sepal-derived petaloids relies on the delay of calyx initiation, which in an extreme situation could lead to the incorporation of calyx parts in the androecium, maybe due to lack of space as the androecium initiation is very fast and occurs centrifugally. A similar situation has been recorded for Portulaca L., where there is also a delayed growth of perianth parts (dos Santos 2011 unpublished MSc thesis), showing that this situation is likely to happen also in Claytonia. Another evidence for this theory is the quincuncial aestivation pattern found in Claytonia. Although there is no evidence of a typical 2/5 calyx initiation due to the simultaneous arising of the petaloids, these show a quincuncial aestivation in maturity. This suggests that the calyx (or the perigone) is being expressed in androecium tissue due to a delay of growth of the petaloids. This can lead to a misinterpretation of the identity of perianth parts, appearing as stamen appendages.

Further evidence could be given by arguing that, for example, in Montia fontana L., a species belonging to a sister genus of Claytonia, there is sometimes a reduction in the number of stamens (3 instead of 5) but not in the number of petaloids (Hofmann 1993), however rudimentary stamens in the single petaloids were not reported to be found. This might give evidence for the theory that the petaloids are not part of the androecium in Claytonia, although further investigation of these species should be carried out. Hofmann (1993) described these organs as petals (stapetals) with a delayed growth, assuming also the total loss of sepals in this genus but not giving an explanation for the emergence of petals, lacking in the rest of the group. A new gain of petals in Claytonia, as assumed by Hofmann (1993) is very unlikely and these are rather other misinterpreted organs of the flower that should be worth of a detailed morphological study in future.

The delay of calyx growth and subsequent incorporation in the androecium tissue seems to be the most suitable interpretation of our results; however molecular studies are essential to support this hypothesis.

Acknowledgements

This study was financially supported by the National Environment Research Council (NERC).

Many thanks to an anonymous reviewer for the helpful comments.

25

References

Brockington S.F., Alexandre R., Ramdial J., Moore M.J., Crawley S., Dhingra A., Hilu K., Soltis D.E., Soltis P.S. 2009. Phylogeny of the Caryophyllales sensu lato: revisiting hypotheses on pollination biology and perianth differentiation in the Core Caryophyllales. Int. J. Pl. Sci. 170 (5): 627643.

Dos Santos P. 2011. The Cryptic Evolution of Petaloids in the Portulacineae (Caryophyllales). MSc thesis unpublished data. University of Edinburgh, Royal Botanic Gardens Edinburgh & the University of Cambridge.

Endress P.K. 1996. Diversity and Evolutionary Biology of Tropical Flowers. Cambridge University Press.

Hofmann U. 1993. Flower morphology and ontogeny. in: Behnke H.-D. & Mabry T.J. (eds). Caryophyllales Evolution and Systematics: 123166. Springer-Verlag Berlin-Heidelberg.

Milby T.H. 1980. Studies in the floral anatomy of Claytonia (Portulacaceae). Am. J. Bot. 6 (7): 10461050.

Nyffeler R., Eggli U. 2010. Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59 (1): 227240.

Ronse de Craene L. 2008. Homology and Evolution of Petals in the Core Eudicots. Systematic Botany 33 (2): 301325.

Fig. 1. Claytonia sibirica (A-F and i) and Claytonia perfoliata (G-H) SEM figure plate. A-B inflorescence; C-D stamen and carpel early development; E-G petaloid initiation from the base of the stamens; H-i growth of petaloids and full developed flower. (White bar Ai = 100m). Br = bracteoles; = petaloids; = stamens and stamen primordia; G = gynoecium and gynoecium primordium.

dos Santos P., Brockington S., Glover B. & Ronse de Craene L.P.


VASCULAR ANATOMY AND MORPHOLOGY OF THE FLOWERIN FRitillaRia montana HOPPE (LILIACEAE)

Andrew V. Novikoff 1 & Maria A. Kazemirska 2

Abstract. in the first time structural type of gynoecium and vascular system organization in Fritillaria montana Hoppe flower were described in details. We ascertain that gynoecium is divided in three vertical zones viz synascidiate, symplicate and apocarpous. Therefore the gynoecium was described as syncarpous in wide sense and typified as Fritillaria-type. in the other hand, for the vascular system of the flower six main groups of bundles were described, e.g. recurrent bundles in central column of ovary were observed. it was ascertained that ovules supply by the complex of ventral and dorsal as soon as recurrent bundles. A little difference in the innervation of inner and outer tepals was ascertained too.

Key words: Fritillaria montana, flower, gynoecium, morphology, vascular anatomy

1 State Natural History Museum NAS Ukraine, Department of Evolution and Biosystematics, Teatralna Str., 18, Lviv, 79008, Ukraine;[email protected] Yuriy Fedkovich Chernivtsi National University, Faculty of Biology, Ecology and Biotechnology, Department of Botany and Nature Protection, Fedkovycha Str., 11, Chernivtsi, 58022, Ukraine; [email protected]

to W. Leinfellner & H. Baum

Introduction

Fritillaria montana Hoppe is an endangered species listed in the third edition of the Red Book of Ukraine (Chorney et al. 2009) and Convention on the Conservation of European Wildlife and Natural Habitats (Bern Convention 2002).

it is south-european-Balcanian species found in disjunctive areas and its general natural range covers Central and Southern Europe (italy, France, Bulgaria), South Hungary, Austria and Balcanian penninsula. Also it can be found in Moldova and Romania. in Ukraine F. montana occurs in the northeastern limit of its natural range, and is reported in Khmelnytska, Chernivtsi and Odessa regions.

At the present time, in Chernivtsi Region there are 9 habitats (sites) of F. montana, which can be found mainly in communities of Querco-Fagetea Br.-Bl. et Vlieg. in Vlieg. 1937, Molinio-Arrhenatheretea Tx. 1937 and Trifolio-Geranietea sanguinei Th. Mull. 1962 classes (Kazemirska & Chorney 2010a, 2010b; Kazemirska 2011).

Previous investigations on Fritillaria mainly were attached to the morphology and phenetic studies (Zaharof 1988; Mohammadi-Dehcheshmeh et al. 2008; Bartolucci et al. 2009) and only particularly to the anatomy of vegetative organs (Corneanu & Popescu 1981; Alan 2008). There no profound investigations on the floral anatomy or gynoecium

morphology of Fritillaria because the most important taxonomical features in the flower organization in this genus are: perigonium colour, flower and tepal shape, shape of the nectaries (Rix 1974, 1975, 1978, 1980; Bartolucci et al. 2009). Although the studies on gynoecium vertical zonality and organization of floral vascular system are not less interesting features for comparative morphological and taxonomical research.


Flowers of Fritillaria montana were collected by M. Kazemirska at the different stages of ontogenesis during 2011 in: 1) Botanical garden of Yuriy Fedkovich Chernivtsi National University; 2) environs of Kaplivka village, Chernivetsi region; 3) environs of Podviryivka village, Chernivetsi region; 4) environs of Zelene village, Chernivetsi region. Flowers were fixed in 70% alcohol, dehydrated with chloroform and then embedded in paraffin mixture (Gerlach 1984). After that flowers have been cut into 15 m cross-sections by rotary microtome MS-2 (USSR), stained by 1% safranin and 0,5% methylene blue and embedded in Canadian Balm. Figures of cross-sections were drawn by the using of Carl Zeiss Q1 microscope and camera lucida Lomo RA-4 (USSR).

The total ovary height (TOH) was calculated as a distance from the ovary loculi bottom up to the base

A.V. Novikoff, M.A. Kazemirska, 2012


of the style (Fig. 1 E). F. montana has no strict border between the ovary and style, therefore the base of the style we identify by the reduction of mushroom-shaped walls above the level of placentas and by the vanishing of ventral median bundles of carpels. The

length of style and stigma had no calculated because of high variability and intensive elongation during the flower ontogenesis.

Fig. 1. Fritillaria montana Hoppe: A common view of the flower, B outer (left) and inner (right) tepals, C stamens, D flower with detached perigonium, E scheme of longitudinal section of the flower. Gray color congenital fusion, yellow ovules and anthers. Aps apocarpous sterile, Sps symplicate sterile, Spf symplicate fertile, Saf synascidiate fertile and Sas synascidiate sterile zones. TOH total ovary height.

29

Results

Common characteristic of flowerF. montana has hypogynous actinomorphic

flowers with campanulate perigonium divided into 3 outer and 3 inner tepals (Fig. 1). Mature flowers about 2,5-3 cm of length, and about 2-2,7 cm in diameter. Outer tepals more developed, they have massive bases and axipetal tips, while inner tepals have less developed bases and axifugal tips. All the tepals have nectaries in their adaxial surface (Fig.1B). Perigonium purplish-brown, at the base of flower it forms the short hypanthium. Gynoecium is trimerous, represented by 3 fused carpels which are situated on the radii of outer tepals, without clear borders between carpels. Style elongated, rounded-triangular on the cross sections. Stigma consists of 3 separated narrow upward directed lobes (Fig.1 D). The total ovary length varies in different flowers and considerably increases with maturity. in immature flowers ovary length is about 0,3-0,5 cm. Androecium consist of 6 stamens (Fig. 1 C). Stamens basifixed, epipetalous at the base, they have linear filaments and developed anthers, their connectives are not elongated. Stamen length is about 1,5 cm.

ovary structureTepals and stamens are accreted together in short

hypanthium, which fuses with receptacle below the ovary. Above the level of hypanthium fusion, after short receptacle stretch, the ovary forms 3 small congenital isolated loculi on the radii of outer tepals level of separated sterile loculi (Fig. 2 D). in the same time, outer tepals begin the detaching of their margins from hypanthium wall (Fig. 1 E; Fig. 2 C-F).

in the next each loculus become fertile and contains two rows of axile placentas (Fig. 2 E). Ovules are anatropous, with short funiculi. The part of ovary with 3 separated fertile loculi make about 40,1% from TOH. in immature flowers share of this part is about 33,5% and it increases with

maturity up to 46,8%. As well as in the middle of the level of separated fertile loculi all the stamens become detaching from hypanthium wall (Fig. 2 F). Therefore, in the top of this level hypanthium break up into 6 (3+3) separated

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