Relationship of animals to the cyclamen Cyclamen fatrense Halda et Soják: pollinators, consumers and occasional visitors Peter Turis1 & Ľubomír Vidlička2,3 1
The administration of the Low Tatras National Park, Lazovná 10, SK-97401 Banská Bystrica, Slovakia; e-mail: [email protected] 2 Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, SK-84506 Bratislava, Slovakia and 3 Department of Biology, Faculty of Education, Comenius University, Moskovská 2, Bratislava, Slovakia; e-mail: [email protected]
Abstract: The West-Carpathian endemic Cyclamen fatrense is a forests species. Its distribution area lies at the northern margin of distribution area of the Cyclamen genus. Several studies have focused on the pollinators and consumers of representatives of this genus, but they mention only some more important animal species. The aim of our study was to determine the species spectrum of animals having a relationship to C. fatrense in its natural environment and to deﬁne mutual relationships between this plant and animals based on comparison of phenologic manifestations of cyclamen and known ecologic data about its visitors. In course of diﬀerent phenophases of C. fatrense we recorded 127 invertebrate taxa on this plant. Among them 7 taxa were identiﬁed as consumers, 29 taxa as potential consumers, 5 taxa as potential pollinators and 91 had an indiﬀerent relationship to the cyclamen. Damaging or direct consummation of cyclamens by vertebrates has not been observed. Key words: Cyclamen fatrense; pollinators; consumers
Introduction Species of Cyclamen L. (Primulaceae) are distributed predominantly in the Mediterranean area. From among 20 known species (Davis et al. 1999) the northernmost occurring species are Cyclamen purpurascens Mill. and Cyclamen fatrense Halda et Soják. While C. purpurascens grows on a wide territory from the Swiss Jura and West Alps to Balkan Mts (Meusel et al. 1978), C. fatrense is considered as a subendemic of the small mountain range Veľká Fatra in West Carpathians in Slovakia (Kliment 1999). Both species have very similar shape and diﬀer from each other mainly by form and color of leaves (Halda & Soják 1971). Relationships of animals to the Cyclamen species are presented on several levels. Most authors mention herbivory of molluscs, Diptera and caterpillars, which eat leaves, fruits or tubers of cyclamens in natural habitats (Hildebrand 1898; Hering 1957; Aﬀre et al. 1995; Reiprich 2001). L¨ udi (1975) mentions damaging of buds, leaves, tubers and roots of cultivated individuals by aphids, caterpillars, thrips, beetle larvae, nematodes, molluscs and rodents. Other animals are reported in connection with the Cyclamen genus as pollinators in relation with their reproduction. In wild plants, this role is only rarely played by bees, bumblebees and ants. Hildebrand (1898) observed these hymenopterans on ﬂowers of the cultivated cyclamens. More frequently the cyclamens are pollinated by night moths, thrips or hoverﬂies (L¨ udi 1975; c 2013 Institute of Zoology, Slovak Academy of Sciences
Aﬀre et al. 1995; Aﬀre & Thompson 1997; SchwartzTzachor et al. 2006). Several authors report on animals participating on spreading fruits and seeds of the cyclamen. According to Kerner (1898), the spiral-like rolled stalks of fruits are, at occasional attaching, spread by some vertebrates. In connection with seed propagation, the myrmecochoria is mostly mentioned (Hildebrand 1898; Nordhagen 1932; Kovanda 1992). Aﬀre et al. (1995) even discovered one ant species participating on seed transport. In connection with occurrence of animals on C. fatrense only a ﬁnding of the weevil Phyllobius argentatus (L., 1758) (Coleoptera: Curculionidae) in an empty capsule has been published (Turis 2008). In the framework of the running ecobiological investigation of C. fatrense, it is important to know as wide as possible spectrum of animals visiting this plants in order to deﬁne: (i) occasional visitors without obvious consequences for the plant, (ii) visitors with more signiﬁcant consequences for the plant (phytophages damaging vegetative or generative organs), (iii) pollinators. Brief description of Cyclamen fatrense Cyclamen fatrense is a hemicryptophyte or geophyte bound to forests of submountain and mountain zone. Usually a single stem reaching to the layer of leaves fallen from the surrounding trees, grows from the tuber. From the terminal part of stem the permanent leaves
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grow and, in summer, striking, moderately smelling ﬂowers in various shade of rose color. Flowering lasts approximately from July until late October. Peduncle of fertilized ﬂowers spiral-like rolls on the ground surface and the fruits ripe in next summer. Absence of elaiosome on seed is important for potential relationships with animals. Production of nectar in ﬂowers has not been observed. In Cyclamen repandum Sm., Cyclamen hederifolium Aiton and Cyclamen persicum Mill. nectar is not produced (Aﬀre & Thompson 1997; SchwartzTzachor et al. 2006), while Aﬀre et al. (1995) mentioned its absence in the whole Cyclamen genus. Fig. 1. Localization of the research area (grey circle – wider surrounding of the Motyčky village).
Material and methods Invertebrates on Cyclamen fatrense were collected individually in 2006–2007. Observations and catches were carried out in the snow-free period (approximately March – November), usually once a week. The invertebrates were sampled (leg. P. Turis) from the above-ground parts of plants during light period of the day and were preserved in alcohol and distributed to specialists for identiﬁcation. Relationships of vertebrates (forest rodents, wild boars, red deer, roe deer etc.) to cyclamens were evaluated on base of visual observations in 2006–2008. Nomenclature of invertebrate taxa and their systematic arrangement in the Table 1 was used according to Fauna Europaea (de Jong 2010). Trophical characteristics of individual taxa are based on oral information from the specialists and on published data (Buchar et al. 1995; Buchar & Kůrka 1998; Doskočil 1977; Jasič 1984, Kočárek et al. 2005; Pﬂeger 1988; Zahradník 1987). Relations of the recorded animals to C. fatrense given in Table 1 are characterized according to trophic relation of the caught developmental stages and on direct ﬁeld observations. We consider the zoophages, parasites and saprophages to have an indiﬀerent relation to cyclamens, similarly as phytophages obviously specialized on other plant species (except for cyclamen). Other phytophages are taken as potential consumers, while the phytophages directly observed to eat leaves or ﬂowers of cyclamens are taken as consumers.
When evaluating signiﬁcance of melliphages for pollination we took in consideration a period of their occurrence in relation to the period of cyclamen ﬂowering. Study area Observation and collection of animals were carried out mainly in the eastern part of the species area in the Veľká Fatra Mts and the Starohorské vrchy Mts. The studied territory has an area of 1,763 ha and lies at altitudes of 450– 1100 m (Turis & Žiačik 2008). The broadleaved and mixed forests belonging phytocenologically into association Carici albae-Fagetum Moor 1952 (Kanka et al. 2008) are dominant. The major part of research was performed in a wider surrounding of the Motyčky village (48◦ 50–53 N, 19◦ 7–12 E, Central Slovakia, Banská Bystrica district, 680 m a.s.l.; Fig. 1).
Results During the two-year investigation we recorded 127 taxa of invertebrates (107 species and representatives of further 12 genera, 7 families and 1 order, which could not be precisely identiﬁed) (Table 1). We recorded
Fig. 2. Survey of orders and number of invertebrate taxa recorded on Cyclamen fatrense in years 2006–2007.
Animals on Cyclamen fatrense
Table 1. Systematic survey of invertebrates recorded on Cyclamen fatrense in 2006–2007. Recorded on
Abbreviations: L – leaves, Fl – ﬂowers, Fr – fruits, FP – ﬂowering period, EFP – extra ﬂowering period, TR – trophic relations, RP – relation to the plant, P – phytophage, Z – zoophage, S – saprophage, M – melliphage, Po – polyphage, Par – parasitoid, PS – phytosaprophage, A – aphidophage, PC – potential consumer, C – consumer, I – indiﬀerent, PP – potential pollinator, ? – indeterminate classiﬁcation.
Fig. 3. Number of invertebrate taxa recorded on Cyclamen fatrense in 2006, 2007 and number of taxa common for both studied years.
mainly consumption of tubers, leaves and ﬂowers. The richest in species were Diptera (21 taxa), Lepidoptera (17 taxa), Collembola and Pulmonata (16 taxa each) (Fig. 2). In both years of investigation approximately equal number of taxa were recorded (Fig. 3). A considerably higher number of taxa were found on leaves than on ﬂowers and fruits (Fig. 4). This is obviously connected with diﬀerent lasting of individual parts of the plant and their spatial distribution. During ﬂowering and extra ﬂowering period almost equal number of taxa were recorded (Fig. 5).
Consumption of leaves and ﬂowers of C. fatrense by invertebrates was observed only to a limited extent. We consider only 7 among all 127 taxa as consumers. The leaves were eaten by caterpillars of Xestia xanthographa and slugs of the genus Deroceras Raﬁnesque, 1820, after which excrement pieces remained on the leaf blade. Flower corollas were consumed by caterpillars of Eupithecia subfuscata, Eupithecia vulgata and Idaea aversata. Flower stamens were eaten by caterpillars of Clepsis senecionana. We consider the melliphagous beetle Meligethes denticulatus found in the ﬂower as a con-
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Fig. 4. Number of invertebrate taxa recorded on leaves, ﬂowers and fruits of Cyclamen fatrense in 2006–2007.
Fig. 5. Number of invertebrate taxa recorded during ﬂowering period, extra ﬂowering period and during both periods together on Cyclamen fatrense in 2006–2007.
sumer of pollen grains. Further 29 taxa we classiﬁed as potential consumers. This group consists of phytophagous species, which have not been observed to feed on a part of cyclamen, as well as the melliphagous psocid Caecilius despaxi and moth Micropterix calthella caught out of the cyclamen ﬂowering period. Five recorded taxa are classiﬁed as potential pollinators (the investigation was not focused on experimental veriﬁcation of pollination). First of all they are represented by the thrips Frankliniella intonsa and Taeniothrips picipes frequently and abundantly occurring on the ﬂowers, as well as the sap beetle Meligethes denticulatus. Two recorded melliphagous species, the psocid Caecilius despaxi and moth Micropterix calthella, also might participate on cyclamen pollination in the case of occurrence in the ﬂowering period of this plant.
Further development of caterpillars of the moths Clepsis senecionana, Philedonides rhombicana, Pseudoips prasinana and Syndemis musculana was also connected with cyclamen. Their pupation in the tube-like rolled leaf blades cause damages of assimilation organs of cyclamens, but only in a negligible measure. The cultivated seedlings and juvenile individuals of C. fatrense were strongly eaten by slugs Arion rufus (O.F. M¨ uller, 1774). This synanthropic slug has not been found in cyclamen populations in the natural habitats. Trophical relations of vertebrates to C. fatrense are occasional. Eating of leaf blades or tubers by larger herbivores (red deer, roe deer, hares, wild boars etc.) has not been observed at all. Consumption of leaf blades on some individuals, observed in a single case, probably originated from forest rodents. In the soil we found
Animals on Cyclamen fatrense access galleries leading up to the tuber, which were browsed by the rodents. Discussion Studies of other authors dealing with relationships of animals to diﬀerent species of cyclamens in natural habitats do not present rich species spectra of the taxa recorded. Aﬀre et al. (1995) consider slugs as significant consumers of ﬂowers and fruits of the endemic species Cyclamen balearicum Willk. In C. fatrense, the slugs ate stronger only leaves, namely in the period immediately after snow melting. It is probably caused by shortage in oﬀer of over-wintered green leaves in forests habitats at that time, when leaves of few broadleaved herbs (e.g., Asarum europaeum, C. fatrense, Viola reichenbachiana, locally Soldanella carpatica) are available. In the growing season, predominantly moth caterpillars participate on consumption and damaging of the above-ground plant parts of cyclamens. Hering (1957) and Reiprich (2001) also consider caterpillars of Cnephasia chrysantheana (Duponchel, 1843), Cnephasia incertana (Treitschke, 1835), Cnephasia stephensiana (Doubleday, 1849) and Cnephasia terebrana Amsel, 1935 as consumers of cyclamen leaves. We do not consider the presence of thrips on cyclamen as occasional and suppose them to suck the pollen grains. The pollen grains can attach on bodies of these less mobile animals when they are mowing in open ﬂowers and searching for food (Annand 1926). It has doubtless a great signiﬁcance for reproduction of the cyclamens. Ability of thrips to move on relatively large distances by means of anemochory (Rhainds & Shipp 2003; Fedor et al. 2008) makes possible transfer of pollen grains between several individuals in cyclamen’s populations and helps to pollinate them. The sap beetle Meligethes denticulatus also can have such ecological signiﬁcance for C. fatrense, but its occurrence frequency on cyclamens is negligible in comparison with the thrips. Schwartz-Tzachor et al. (2006) also considers thrips to be the commonest pollinators of the Cyclamen persicum populations in Israel (thripophily). In contrast to it, the beetles have not been mentioned as pollinators of the Cyclamen genus. Schwartz-Tzachor et al. (2006) consider the little moth Micropterix elegans Stainton, 1867 as the speciﬁc pollinator of C. persicum. In C. fatrense, we recorded the congeneric melliphagous species Micropterix calthella, but we classify it among the potential consumers and pollinators, because of its occurrence out of the ﬂowering period. In spite of the relatively large, strikingly colored and smelling ﬂowers of C. fatrense we did not observe any bees or bumblebees visiting its ﬂowers. This is more or less in accordance with results of other authors studying pollinators of diﬀerent cyclamen species (Aﬀre et al. 1995; Aﬀre & Thompson 1997; SchwartzTzachor et al. 2006). The presupposed lost of attractiveness of cyclamen ﬂowers for these pollinators is usually explained by historical causes (climatic changes,
523 glaciations, tectonic movements – Schwartz-Tzachor et al. 2006). In beach forests with C. fatrense, the bees and bumblebees forage only during springtime, when majority of herbs (Ajuga genevensis, Dentaria bulbifera, D. enneaphyllos, Fragaria vesca, Lathyrus vernus, Mercurialis perennis, Soldanella carpatica, Valeriana tripteris, Viola reichenbachiana etc.) are blooming (Hegi 1924, 1927; Beattie 1972; Blažyt˙e-Čereškien˙e et al. 2012). In summertime these pollinators ﬁnd food on meadowland, where quantity of ﬂowers is blooming at that time. During gradual blooming of the cyclamen (from July to September), beach forest does not attract these insects because of deﬁciency of food. The cyclamen allure of potential pollinators by pollen productivity. Therefore the transport of pollen grains is provided mostly by melliphages. Probably for the same reason we did not record the presence of adult butterﬂies feeding usually on nectar. Eating of cyclamen tuber by wild boars (Sus scrofa L., 1758) has been mentioned already in the older period. Some authors (e.g., Hildebrand 1898; Polívka 1901) probably based this statement on using of tubers of Cyclamen graecum Link., which was called by ancient Greek physicians as „bread for pigs“ (L¨ udi 1975). Translation of this name is also reﬂected in the popular names of cyclamens in Germans (“Saubrot”), English (“Sowbread”), Czech (“sviňský chléb”) or Slovak (“svinský chlieb”). However, we did not at all record digging out and consumption of the tubers of C. fatrense. We did not record the consumption of cyclamen by other large vertebrates. Damage of the tubers by rodents was an exception. The cyclamens are probably protected against consumption by most animals by production of diﬀerent types of poisonous glycosides. These toxins have been detected also in the tubers (Reznicek et al. 1989). The survey of animals recorded on C. fatrense includes only the species observed by light period of day. Occurrence of some specialized night species is not probable. Aﬀre et al. (1995) have studied insects associated with Cyclamen balearicum over the whole day but did not record any night species. Conclusions The animals recorded on C. fatrense in the eastern part of its distribution area attack only a small part of individuals in its populations and do not cause extensive damages with lethal consequences. The recorded consumers do not represent a serious danger. Lose of assimilation organs caused by eating of leaf stalks by animals occurs only sporadically, but the plants damaged in this way can survive the whole growing season in dormancy due to the tubers longevity. Damaging of seedlings or tubers of very small juvenile plants could have a negative impact. Only few animal species recorded on C. fatrense have a non-occasional relationship to this plant species. Among 127 taxa, the consumers represented 5.5%, po-
524 tential consumers 22.8% and potential pollinators only 3.9%. Other animals (zoophages, saprophages and specialized phytophages) have an indiﬀerent relation to this plant. Acknowledgements We thank the following specialists for identiﬁcation of animals: P. Bitušík (Diptera: Chironomidae), J. Čarnogurský (Collembola), T. Čejka (Mollusca), P. Fedor (Thysanoptera), P. Gajdoš (a part of Araneae, Opiliones), M. Holecová (Hymenoptera: Formicidae), J. Holman (Sternorrhyncha), O. Holuša (Psocoptera), J. Chrystophoryová (a part of Pseudoscorpiones), V. Janský (Auchenorrhyncha), S. Kalúz (Mesostigmata, Oribatida, Prostigmata), A. Krištín (Orthoptera), I. Krno (Plecoptera), Z. Krumpálová (a part of Araneae, Opiliones), M. Krumpál (a part of Pseudoscorpiones), J. Lukáš (Hymenoptera: Braconidae, Diapriidae, Chalcididae), O. Majzlan (a part of Coleoptera), A. Mock (Diplopoda, Isopoda), J. Patočka (Lepidoptera), M. Roháčová (Heteroptera), L. Roller (Hymenoptera: Symphyta), I. Rychlík (a part of Coleoptera) and V. Straka (Diptera). References Aﬀre L. & Thompson J.D. 1997. Variation in the population genetic structure of two Cyclamen species on the island of Corsica. Heredity 78: 205–214. DOI: 10.1038/hdy.1997.28 Aﬀre L., Thompson J.D. & Debussche M. 1995. The reproductive biology of the Mediterranean endemic Cyclamen balearicum Willk. (Primulaceae). Bot. J. Linn. Soc. 118 (4): 309–330. DOI: 10.1111/j.1095-8339.1995.tb00474.x Annand P.N. 1926. Thysanoptera and pollination of ﬂowers. Amer. Nat. 60 (667): 177–182. Beattie A.J. 1972. The pollination ecology of Viola. 2, Pollen loads of insect-visitors. Watsonia 9 (1): 13–25. http:// archive.bsbi.org.uk/Wats9p13.pdf (accessed 15.06.2012) Blažyt˙e-Čereškien˙e L., B¯ uda V. & Bagdonait˙e E. 2012. Three wild Lithuanian strawberry species and their pollinators. Plant Syst. Evol. 298 (4): 819–826. DOI: 10.1007/s00606-012-05939 Buchar J., Ducháč V., Hůrka K. & Lellák J. 1995. Klíč k určování bezobratlých. Pedagogické nakladatelství Scientia, Praha, 287 pp. ISBN: 8085827816, 9788085827811 Buchar J. & Kůrka A. 1998. Naši pavouci. Academia, Praha, 155 pp. ISBN: 8020003312, 9788020003317 Davis A.P., McGough N., Mathew B. & Grey-Wilson Ch. 1999. CITES Bulb Checklist for the genera: Cyclamen, Galanthus and Sternbergia. Royal Botanic Garden, Kew, 135 pp. ISBN10: 1900347393, ISBN-13: 978-1900347396 de Jong H. (ed.) 2010. Fauna Europaea (version 2.2.). http:// www.faunaeur.org/ (accessed 03.10.2011) Doskočil J. (ed.) 1977. Klíč zvířeny ČSSR. Díl V. Dvoukřídlí. Československá akadémia vied, Praha, 376 pp. Fedor P., Sierka W. & Országhová Z. 2008. Thrips (Thysanoptera) in nests of the tree sparrow, Passer montanus (L.). Acta Phytopathol. Entomol. Hung. 43 (2): 277–282. DOI: 10.1556/APhyt.43.2008.2.9 Halda J. & Soják J. 1971. Cyclamen fatrense sp. n., nový druh západokarpatské květeny. Čas. Nár. Mus., odd. Přírodověd. (Praha) 140 (1/2): 63–65. Hegi G. 1924. Illustrierte Flora von Mittel-Europa mit Beson¨ derer Ber¨ ucksichtigung von Deutschland, Osterreich und der Schweiz zum Gebrauche in den Schulen und zum Selbstunterricht. Bd. 4. T. 3 (Leguminosae bis Tropaeolaceae), J.F. Lehmann, M¨ unchen, pp. 1113–1748. Hegi G. 1927. Illustrierte Flora von Mittel-Europa mit Beson¨ derer Ber¨ ucksichtigung von Deutschland, Osterreich und der Schweiz zum Gebrauche in den Schulen und zum Selbstunterricht. Dicotyledones. Bd. 5, T. 4, pp. 2253–2631.
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