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Chromosome numbers in Pinguicula (Lentibulariaceae): survey, atlas, and taxonomic conclusions

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Abstract

Our study (survey, atlas of 136 microphotographs and 67 drawings) points out the actual chromosome numbers of 82 taxa of the genus Pinguicula L. They were gathered from literature and critically examined. In addition, numerous counts are published for the first time. They represent about 80% of all the taxa known. The basic chromosome numbers are x = 6, 8, 9, 11, and 14; the ploidy levels are 2n (diploid), 4n (tetraploid), 8n (octoploid) and 16n (hexadecaploid). The basic number x = 6 is a one-off, x = 8 and 11 are the most frequent in the genus; x = 14 indicates a hybridogenous differentiation process in the past. The caryological differentiation—chromosome numbers and ploidy level—is discussed with regard to distribution pattern, growth type, and infrageneric classification (at the level of sections).

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Notes

  1. In some cases the present taxonomic practise is not in full accordance with the Code. Nevertheless, in our study we are using the names especially for the sections as given in relevant literature.

  2. About 95% of current taxonomic studies in Pinguicula refer to Casper (1966b) apparently, because an actual comprehensive taxonomic treatment is lacking. Legendre (2000) and especially Cieslack et al. (2005) present the most recent example of a modern approach to a phylogenetic classification. It is not discussed in our presentation.

  3. The description of the new species from the Apuan Alps is in preparation (see also footnote 10).

  4. Personal communication by M. Studnička (Liberec). In 1990, he found a single specimen near Mělnická Vrutice (Polabská černava) and succeeded in cultivation. This is the last existing sample of P. bohemica which originates from the type locality (river Elbe area).

  5. M. Studnička showed the locality to the senior author and provided us with specimens from his cultures in BG Liberec.

  6. As subspecies: Pinguicula vulgaris subsp. bohemica (Krajina) Domín (Kubát 2002, p. 572). In Čeřovský et al. (1999, p. 282) Dostál is quoted as author of the subspecies: P. vulgaris subsp. bohemica (Krajina) Dostál.

  7. In his monograph on Pinguicula (Casper 1966a, b, p. 137, Fig. 38; 138) the senior author followed McVaugh and Mickel (1963) who had identified the specimens collected by G.B. Hinton et al. between 1937 and 1939 in Guerrero (nos. 10682, 14442) and Michoacan (no. 15107) as P. colimensis. This identification was corrected by Zamudio Ruiz (2001), who ranked the Hinton collections as P. zecheri.

  8. We did not find any adequate term in English; perhaps “strap-shaped upright sticky leaves” as one of the reviewers proposed.

  9. Pinguicula mariae spec. nov. will be published later on in 2008. We are aware that a name published in anticipation of further acceptance is no standard practise. However, in our context we break the rule for a better understanding.

  10. 32.1 BGJ is a sample that has been cultivated without any interruption in the botanical garden of Jena from the beginning of the twentieth century. Collection locality and source are not known. Therefore it remains doubtful whether the voucher specimens are identical with the cultivated specimens studied by Kondo.

  11. The hexadecaploid chromosome number 2n = 128 could not be counted exactly; it is estimated (tolerance +2/-3). It was not found in all specimens studied. There seems to be a tendency to polyploidy like in other taxa of the Apennine area said to be similar to P. vulgaris.

  12. In our context we will not discuss the taxonomic rank of the taxa in question in detail (see Conti and Peruzzi 2006 for the Apennine populations).

  13. If we followed Zamudio Ruiz (2001, p. 213) we should place P. debbertiana with its 2n = 22 caryotype into section Pinguicula. However, there are not really compelling reasons for such a taxonomic step.

  14. We do not understand Peruzzi’s (2004, p. 103, 108) report of 2n = 44 for P. balcanica. He discusses two basic numbers (x = 8 and the “neo-basic” x = 11), the possibility of “misidentification” (however, see Shuka et al. 2007), and arranges the endemic (Balcan Peninsula) European taxon partially into the New World Orcheosanthus-group (Peruzzi 2004, p. 108). That’s a clear misinterpretation of the known facts.

  15. Therefore, P. dertosensis can no longer be arranged as subspecies of P. longifolia as Schlauer [1974, p. 67: P. longifolia ssp. dertosensis (Cañigueral) J. Schlauer] did.

  16. It is possible, that the Portuguese P. vulgaris-like population from Gerez (in Casper 1966b, on “Serra do Geres”), belongs to this group, too. The chromosome number is unknown.

  17. Formerly, Casper (1974b) believed the Pinguicula inhabiting NE Turkey and the Caucasus to be a subspecies of P. balcanica. However, our recent chromosome counts show that the populations of this region should be seen as a separate species (=P. pontica spec. inedit.).

  18. Pinguicula corsica is a representative of the Pinguicula corsica-Trichophorum caespitosum—association which inhabits—together with Narthecium reverchonii, Carex fusca, Nardus stricta, Ranunculus cordigerus, Poa supina, and Drosera rotundifolia—water-rinsed sinks of pozzines. These humid montane-subalpine meadows are developed on the bottom of desiccated glacial lakes (Gamisans 1976, 1977, 1978; Mayer 1990). They originated after the quaternary glaciations (Conchon 1986). This statement does not really contradict the view of Steiger (1998) who put forward the opinion that Corsica remained more or less (i.e. compared with the situation in the Alps—added by Casper) untouched by several ice ages.

  19. As P. villosa var. ramosa (Miyoshi) Tamura (see Kadono 1993, p. 400).

  20. From cultivation it is likely that the taxa are short lived perennials. We could not check the chromosome number of P. elongata which is said to be a heterophyllous species.

  21. The caryotype formulas discussed in this paper should be regarded with reservation because of the inadequate methods used (see Footnote 20, too) and of the small size of the chromosomes (Kondo and Shimai 2006; see also Greilhuber et al. 2006).

  22. In this context we remind about the statement of Ledebour (1847) who compared his P. spathulata (=P. variegata) from Transbaikal (Russian Federation) with P. lutea, a typical representative of section Isoloba.

  23. Our method used for counting of the chromosomes is not suited to solve structural problems of the caryotype. The individual chromosomes are difficult to classify as to L, M. or S group. Nevertheless, in the case of P. primuliflora we can see that in the complement two unusual large chromosomes exist which can be interpreted as the result of a fusion process of two S-chromosomes at the haploid level. Naturally, any genetic proof is lacking.

  24. The plants we studied are euploid, i.e. there is no intra individual variation in chromosome number.

  25. Speta and Fuchs (1982, p. 118) put forward the argument that Casper’s (1966a, b) division of the genus into three subgenera contradicted the caryological data especially regarding the taxonomic position of section Orcheosanthus. In principle that is right. They could not know that Casper’s creation of subgenera was enforced by external influence and did not express the actual opinion of the author. They realised that later on Casper (for instance 1974b) quoted only sections instead of subgenera, however, they could not draw appropriate conclusions.

  26. The octoploid P. macroceras and P. vulgaris inhabit partly the northern latitudes of the New World however, are of Old World origin.

  27. In this context the view of Deil (1989) should be mentioned who described the “Balmenflora” (“Felsstirn-Halbhöhlen-Catena”; Quézel 1968), i.e. the Adiantetea communities with Pinguicula, in the mediterranean area (Iberian, Italian, and Balcan peninsula), phytosociologically as Coeno-Pinguiculion. He came to the conclusion that the taxa involved should be thought as highly (paleo-) endemic vicarious species.

  28. The available molecular data (Schmidt 1998; Jobson et al. 2003; Müller et al. 2004; Cieslack et al. 2005; Degtjareva et al. 2006; Müller et al. 2006; Kondo and Shimai 2006) will be discussed in detail in a forthcoming separate paper dealing with our own findings.

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Acknowledgments

We got living material for our chromosome studies from Dr. Maria Ansaldi (Pisa, Italy), P. Debbert (Munich, Germany), O. Gluch (Harthausen, Germany). M. Gube (Jena, Germany), P. Harbarth (Dossenheim, Germany), Prof. Dr. L. Kashta (Tirana, Albania), Prof. Dr. E. Köhler (Berlin, Germany), J. Lautner, (BG Göttingen, Germany), R. Mangelsdorff (Frankfurt/Main, Germany), Dr. A. Schmidt (Berlin, Germany), Dr. M. Studnička (Liberec, Czech Republic), F. Fuchs (BG Linz, Austria), Dr. L. Shuka and Prof. Dr. Xhulaj (both Tirana, Albania), Prof. Dr. J. Steiger (Bern, Switzerland), M. Welge (Barsinghausen, Germany), PD Dr. K. Zoglauer (Berlin, Germany). To all of them we are greatly indebted. The specimens were professionally cultivated by Mr. Wolf from BG Jena, who committed successfully large amount of his time to provide us with living specimens and to find out new and suited culture methods. Our work was critically accompanied by Prof. Dr. F. Hellwig, Dr. H. Manitz (both of Jena, Germany), Dr. B. Rice (Davis, USA), Dr. W. Spanowsky (Dresden, Germany), PD Dr. J. Schlauer (Frankfurt/Main, Germany—to whom we are much indebted for correcting our poor English), and Prof. Dr. J. Steiger (Bern, Switzerland).

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    S. Jost Casper & Rosemarie Stimper

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Casper, S.J., Stimper, R. Chromosome numbers in Pinguicula (Lentibulariaceae): survey, atlas, and taxonomic conclusions. Plant Syst Evol 277, 21–60 (2009). https://doi.org/10.1007/s00606-008-0097-9

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