Research eco.mont – Volume 9, Number 2, July 2017 16 ISSN 2073-106X print version – ISSN 2073-1558 online version: http://epub.oeaw.ac.at/eco.mont https://dx.doi.org/10.1553/eco.mont-9-2s16
Euphrasia’s hide-and-seek: periodicity of occurrence in some high-mountain spe- cies of Euphrasia (Orobanchaceae) and its consequences for botanical research on protected areas
Ewa Posz
Keywords: Euphrasia, periodicity, protected areas, botanical research
Abstract The paper presents the results of long-term observations made on selected populations of two high-mountain Euphra- sia species. The monitoring was conducted during 2002–2016 in ranges of the Polish parts of the Carpathians and Sudetes. Field studies revealed periodical occurrence and disappearance of populations in the localities for one or more seasons, which may be connected with the life cycle of annual plants. The phenomena of periodicity occurrence and seed dormancy and their consequences are discussed.
Introduction
The genus Euphrasia (eyebright) is represented by ca. 350 species worldwide (Fischer 2004). High-moun- tain eyebright species occur at altitudes up to 4 450 m a.s.l. in the Himalayas (Li 1953). In Europe, eyebright species have been found in almost all mountain ranges at altitudes as high as 2 700 m a.s.l. (in the Alps) (Yeo 1978). There are 11 species of eyebright in Poland (Posz 2014, 2016), three of which, E. minima Jacq., E. picta Wimm. and E. corcontica (Smejkal) Smejkal et are typical for siliceous bedrock. The range of E. salis- burgensis Funck is also limited to mountain areas, but it prefers calcareous bedrock. European eyebrights are small, semi-parasitic her- baceous annuals, which range in height from several centimetres to nearly twenty centimetres. The dou- (though occasionally yellow or blue), with a yellow spot on the lower petal, with individual leaves situated radially around the stem (Figure 1). Eyebrights are therophytes, meaning that their entire life cycle from Figure 1 – Euphrasia picta Wimm. in the Tatra National germination to producing their own seeds takes place Park. © E. Posz within a single growing season. After producing seeds, the plants die off, creating a soil seed bank. Given the high number of differences found within annual life cycle and seed dormancy in the soil seed individual species, the high number of hybrids, and bank. The possibility of germination (particularly their overlapping ranges, it should be said that Euphra- for therophytes) has been investigated for different sia ! \ # ! species by many scientists (Symonides 1989; Fenner atypical forms, probably as a result of semi-parasit- ' *++/% : # ism, creates additional problems. Their morphological on germination, such as soil temperature, moisture $ % level, the impact of the surrounding plant commu- the presence or absence of host plants (Crosby-Brown nities, microbes or fungi (Rees & Long 1992; Egley 1950; Philipson 1959; Wilkins 1963; Yeo 1961, 1964). 1995; Fenner & Thompson 2005). Observations of & \ - seed dormancy and germination behaviour of the ge- versity even within one population. nus Euphrasia have been made, but only with regard Field studies over many years revealed the periodi- to seeds collected from nature and sowed in green- cal occurrence and disappearance, for one or several houses, gardens, pots or Petri dishes (Yeo 1961, 1964; seasons, of certain populations in the localities. This Liebst & Schneller 2008). Periodicity of populations phenomenon was called periodicity (Posz 2014; Pro- \ szkiewicz 2006) and is probably connected with the # \ - Ewa Posz 17
Figure 2 – The areas of local research: protected areas in the Karkonosze National Park, the Tatra National Park, the Babia Góra National Park, the Bieszczady National Park. Prepared by E. Posz. cies had been systematically monitored. Furthermore, es in the forest management of the areas where the the study demonstrated that dormancy (i. e. the seeds species is found, the localities have deteriorated, and lying in the soil, and the plant not appearing in a local- the species is being pushed out by others. It is thus es- ity) may last for several years. sential to discover whether the species is going extinct, The aim of the monitoring programme was to de- or whether the absence of the plant at the locality is a \ - temporary natural phenomenon. lation numbers, including the absence of the plant from E. minima \ a locality, are an isolated anomaly, or whether they occur during the on-going multi-year study at numerous sites regularly and are part of the eyebright’s survival strat- in the Tatras, at many of which, however, it appears egy. The research question was: Does the periodicity irregularly, with breaks of several years. phenomenon affect our understanding of this genus? The study also looked at the duration and frequency Materials and methods of these dormant periods, as well as the number of the # Taxonomical studies and local research on Euphra- they emerged at a locality after a period of absence. sia species have been made in the Polish part of the Two species E. picta and E. minima, especially valu- Carpathians and the Sudetes since 2001 (Posz 2014). able botanically in Poland, were studied. These are, In addition, some sites of E. picta and E. minima have however, typically highland species, and in Poland they been regularly monitored in the Karkonosze, the appear only within a small distribution range and not Bieszczady, the Tatras and on Mt Babia Góra (Fig- in great numbers. As the localities at which these spe- ure 2). The areas of local research are protected areas: cies grow occur within national parks, the species are the Karkonosze National Park (NP), Tatra NP, Babia not under special protection in Poland. They are, how- Góra NP and Bieszczady NP. ever, recognized as a valuable part of the ecosystem Monitoring of the changes in the abundance and and appear on Polish regional lists of rare and endan- disappearance of populations was carried out in the gered plants (Pender 2003). Tatras, the Bieszczady and on Mt Babia Góra (in the E. picta is an exceptionally rare species in Poland, Carpathians) from 2002 to 2015, and from 2008 to \ # # 2016 in the Karkonosze (the Sudetes). It was conduct- Tatra and Bieszczady Mountain ranges. Due to chang- ed every year in late July / early August. Before late Research 18
Table 1 – Monitoring of E. minima in the Tatras: H: high abundance of plants (more than 500 individuals); –: the monitoring was not conducted at this site in this year (in the section Current monitoring), or documentation for the site was not found in her- barium material during research (in the section Herbarium data); X: the site was provided by Herbarium material. No. Locality Herbarium data Current monitoring ski 1875 ski 1875 L L niowski 1886 V niewski 1923 V Kulczy (KRAM 185911) Kulczy (KRAM 185982) Wrze (ZAMU MT 826) Wi (WA 018265) (WA Jasiewicz 1951 (KRAM 434725) & Jasiewicz 1986 Pawlus (KRAM 453093) 2002 Ewa Posz 2003 Ewa Posz 2004 Ewa Posz 2008 Ewa Posz 2012 Ewa Posz 2014 Ewa Posz 2015 Ewa Posz Morskie Oko, nad stawem (the Sea Eye Lake, near 1. X 00148000 the lake) Vci (the Sea Eye Lake, the 2. ––––––180118154738 Peninsula of love) 3. X 0 H 87 H 15 123 190 4. X 0 67 15 0 25 150 43 5. Ciemniak (Mt Ciemniak) X 0 0 H 0 79 – H 6. ! 4ki (the Little Meadow Valley) X 0 0 15 267 0 157 45 Bcz MiBdzy Kopami, Wielka Kopa Królowa (the 7. ––––––0651918726522H Pass between the Kopas) Btusi, miB# $ % & $ 8. B & # X001860 8 0 Kobylarz) ' & Bczy Liliowe (the Sucha Valley 9. ––––––005714004 near the Liliowe Pass) 10. Bcz Liliowe (the Liliowe Pass) ––––––301630–7 11. Kopa Kondracka (Mt Kopa Kondracka) ––––––009 016–28 12. 4' 4czniak) ––––––0H470H–H Hala G4 ' * Lca (G4sienicowa 13. ––––––06826H781800 Hall Mountain, rocks near the Murowaniec shelter) 14. Krzesanica (Mt Krzesanica) ––––––00H065–H Chuda Turnia, Ciemniak (the Skinny Crag, Mt Ciem- 15. ––––––01605H–H niak)
July, in cooler years in the higher reaches of the Tatras, every year. Plants collected from these localities at the the plants often still do not have fully developed seeds beginning of the research for the purpose of species and capsules. On the other hand, in late August, frosts \ or even snowfalls often occur. The experiences of Institute of Botany, Jagiellonian University in Kraków \ # - (Posz 2014; Proszkiewicz 2006). den changes of weather. Historical data pertaining to some localities were @ \ # - gained from research (2001–2013) into the collections od of species population monitoring was applied, as of a total of 30 herbaria (Mirek et al. 1997; Thiers recommended by the Polish Environmental Protec- [continuously updated]). The herbaria referred to are: tion Inspectorate and the Institute of Environmental Herbarium of Anna and Jan Kornasiowie in Cracow Protection of the Polish Academy of Sciences for $@XZ%\ ^ _ U ` C $`U}%\ protected, endangered and rare species (Perzanowska ` C ~ $`~%\ 2010). This method is used successfully in the moni- Agriculture Academy in Bydgoszcz (BYDG); Medical /Q \ - # ~ $~ :@%\ X - toring as part of the Natura 2000 programme (Per- sity in Cracow (KRA); Polish Academy of Sciences zanowska 2010, 2012a, 2012b). – Institute of Botany in Cracow (KRAM); University Studies were conducted at 14 sites for E. picta (Mt of Agriculture in Cracow – Department of Forest Babia Góra, the Bieszczady, the Tatras) and at 17 sites Biodiversity (KRFB); Pedagogical University in Kielce for E. minima (Mt Babia Góra, the Karkonosze, the (KTC); University of Silesia in Katowice (KTU); Tatras). The observations were carried out in order to Maria Curie-Sklodowska University in Lublin (LBL); \ Euphrasia population in a Lódz University (LOD); Upper Silesian Museum in given locality, to determine the population’s abundance Bytom (MGS); Teacher Training College in Olsztyn (counting of individuals), and to analyse the condi- $} %\ : @ # & N tion of individual plants (checking whether plants had of Pharmaceutical Botany (PBMA); Adam Mickie- % U - # & N - graphic documentation of the sites was carried out ment of Plant Taxonomy (POZ); Adam Mickiewicz Ewa Posz 19
Table 2 – Monitoring of E. minima in the Karkonosze Mountains (for the abbreviations, see Table 1). No. Locality Historical data Current monitoring Schube 1896 (SCHUBE 1897) (BIL 423136) 1986 Kusiak (KRAM 423140) Kwiatkowski 1997 1998) (KWIATKOWSKI 2008 Ewa Posz 2010 Ewa Posz 2011 Ewa Posz 2013 Ewa Posz 2014 Ewa Posz 2015 Ewa Posz 2016 Ewa Posz 1. Unieb % ' $ % XXXX11217716027156024 2. Równia pod Uniebk4(the plateau under Mt Uniebka)– – – –HHHHHHH
# & N herbarium collections, although no representatives of Collections Faculty (POZG); Agricultural Academy E. minima were found there before 2004, despite care- & $&`%\ _ & # \ U $_&%\ $} %\ : frequency in different localities varied: some popula- University of Silesia in Katowice (SOSN); University tions disappeared, while some were continuously pre- of Szczecin (SZUB); Nicolaus Copernicus University $ _%\ ~ # $~ @%\ - E. minima was found only at one site in the Karkon- sity of Warsaw (WA); Warsaw Agriculture University $ *% # : # & C # (WAUF); University of Podlasie in Siedlce (WSRP); Z $ # Z % \ - # $_}%\ # scribed in the 19th century by Schube (1897). The next Museum in Zakopane (ZAMU); Polish Academy of locality of the species (two populations) to be discov- Sciences – Institute of Nature Conservation in Zako- *++ _ & C B $ & pane (ZTS). : & C % $& *++% *+Q # - other population was found, 300 m from the Równia Results & C B : Z _ & C B U *+/ \ - Studies on the distribution of E. minima in the : # & C # Z *+Q Tatras carried out in 2002 and 2003 rendered few re- abundance there was relatively high. Plants were ob- sults (Table 1). Only a few localities for this species served in 2016, but the population was not numerous. were discovered, despite the fact that herbarium col- Changes in the population abundance and even the lections (KRA, KRAM) contain a considerable num- disappearance of some of them were also observed ber of sheets for this taxon. However, in 2004, there for another mountain taxon – E. picta. In the Tatras was a mass occurrence of this species on mountain $ % \ # - pastures, rocks and even at the sides of trails. Speci- barium collections (KRA, KRAM) is located on Mt mens were also collected from sites documented in $ _ : %
Table 3 – Monitoring of E. picta in the Tatras (for the abbreviations, see Table 1). No. Locality Historical data Current monitoring dalski 1965 4 Jasiewicz 1953 (KRAM 423147) Jasiewicz 1953 (KRAM 423128) M (KRAM 27958) Jasiewicz 1966 (KRAM 440517) 1981 Pawlus (KRM 450977) Oklejewicz 1975 Czeczott ? (KRAM 254061) ? 1926 (KRAM 334970) 2002 Ewa Posz 2003 Ewa Posz 2004 Ewa Posz 2008 Ewa Posz 2012 Ewa Posz 2014 Ewa Posz 2015 Ewa Posz 1. X X X X 0 0 0 21 15 H H 2. GB ! +B ! ––––––––030176HH Droga nad Reglami (the road above the 3. ––––––––025430–6454 Regles) ; # < ; $ ' = ; 4. # & > < ; = ; X 04000H25 Valley) Dolina Str4byska, wylot (outlet of the 5. X0009000 Str4byska Valley) Dolina KoV' ' ? $ ! 6. (the KoVcieliska Valley, near the Hala X X 01605000 Smytnia) Polana Waksmundzka (the Waksmundzka 7. ––––––––0090350 clearing) Research 20
Table 4 – Monitoring of E. picta in the Bieszczady (for the abbreviations, see Table 1). No. Locality Historical data Current monitoring Jasiewicz 1954 (KRAM 423136) Jasiewicz 1954 (KRAM 423136) Jasiewicz 1954 (KRAM 423136) Jasiewicz 1956 (KRAM 423146) Jasiewicz 1960 (KRAM 423142) Jasiewicz 1968 (KRAM 423125) Zemanek & Winnicki 1993–1997 (Zemanek & Winnicki 1999) 2002 Ewa Posz 2003 Ewa Posz 2004 Ewa Posz 2008 Ewa Posz 2012 Ewa Posz 2014 Ewa Posz 2015 Ewa Posz 1. Tarnica, szczyt (Mt Tarnica, the peak) X H 0 0 0 0 0 0 @ ' ! ' @ ' 2. –––––– –030179689314 rocks below the peak) Tarnica, w drodze na szczyt (Mt Tarnica, in the 3. –––––– –––– ––127253 path on the peak) 4. Bukowe Berdo (the Bukowe Berdo) X X800 05 0 0 5. Krzemie (Mt Krzemie ) X X X000 00 0 0 6. < < X X H 0 0 0 0 0 0 7. Smerek (Mt Smerek) X 0 0 0 0 0 0 0
in the locality of Droga nad Reglami (the road above the high mountain pastures caused by altered land use and Regles) appears and disappears irregularly. Monitoring the abandoning of pastures. carried out in the Bieszczady (Table 4) in 2002–2015 Periodicity is most probably connected with the reported changes in the sites on Mt Tarnica and Bu- creation of soil seed banks by Euphrasia species kowe Berdo (the Bukowe Berdo). The studies did not (Liebst & Schneller 2008), which is associated with the \ # annual life cycle. Many years’ dormancy of seeds in the herbarium collections (KRAM) of Jasiewicz (from the soil was observed earlier, during their experimen- the 1950s and 1960s) or Zemanek and Winnicki (1999). tal cultivation from host plants in pots, greenhouses Mt Tarnica, the highest mountain of the Bieszczady, is and gardens (Vitek 1998; Wilkins 1963; Yeo 1964; a good locality to study population changes. In 2002, Liebst & Schneller 2008). In the high-mountain spe- a large population of E. picta was found at the sum- cies under study, E. minima and E. salisburgensis (Liebst mit, below the viewing platform. During the following & Schneller 2008), the dormancy period lasted ca. years (2003–2015), no plant was found at this locality. three years, while in the low-land species E. stricta D. In 2003, a new population of the species was discov- Wolff. ex F. Lehm. seed germination was observed up ered on rocks, at a different site, in the northwestern \ # $ % part of the summit. The plants at this site were moni- Knowledge about the periodicity phenomenon tored constantly from 2003 to 2015, and noted almost as seen in relation to the eyebright has a special sig- every year. Another population, located slightly lower, \ midway between the pass and the summit, was discov- resources in protected areas (for example NPs or Na- *+Q \ *+/ ture 2000 areas), as described in the European Habitats The occurrence of both E. minima and E. picta on Directive. So far, periodicity has not been taken into Mt Babia Góra is well documented in herbarium col- account in studies of the distribution of eyebright lections (AJK, KRA, KRAM, MGS, WA). However, \ # \ # - gain a fuller and more accurate understanding of Eu- ried out in 2003–2015. phrasia distribution, studies of the same area should # \ Discussion and conclusions years. Floristic 1- or 2-year studies are often insuf- \ Monitoring indicated that huge changes in plant and sometimes population disappearance for several numbers, including the total absence of the species years, can be a cause of incorrect evaluation of the from a locality is a normal phenomenon for highland population’s condition and, consequently, the species’ eyebrights. These dormant periods, when no single status in a given area. Excessive tourism and natural plant of the species is observed at a locality, may be rock erosion of the habitat are usually suggested as quite long, lasting for several years. It has been shown that the condition of the plants on reappearance is very in a population. They were, for example, listed in a # # case of E. minima evaluation for the Karkonoski Park The fact that many of the populations found by Narodowy (Kwiatkowski 1998; Pender 2003; Zemanek and Winnicki (1999) in Bieszczady cannot *++\ B *++Q% ^ Eu- # \ # # phrasia, however, periodicity of occurrence seems to with periodicity but also with natural succession in be just as important a factor. Ewa Posz 21
^ \ - banchaceae) occuring in the Swiss Alps. Botanical Jour- pret such as the disappearance of eyebright localities nal of the Linnean Society 156: 649–656. within the area studied, knowledge about their ecology : } : ' X Polish her- and monitoring of their numbers may allow appropri- baria. Polish Botanical Studies, Guidebook Series. Pol- ate measures to be taken to protect the species. One ish Academy of Sciences, W. Szafer Institute of Bota- X B } ~ ` ny, Kraków, Poland. - Pender, K. 2003. Gatunki endemiczne, reliktowe i Z & $X B } ~ wysokogórskie w Sudetach [Endemic, relict and alpine Bank, The Karkonsze NP [continuously updated]), a # : U ZB $ % programme which has been running for many years, [Endangered \ # Vascular Plants of Lower Silesia]. Instytut Biologii protect and increase the gene pool of different taxa by _ ' # & # vegetative and generative reproduction. One species & # & # # P Q /N/ covered by this programme is a species described in [In Polish] this article, Euphrasia minima. The question of which Perzanowska, J. (ed.) 2010. # - ! " Q 5 % [Monitoring of plant tion of eyebrights’ seed dormancy, which may be quite ~ # & U ~U& - Warszawa. [In Polish] dance, remains open. Perzanowska, J. (ed.) 2012a. Monitoring gatunków ! " Q 5 %% [Monitoring of Acknowledgements plant species. Guidebook of methodology. Part II]. ~U& ¡U & I am grateful to directors of the Babia Góra NP, Perzanowska, J. (ed.) 2012b. Monitoring gatunków the Bieszczady NP, the Karkonosze NP and the Tatra ! " Q 5 %%% [Monitoring of NP for allowing access to the Parks. plant species. Guidebook of methodology. Part III]. This work was partly supported by the Polish State ~U& ¡U & \ _ $Z` Philipson, W.R. 1959. Some observations on root- 2P04G02427 in the years 2004–2006). parasitism in New Zealand (Euphrasia). Transactions and Proceedings of the Royal Society of New Zealand 87: 1–3. References Posz, E. 2010. Euphrasia minima Jacq. (Orobanchaceae) w Karkonoskim Parku Narodowym [Euphrasia minima Crosby-Browne, A.J. 1950. The root parasitism of Jacq in the Karkonosze National Park] – Opera Corcon- Euphrasia salisburgensis Funck. Watsonia. Journal of Bo- tica 47 Suppl. 1: 1–5 (English summary). [In Polish] tanical Society of the British Isles 1(6): 354–355. Posz, E. 2014. & ' *+ / 6 - Egley, G.H. 1995. Seed germination in Soil: Dor- mia i rozmieszczenie. [Genus of Euphrasia L. in Poland. mancy cycles. In: Kigel, J. & G. Galili (eds.), Seed devel- ! # # X opment and germination: 529–544. New York-Basel. raków (English summary). [In Polish] Fenner, M. & K. Thompson 2005. The ecology of Posz, E. 2016. Euphrasia corcontica (Smejkal) Smejkal seeds. Cambridge. # & ¡E. corcontica Fischer, E. 2004. Scrophulariaceae. In: Kadereit, J.W. $ % (ed.), The Families and Genera of Vascular Plants 7: 332– Poland]. 7 N 5 9 432. Berlin. < * => 6 7 - X B } ~ ` Z - tanicznego. [Botany – tradition and modernity, Book of sze National Park [continuously updated]. Avail- abstract of the 57 Congress of Polish Botanical Soci- able at: http://www.bankgenow.kpnmab.pl/ (acessed: ety]. Lublin. [In Polish] 24.01.2017) [In Polish] Posz, E. 2006. 6 ! \ \ Kwiatkowski, P. 1998. Stan aktualny populacji ' *+ / @ ! Q ! ' Euphrasia minima w Karkon- taksonów górskich. Praca doktorska [Taxonomy and phy- oskim Parku Narodowym [The present state of Eu- togeography of Euphrasia L. genus in Poland with a phrasia minima population in the Karkonosze Mts]. special regard of mountain species. PhD Disserta- In: Sarosiek, J. & J. Štursa (eds.), Geoekologiczne Prob- U # ` # X lemy Karkonoszy [Geoecological problems of the Kraków. [In Polish] Z : : # Rees, M. & M.J. Long 1992. Germination biology w Przesiece, 15–18.10.1997: 335–339 (English sum- and the ecology of annual plants. The American Natura- mery). [In Polish] list 139(3): 484–508. Li, H.-L. 1953. Euphrasia in China. Notulae naturae of Schube, T. 1897. Ergebnisse der Durchforschung the Academy of Natural Sciences of Philadelphia 254: 1–6. der Schlesischen Phänerogamen Flora im Jahre 1886. Liebst, B. & J. Schneller 2008. Seed dormancy and Jahrbuch schlesische Gesellschaft vaterländ. Cult 74(2): 39– germination behaviour in two Euphrasia species (Oro- 64. [In German] Research 22
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