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Brief information about the species status of Utricularia cornigera Studnicka PDF

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TechnicalRefereed Contribution Brief information about the species status of Utricularia cornigera Studnicka Miloslav Studnicka • Liberec Botanic Gardens • Purkynova 630/1 • CZ-460 01 Liberec • Czech Republic • [email protected] Keywords: Utricularia cornigera, hybrid, heterosis, apomixis Abstract: The carnivorousplant Utricularia cornigera Studnickawas describedin2009, butauthor- ities ofthe International Carnivorous Plant Societypublishedanopinionthatitis notatrue species, but only a natural hybrid of U. reniformis and U. nelumbifolia. The role ofheterosis is discussed, because U. cornigera is much largerthan boththeoreticalparents. Seedlings, the very characteristic feature ofbladderworts (Utricularia), are different in all the bladderworts described, that is, in the named species and in artificial hybrids of U. nelumbifolia and U. reniformis. No support for the hypothesis supposing ahybrid origin ofU. cornigera was found. Introduction Recentlyahypothesisappearedthat Utriculariacornigera StudnickacouldbeahybridofU. nelum- bifolia Gardn. x U reniformis St.Hil. (Schlauer 2011; Fleischmann 2012). Consequentially, the new specieswasrejectedfromtheCarnivorousPlantDatabase(Schlauer2011).Neverthelessitwasaccepted inthe International PlantName Index (IPNI 2005). This articlepresents the results ofnew experiments withartificialcrossingsofboththeoreticalparentsproposedbythe authors. Themannerofgerminationandspecificallytheappearanceoftheseedlingsarecrucialphenomenain thelife strategyofbladderworts. Inthe Utricularia speciesfromthesectionIperuatherearetwodifferent ways ofgermination: either by floating seedlings (e.g. U. cornigera, U. nelumbifolia), or by terrestrial U seedlings(e.g. U. geminiloba, U. nephrophylla, andthetrue reniformis). ThedifferencewasdescribedandpicturedasaveryimportantdiacriticalattributeofU. cornigeraas comparedwith U. reniformis (Studnicka 2009).As mentionedintheprevious paper, U. reniformis does nothavewhorl-shapedseedlings anditcannotgerminate inwater, but U. cornigera germinates inwater. Thefloatingwhorl-shapedseedlingsofU cornigeraarerathersimilartothealsowhorl-shapedseedlings ofthe Brazilian U. nelumbifolia andtheVenezuelan U. humboldtii Schomb. These species canusephy- totelmeswithinthe leafrosettesofcertainhostplantstogerminate(Taylor 1989; Studnicka2011). Methods Cultivatedplantswereusedformanualhybridization,whichwascarriedoutduringJulyandAugust 2012 inLiberecBotanicGardens. SpecificidentityofU. reniformisisgrantedbyapreviousstudymade usingthe same material(Studnicka2009). Specific identity ofU. nelumbifolia is clearbecausethe sto- loniferous plants distributed in European botanic gardens are ofnatural origin. Numerous fresh seeds (thewholecontentofatleast2fullseedcapsulesofeachspeciesorhybrid)wereputintowaterinaglass and cultivated in a greenhouse. The seeds of U. reniformis, which couldnot germinate in such condi- tions, werealsoplacedonawetsoilmixofpeatanddriedSphagnummoss 1:1 andwere alsocultivated inthe greenhouse. Justthe soil surfaceprovides suitable conditions for U. reniformisto germinate. Volume 42 March 2013 15 The seedlings ofU. reniformis, U. nelumbifolia, U. nelumbifolia x U. reniformis, and U. reniformis mm x U. nelumbifoliawere documentedbyphotographstakenusingaPentax35 macrolens.The float- ing hybrid seedlings were compared with the seedlings of U. cornigera formerly used to describe the species (Studnicka2009). Possible heterosis was evaluated according to the maximal size ofthe leafblades found. The blad- derwort U. cornigera was measured in natural localities as well as in cultivated specimens, both giant (Studnicka 2009). The species U. nelumbifolia found thanks to F. Rivadavia nearTeresopolis in Serra dos Orgaoswasmuchsmallerthanthe specimens cultivatedinLiberec Botanic Gardens, whichiswhy Ipresent a measurement ofa cultivatedplanthere. The leafofthe flowering cultivated specimen ofU. reniformisusedinthis researchwas also measuredandcomparedwiththe largest leaffoundinnatural conditions (cf Studnicka2009). Results The question is whether the prospective hy- bridtheoreticallyresulting in U. cornigera would be an F generation or a descendant ofmultiple 1 breeding. To assess this problem measurements ofthe largestknown leaves in allthree species in questionareneeded. Icangivethesedimensions: U cornigera 24.4 x 20.7 cm (cultivated speci- men), but also up to 17.5 x 13.0 cm in natural conditions; U. reniformis9.2 x7.5 cm(cultivated specimen),buteven 10.5 x8.5 cminnaturalcon- ditions; U. nelumbifolia 9.9 x 9.6 cm (cultivated specimen), butdistinctly smallerspecimenswere seen in natural conditions. The preserved leafof U. cornigeraphotographedinthis article is locat- ed in the PRC herbarium; the leaves ofthe other species are found in Liberec Botanic Gardens (www.botaniliberec.cz). It was documented that theleaflaminasofthetheoreticalparentalspecies arebothabouthalfthesizeofthoseofU. cornige- ra (Fig. 1-3). IfU. nelumbifolia and U. reniformis weretheparents ofU. cornigera the giant size of , its leaflaminawouldbe asignofheterosis. The seedlings of all three species are very characteristic. Are the juvenile hybrids of U. nelumbifolia x U. reniformis (and conversely) similar to U. cornigera? The results of crossing experiments are documentedbelow(Fig. 4-8). Discussion Figures 1-3: Leaves of luxurious specimens The bladderwort U. cornigera is much more of cultivated bladderworts. The scale is in vigorous than both U. nelumbifolia and U. reni- centimeters. 16 Carnivorous Plant Newsletter Figures 4-5: Floating seedlings of Utricularia nelumbifolia (left) and terrestrial seedlings of U. reniformis (right). These theoretical parents of U. cornigera were crossed artificially. (See the following figures.) ' A /> -7* . x ' ri * A V v-W a * 9 Xx, V r Figures 6-7: Seedlings of the hybrid Utricularia nelumbifolia 9 x U. reniformiswith 4-5 leaf primordia (left) and of the hybrid U. reniformis 9 x U. nelumbifolia with 3-4 leaf primordia (right). formis. Therefore, the heterosis effect in the FI crossbredgenerationshouldbenoted, ifU. corni- gem were to be considered as a hybrid. The het- erosis effect is, however, unsustainable in subse- quentgenerations. The gigantism in U. cornigera (considered a hybrid) would be stable solely by two prerequisites: 1. It is a primary hybrid; 2. It propagates onlyasexually. U. cornigera produces viable seeds, but they could theoretically be a result ofapomixis. Nev- Figure 8: Seedlings of Utricularia cornigera ertheless, research of seedlings made by cross- with 6-8 leaf primordia are dissimilar to the breeding experiments between U. nelumbifolia hybrids pictured above. and U. reniformis indicates lowernumbers ofleaf primordia in these hybrids in comparison with U. cornigera. This is clear evidence that the seedlings of U. cornigera cannot be an apomictic derivate ofthe hybrids U. nelumbifolia x U. reniformis or U. reniformis x U. nelumbifolia. Experience with a cultivated U. cornigera indicates thatmanual pollina- tionisnecessaryto obtainseedsandseedlings.Thatis also a signofnormal sexualpropagation.Asthe aforementioned prerequisites are not fulfilled, the idea that U. cornigera originated as a hybrid is not supported. According to this preliminary result, U. cornigera arose due to reproductive isolation, like other endemic plants restricted to the same area, ratherthan from ahybrid origin. The future study of Volume 42 March 2013 17 traps, and or fertility/sterility in mature hybrids would also be interesting corroboration ofthe species status ofU. cornigera. Conclusions 1. The seedlings originating from both crossings of U. nelumbifolia and U. reniformis are dis- similarto seedlings ofU. cornigera. 2. This fact excludes the possibility that U. cornigera couldbe an apomictic primary hybrid. 3. The bladderwort U. cornigera is the most vigorous species ofthe three named bladderworts, but no heterosis could survive inmany generations ofthe sexuallymultiplying bladderwort. That is why it canhardly be a stabilizedhybrid ofmuch smallerparents. Acknowledgement: I wish to thank Professor R. J. Valka Alves, University Federal do Rio de Ja- neiro, forhis helpful suggestions. References Fleischmann,A. 2012.Thenew Utricularia speciesdescribedsincePeterTaylor’smonograph. Camiv. PI. Newslett. 41: 67-76. IPNI. 2005. The International PlantNames Index, http://www.ipni.org/ipni/plantnamesearchpage.do, accessed 17 July2012. Schlauer, J. 2012. Carnivorous Plant Database, http://www.omnisterra.com/bot/cp_home.cgi, ac- cessed 17 July2012. Studnicka, M. 2009. Brazilian bladderwort Utricularia reniformis is a blend oftwo species. Thaiszia -J.Bot. 19: 131-143. Studnicka, M. 2011. Surprising phenomena in the life strategy of Utricularia reniformis in Brazil. Thaiszia-J. Bot. 21: 37-43. Taylor, P. 1989. The genus Utricularia - a taxonomic monograph. Kew Bull. Additional Ser. XIV, Royal Bot. Gardens, Kew. 723 p. The Largest Most Famous CP Nurseryin North America Home ofthelargest collection of CP in the worldon display Thurs. - Mon. 10 Am -4 Pm We ship only thehighest quality plantsin ourdeluxe pots to the entireU.S. Owned by Peter D'Amato, author of"The Savage Garden" 707-824-0433 www.Californiacarnivores.com [email protected] Ituikli 18 Carnivorous Plant Newsletter

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