Roczniki Akademii Rolniczej w Poznaniu CCCLXXXVIII

Botanika – Steciana , , - www.up.poznan.pl/steciana ISSN -

MORPHOLOGICAL DIFFERENTIATION OF POLLEN OF VULGARIS L. FROM POLAND

DOROTA WROŃSKA-PILAREK, MARIA MOROZOWSKA

D. Wrońska-Pilarek, Department of Forestry Natural Foundations, Poznań University of Life Sciences, Wojska Polskiego D, - Poznań, Poland, e-mail: [email protected] M. Morozowska, Department of Botany, Poznań University of Life Sciences, Wojska Polskiego C, - Poznań, Poland, e-mail: [email protected]

(Received: March , . Accepted: June , )

ABSTRACT. The morphology and variability of pollen of Lysimachia vulgaris L. were studied on the base of the material coming from native localities. pollen grains were examined. The diagnostic features of pollen grains of studied species are: exine sculpture irregularly reticulate with lumina of dif- ferent diameters, presence of many minute granules of diff erent size inside lumina (a new feature for the examined species), muri with medium thickness, pollen shape most often subprolate, rarely prolate. Ob- tained results do not fully correspond with the available L. vulgaris pollen descriptions. Statistical analysis of nine quantitative grain characteristics showed their variability to be very diff erentiated. The highest variability was found to occur in trait the Exe/E (the distance between the apices of two ectocolpi). No de- pendence in pollen morphology diff erentiation due to the localization of natural populations was observed.

KEY WORDS: Lysimachia vulgaris, pollen morphology, statistical analyse, LM, SEM

INTRODUCTION of Lysimachia done by BENNELL and HU () resulted in the recognition of ten major pollen-types and four Genus Lysimachia L. consists of ca. species of subtypes. Lysimachia vulgaris pollen was classifi ed by perennial, biennial or annual herbs with almost all over these authors into Fraseri-pollen type, together with the world distribution (STÅHL and ANDERBERG ). several other species e.g. L. davurica, L. fraseri, L. ter- Lysimachia is traditionally placed in the tribe Lysima- restris, L. thyrsifl ora. According to them, the general chieae of the family (CRONQUIST , form of pollen in Lysimachia is constant, the grains are TAKHTAJAN ) and is one of the largest genera in isopolar, tricolporate and they vary in size from smallest this family. Results of the recent phylogenetic studies of (. × . μm in L. capillepsis) to medium (largest . primuloid families based on molecular and morphologi- × . μm in L. franchetii). cal data support the transfer of the genus Lysimachia to We have examined pollen morphology of Lysimachia Myrsinaceae (KÄLLERSJÖ et AL. , APG , MAR- vulgaris from natural habitats distributed all over Po- TINS et AL. , OH et AL. ). Genus Lysimachia land and from the cultivated population with the aim is divided into fi ve-six subgenera and many subgeneric of () the characteristic of L. vulgaris pollen morphology sections (CHEN and HU , BENNEL and HU , AN- on the base of a large sample and () the comparative DERBERG et AL. ). Lysimachia vulgaris belongs to the statistical analysis of the examined pollen traits in order subgenus Lysimachia and section Lysimachia (BENNEL and HU , HAO et AL. ). to evaluate L. vulgaris pollen grains diff erentiation. Lysimachia vulgaris is distributed in Europe, N Amer- ica, SW Asia and NW Africa (FERGUSON , INTER- NET http://www.eol.org). In Poland it is a common herb MATERIAL AND METHODS (ATLAS... ) occurring in moist habitats, such as fens, wet woods, lake shores, and river banks. It is a medicinal The examined pollen grains were collected from plant and the latest results of chemical studies show its natural Lysimachia vulgaris populations distributed all potential cytostatic and anticancer activity (UMEK , over Poland and from one cultivated population, which ŁUCZAK et AL. , JANIK et AL. , PODOLAK et AL. was treated by us as a preliminary attempt to fi nd out , PODOLAK and STRZAŁKA ). if there are diff erences in pollen morphology in de- Morphologically, Lysimachia is very heterogenous pendence of the growing conditions (Table ). with the variation in infl orescence structure, fl oral The cultivated population is located in the garden of morphology, pollination adaptation and growths habit. the Department of Botany of Poznań University of Life Studies of pollen morphology of species and varieties Sciences. It was started in from seeds collected D. Wrońska-Pilarek, M. Morozowska

TABLE . Location of studied pollen samples of Lysimachia vulgaris

Population number Localities Latitude (N) and longitude (E) Year of collection, herbarium

Antoninek °’, °’ , POZNB

Poznań°’, °’ , POZNB

Kamińsko °’, °’ , POZNB

near Trzciel °’, °’ , POZNB

near Świebodzin °’, °’ , POZNA

Oława ° , °’ , POZNA

near Sępolno °’, °’ , POZNA

Mielno °’, °’ , POZNA

near Wejherowo °’, °’ , POZNA

Wilczkowo Lake °’, °’ , POZNA

Puszcza Knyszyńska °’, °’ , POZNA

Puszcza Bukowa near Szczecin °’, °’ , POZNA

Małe Oborowo Lake °’, ° ’ , POZNA

near Chojnice °’, °’ , POZNA

Bługowo °’, °’ , POZNA

near Konin °’, °’ , POZNA

Stawek Lake °’, °’ , POZNA

near Zielona Góra °’, °’ , POZNA

Pępowo °’, °’ , POZNA

cultivated population

Potasze °’ , °’ , POZNB

PZNB – Herbarium of Department of Botany of Poznań University of Life Sciences, PZNA – Herbarium of Adam Mickiewicz University in Poznań.

from one of the natural populations examined in our TABLE . Pollen grains traits analysed present work. Each pollen sample from herbaria collections con- No Traits tained pollen grains sampled from few fl owers of two- -three plants, in dependence on the plant material avail- Length of polar axis (P) ability. Pollen samples from existing natural populations Length of equatorial axis (E) and from the cultivated one were collected every time from fi ve fl owers of plants growing at least two me- Thickness of exine along polar axis (Exp) ters one from another, in order to avoid sampling the Thickness of exine along equatorial axis (Exe) same individual. Plants in cultivation grew on podosolic soil with the humus level of .% and pH = . In the year Length of ectocolpi (Le) before the cultivation on the experimental plot lupine P/E ratio was sown in order to plough back. During the vegetation period plants were fertilized twice with NPK in stand- Relative thickness of exine (P/Exp ratio) ard doses recommended for ornamental species of Lysi- Relative thickness of exine (E/Exe ratio) machia and watered every second day. Each sample was represented by pollen grains and pollen grains Distance between the apices of two ectocolpi (d) were examined in total. They were analysed for nine quantitative features and the following qualitative ones: shape outline and type of exine sculpture (Table ). Only All samples were acetolysed according to ERDTMAN’S grains that were unbroken, symmetrical, and fully ex- () method, with insignifi cant modifi cations (WROŃ- panded in equatorial view were used. SKA-PILAREK ). The terminology follows ERDTMAN Morphological diff erentiation of pollen of Lysimachia vulgaris L. from Poland

(), REITSMA (), PUNT et AL. () and HAL- BRITTER et AL. (). Relative exine thickness is the ra- tio of exine thickness measured along the P (Exp) axis to the length of the P axis, and along the E (Exe) axis to the length of the E axis. Pollen grains were mounted in glyc- erol jelly for LM (Biolar ) studies. Surface sculpture of the examined pollen grains was characterised with the use of SEM technique. The microphotographs were taken on a Zeiss EVO electron microscope (Labora- tory of Electron Microscopy of Adam Mickiewicz Uni- versity in Poznań). The empirical data from grain measurements em- braced quantitative features listed in Table . Their analysis included descriptive statistics and correlation coeffi cients, univariate analysis of variance and Tukey’s procedure, agglomerative grouping by Ward’s method – dendrogram (CALIŃSKI and KACZMAREK , KAROŃ- SKI and CALIŃSKI ).

RESULTS

General characteristic A description of the pollen grain morphology of the studied species is given below and illustrated with a few SEM photographs (Figs -). The morphological observa- tions are summarised in Table . The pollen of Lysimachia vulgaris is three zonocolpo- rate, occasionally four zonocolporate. Pollen size usually FIG. . SEM. Pollen grain in equatorial view; reticulate exine medium (.%), rarely small (.%). Average length of sculpture and ectocolpus visible (× ) polar axis (P) . μm (.-, μm) and equatorial

FIG. . SEM. Pollen grain in equatorial view; pollen shape, FIG. . SEM. Elongated, narrow ectocolpus with colpus mar- exine sculpture and two ectocolpi visible (× ) gins and opened endopore visible (× ) D. Wrońska-Pilarek, M. Morozowska ...... (%) ance ance vari- coef- cient cient fi ...... mum maxi- ...... mean metic arith- ...... mum mini- ...... (%) ance ance vari- coef- cient cient fi ...... mum maxi- ...... mean metic arith- ...... mum mini- ...... (%) ance ance vari- coef- cient cient fi ...... mum maxi- Traits ...... mean metic arith- ...... mum mini- ...... (%) ance ance vari- coef- cient cient fi ...... mum maxi- ...... mean metic arith- ...... mum mini- ...... (%) ance ance vari- coef- cient cient fi ...... mum maxi- PEP/EExpExe ...... mean metic arith- ...... mum mini- . Numeral characteristics of traits of studied pollen grains

ties ABLE Locali- T Morphological diff erentiation of pollen of Lysimachia vulgaris L. from Poland - ...... ffi . . coe variance variance cient (%) ...... mum maxi- ...... tic mean tic arithme- ...... mum mini- - ...... ffi . coe variance variance cient (%) ...... mum maxi- ...... tic mean tic arithme- ...... mum mini- Traits - ...... ffi coe variance variance cient (%) ...... mum maxi- ...... tic mean tic arithme- ...... mum mini- - ...... ffi coe variance variance cient (%) ...... mum maxi- Exp/P Exe/E Le d ...... tic mean tic arithme- ...... mum mini- ties Locali- D. Wrońska-Pilarek, M. Morozowska

FIG. . SEM. Reticulate exine sculpture with lumina with small FIG. . SEM. Reticulate exine sculpture with lumina with small granules and muri visible (× ) granules and muri visible (× ) axis (E) . μm (.-. μm). Average P/E ratio ., Intraspecifi c variability with the range from . to .. Pollen shape spheroi- Features P, E and P/E ratio showed low variability dal to prolate, most often subprolate (.%) or prolate (P – .%, E – .% and P/E – .%). The features Exp, (.%), rarely prolate-spheroidal (.%) and spheroidal Exs, Le, d and Exp/P ratio were characterised by moder- (.%, Table ). Outline in polar view mostly circular, ate level of variability (Exp – .%, Exe – .%, Le – more rarely elliptic and in equatorial view mostly el- .%, d – .% and Exp/P – .%) and for the Exe/E liptic, very rarely circular. ratio variability was found to be the highest (.%). Exine . (.-.) μm thick, with a partial tec- On the basis of an analysis of the diagonal elements tum. Exine is two-layered, with ectexine and endexine of the inverse of the R correlation matrix for all the of about the same thickness. The relative thickness of characteristics, those selected for further study were the exine measured along P (Exp/P ratio) averaged . ones with low values of the diagonal element, which (.-.), and along E (Exe/E ratio) . (.-.). is indicative of a weak correlation between the given The value and range of the coeffi cients of the relative thickness of the exine measured along P (Exp) and along characteristic and the others. In this way the following E (Exe) similar, what indicates that thickness of exine arrangement of traits was chosen: P with the diagonal measured along both axes similar. element value of ., E – ., Exp/P – ., Exe/E – ., Exine sculpture is reticulate. Lumina irregular, with Le – . and d – .. diff erent diameter (.-. μm; average -. μm) de- To verify the hypothesis about the eff ect of the loca- creasing towards colpi. In the -% of studied pollen tion of analysed populations on pollen grain traits grains, inside lumina, minute granules of diff erent size under study, a univariate analysis of variance was car- are present. Slightly thickened muri (average thicknes .- ried out. The assumption of distribution normality was -. μm), usually twice thinner than the lumina diameter. verifi ed by Shapiro-Wilk test, and of variance uniform- Apertures rd (occasionally th), both ectocolpi and ity, by Levene’s test. While analysing the empirical value endopores. Ectocolpi arranged meridionally, regularly, of F statistic and probability p, for each feature under more or less evenly spaced. Long, narrow, elliptic in study the diff erences were found to be signifi cant at the outline, sunken, with distinct margins, ends acute with % signifi cance level (feature P – F = . and p = margo and costae. Mean length . (.-.) μm; calc ., E – F = . and p = ., Exp/P ratio – F = ectocolpi constitute .% of length of the polar axis calc calc . and p = ., Exe/E ratio – F = . and p = ., (P). Ectocolpus sometimes bridged at the equator. Col- calc pus margins present. Endopores equatorially elongated, Le – Fcalc = . and p = . and d – Fcalc = . and p = forming an almost continuous oral zone. Usually located .). In analysing the particular localities a lack of in the middle of ectocolpi, rarely asymmetrically, usually signifi cant diff erences was marked by two points joined one. Fastigium present, sometimes indistinct. with a dashed line (Fig. ). Morphological diff erentiation of pollen of Lysimachia vulgaris L. from Poland . . . ● ● ● ● ● . ● . . ------ . . . ● ● . ● ● . . ------ . . . ● ● ● ● ● . . . . . . ● ● ● . ● . . ------ . . . ● ● ● . ● ● . . ------ . . . ● ● . . . ------ . . . . ● . . ------ . . . ● ● . ● . . ------ . . . ● ● ● . . . . . . . . . ------ . . . ● ● ● . . . ------ . . . ● ● . . . ------ . . . ● . . . ------ . . . ● ● . . . ------ . ● . . ● ● . ● . . ------ . . ● . ● . ● . . ------ . . . ● . . . ------ ● . . ● ● . . . . ------ ● . . ● . ● . . . ------ ● ● . ● . . ● . . . ------ ------● . ● . . ● ● . ● . . . Multiple comparisons procedure by Tukey’s trait . Mean Mean Mean Mean of Mean of of d trait of E trait of P trait IG Localities Localities Localities Localities Localities Localities Exe/E trait Exp/P trait Mean of Le F D. Wrońska-Pilarek, M. Morozowska

The highest mean values of P, E, Le and d were no- The key to pollen types in genus Lysimachia given by ticed for pollen grains from sample no , and the lowest BENNEL and HU () classifi es Lysimachia vulgaris pol- for sample no . In sample no the value of the Exp/P len grains into Fraseri-type by, among others things, the ratio was highest, and Exe/E ratio was highest in the reticulate exine sculpture with lumina < muri, the size of sample no . The lowest values of these features were lumina > . μm and the small pollen grains size. BEUG noticed for samples nos and . () informs, that lumina (brochi) measure - μm In order to distinguish the most homogenous clus- (near colpus only μm), and muri ca. . μm. Accord- ters of pollen grains collected in examined populations ing to our observations lumina are of irregular shape by the criterion of the features under study (P, E, Exp/P, with the medium size .-. μm and not expanding Exe/E, Le and d), an agglomerative grouping by Ward’s . μm, and muri are slightly thickened up to . μm. method was carried out to obtain a dendrogram (Fig. ). It is in agreement with the results of HUYNH (), The grouping revealed that pollen samples from all ex- who reported that size of lumina for L. vulgaris pollen amined localities formed three groups. The greatest was maximally . μm. NOVICKE and SKVARLA () diff erences occurred between all three of them and the found out that for L. terrestris and L. thyrsifl ora, the two cultivated population (no ). The fi rst group comprised species belonging to the same pollen type as L. vulgaris, localities nos , , , , and locality no showed the surface of pollen grains was fi nely reticulate or irreg- similarity to them. The second group included localities ularly perforate and the tectal perforations were more nos , , , , , and the third one localities nos , pronounced in the mesocolpial regions. According to , , , , , , . The grouping revealed also similar- our observations L. vulgaris pollen grains show similar ity to hold among the grains from the latest two groups features. BENNEL and HU () accounted that L. thyr- and the locality no . sifl ora pollen characterises coarse reticulum. In our in- vestigations it was found out that inside the lumina of 40 L. vulgaris pollen grains minute granules of diff erent size are present. Observation of this new feature allows 35 us to suggests that, on the base of pollen morphology, L. vulgaris is more alike L. thyrsifl ora than L. terrestris 30 and it confi rms the attachment of these two species to

25 the same pollen type group. It is also congruent with the results of the latest investigations on phylogenetic 20 relations within genus Lysimachia based both on the Linkage distance molecular as well as morphological data (ANDERBERG 15 et AL. , OH et AL. ). Experiments on Lysimachia vulgaris pollen variabil- 10 ity were not carried till now. Examined grains showed

5 diff erent variation level. The lowest variation level was 20 11 18 13 19 6 4 2 21 8 14 12 5 16 9 15 3 7 10 17 1 found for the most precisely measured length and thick- Number of locality ness of grains (features P, E and P/E ratio). More dif- FIG. . Dendrogram of localities constructed on the basis fi cult to exact measurements, and therefore measured of Ward’s method roughly, exine thickness, distance between colpis and their lengths showed moderate variability. High level of variation may be correlated with the high value of the DISCUSSION one of exine measurements. The statistical analysis showed that pollen morpho- According to PUNT et AL. (), BENNELL and HU logical features of one species may be distinctly diff eren- () and FAEGRI and IVERSEN (), MOORE et AL. tiated. Such a situation was observed concerning pollen (), BEUG () pollen grains from Lysimachia vul- samples no and no . In the sample no the mean garis are tricolporate. BENNELL and HU () described value of the feature P is . μm, and in the sample no them as small (- × - μm), prolate-spheroidal to its value is . μm. Furthermore, in sample no the subprolate. According to our measurements L. vulgaris min. and max. values of the feature P diff er for μm. pollen grains are medium sized and they are most of- It results from these observations that the number of ten subprolate or prolate, very rarely spheroidal. PUNT measured pollen grains is signifi cantly important in et AL. () includes L. vulgaris into L. vulgaris – pol- obtaining the precise pollen morphology characteristic len type together with such species as L. nummularia, for the given species. L. terrestris, L. thyrsifl ora and L. punctata. According The performed statistical analysis showed no simi- to BENNELL and HU () L. vulgaris belongs to the larity in the examined pollen features in dependence on Fraseri-pollen type together with L. davurica, L. fraseri, the population locality. According to that, grains col- L. quadrifolia, L. serulata, L. terrestris and L. thyrsifl ora. lected from plants growing near Zielona Góra (lubelskie They suggested a closer relationship between L. thyr- province), Gostyń (Wielkopolskie province) or Oława sifl ora and L. terrestris based on pollen morphology. (dolnośląskie province) were very similar with each Results obtained in the presented work are generally other, while signifi cant diff erences have been found in agreement with already published descriptions, but between pollen morphology of grains collected from in few cases they are new or not fully congruent with very closely located populations. For example, grains them. from samples no and collected in Poznań and its Morphological diff erentiation of pollen of Lysimachia vulgaris L. from Poland surroundings belonged to diff erent groups according to fort he orders and families of fl owering plants: APG the obtained dendrogram. II. Bot. J. Linn. Soc. : -. ATLAS rozmieszczenia roślin naczyniowych w Polsce. (). Eds A. Zając, M. Zając. Pracownia Chorologii CONCLUSIONS Komputerowej Instytutu Botaniki UJ i Fundacja dla UJ, Kraków. . The most crucial traits of the pollen grains of the BENNELL A.P., HU C.M. (): The pollen morphology studied species: exine sculpture irregularly reticulate and of Lysimachia. Not. Roy. Bot. Gard. with lumina of diff erent diameters, inside lumina many Edinb. : –. minute granules of diff erent size (a new feature for the BEUG H.J. (): Leitfaden der Pollenbestimmung für examined species), muri with medium thickness, pollen Mitteleuropa und angrenzende Gebiete. Verlag Dr. shape most often subprolate, rarely prolate. F. Pfeil, München. . Observed features of Lysimachia vulgaris pollen CALIŃSKI T., KACZMAREK Z. (): Metody komplekso- do not allow to classify it into given by BENNEL and HU wej analizy doświadczenia wielocechowego. Wydz. () Fraseri-pollen type. V PAN i PTB, Warszawa. . Statistical analysis showed that the variability of CHEN F.H., HU C.M. (): Taxonomic and phytogeo- the quantitative features of Lysimachia vulgaris pollen graphic studies on Chinese species of Lysimachia. grains was moderate. The highest level of variability was Acta Phytotaxon. Sin. : -. found for the trait Exe/E, moderate variability level for CRONQUIST A. (): An integrated system of classifi ca- the features Exp, Exe, Le, d and Exp/P ratio, the rest of tion of fl owering plants. Columbia University Press, the features (P, E and P/E) were characterised by law New York. variability. DOBOSZ. M. (): Wspomagana komputerowo staty- . It is necessary to measure large number of pollen, styczna analiza wyników badań. Akademicka Ofi cy- at least grains in each sample, to obtain precise bio- na Wydawnicza Exit, Warszawa. metrical data. ERDTMAN G. (): Pollen morphology and plant taxon- . An agglomerative grouping by Ward’s method omy. Angiosperms. Almqvist & Wiksell, Stockholm. FAEGRI K., IVERSEN J. (): Bestimmungsschlüssel für showed that pollen grains collected from the cultivated die nordwesteuropäische Pollenfl ora. Fischer, Jena. population diff er from all other pollen samples collected FERGUSON L.F. (): Lysimachia L. In: Flora Europaea. from natural localities. No dependence on population’s Vol. . Diapensiaceace to Myoporaceae. Eds T.G. Tu- localization and pollen similarity was observed. tin, V.H. Heywood, N.A. Burges, D.M. Moore, D.H. . To confi rm the observed Lysimachia vulgaris pol- Valentine, D.A. Walters. Cambridge University Press, len morphology diff erentiation between grains coming Cambridge: -. from natural populations and from the cultivated one HALBRITTER H., WEBER M., ZETTER R., FROSCH-RADIVO it is necessary to examine more pollen samples from A., BUCHNER R., HESSE M. (): Pal dat-illustrated higher number of cultivated populations. handbook on pollen terminology. Vienna. . Obtained results confi rm resemblance of Lysi- HAO G., YUAN Y.-M., HU CH.-M., GE X.-J., ZHAO N.-X. machia vulgaris pollen morphological features with (): Molecular phylogeny of Lysimachia (Myrsi- L. thyrsifl ora and L. terrestris, which are, according to naceae) based on chloroplast trnL-F and nuclear ri- recently published phylogenetical relations within genus bosomal ITS sequences. Mol. Phyl. Evol. : -. Lysimachia, the closest relatives of the examined spe- HUYNH K.-L. (): Le pollen et la systématique chez le cies. The new observed feature, the presence of minute genre Lysimachia (Primulaceae). Part . Morphologie granules in lumina, allows to accept L. vulgaris as more générale du pollen et palynotaxonomie. Candollea alike L. thyrsifl ora than L. therrestris. : -. INTERNET http://www.eol.org .. Acknowledgements JANIK I., JANECZKO Z., SENDRA J., BURCZYK J. (): The authors would like to thank Mirosław Dziurla Benzoquinone pigment from Lysimachia vulgaris. (Department of Forestry Natural Foundations, Poznań Fitoterapia , : . University of Life Sciences) for his technical assistance KÄLLERSJÖ M., BERGVIST G., ANDERBERG A.A. (): in preparing micrographs and Dr. Jarosław Lira (De- Generic realignment in primuloid families of the partment of Food and Agricultural Economics, Poznań s.l.: a phylogenetic analysis based on DNA University of Life Sciences) for carrying out statistical sequences from three chloroplast genes and mor- data analysis. phology. Amer. J. Bot. : -. KAROŃSKI M., CALIŃSKI T. (): Grouping in multivar- iate populations on the basis of Euclidean distances. REFERENCES Algor. Biometr. Statyst. : -. ŁUCZAK S., ŚWIĄTEK L., DANIELEWSKI M. (): Phe- ANDERBERG A.A., MANNS U., KÄLLERSJO M. (): nolic acids in herbs Lysimachia nummularia L. and Phylogeny and fl oral evolution of the Lysimachieae L. vulgaris L. Acta Pol. Pharm. , : -. (Ericales, Myrsinaceae): evidence from ndhF sequence MARTINS L., OBERPRIELER C., HELLWIG F.H. (): data. Willdenowia : -. A phylogenetic analysis of Primulaceae s.l. based on APG (Angiosperm Phylogeny Group) (): An update internal transcribes spacer (ITS) DNA sequence data. of the Angiosperm Phylogeny Group classifi cation Plant Syst. Evol. : -. D. Wrońska-Pilarek, M. Morozowska

MOORE P.D., WEBB J.A., COLLINSON M.E. (): Pollen European Pollen Flora . Ed. W. Punt. Elsevier, Am- analysis. Blackwell Scientifi c Publications, London. sterdam: -. NOWICKE J.W., SKVARLA J.J. (): Pollen morphology REITSMA T.J. (): Suggestions towards unifi cation of and the relationship of the Plumbaginaceae, Polygo- descriptive terminology of angiosperm pollen grains. naceae, and Primulaceae to the Order Centrosper- Rev. Paleobot. Palynol. : -. mae. Smithsonian Institute. Press, Washington: STÅHL B., ANDERBERG A.A. (): Myrsinaceae. In: The -. families and genera of vascular plants. Flowering OH I.C., ANDERBERG A.L., SCHÖNENBERG J., ANDERBERG plants-dicityledons. Celastales, Oxalidales, Rosales, A.A. (): Comparative seed morphology and Cornales, Ericales. Ed. K. Kubitzki. Springer, Berlin: character evolution in the genus Lysimachia (Myrsi- -. naceae) and related taxa. Plant Syst. Evol. , -: TAKHTAJAN A.L. (): Diversity and classifi cation of -. fl owering plants. Columbia University Press, New PODOLAK I., ELAS M., CIESZKA K. (): In vitro antifu- York. gal and cytotoxic activity of triterpene saponosides UMEK A. (): Saponins and sterols of yellow loos- and quinoid pigments from Lysimachia vulgaris L. estrife Lysimachia vulgaris L. Farm. Vestn. (Ljublja- Physiol. Res. :-. na) : -l. PODOLAK I., STRZAŁKA M. (): Qualitative and quan- WROŃSKA-PILAREK D. (): Pollen morphology of the titative LC profi le of embelin and rapanone in se- polish species of the genus Ribes L. Acta Soc. Bot. lected Lysimachia species. Chromatographia (S): Pol. , -: -. -. PUNT W. (): The northwest European pollen fl ora. Vol. , Parts -. Elsevier, Amsterdam. For citation: Wrońska-Pilarek D., Morozowska M. (): PUNT W., BLACKMORE S., NILSSON S., LE THOMAS A. Morphological diff erentiation of pollen of Lysimachia vul- (): Glossary of pollen and spore terminology. garis L. from Poland. Rocz. AR Pozn. , Bot.-Stec. : http//www.bio.uu.nl /~palaeo/glossary/glos-int.htm. -. PUNT W., DE LEEUW VAN WEENEN J.S., VAN OOSTRUM W.A.P. (): Primulaceae. In: The Northwest