ANNALES HISTORICO-NATURALES MUSEI NATIONALIS HUNGARICI Volume 96 Budapest, 2004 pp. 51-62.

Some differences in morphological structure of the vegetative parts (herbage) of heufleriana Schur and Sesleria hungarica Újhelyi ()

D. KOVATS

Department of Botany, Hungarian Natural History Museum H-1087 Budapest, Könyves Kálmán krt. 40. Hungary E-mail: [email protected]

Abstract - Comparative morphological measurements were carried out on the culms and leaves of Sesleria heufleriana SCHUR and Sesleria hungarica ÚJHELYI. On the basis of observations and mea­ surements it can be stated that differences occur in numbers and length values of internodes. The length dimension (ratio) of leaf blades and leaf sheets changes on another level of the culms in Sesleria heufleriana as in Sesleria hungarica. The investigations were carried out on collected from original habitats and on those planted in an experimental garden. With 5 figures and 4 tables.

Key words - Sesleria heufleriana, Sesleria hungarica.

INTRODUCTION

FERDINAND SCHUR (1799-1878) - who had a hard life as a pharmacist, chemist and botanist - during his botanical round trip in Transylvania between 5th luly and 15th August in 1853 discovered a lot of new taxa, among them a new spe­ cies of Sesleria, and described it as Sesleria heufleriana SCHUR (SCHUR 1853). Another outstanding botanist, taxonomist in Hungary several decades later was JÓZSEF ÚJHELYI (1910-1979), who had always been a paternal friend to this author. He was the best specialist of the European of Koeleria, Sesleria, Achillea, and Lotus. ÚJHELYI described Sesleria hungarica ÚJHELYI as a new spe­ cies of Poaceae (ÚJHELYI 1959). These two Sesleria are closely related taxa. Ac­ cording to ÚJHELYI, S. hungarica is octoploid 2n=56 and a much more robust than the tetraploid 2n=28 S. heufleriana SCHUR (ÚJHELYI 1959). In this paper the author sketches out the morphological structure of the vege­ tative parts of culms, makes comparision with culms, internodes, leaf blades, leaf sheaths, the measures of lengths, and their ratios. The author tries to find more morphological differences between these two closely related taxa, because most of the scientific literature does not accept S. hungarica as an independent species. In most of botanical literature Sesleria hungarica ÚJHELYI is considered to be a subspecies of Sesleria heufleriana (DEYL 1978, 1980, 1989, 1991, 1996, 2000, SOÓ 1980, PRISZTER 1985, SIMON 2000). Only in few instances Hungarian litera­ ture accepts Sesleria hungarica as a species (SOÓ 1973, SIMON 1992). In the floristic work of the Hungarian Bükk Mountains (VOJTKÓ 2001) and in protected plants of Hungary (LESS 1999), Sesleria hungarica is dealt with as an independent species.

MATERIALS AND METHODS

The research material of both Sesleria taxa amounting to 320 specimens, from 13 populations (Table 1) was collected at different times (see below), and in different places of the northeastern part of Hungary partly from original habitats, and partly from an experimental garden of identical condi­ tions (100 specimens from 4 populations). They were side by side at a distance of about 40 cm from one another. The sampling for measurement in the case of samples from the original habitats were 220 specimens from 9 populations (Table 1) gathered in May, immediately after the blooming period, at the beginning of the fruiting stage (see below). The collection of samples from the experimental garden for measurement took place about a month earlier, in April, in the blooming period. In this lat­ ter case the lengths of culms and those of the internodes of specimens from the original habitats can­ not be compared with the specimens from the experimental garden. As is known, the culms and the leaves of grasses further increase in length not only until the blooming period but also after that. The research material was collected from the following habitats. Sesleria heufleriana SCHUR, 1853 - Population 1: Comit. Cibin. In lapidosis montis "Verfu Magura" prope pagum Nagy Talmács. Solo arenoso saxoso, alt. c. 250 m.s.m. Loco classico! Leg. Dr. ÁRPÁD DE DEGEN 05.05. 1903. Gramina Hungarica exsiccata. Topotypus DEZSŐ KovÁTS, 09.2002. Population 2: In rupibus apud Szarvaskő, mt. Bükk. Legit: 05.05.1949. ANDRÁS HORÁNSZKY. Population 3: Material from the experimental garden originated from: 10-11. Comit. Heves, prope pagum Szarvaskő. In rupibus gabrosis, in monte "Várhegy", in "Bükki Nemzeti Park", alt. cca. 300 m.s.m. 06.06.1983. Leg. et det. TIBOR SZERDAHELYI. Taken out of the experimental garden: 16.04.1986. Population 4: Comit. Abauj-Torna. In rupibus calcar. sept, silvat. montis Szádvár prope Szögliget. Alt. cca. 450 m.s.m. 01.05.1953. Leg. ÁDÁM BOROS.

Table 1. Sesleria research material from original from experimental sum habitat garden number of number of number of number of number of number of specimens populations specimens populations specimens populations S. heufleriana 110 4 30 1 140 5 S. hungarica 110 5 70 3 180 8 Sum 220 9 100 4 320 13 Population 5: Hungária sept.-orientalis. In montibus Tornai Karszt, in monte Nagyoldal exp. merid. in ass. Seslerietum heuflerianae subcarpaticum, 400 m.s.m. prope pag. Jósvafő. 18. 05. 1958. Leg. et det. JÓZSEF ÚJHELYI. Sesleria hungarica ÚJHELYI, 1959 - Population 1: Comit, Borsod. Catena Bükk: in saxosis herbosis montis "Tarkő". Sol. calc.; alt. c. 900 m.s.m. Legit: 25.05.1916. JÁNOS HULJÁK. Population 2: Hungária sept.-orientalis. In montibus Bükk-hegység. In caespitibus saxi montis Szentléleki Látókő supra casam Szentlélek ad pag. Omassa prope opp. Miskolc, alt. 700 m.s.m. Exp. NO. Solo calcareo 2n=56! 21.05.1959. leg. JÓZSEF ÚJHELYI et TERÉZ ASBÓTH. Population 3: Hungária sept.-orientalis. In montibus Bükk hegység. In saxosis sylvis fagetis valle Sebes völgy, ad. pag. Ómassa, prope opp. Miskolc, alt. 600 m.s.m. Exp. N. Solo calcareo. 2n=56! 21.05.1959. Leg.: JÓZSEF ÚJHELYI et TERÉZ ASBÓTH, det.: JÓZSEF ÚJHELYI. Population 4: Material from the experimental garden originated from: Mts. Bükk, Bükk Na­ tional Park, prope opp. Felső-Hámor, in saxi montis "Puskaporos" 15.04.1983. Leg. et det.: DEZSŐ KOVATS took it out of the experimental garden: 18.04.1998. Population 5: Comit. Heves, prope pag. Bánkút, prope opp. Nagyvisnyó, in rupibus calcareis Ablakoskő, cca. 750 m.s.m. 18.05.1983. Leg. DEZSŐ KOVATS etTlBOR SZERDAHELYI, det.: DEZSŐ KOVÁTS. Population 6: Originated as before. Taken out of the experimental garden: 17.04.1986 Population 7: In montibus Bükk hegység, prope opp. Felsőtárkány, Csákpilis 19.05.1983. Leg. et det.: DEZSŐ KOVÁTS. Population 8: Material from the experimental garden originated from: Comit. Borsod-Abaúj- Zemplén, in montibus Bükk hegység, in monte Uppony prope opp. Dédestapolcsány, ad limites lacuum Lázbérc, 05.10.1982. Leg. et det.: DEZSŐ KOVÁTS. Taken out of the experimental garden: 14.04.1986.

RESULTS AND DISCUSSION

Distribution of culms according to internode numbers

The author made a comparision of the internode numbers of culms bearing panicles on 320 specimens from 13 populations. The comparision between the two Sesleria taxa was made with experimental probability (frequency), and the differ­ ences between the specimens collected from the original habitats (220 specimens from 9 populations) and those grown in the experimental garden (100 specimens from 4 populations), (Fig. 1, Tables 1-2). As regards the specimens grown in the original habitats the double internoded culms (culms with 2 internodes) are the most frequent and, in an interesting way, of the same rate, 64% in both Sesleria taxa. After that follow the culms with 3 internodes, also of the same percentrage in both Sesleria taxa: 27% level. Differ­ ence beetween the two closely related taxa is found at 9-9%. That means culms without node - culms with 1 internode - at Sesleria heufleriana (9%) and culms with 4 internodes at Sesleria hungarica (9%). Table 2. Distribution of culms by internode numbers

Sesleria heufl zriana Sesleria h ungarica number of from original from experimental from original from experimental internodes habitats garden habitats garden 1 10/110 = 9% 10/30 = 33% 0 0 2 70/1 10 = 64% 20/30 = 67% 70/110 = 64% 0 3 30/1 10 = 27% 0 30/1 l() = 27(/r 50/70 = 72% 4 0 0 10/110 = 9% 10/70= 14% 5 0 0 0 10/70= 14% sum % 100 100 100 100

At experimental garden conditions there are significant differences in the number of internodes between the two taxa (measuring 100 specimens from 4 pop­ ulations). Here in the case of Sesleria heufleriana 33% of the culms have not nodes; 67% of the culms have 2 internodes. In the case of Sesleria hungarica 72% of the culms have 3 internodes, 14% 4 internodes, and 14% even 5 internodes (Fig. 1). In this way at garden conditions there remain culms with 2 internodes in most cases at Sesleria heufleriana, but in contrast with this no culms with 2 internodes or culms without node can be found at S. hungarica. At this taxon culms with 3 internodes are the most frequent, even culms with 4 or 5 internodes are not rare. According to the foregoing observations and measurements it seems that a higher number of internodes is characteristic of Sesleria hungarica especially at garden conditions where the ecological differences are basically eliminated (Fig. 1). In a previous work, similarly to Sesleria, on two taxa of Phleum {Ph. hertolonii DE CANDOLLE = Ph. hubbardii D. KOVÁTS and Ph. pratense LINNAEUS) I found sig­ nificant differences in the number of internodes especially at garden conditions (KOVÁTS 1976, 1977).

Fig. 1. Distribution of culms by internode numbers Distribution of culms according to lengths and their internode lengths

Measuring 220 specimens from 9 populations from the original habitats shows that the fertile culms of Sesleria hungarica are ca. 10-15-20 cm higher than the culms of Sesleria heufleriana (Fig. 2, Tables 3-4). In accordance with this dif­ ference of lengths in culms, the lengths of internodes are also different at each level of the culms. The panicles of Sesleria hungarica are also longer than those of Sesleria heufleriana. In order to comparable the internodes at the proper levels, the culms with 2, 3, and 4 internodes are to be separately matched. In each case the lowest internodes of Sesleria hungarica are twice longer than those of Sesleria heufleriana. The upper­ most (longest) internodes of the culms, below the panicles, are 10-15 cm longer than in the case of Sesleria hungarica. The differences in the full lengths of culms - at both taxa - approximately correspond to the length differences and their ratio of the uppermost (longest) intednodes below the panicles (Fig. 2, Tables 3-4).

Table 3. Average data of panicle, culm, and internode lengths, ratio of the lengths of culms and uppermost internodes, ratio of the lengths of leaf blades and leaf sheats of Sesleria heufleriana average internode average cumulative uppermost ratio of leaves ratio of width of serial lengths of lengths of internodes the aver- serial the aver­ panicles numbers 1. internodes internodes free from age num- age lowest 2.3. leaf lengths of bers lengths uppermost sheaths culms and of leaf internodes uppermost blades internodes and leaf sheaths 0.8 cm 1 3 cm 3 cm 29 cm 46/43 cm 1 5 2.5/14 cm up­ permost leaves 2 4 8/6.5 cm 2 43 cm 46 cm 3 3 16+7/5 cm panicles 1.8 cm 1.8 cm 46/42 cm 4 2 22+7/4.5 cm 5 1 28+7/4.5 cm low­ est leaves 3 42 cm 46 cm 2 3.5 cm 4 cm 1 0.5 cm 0.5 cm I have found intravaginal innovation in both taxa with the exception of one population of Sesleria heufleriana at Szarvaskő (in population 2), in which I have found extravaginal innovation too.

Table 4. Average data of panicle, culm and internode lengths, ratio of the lengths of culms and uppermost internodes, ratio of the lengths of leaf blades and leaf sheats of Sesleria hungarica

average internode average cumula­ upper­ ratio of the leaves ratio of width of serial lengths of tive most average serial the aver­ panicles numbers internodes lengths of inter­ lengths of numbers age 1. lowest internodes nodes culms and lengths 2.3.upper­ free from uppermost of leaf most leaf internodes blades internodes sheaths and leaf sheaths

1 cm I 4 cm 4 cm 39 cm 57/53 cm 1 7 3/16 cm upper­ most leaves

2 6 4/9 cm 2 53 cm 57 cm 3 5 19/7 cm

panicles 2.2 cm 2.2 cm 4 4 25+'?/7 cm

62.5 /57 5 3 31+7/6 cm 3 57 cm 62.5 cm 6 2 40+7/5 cm

2 4.5 cm 5.5 cm 7 1 21+7/5 cm low­ est leaves

1 1 cm 1 cm 67.5/53 cm

4 53 cm 67.5 cm

3 8 cm 14.5 cm

2 5 cm 6.5 cm

1 1.5 cm 1.5 cm The number of leaves

The number of leaves measurable in the blooming period and immediately af­ ter that is usually 6 or 7. The greatest number of leaves is 9 among the investigated populaions, but occurring also only at garden conditions. In most of the cases the lowest 3 or 4 senile leaf blades are split, leaf sheaths reduced to fibres (DEYL 1946). According to my observations the upper (young) 1 to 3 leaves remain un­ broken and mesurable.

The ratio of lengths of leaf blades and leaf sheaths at different levels of the culm

Regarding the average data of leaf blade and leaf sheath lengths in both spe­ cies the lengths of leaf blades decrease at the culm from below to upwards, from senile leaves to juvenile leaves (Fig. 3, Tables 3-4). In contrast with this the lengths of leaf sheats change the other way round and consequently there is in­ crease at the culm from below to upwards, the sheaths of senile lower leaves are shorther, then they continually increase in lengths at a higher level, finally the leaf sheats of the youngest leaves are the longest, as usual in most grasses. The sheaths of the youngest, upermost leaves are on the average 5 times lon­ ger than the leaf blades (16/3 cm, or 14/2.5 cm) on average at both Sesleria taxa. This ratio turns to the contrary already at the second, youngest leaves from above

Fig. 2. Distribution of internode lengths by levels of culms (empty bar = S. hungarica, filled bar = S. heufleriana ) the culms of Sesleria heufleriana. Here the leaf blades are longer ( 1.5 cm) than the leaf sheaths (8/6.5 cm). Whereas at Sesleria hungarica on this level the lengths of leaf blades are not yet half the lengths of the leaf sheaths (4/9 cm). The change of ratio in lengths of leaf blades and sheaths takes place only at the third youngest leaves from above the culms of Sesleria hungarica (in ratio 19/7 cm), (Fig. 3, Ta­ bles 3-4). This change of ratio is already significant at the fourth youngest leaves from above the culms of both Sesleria taxa. At this level of culm the leaf blades are about 5 times longer than the leaf sheths (22+?/4.6cm, or 25+?/7cm). The lengths of leaf blades in most cases are uncertain because they are usually damaged, frag­ mentary. In this state of development, the lengths of culms are not completely mea­ surable, as has been mentioned above. The lengths of the further lower leaf sheaths do not decrease yet, or only to a lesser degree, and on the other hand the leaf blades still further increase in lengths rather than decrease at the bottom of the culms while they are measurable (Fig. 3, Tables 3-4).

Some other morphological observations and remarks

Though not in keeping with the average data, certain population and sample differences are observable from the data of the average lengths of the leaf blades and sheaths, and their ratio, first of all at Sesleria heufleriana. In the foregoing demonstrated relations, the data of the leaf blades and sheaths are different in each case at the different levels of the culms. For example, at two populations of Sesleria heufleriana collected at Szarvaskő (population 2) and Szádvár (popula-

Fig. 3. Ratio of the average length of leaf blades and leaf sheaths by different levels of culms tion 4) a few (second from above) youngest leaf blades and the next (third from above) youngest leaf blades leave the culms at the same level. At the population collected at Szádvár (population 4) a few third and sixth leaf blades leave the culms at the same level, consequently the lenghts of the leaf sheaths are the same. A few fifth leaf blades leave the culms higher than the blades of the third and sixth leaves, and though they are older than the third and sixth leaves, their leaf sheaths are lon­ ger. Such exceptions can be found as compared with the average data. In the foregoing already mentioned population (population 2) from Szarvaskő at Sesleria heufleriana a few older fourth leaves have longer sheaths than the youn­ ger third leaves have. Consequently a few older, lower leaf blades leave the culms at a higher level than the younger ones do. Naturally in this case the leaf sheaths of the fourth leaves open for the leaf blades of the third leaves, in order to be able to leave the culms According to DEYL (1946: 8): "Most characteristic is the length of the last culm leaf." I have not found significant differences in the lengths of the last culm leaves between the two closely related Sesleria taxa. According to my measure­ ments the lengths of the blades of the topmost (youngest) leaves are only slightly shorter (2.5 cm) at Sesleria heufleriana than at Sesleria hungarica (3 cm). The same aplies to the leaf sheaths, their lengths being 14 cm and 16 cm on the average at Sesleria heufleriana and Sesleria hungarica respectively.

Fig. 4. Morphological sructure of Sesleria heufleriana SCHUR with 2 internodes from original habitat At both Sesleria taxa there sprout young intravaginal branches (shoots) - even 3 or 4 on a culm - from the 2nd and/or the 3rd leaf sheaths from above of the culms during and immediately after the blooming period. All the senile, i.e. lower, leaves sprout from the stock at both Sesleria taxa. The uppermost - the youngest - leaf springs up from a node, from the top one if there are more than one, the leaf below from the node below, and when the nodes run out all the other leaves sprout from the stock. At Sesleria hungarica the breadth of senile leaves is about the double ofthat at Sesleria heufleriana, in the case of the former about 6 or 7 mm, in that of the lat­ ter 2.5 or 3 mm, rarely even 4 mm, as was the case in population 5 in montibus Tornai Karszt. The differences between the two Sesleria taxa are shown in two figures (Figs 4-5). To ensure comparability, all the materials have been taken from their original habitats, three populations having two internodes alike. The differences in mea­ surement are marked, and in addition, the length ratio of the leaf blades and sheaths at Sesleria heufleriana changes in the opposite direction at the second level from above of the youngest leaves, whereas at Sesleria hungarica it takes places only at the third level, as has been mentioned above. The figures also indicate - as has been experienced in practice - that in this stage of development there are two more - partly still measurable - senile leaf vestiges to be found at Sesleria hungarica than in the case of Sesleria heufleriana.

Fig. 5. Morphological structure of Sesleria hungarica ÚJHELYI with 2 internodes from original habitat SUMMARY

Besides the differences in measurement, a few less marked differences have also been found in the morphology of the two Sesleria taxa (S. heufleriana SCHUR and S. hungarica ÚJHELYI). Observation and measurement show that there is difference in the number of the internodes of the culms, first of all apparent in specimens grown under similar circumstances in an experimental garden, where the culms have had only 1 and (mostly) 2 internodes at Sesleria heufleriana, whereas at S. hungarica the culms have had mostly 3, but occasionally 4 or 5 internodes. Consequently there is not even overlaping in the number of internodes between the two taxa. In the case of the specimens that grew in their original habitats there is complete agreement in the percentage of the number of the culms with 2 and 3 internodes between the two taxa. Difference is manifest between them at 9 per cent: to that tune the specimens of Sesleria heufleriana have no nodes as opposed to S. hungarica in whose case culms with 4 internodes can be found to that amount. Sesleria hungarica is a more robust and taller plant than Sesleria heufleriana, as regards both the culm and its internodes, sometimes double in length. Compared with the average height of the specimens that grew in their original habitats, this difference in measurment is 10 to 15 cm, and in proportional to the height (length) of the uppermost internodes. Contrary to the average data, the length rates of the leaf sheaths and leaf blades change at different levels of the culm in the case of the two taxa. At Sesleria heufle­ riana the blades of the youngest leaves second from above are already longer than the leaf sheaths, whereas at Sesleria hungarica it takes place only in the case of the youngest leaves third from above. Though the average data have been taken into consideration, no marked dif­ ference has been found in the length of the last culm leaf (DEYL 1946: 8), nor in the length of the leaf blades, nor in that of the leaf sheaths. Through the above observations, measurements, and their comparison, au­ thor has intended to contribute to the separation of the two Sesleria taxa.

Acknowledgements - I gratefully acknowledge my indebtedness to my sons DEZSŐ KRISTÓF KOVÁTS and SZABOLCS MIKLÓS KOVÁTS, who rendered me invaluable help with computer editing, especilly with the tables and graphs. REFERENCES

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