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Roczniki Akademii Rolniczej w Poznaniu CCCLXXXVIII www.up.poznan.pl/steciana ISSN ͻ-ͻͺ Botanika – Steciana ͻͽ, ͼͺͺ, ͻ-ͻͺ INTRODUCTION The genus Linaria Mill. comprises ͻͿͺ species dis- tributed throughout the northern hemisphere. Its main centre of distribution is located in the Mediterranean basin. Linaria together with few other genera tradi- tionally placed in family Scrophulariaceae was recently added to the expanded Plantaginaceae (APG II ͼͺͺͽ, ALBACH et AL. ͼͺͺ;). The seed characters have proved to be useful in many morphological studies of the Linaria. For instance SUTTON ( ͻ) divided the genus into ve sections (Li- naria, Supinae, Pelisseriannae, Versicolores and Diusae ) on the base of seed morphology. The SEM studies of seeds were carried out in the identication of the most species of Linaria from south-western Iberian Penisula (e.g. JUAN et AL. ͻ , SEGARRA and MATEU ͼͺͺͻ a, b, EZ and CRESPO ͼͺͺͿ). An interesting contribution to the taxonomy of Plantaginaceae sensu lato based on ul- trastructure of seed coat was recently given by SHEHATA and LOUTFY (ͼͺͺ). VIANO ( ͻ) indicated the potential signi cance of pollen morphology in the systematics of the genus Linaria. Author ascertained important range of varia- tions between the grains of ͽͽ species, originated from the dierent regions of the Mediterranean basin. Pol- len grains of some other species of the genus were also described in the works of OLSON ( ͻ;), ELISENS ( ͻ) and HALBRITTER (ͼͺͺͺ). From among Ϳͽ species of Linaria growing in Eu- rope almost ͺ% are endemic to the continent. Lina- ria loeselii is an endemic species of the Baltic Sea coast which occurrence is restricted to its south-eastern part (PANKHURST ͼͺͺ). In Poland it grows only on coastal dunes between the village Unieście in the west and Vistula Bar in the east. At present the most numerous populations exist in the Słowiński National Park (PIO- TROWSKA et AL. ͻ , PRZYRODA... ͻ). The species is listed in Bern Convention on the Conservation of Euro- pean Widelife and Natural Habitats and it is protected by law in Poland. Linaria loeselii is also considered to be vulnerable within Polish vascular ora (KAŹMIERCZA- KOWA and ZARZYCKI ͼͺͺͻ, FREY ͼͺͺ;, ZARZYCKI and SZEL ĄG ͼͺͺ). Linaria loeselii is sometimes treated as subspecies of L. odora [L. odora (M. Bieb.) Fisch. subsp. loeselii (Schweigg.) Hartl] or even, both mentioned names of species are used as synonyms (STASIAK ͻ, KAŹMIER- CZAKOWA and ZARZYCKI ͼͺͺͻ, MIREK et AL. ͼͺͺͼ, PAN- KHURST ͼͺͺ). The aim of the presented work was to investigate the pollen morphology and the ultrastructural traits of seed-coat of this interesting, rare species. MATERIAL AND METHODS In ͼͺͺ the owers and the seeds of L. loeselii were collected from one rich population on the Łeba Sand- bar in the Słowiński National Park (E ͼͺ°ͻ’, N ;°ͼ’). The sample of pollen grains (ripe anthers were taken only) was acetolysed according to ERDTMANS method ( ͻͿͼ), with some insignicant modications (TOMLIK- -WYREMBLEWSKA ͻͿ). The sample of ͽͺ well devel- oped and ripe seeds was not prepared, but only cleaned. The using of scanning electron microscopes (JEOL JSM MORPHOLOGY OF POLLEN GRAINS AND SEEDS OF THE BALTIC SEA COAST ENDEMIC SPECIES LINARIA LOESELII SCHWEIGG. LESZEK BEDNORZ, IRMINA MACIEJEWSKA-RUTKOWSKA L. Bednorz, Department of Botany, Poznań University of Life Sciences, Wojska Polskiego ΅Ϳ C, ΄;- ΄ Poznań, Poland, e-mail: [email protected] I. Maciejewska-Rutkowska, Department of Natural Forestry Foundation, Poznań University of Life Sciences, Wojska Polskiego ΅Ϳ D, ΄;- ΄ Poznań, Poland (Received: February ͼͿ, ͼͺͺ. Accepted: July ͼͿ, ͼͺͺ) ABSTRACT. Linaria loeselii Schweigg. is an endemic species of the Baltic Sea coast which occurrence is restricted to its south-eastern part. The aim of the presented work was to investigate its pollen morphol- ogy as well as ultrastructural traits of its seed-coat. Pollen grains of this species are small, spheroidal or prolate spheroidal, ͽ-zono-colporate, with microreticulate to foveolate ornamentation. The seeds are winged, concolourous, black to golden brown, metallic shiny. The seed-coat ornamentation of L. loeselii are of verruculate to rugulate type. KEY WORDS: Linaria loeselii, the Baltic Sea coast endemic species, pollen, seeds, SEM

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Page 1: Botanika – Stecianamerlin.up.poznan.pl/steciana/wp-content/uploads/2014/03/13_Stec_20... · Pollen grains of this species are small, ... The seeds are winged, concolourous, black

Roczniki Akademii Rolniczej w Poznaniu CCCLXXXVIII

www.up.poznan.pl/steciana ISSN -

Botanika – Steciana , , -

INTRODUCTION

The genus Linaria Mill. comprises species dis-tributed throughout the northern hemisphere. Its main centre of distribution is located in the Mediterranean basin. Linaria together with few other genera tradi-tionally placed in family Scrophulariaceae was recently added to the expanded Plantaginaceae (APG II , ALBACH et AL. ).

The seed characters have proved to be useful in many morphological studies of the Linaria. For instance SUTTON ( ) divided the genus into fi ve sections (Li-naria, Supinae, Pelisseriannae, Versicolores and Diff usae) on the base of seed morphology. The SEM studies of seeds were carried out in the identifi cation of the most species of Linaria from south-western Iberian Penisula (e.g. JUAN et AL. , SEGARRA and MATEU a, b, SÁEZ and CRESPO ). An interesting contribution to the taxonomy of Plantaginaceae sensu lato based on ul-trastructure of seed coat was recently given by SHEHATA and LOUTFY ( ).

VIANO ( ) indicated the potential signifi cance of pollen morphology in the systematics of the genus Linaria. Author ascertained important range of varia-tions between the grains of species, originated from the diff erent regions of the Mediterranean basin. Pol-len grains of some other species of the genus were also described in the works of OLSON ( ), ELISENS ( ) and HALBRITTER ( ).

From among species of Linaria growing in Eu-rope almost % are endemic to the continent. Lina-ria loeselii is an endemic species of the Baltic Sea coast which occurrence is restricted to its south-eastern part

(PANKHURST ). In Poland it grows only on coastal dunes between the village Unieście in the west and Vistula Bar in the east. At present the most numerous populations exist in the Słowiński National Park (PIO-TROWSKA et AL. , PRZYRODA... ). The species is listed in Bern Convention on the Conservation of Euro-pean Widelife and Natural Habitats and it is protected by law in Poland. Linaria loeselii is also considered to be vulnerable within Polish vascular fl ora (KAŹMIERCZA-KOWA and ZARZYCKI , FREY , ZARZYCKI and SZELĄG ).

Linaria loeselii is sometimes treated as subspecies of L. odora [L. odora (M. Bieb.) Fisch. subsp. loeselii (Schweigg.) Hartl] or even, both mentioned names of species are used as synonyms (STASIAK , KAŹMIER-CZAKOWA and ZARZYCKI , MIREK et AL. , PAN-KHURST ).

The aim of the presented work was to investigate the pollen morphology and the ultrastructural traits of seed-coat of this interesting, rare species.

MATERIAL AND METHODS

In the fl owers and the seeds of L. loeselii were collected from one rich population on the Łeba Sand-bar in the Słowiński National Park (E ° ’, N ° ’). The sample of pollen grains (ripe anthers were taken only) was acetolysed according to ERDTMAN’S method ( ), with some insignifi cant modifi cations (TOMLIK--WYREMBLEWSKA ). The sample of well devel-oped and ripe seeds was not prepared, but only cleaned. The using of scanning electron microscopes (JEOL JSM

MORPHOLOGY OF POLLEN GRAINS AND SEEDS OF THE BALTIC SEA COAST ENDEMIC SPECIES LINARIA LOESELII SCHWEIGG.

LESZEK BEDNORZ, IRMINA MACIEJEWSKA-RUTKOWSKA

L. Bednorz, Department of Botany, Poznań University of Life Sciences, Wojska Polskiego C, - Poznań, Poland, e-mail: [email protected]

I. Maciejewska-Rutkowska, Department of Natural Forestry Foundation, Poznań University of Life Sciences, Wojska Polskiego D, - Poznań, Poland

(Received: February , . Accepted: July , )

ABSTRACT. Linaria loeselii Schweigg. is an endemic species of the Baltic Sea coast which occurrence is restricted to its south-eastern part. The aim of the presented work was to investigate its pollen morphol-ogy as well as ultrastructural traits of its seed-coat. Pollen grains of this species are small, spheroidal or prolate spheroidal, -zono-colporate, with microreticulate to foveolate ornamentation. The seeds are winged, concolourous, black to golden brown, metallic shiny. The seed-coat ornamentation of L. loeselii are of verruculate to rugulate type.

KEY WORDS: Linaria loeselii, the Baltic Sea coast endemic species, pollen, seeds, SEM

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L. Bednorz, I. Maciejewska-Rutkowska

and LEO VP) allowed to investigate the details of exine sculpture and the seed ultrastructure. Light mi-croscopy (BIOLAR ) was mainly applied to study the exine structure and quantitative characters of pollen grains. In all pollen grains were observed. The pollen terminology follows PUNT et AL. ( ). With regard to the seed surface, mainly terminology of BARTHLOTT ( ) was applied.

RESULTS

Pollen morphology (Fig. )Pollen grains of L. loeselii are small in size; length

of polar axis (P) is . μm (with the whole range . -- . μm) and equatorial axis (E) . μm (with the

whole range . - . μm). P/E value oscillates from . to . , with the mean value . . They are most of-

ten spheroidal or prolate-spheroial, with circular equa-torial outline. Pollen grains are three-zono-colporate. Ectocolpi are narrowly oblong, with parallel margins and rounded or truncate ends, about ¾ length of po-lar axis. The colpus membranes are without perfora-tions, covered sparcely by minute granules. They are raised and connected with each other in the equato-rial region, forming a bridge over the ectoaperture. The endopori are equatorial. The ornamentation of exine is microreticulate to foveolate. Lumina or perforations of variable outline and size; circular to prolate and irregu-lar; diameter (or length) . - . μm. Muri with smooth and obtuse ridges. The ornamentation of apocolpi and mesocolpi is similar, or depressions of the polar area

FIG. . Pollen morphology of Linaria loeselii in SEM ( - ) and LM ( - ): - – equatorial view, colpi and microreticulate ornamentation of mesocolpi visible, – outline in polar view, microreticulate ornamentation of apocolpus visible, – details of reticulate ornamentation, - – outline in equato-rial view, – outline in polar view, exine structure visible

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Morphology of pollen grains and seeds of the Baltic Sea coast endemic species Linaria loeselii Schweigg.

are slightly smaller than of the ones of equatorial areas. Exine - μm thick, remaining the same thickness from mesocolpia to poles; endoexine of the same thickness as endoexine.

Seed morphology (Fig. )The seeds of L. loeselii are winged, consisting of two

parts: disc (seed body) and wing. Seeds are concolourous (seed body and wing are similar in colour, disc a little bit darker), black to golden brown, metallic shiny. The examined seeds are . - . mm long and . - . mm wide. Wing is broad, . - . mm wide, usually fl at or concave-convex, membranous. The seed-coat surface is smooth and formed by irregularly polygonal cells. They are more or less isodiametric on disc ( - μm in diam-eter), and prosenchymatic-elongated on wing ( - μm long) orientated ± longitudinally to seed radial. The cells of seed-coat have shallow or obscure, concave anticlinal walls and raised, verruculate to rugulate, and convex periclinal walls.

DISCUSSION

VIANO ( ) determined the ranges of variation of pollen size between the diff erent species of Linaria and at the same time he observed the relationship between pollen size and sections of the genus. ELISENS ( ) stated that pollen dimension was mainly correlated with the length of fl ower style within the tribe Antirrhineae. Unfortunately both cited authors did not investigated the species from section Linaria (to which belongs L. lo-eselii). We noticed, that pollen morphology of L. loeselii was generally typical also for the other species of genus,

that had been investigated so far (OLSON , VIANO l , ELISENS , HALBRITTER , HAMDI and AS-SADI ). However, observed some diff erences were in exine ornamentation of L. loeselii pollen. According to the authors, cited above, the grains of the genus are mi-croreticulate. It means network-like pattern, consisting of lumina smaller than μm, delimited by signifi cantly narrower muri (PANKHURST ). The pollen grains of L. loeselii was observed with the lumina wider than muri, forming microreticulate ornamentation, as well as shorter than the distance between adjacent depressions, forming foveolate ornamentation. Besides, both types of sculpture occurred nearly always together, forming a system of lumina and foveolae, being mixed with each other.

The seed features of Linaria species have been tradi-tionally used in its intrageneric classifi cation, but there is lack of detailed information of the most species of the Linaria section. On the other hand SEGARRA and MATEU ( a) stated the variability in seed micromorphology of the species of section Supinae required using of other plant features to identify particular taxa or to determine taxonomic relationships in this group.

As all species of the section Linaria, L. loeselii have the seeds with a marginal encircling wing (CHATER et AL. , SUTTON ). Its seed-coat microsculpture is of verruculate to rugulate type. In this respect it re-sembles the seed-coat surface of L. pedunculata and of L. tursica (section Supinae), but both species have the seeds without wing. Besides, the anticlinal walls of seeds of L. loeselii are slightly concave, while till now only the raised anticlinal ones have been observed (for Linaria species of sections: Supinae, Diff usae, Speciosae and Versicolores; JUAN et AL. , SEGARRA and MATEU

FIG. . Ultrastructure of seed of Linaria loeselii: – outline of the whole seed, – close up of seed wing, – ornamentation of seed body (disc) coat, – ornamentation of wing cells

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L. Bednorz, I. Maciejewska-Rutkowska

a, b, SÁEZ and CRESPO , SHEHATA and LOUTFY ). It seems to be a rare feature of Linaria seeds or

it may be a characteristic one for the seeds of Linaria sections.

AcknowledgementsWe would like to thank Dr. Toshiyuki Fujiki (Nagoya

University) for taking microphotographs of pollen grains and Dr. Hanna Jackowiak (Poznań University of Life Sci-ences) for taking microphotographs of seeds.

REFERENCES

ALBACH D.C., MARTINEZ-OETEGA M.M., FISHER M.A., CHASE M.W. ( ): A new classifi cation of the tribe Veroniceae, problems and a possible solution. Taxon

: - .APG (ANGIOSPERM PHYLOGENY GROUP) ( ): An up-

date of the Angiosperm Phylogeny Group classifi ca-tion for the orders and families of fl owering plants: APG II. Bot. J. Linn. Soc. : - .

BARTHLOTT W. ( ): Epidermal and seed surface char-acters of plants: systematic applicability and some evolutionary aspects. Nord. J. Bot. : - .

CHATER A.O., VALDÉS B., WEBB D.A. ( ): Linaria. In: Flora Europaea. Vol. . Eds T.G. Tutin, V.H. Heywood, N.A Burges, D.M. Moore, D.H. Valentine, S.M. Wal-ters, D.A. Webb. Cambridge University Press, Cam-bridge: - .

ELISENS W. ( ): Pollen morphology and systemat-ic relationships among new world species in tribe Antirrhineae (Scrophulariaceae). Am. J. Bot. , :

- .ERDTMAN G. ( ): Pollen morphology and plant tax-

onomy. Angiosperms. An introduction to palynology . Almqvist & Wiksell, Stockholm.

FREY L. ( ): Linaria odora (M. Bieb.) Fisch. Lnica wonna. In: Poradniki ochrony siedlisk i gatunków Natura – podręcznik metodyczny. Gatunki roś-lin . Eds B. Sudnik-Wójcicka, H. Werblan-Jakubiec. Ministerstwo Środowiska, Warszawa: - .

HALBRITTER H. ( ): Linaria genistifolia, Linaria vul-garis. In: Paldat – a palynological database: Descrip-tions, illustrations identifi cation and information retrieval. Eds R. Buchner, M. Weber. http://www.paldat.org.

JUAN R., PASTOR J., FERNANDEZ I. ( ): Morphologi-cal and anatomical studies of Linaria species from south-west Spain: seeds. Ann. Bot. : - .

KAŹMIERCZAKOWA R., ZARZYCKI K. ( ): Polska czer-wona księga roślin. Paprotniki i rośliny kwiatowe. Instytut Botaniki im. W. Szafera, Polska Akademia Nauk, Kraków.

MIREK Z., PIĘKOŚ-MIRKOWA H., ZAJĄC A., ZAJĄC M. ( ): Flowering plants and pteridophytes of

Poland. A checklist. Vol. . Biodiversity of Poland. – Krytyczna lista roślin naczyniowych. T. . Różnorod-ność biologiczna Polski. W. Szafer Institute of Botany Polish Academy of Sciences, Kraków.

OLSON V. ( ): A biometric study of the pollen mor-phology of Linaria vulgaris (L.) Miller. (Schrophu-lariaceae) and their hybrid progeny in F and F generations. Grana , / : - .

PANKHURST R. ( ): PANDORA taxonomic data base system at the Royal Botanic Garden Edinburgh. http://www.rbg web .rbge.org.uk/FE/fe.html.

PIOTROWSKA H., ŻUKOWSKI W., JACKOWIAK B. ( ): Rośliny naczyniowe Słowińskiego Parku Narodowe-go. Pr. Zakł. Takson. Rośl. UAM Pozn. . Bogucki Wyd. Nauk., Poznań.

PRZYRODA Słowińskiego Parku Narodowego. ( ). Ed. H. Piotrowska. Bogucki Wyd. Nauk., Poznań.

PUNT W., HOEN P.P., BLACKMORE S., NILSSON S., LE THOMAS A. ( ): Glossary of pollen and spore terminology. Rev. Palaeobot. Palynol. : - .

SÁEZ L., CRESPO M.B. ( ): A taxonomic revision of the Linaria verticillata group (Antirrhineae, Scrophu-lariaceae). Bot. J. Linn. Soc. : - .

SEGARRA J.G., MATEU I. ( a): Seed morphology of Linaria species from eastern Spain: identifi cations of species and taxonomic implications. Bot. J. Linn. Soc. : - .

SEGARRA J.G., MATEU I. ( b): Taxonomic study of Li-naria depauperata and L. supina complexes in eastern Spain. Ann. Bot. : - .

SHEHATA A.A., LOUTFY M.H.A. ( ): On the taxono-my of Plantaginaceae Juss. Sensu Lato: evidence from SEM of the seed coat. Turk. J. Bot. : - .

STASIAK J. ( ): Materials for knowledge of the rare and endangered species of Poland . Linaria odora ssp. loeselii Schweigg. Hartl. Fragm. Florist. Geobot.

, - : - .SUTTON D.A. ( ): A revision of the tribe Antirrhineae.

Oxford University Press, London.TOMLIK-WYREMBLEWSKA A. ( ): Pollen morphology

of genus Rubus L. . Introductory studies of the Eu-ropean representatives of the subgenus Rubus L. Acta Soc. Bot. Pol. , : - .

VIANO J. ( ): Recherches palynologiques sur le genre Linaria. Anal. Inst. Bot. Cavanilles : - .

ZARZYCKI K., SZELĄG Z. ( ): Red list of vascular plants in Poland. In: Red lists of plants and fungi in Poland. Eds Z. Mirek, K. Zarzycki, W. Wojewoda, Z. Szeląg. W. Szafer Institute of Botany, Polish Acad-emy of Sciences, Kraków: - .

For citation: Bednorz L., Maciejewska-Rutkowska I. ( ): Morphology of pollen grains and seeds of the Baltic Sea coast endemic species Linaria loeselii Schweigg. Rocz. AR Pozn. , Bot.-Stec. : - .