Studia bot. hung. 34, pp. J 2 J-/26, 2003
T H O U S A N D SEED W E I G H T RECORDS OF SPECIES F R O M
T H E F L O R A OF H U N G A R Y , I . M O N O C O T Y L E D O N O P S I D A
P . C S O N T O S1, J. T A M Á S2 and L . B A L O G H3
'MTA-ELTE Research Group on Theoretical Biology and Ecology
Eötvös Loránd University, Pázmány Péter sétány. 1/c, Budapest, H-I1J7, Hungary
2Department of Botany, Hungarian Natural History Museum
H-1476 Budapest, Pf 222, Hungary 'Department of Natu red History, Savaria Museum Kisfaludy S. it. 9., Szombathely, H-9700, Hungary
New thousand seed weight records are reported for 38 members o f the Hungarian monocotyledonous flora. Seed samples were harvested from natural populations, kept dry t i l l the end o f the vegetation period, then their seed mass were determined. One to three hundred-seed lots were used for measure ments w i t h accuracy o f 0.1 mg (up to 30 g) and 0.1 g (above 30 g). M o s t o f the new records corre spond to already k n o w n rules o f seed m o r p h o l o g y and ecology. I n some cases the new records re quired i n d i v i d u a l interpretation as they d i d not support general trends. W h e n thousand seed weight records were plotted against naturalness o f species a slight positive correlation occurred indicating that monocots w i t h larger seed mass values often received higher rank o f naturalness values. W i t h one table and one figure.
Key words: diaspore weight, Hungarian flora, Monocotyledonopsida, naturalness value, seed weight
I N T R O D U C T I O N
The role of seeds and fruits in the dispersal and establishment of species and
the recognition of their importance in vegetation restoration resulted an increasing
interest on the reproductive propagules of higher plants( H O W E
1986,B A K K E R
al. 1996,L E V E Y
et al. 2002). Vegetation scientists reported new ecological rules
derived from size and shape of seeds and fruits, that requires an extensive database
of raw data of species' reproductive propagules. Also for appropriate operations by
environment engineers in vegetation restoration projects these databases offer use
ful information. T o serve these challenges seed morphological and seed ecological
databases were established in the last decades( G R I M E
et al. 1981,T H O M P S O N
al. 1997,V A N S L Á G E R E N
T o j ó i n the international trend completion of a seed size and shape database of
the Hungarian flora was decided( C S O N T O S
1998). For filling up the database we
used three types of data sources: data published in the literature, measurements of
seeds received from botanical gardens and measurements of freshly collected
seeds originated from spontaneous populations. Evaluation of the seed weight da
tabase w i t h application examples were given byC S O N T O S
andT A M Á S
(2001 ) and
C S O N T O S
(2002). In this paper we publish data o f 38 monocotyledonous species o f
the Hungarian flora for what thousand-seed-weights are first reported.
M A T E R I A L S A N D M E T H O D S
Seeds or fruits o f species were collected in the f i e l d at mature stage. For species w i t h easy-to-harvest seeds several hundreds were collected (3 hundreds being the m i n i m u m ) , otherwise at least 100 seeds were taken. Whenever it was possible, attention was paid to collect pooled seed sam ples from several mother plants o f the same stand to avoid bias caused by maternal effects ( K a z i n c z i and M a g y a r 2003). In case o f fleshy fruits the seeds were removed in the lab d u r i n g the subsequent days then cleaned and d r i e d at r o o m temperature. A l l seed samples were put i n labelled paper bags and stored at room temperature t i l l the end o f the vegetation period (late October - early N o v e m b e r ) . For measurements hundred-seed lots (or hundred-diaspore lots) were prepared. Damaged seeds were not considered. Three lots were measured for the majority o f species, i n some cases t w o or one. U p to 30 g the accuracy was 0.1 m g (using K E R N - 4 1 0 automatic chemical balance), whereas above 30 g it was 0.1 g ( N A G E M A digital weighting machine). Thousand seed weight o f a species were calculated from the average weight o f the seed lots. I n case o f six species seeds were collected from spontaneous populations by the Botanical Garden o f the Ecological and Botanical Research Institute o f the H u n garian Academy o f Sciences ( V á c r á t ó t ) . These samples were subjected to the same operations as de scribed above.
As an example for seed mass data application in botanical research a pilot test was made re garding the relationship o f seed mass and naturalness values (sensu B O R H I D I 1993) o f the 38 species discussed in this paper. For statistical evaluation the ordinal scale o f naturalness values was d i v i d e d into the f o l l o w i n g three parts: " l o w " , from - 2 to + 2 ; " m e d i u m " , from +3 to + 5 ; " h i g h " , from +6 to + 10; then the seed mass data o f the " l o w " and " h i g h " groups were compared i n t-test ( S v á b 1981).
RESULTS A N D D I S C U S S I O N
New thousand seed weight records are given for 38 monocotyledonous spe
cies o f the Hungarian flora representing the following taxonomic groups: Poaceae:
12 spp., Cyperaceae: 9 spp., Liliaceae (sensu lato): 7 spp., other monocots: 10 spp.
(Table 1). Seed samples are stored in the seed collection of the Botanical Depart
ment of the Hungarian Natural History Museum (Collectio seminum et fructum, BP).
Most of the new records correspond to already known trends o f seed ecology.
Seed mass of three rush species (Juncus atratus, J. conglomeratus and J. gerardii)
proved to be extreme low (Table 1) similar to most o f their congeners. Several
small seeded Juncus species including J. conglomeratus are reported to establish
long term persistent seed bank( T H O M P S O N
et al. 1997) and / . gerardii was already
classified as having well developed seed bank w i t h considerable persistence in
various European vegetation types( E R K K I L A
1998,W O L T E R S
andB A K K E R
Therefore we can assume a similar soil seed bank behaviour f o r i , atratus as well.
T a b l e 1. Seed weight records first published to 38 species o f the Hungarian flora. S N = number o f seeds measured, W = thousand seed w e i g h t in grams
Species name O r i g i n o f the seeds Date S N W ( g )
Potamogeton pectinatus L . D i n n y é s , D i n n y é s K a j t o r i
30.09.2000 100 6.280
Allium rotundum L . subsp. V á c r á t ó t 2000 300 0.574
waldsteinii (G. D o n ) Richter
Allium sphaerocephalon L . N a g y k o v á c s i , N a g y - S z é n á s ,
1995 300 0.766
Hemerocallis lilio-asphodelus L . Vas county, Nárai 1996 200 52.4
Ornithogalum comosum L . C s i k i M t s , T ö r ö k u g r a t ó 18.07.1996 300 3.511
Polygonatum latifolium (Jacq.) Desf. C s i k i M t s ( K F K I ) 20.09.1998 300 23.90
Scilla autumnalis L . V á c r á t ó t 2000 100 2.090
Scilla vindobonensis Speta K u p , K u p i - e r d ő1 23.05.1996 300 5.056
Galanthus nivalis L . K u p , K u p i - e r d ő1 23.05.1996 300 6.1 12
Crocus heuffelianus Herbert K u p , K u p i - e r d ő1 10.06.1996 300 5.664
Crocus reticulatus Stev. K o c s1 23.05.1996 200 4.921
Iris sibirica L . K ő s z e g , A l s ó - r é t e k 05.09.1995 60 15.80
Juncus atratus K r o c k e r V á c r á t ó t 2000 100 0.032
Juncus conglomeratus L . Dunaremete, Sorjás pasture 1995 100 0.01 1
Juncus gerardii L o i s . S á r k e r e s z t ú r , Lake S á r k á n y 22.08.1996 300 0.044
Limodorum abortivum ( L . ) Sw. Budapest, Ö r d ö g - o r o m 02.07.2000 100 0.002
Neottia nidus-avis ( L . ) Rich. H o l l ó k ő , near F ö l d v á r 12.09.1998 300 0.008
Carex alba Scop.2 Buda M t s , K i s - S z é n á s 08.08.1999 200 3.271
Carex cuprina ( S á n d o r ) Nendtvich S z á z h a l o m b a t t a , fish-ponds 1995 300 1.218
Carex elata A l l . S z á z h a l o m b a t t a 1995 300 0.414
Carex hallerana Asso V á c r á t ó t 2000 200 2.702
Carex liparicarpos Gaud. Budapest, T ü n d é r - s z i k l a 15.08.1996 300 3.257
Holoschoenus romanus ( L . ) Fritsch V á c r á t ó t 2000 300 0.090
subsp. holoschoenus ( L . ) Greuter
Schoenoplectus litoralis (Schrad.) Velence 1995 300 0.974
Schoenoplectus triqueter ( L . ) Palla Velence 1995 300 3.012
Scirpus syivaticus L . F e l s ő s z ö l n ö k , wet meadow 1995 300 0.089
Bromus ramosus Huds. B ö r z s ö n y M t s 02.07.1998 200 4.36
Calamagrostis epigeios ( L . ) Roth near village N a g y k o v á c s i 10.10.1998 200 0.064
Cenchrus incertus M . A . C u r t i s3 V á c r á t ó t 2000 100 44.4
Eleusine indica ( L . ) G ä r t n . Budapest, I X . district 22.10.1996 300 0.363
T a b l e 1 (continued)
Species name O r i g i n o f the seeds Dale S N W (g)
Hordelymus euwpaeus ( L . ) Jessen Beech forest above 05.08.1998 300 7.32
P i l i s s z e n t l á s z l ó
Phleum phleoides ( L . ) Karsten Gerecse M t s , Nagy-Gete 15.08.1998 200 0.075
Poa nemoralis L . B ö r z s ö n y Mts 11.08.1998 100 0.13
Puccinellia limosa (Schur) Holmberg H o r t o b á g y 1995 300 0.132
Stipa eriocaidis Borb. Budapest, S z é c h e n y i - h e g y 13.06.1999 100 25.30
Tragus racemosus ( L . ) Desf. P o r p á c , railway station 28.08.1998 100 0.429
Triticum monococcum L . Szombathely, Beithe garden 27.08.1998 150 27.13
1 = collected by S. B a r a b á s ; 2 = grain crop w i t h silicle (utriculus); 3 = i n c l u d i n g spiny grain coat.
The two Scilla species can be mentioned as good example for seed mass dif
ference of congeners preferring habitats with contrasting light climate (CSONTOS
1998). In accordance w i t h earlier observations, here again the heliophilous mem
ber (Scilla autumnalis) has the smaller seed mass.
Some of the new records do not support the general rules and require special
interpretation. The relatively large seed mass of the small-sized sedge, Carex
liparicarpos contradict to the sunny habitats where this species is a member o f sev
eral plant communities: Asplenia rutae-murariae-Melicetum ciliatae,
Cleistoge-no-Festucetum rupicolae, Astragale? aiistriacae-Festucetum sulcatae, Stipe?
erio-cauli-Festucetum pallentis, Seseli leucospermi-Festucetum pallentis (PENKSZA et
al. 1995, BORHIDI and SÁNTA 1999). This can probably be explained by extreme
high level of abiotic stress - mainly drought - in these habitats, that may support
larger seed mass governed by higher rate of seedling survival. Weedy species are
known to have relatively small seeds that are produced in large numbers to serve
the "colonisation strategy" of their seed dispersal behaviour (cf. HOWE 1986). In
this paper we report considerable diaspore mass for Cenchrus incertus (see Table
1) an alien graminoid weed of the Hungarian flora. A possible interpretation of it is
the epizoochorous dispersal of Cenchrus what may permit heavy diaspores, while
ability for long distance dispersal is still retained.
Finally, we made a pilot test on the relationship of seed mass and nature con
servation status of the 38 species treated in the paper.
The ranking system of BORHIDI (1993) was used for classifying species ac
cording to their naturalness values. In Borhidi's system the lowest ranked "aggres
sive alien species" (invadors) score - 3 , whereas the highest score, +10, was given
to rare specialist species that are represented in the Hungarian flora with 1 to 5 pop
ulations only. (In our case the naturalness values varied between - 2 and +10.) The
seed mass records were log-transformed to ensure normal distribution o f data.
When the logarithm of thousand seed weight data were plotted against natu
ralness values (Fig. 1), a slight positive correlation occurred between the two
vari-O 05 O <U TJ CD CD CO TJ c (Ö CO o -2 -1 0 1 2 3 4 5 6 7
Naturalness value of species
F i g . 1. Seed mass and naturalness value o f 38 monocotyledonous species o f the Hungarian flora. E m p t y squares indicate posititions o f Orchidaceae species.
ables. (Orchids, though indicated, were not encountered because their seeds have
different anatomy with lack o f endosperm, thus they should not be discussed to
gether with the "normal" seeded species.) The statistical evaluation by r-test con
firmed a significantly higher seed mass o f species w i t h " h i g h " naturalness values
compared to that of " l o w " naturalness values ( n( l o w )
= 6; n( h i g h )
= 11; p < 0.1 ; critical
t '-value = 2.006; calculated f-value = 2.358). We found the results o f the pilot test
interesting and thought provoking, that may reflect to both underlying ecological
phenomena and human attitude in evaluating species. Larger seed size indicate
more pronounced K-strategist behaviour of a species and it is often linked with
higher sensitivity to habitat disturbances that may cause the rarity of those species
thus involving a higher value given by botanists. For better interpretation o f the re
lationship between seed mass, naturalness and rarity o f a species further studies are
required using larger set o f species w i t h a detailed analysis of their function in the
* * *
Acknowledgements - W e are indebted to several colleagues for their contribution i n collecting seed samples. M a n y thanks are due to H e d v i g Punka for the accurate handling o f the samples, and to E r z s é b e t Szurdoki and I s t v á n R á c z for their comments on the manuscript. Financial support f r o m the Hungarian Scientific Research Fund ( O T K A T025350 and T 0 3 7 7 3 2 ) , and S z é c h e n y i I s t v á n Fellow ships grant ( S Z Ö 372/2002, given to the first author) is greatly acknowledged.
B A K K E R , J . P . , P O S C H L O D , P., S T R Y K S T R A , R. J., B E K K E R , R. M . and T H O M P S O N , K . ( 1 9 9 6 ) : Seed banks and seed dispersal: important topics i n restoration ecology. - Acta Bot. Neerl. 45(4): 4 6 1 - 4 9 0 .
B O R H I D I , A . (1993): A magyar flóra szociális magatartási típusai, természetességi és relatív ökológiai énekszámai. [Social behaviour types o f the Hungarian flora, its naturalness and relativ ecological indicator values.] - A K T M T e r m é s z e t v é d e l m i Hivatala é s a Janus Pannonius T u d o m á n y e g y e t e m k i a d v á n y a , P é c s .
B O R H I D I , A . and S Á N T A , A . (eds) ( 1999): Vörös könyv Magyarország növénytársulásairól 1-2. [Red data book o f the Hungarian plant communities 1-2.] - A K Ö M T e r m é s z e t v é d e l m i H i v a t a l á n a k T a n u l m á n y k ö t e t e i 6., T e r m é s z e t B Ú V Á R A l a p í t v á n y K i a d ó , Budapest.
C S O N T O S , P. (1998): The applicability o f a seed ecological database ( S E E D ) in botanical research. -Seed Science Research 8(1): 4 7 - 5 1 .
C S O N T O S , P. (2002): A magökológiai adatbázis és alkalmazhatósága a botanikai kutatásban. [The seed ecological database and its applicability i n botanical research.] - I n : P A P P , L . and B O R H I D I , A . (eds): S z u p r a i n d i v i d u á l i s B i o l ó g i a i K u t a t á s o k . [Studies in supraindividual b i o l ogy.] M T A - Ö B K I , V á c r á t ó t , pp. 2 9 - 3 6 .
C S O N T O S , P. and T A M Á S , J. (2001): Alkalmazási példák az adatbázisok használatára. [ A p p l i c a t i o n examples for the use o f databases.] - I n : C S O N T O S , P. (ed.): A t e r m é s z e t e s magbank k u t a t á s á n a k m ó d s z e r e i . [Methods in natural seed bank studies.] Scientia K i a d ó , Budapest, pp. 7 7 - 8 1 .
E R K K I L A , H . M . J. B . (1998): Effect of different treatments on the seed bank o f grazed and ungrazed Baltic seashore meadows. - Can. J. Bot. 76(7): 1188-1 197.
G R I M E , J. P., M A S O N , G . , C U R T I S , A . v . , R O D M A N , J., B A N D , S. R., M O W F O R T H , M . A . G . , N E A L , A . M . and S H A W , S. (1981): A comparative study o f germination characteristics in a local Bora. -J. Ecol. 69: 1017-1059.
H O W E , H . F. (1986): Seed dispersed by fruit-eating birds and mammals. - I n : M U R R A Y , D . R. (ed.): Seed dispersal. Academic Press, N e w Y o r k , pp. 123-189.
Kazinczi, G . and Magyar, L . (2003): A g y o m m a g v a k nyugalmi á l l a p o t a és c s í r á z á s ö k o l ó g i á j a I I . A gyommagvak s z á n t ó f ö l d i c s í r á z á s a és a dormancia fajon belüli e l t é r é s e . [ D o r m a n c y and germi nation ecology o f weed seeds I I . Field germination and variations w i t h i n a species.] - Magyar gyomkutatcis és technológia 4(1): 3 - 1 7 .
L E V E Y , D . J., S I L V A , W . R. and G A L E T T E M . (eds) (2002): Seed dispersal and frugivory: ecology, evolution and conservation. - C A B I Publishing, W a l l i n g f o r d .
P E N K S Z A , K . , B E N Y O V S Z K Y , B . M . , Ö T V Ö S , E. and A S Z T A L O S , J. ( 1 9 9 5 ) : P h y t o s o c i o l o g i c a l studies o f the c l i f f F e h é r - s z i r t , near K e s z t ö l c , Hungary. - Acta Bot. Hung. 3 9 ( 1 - 2 ) : 7 1 - 9 5 .
SVÁB J. (1981): Biometriai módszerek a kutatásban. [ B i o m e t r i c a l methods in research w o r k . ] - 3rd ed., M e z ő g a z d a s á g i K i a d ó , Budapest, 557 pp.
T H O M P S O N , K . , B A K K E R , J. P. and B E K K E R , R. M . (1997): The soil seed banks of North West Eu rope: methodology, density and longevity. - Cambridge University Press, Cambridge. V A N S L Á G E R E N , M . W . (2003): The M i l l e n n i u m Seed Bank: b u i l d i n g partnerships in arid regions for
the conservation o f w i l d species. - J. Arid Environm. 54(1): 195-201.
W O L T E R S , M . and B A R K E R , J. P. (2002): Soil seed bank and driftline composition along a successional gradient on a temperate salt marsh. - Appl. Veg. Sei. 5(1): 5 5 - 6 2 .