Bibliotheca Botanica
ISSN 0067-7892 Bibliotheca Botanica
Original Contributions to Botany, founded in 1886 Edited by H. W. Lack, Berlin, P. Leins, Heidelberg and S. Porembski, Rostock
Volume 159
Arno Wörz Revision of Eryngium L. (Apiaceae-Saniculoideae): General part and Palaearctic species pages
Sample
E Schweizerbart Science Publishers
eschweizerbart_XXX BIBLIOTHECA BOTANICA, VOLUME 159
Sample pages BIBLIOTHECA BOTANICA Original Contributions to Botany
Edited by H. W. Lack, Berlin; P. Leins, Heidelberg and S. Porembski, Rostock
Volume 159
Arno Wörz
Revision of Eryngium L. (Apiaceae-Saniculoideae): General part and Palaearctic species
with 84 text-figures, 41 tables and 12 plates Sample pages
Schweizerbart Science Publishers Stuttgart • 2011 Contents 1
Contents
Preface...... 5 Abstract...... 6 I General part: Taxonomy, Biogeography and Evolution (6) ...... 9 1. Introduction ...... 9 2. History of Illustration and Research in Eryngium ...... 10 3. Material and Methods ...... 14 3.1 Nomenclature and Herbarium Studies ...... 14 3.2 Phytosociological Methods ...... 14 3.3DistributionMaps...... 14 3.4 Chromosome Numbers ...... 15 3.5 Fruit Anatomical and Petal Morphological Studies ...... 15 3.6 Cladistic and Systematic Methods ...... 19 4. Reproductive biology ...... 32 5. The Characters and their Taxonomic Values ...... 33 5.1 Morphological and Anatomical Characters ...... 33 5.1.1 The Habits and Life Forms ...... 33 5.1.2 Rhizomes and Tubers ...... 33 5.1.3 Aerial Stems ...... 33 5.1.4 Basal Leaves ...... 33 5.1.5 Cauline Leaves ...... 36 5.1.6 Leaf Sheaths ...... 37 5.1.7 Stomata Types ...... 37 5.1.8 Collenchyma ...... 37 5.1.9 Capitula, Inflorescences, and Synflorescences ...... 37 5.1.10 Involucral leaves ...... 37 5.1.11 Bracts ...... 38 5.1.12 Sepals ...... 38 5.1.13 Petals ...... 39 5.1.14 Nectarium ...... 39 5.1.15 Pollen ...... 44 5.1.16 Mericarps ...... 44 5.1.16.1 General ...... 44 5.1.16.2 Carpell layers and structures ...... 44 5.1.16.3 Oil Ducts and Bundles ...... 44 5.1.16.4 Inclusions ...... 47 5.1.16.5Sample Relictual Carpophor ...... pages 47 5.1.16.6 Wings ...... 47 5.1.16.7 Scales ...... 47 5.1.16.8 Lignified Mesocarp ...... 48 5.1.16.9 Classification of Eryngium byFruitAnatomy...... 49 5.1.17 Seedlings ...... 51 5.2 Chemical Characteristics ...... 51 5.3 Cytological Characteristics ...... 56 5.3.1 Results and General Remarks ...... 56 5.3.2 Discussion and Evolutionary Trends ...... 56 5.4 Ecological Characteristics ...... 67 5.4.1 General ...... 67 2 Contents
5.4.2 The Habitat Types ...... 67 5.4.3 Discussion of the ecological characteristics ...... 74 6. Nomenclature and Description of the genus Eryngium L...... 76 7. Eryngium: Classification and Evolution ...... 77 7.1 The Position of Eryngium withintheSaniculoideae ...... 77 7.2 The Subgeneric Classification ...... 77 7.3CladisticAnalysis...... 79 7.3.1 General ...... 79 7.3.2 Cladistic analysis of Subg. Eryngium ...... 81 7.3.3 The clade of Subg. Ilicifolia ...... 81 7.3.4 The cladistic analysis of subgg Foetida, Monocotyloidea, and Semiaquatica ...... 81 7.4 Comparison with Molecular Results ...... 86 7.5 Fossil Records ...... 88 7.6 Biogeography ...... 88 8. Sectional classification and keys ...... 91 8.1 Division of Eryngium into subgenera ...... 91 8.2 Division of Eryngium subg. Eryngium into sections ...... 91 8.3 Division of the sections of Eryngium subg. Eryngium into species ...... 92 8.4 Description of the anomalous Eryngium sect. Aquifolia and the division of this section into species ...... 96 8.5 Description of Eryngium subg. Ilicifolia and the division of this subgenus into species ...... 97 8.6 Division of Eryngium subg. Semiaquatica into species ...... 97 8.7 Division of the European and North African species of Eryngium subg. Semiaquatica ...... 98 II Systematic treatment of the Eurasian and North African Species ...... 99 9. Eryngium subg. Eryngium ...... 99 E. alpinum L...... 99 E. amethystinum L...... 112 var. tenuifolium Boiss.&Heldr.inBoiss...... 114 var. transiens (Halácsy) A.Wörz...... 114 E. amorginum Rech.f...... 137 E. antiatlanticum Jury ...... 140 E. aquifolium Cav...... 143 var. barbarum Jahand.&Maire ...... 145 E. billardierei F.Delaroche ...... 148 E. bithynicum Boiss...... 155 E. bornmuellerii Nábělek ...... 159 E. bourgatii Gouan...... 161 var. bourgatii ...... 162 var. atlanticumSampleBall in Hook.f...... pages 162 E. bungei Boiss...... 176 E. caeruleum M.Bieb...... 180 E. caespitiferum FontQuer&Pau...... 189 E. campestre L...... 189 var. virens (Link)Weiss...... 192 E. carlinoides Boiss...... 214 E. creticum Lam...... 217 E. davisii Kit Tan & Yıldız ...... 226 E. desertorum Zohary...... 229 E. dichotomum Desf...... 231 E. dilatatum Lam...... 235 Contents 3
E. duriaei GayexBoiss...... 242 subsp. juresianum (Lainz) Lainz ...... 242 E. falcatum F.Delaroche ...... 246 E. giganteum M.Bieb...... 253 E. glaciale Boiss...... 257 E. glomeratum Lam...... 261 subsp. elongatum A.Wörz...... 263 E. grosii FontQuer...... 267 E. hainesii C.C. Towns...... 270 E. heldreichii Boiss...... 273 var. fallax Bornm...... 273 E. huteri Porta...... 276 E. ilex P.H.Davis ...... 279 E. isauricum Contandr. & Quézel...... 281 E. karatavicum Iljin ...... 283 E. kotschyi Boiss...... 286 E. macrocalyx SchrenkinFisch.&C.A.Mey...... 289 E. maritimum L...... 292 E. marocanum Pit...... 304 E. octophyllum Korovin ...... 308 E. palmatum Pančić & Vis. in Vis. & Pančić ...... 310 E. palmito Boiss.&Heldr...... 316 E. planum L...... 318 var. armatum Csató ex Simonk...... 320 E. polycephalum Hausskn.exH.WolffinEngl...... 326 E. pseudothorifolium Contandr. & Quézel...... 329 E. pyramidale Boiss.&Hausskn.exBoiss...... 332 E. serbicum Pančić ...... 334 E. spinalba Vill...... 337 E. ternatum Poir...... 341 var. subinerme Halácsy...... 343 E. thorifolium Boiss...... 345 E. thyrsoideum Boiss...... 349 E. tricuspidatum L...... 352 subsp. occidentalis A.Wörz...... 354 subsp. bovei (Boiss.) Batt. in Batt. & Trab...... 354 subsp. mauritanicum (Pomel) Batt. in Batt. & Trab...... 354 E. triquetrum Vahl ...... 369 subsp. xauense (Pau)Jovet&Sauvage...... 370 Sampleö pages E. trisectum A. W rz&H.Duman...... 375 E. variifolium Coss...... 378 E. wanaturi Woronow ...... 384 E. wiegandii Adamović ...... 385 10. Eryngium subg. Ilicifolia ...... 391 E. argyreum Maire ...... 391 E. ilicifolium Lam...... 394 11. Eryngium tenue ...... 405 E. tenue Lam...... 405 12. Eryngium subg. Semiaquatica ...... 416 E. atlanticum Batt. & Pit. in Pit...... 416 4 Contents
E. corniculatum Lam...... 419 E. galioides Lam...... 424 var. leiocarpum H.WolffinEngl...... 426 var. trachycarpum Gay ...... 426 E. pusillum L...... 431 E. viviparum Gay...... 435 13. Hybrids of Eryngium ...... 441 E. x chevalieri Sennen ...... 441 E. x embergeri Sennen ex Sennen & Mauricio ...... 441 E. x heteracanthum Teyber ...... 442 E. x kalotaszegense J. Papp & Ujvárosi in Ujvárosi ...... 442 E. x microcephalum Sieber...... 442 E. x mohamedanii FontQuer&Pau ...... 443 E. x oliverianum F.Delaroche ...... 443 E. x rocheri L.Corb. in Guétrot ...... 445 E. x tripartitum Desf...... 445 E. x visianii Teyber ...... 445 E. x zabelii H.Christ ...... 446 14. Nomina illegitima, nomina nuda ...... 447 14.1. Nomina illegitima ...... 447 14.2. Nomina nuda ...... 447 15. Acknowledgements ...... 448 16.References...... 450 17. Index nominum ...... 471
Sample pages Preface 5
Preface
This volume, the result of a 12-year study, is an attempt tion. Systematic evaluations and cladistic trees may to understand Eryngium as a small part of global biodi- thus lead to misinterpretations if considered uncritical- versity. For this, a variety of character types have been ly or without a thorough understanding of the taxa. studied: they comprise the usual morphological, chem- Against this backdrop, my own cladistic studies otaxonomic and genetic characters, chromosome num- are presented in this volume and compared with the re- bers as well as ecological characteristics. Every spe- sults of molecular systematic research results published cies, every individual, is embedded in its environment; by others – as is standard in modern science. Both the conditions of a habitat, biological and non-biologi- methods deliver new results to be interpreted critically, cal, form the foundations for an organism’s existence and a deeper understanding of these results is possible and the challenges an organism faces. The environment by giving a closer, more thorough look at each taxon. stimulates cooperation and competition – all in all, the This may seem old fashioned, differing as it does from “struggle for existence”. These external factors are the many recent publications with their plethora of authors driving forces of evolution. often lacking in background knowledge. This field does These external influences undergo fluctuations and, not offer an opportunity to produce a long publication occasionally, dramatic shifts, as we can see, or certainly list or to make a fast career. The still excellent working will see, in the recent climatic changes. Evolution fol- conditions in a natural history museum (where I am lows with adaptations to these unpredictable and more employed) seem to present one of the rare opportuni- or less chaotic fluctuations. Necessarily, evolution over ties today to undertake such a study. time and over generations is also a chaotic process. It is I hope I will pique some interest in the work I not predictable and not really reconstructable; it does present here. not, for example, follow parsimony principles. System- Stuttgart, October 2009 atic evaluations and cladistic trees are thus approxima- Arno Wörz tions that do not reflect the actual pathways of- evolu
Sample pages 6 Abstract
Abstract
A phylogenetic and biogeographic analysis of Eryn- cies of subg. Eryngium grow. The vernal pool species gium L. (Apiaceae-Saniculoideae) and a systematic are uniform in their habitat requirements, and so the treatment of its Eurasian and North African species are radiation of this group in based exclusively on a geo- presented. In the general part, results of classic meth- graphic differentiation. Subg. Eryngium is much more ods have been used for a cladistic analysis and for com- variable in its habitats, suggesting an ecological as well parisons with molecular results. The classic methods as a geographical radiation. The habitats range from support the earlier classification of the genus into five (mostly) dry to moderately moist sites with (mostly) subgenera (subg. Eryngium, Monocotyloidea, Semi- calcareous to occasionally crystalline or serpentine aquatica, Foetida, and Lessonia) and the description of soils. Subg. Ilicifolia prefers ephemeral vegetation a small new subgenus, E. subg. Ilicifolia. types and habitats in mainly dry, semi-desert condi- A number of characters and character groups have tions. been used for this study: Cladistic analysis: All available results have been Morphological characters: The most important included in a cladistic analysis, which roughly supports morphological character is the structure of the basal the subgeneric classification mentioned above. Subg. leaves, which is palmate in subg. Eryngium, undivided Ilicifolia is located at the base of the New World spe- and parallel veined in subg. Monocotyloidea, arcuate, cies in this analysis. The differences between the Old parallel to pinnate in subg. Semiaquatica, and more or World subg. Eryngium and the other subgenera are vis- less pinnate or pinnately veined in subgg. Foetida and ible, but the delimitation of subg. Foetida and subg. Ilicifolia. This formal grouping is roughly supported by Semiaquatica is in some ways doubtful. other morphological characters such as petal and fruit Comparisons with molecular results: Not all mo- morphology. lecular data point to Eryngium being monophyletic, Fruit anatomy: The oil ducts are dorsally and later- and the results published in the literature are contradic- ally located in subg. Eryngium, dorsally and ventrally tory. Molecular results in part support the morphology- in the mainly New World subgenera, and in the fruit based classification. The difference between subg. Er- ribs in subg. Ilicifolia. Fruit wings are present in subg. yngium and the other subgenera is clearly evident. Eryngium; they include the outer fringes of the ligni- Subg. Monocotyloidea is largely supported as a con- fied ventral band typical for this subgenus. Insubg. sistent group. Subgg. Foetida and Semiaquatica are Monocotyloidea, wings are formed by exocarp cell lay- only in part confirmed by the molecular data and re- ers only. Subgg. Foetida and Semiaquatica lack wings quire further study. Contrary to the cladistic results, entirely. subg. Ilicifolia turns out to be a sister group to subg. Chromosome numbers: The species of subg. Er- Eryngium. This is plausible, as both groups grow sym- yngium are diploid (with few exceptions) with a chro- patrically in the Old World. The differences between mosome base number of x=8 or x=7, the latter concen- the classical and the molecular approach are dis- trated in sect. Amethystina. Subg. Monocotyloidea cussed. comprises some diploids, mainly in SE South America Classical and molecular results suggest two main (indicating a primary centre of diversity there), with a lineages: (1) subg. Eryngium with palmate basal leaves vast majority of polyploidsSample dispersed over North and is limited pages to the Eurasian and North African region, and South America with a secondary centre of diversity in (2) the remainder of the subgenera (subgg. Monocoty- Mexico. In subg. Semiaquatica, both diploids and poly- loidea, Semiaquatica, Foetida, Lessonia) to the New ploids are common, with diploids in the western Medi- World, Australia, and (with 5 species in subg. Semi- terranean and polyploids in the western part of North aquatica) in the Mediterranean. A small isolated group America. Subg. Foetida consists of diploids, some of (subg. Ilicifolia) occurs in the western part of the Med- them with a reduced chromosome base number (down iterranean, indicating the importance of this region for to x=5). the evolution of the genus. Morphological and fruit- Ecology: In the Eurasian and North African species anatomical results and chromosome numbers suggest presented here, the most conspicuous difference in the long-range dispersal events from the western Mediter- ecology is the vernal pool habitats of subg. Semiaquat- ranean to the New World, and – within the Eurasian ica, and the more or less dry conditions where the spe- and North African regions – a radiation in an easterly Abstract 7
direction (subg. Eryngium). Molecular results support ter common to all Eryngium species is the form of the this view. The colonization of the New World happened capitulum. The homology of this character has not yet within subg. Semiaquatica; a second event may have been examined. occurred, resulting in the subgg. Foetida and Mono- In the special part of this revision, 61 Eurasian and cotyloidea. North African species are treated, descriptions, distri- In all character groups, large and conspicuous dif- bution maps, and illustrations are provided and keys for ferences are visible between subg. Eryngium and all the identification included. Herbarium studies form -the ba other subgenera. This may justify a separation of subg. sis of this work. Phytosociological data serve as indica- Eryngium as a genus in its own right. The only charac- tors for ecological requirements.
Sample pages 10 General Part: Taxonomy, Biogeography and Evolution
2 History of Illustration and Research in Eryngium
Eryngium was already mentioned by Dioscurides in the (1581a: 24–26) as Eryngium pusillum planum Moutoni 1st century (see Berendes 1902: 275), who indicated the and Eryngium pumilum hispanicum. Clusius (1601: use of the young leaves as vegetables. Dioscurides was CLVIII–CLIX) later copied them. At about the same probably referring to E. campestre L. He also men- time, other Eryngium species were illustrated for the tioned medicinal uses: as a diuretic, for intestinal and first time (according to Baumann 1998) by Clusius liver afflictions, for the bites of poisonous animals, for (1576, E. amethystinum), Turner (1551, E. maritimum), epilepsy, and as a remedy for healing swellings when and Matthioli (1565, E. planum). worn as an amulet. Theophrastus (372–287 BC) notes The first to report on the New World Eryngium Eryngium in his Historia Plantarum as a species with species was probably Francisco Hernandez (1517– thorny leaves (Theophrastus VIth book, chap. 1/3; see 1587), who recorded and illustrated Aztec medicinal also Sprengel 1822 v. 1: 217, v. 2: 396). It is, however, plants from Mexico, including two Eryngiums (Con- not clear, which species Theophrastus meant with his stance 1978: 9). This herbal was not published until ήρύγγιον. 1651 (Hernandez 1651: 143, 222). Constance identi- Some species of Eryngium were illustrated by me- fied the plant called “Chichica Hoazton” or- “Cohay dieval and early Renaissance painters. The most fa- elli” as E. serratum Cav. (Fig. 2) and “Ocopiaztli” as E. mous is probably Albrecht Dürer (1471–1528). He in- deppeanum Schlechtendahl & Chamisso (Fig. 3). Paul cluded simplified Eryngium plants in several of his Hermann (1646–1695) was (according to Constance works, for the first time in his self-portrait ofE. 1493( 1978: 9) the first to present a description and illustra- amethystinum), which is today in the Louvre, Paris, and tion of a South American species: E. foetidum L. from later in his Allegorisches Wappen der Trägheit [Alle- Surinam (“Eryngium Americanum foetidum”; see Her- goric Emblem of Idleness], Die Heilige Familie mit der mann 1687: 236). An early illustration of E. yuccifo- Heuschrecke [The Holy Family with the Locust], and lium Michx. is presented in Plukenet’s Phytographia Liebespaar zu Pferde [Lovers on a Horse]. Apart from (Plukenet 1692: pl. 175), based on specimens collected an erotic symbol, it may also be interpreted as a Chris- by the missionary John Banister (1650–1692) (Con- tian allegory on the pains and suffering of Jesus Christ, stance 1978: 9). Caspar Bauhin (1560–1624) men- as is the thistle (Grote 1965: 156ff.). In this context, tioned in his Pinax (Bauhin 1623: 386) nine species, all Eryngium is often miscalled “a kind of thistle” (Grote from the Old World, some of which may in actuality 1965). The same is true for a very nice drawing of E. belong to the Compositae family. Joseph Pitton de alpinum by Nicolas Robert (Haarlem, Teylers Museum, Tournefort (1656–1708) lists 20 names (Tournefort inv. no. T 86), entitled The Blue Thistle. 1700: 327), including one for an American species, Apart from works of art, the genus Eryngium is which is probably E. foetidum (“Eryngium America- mentioned and illustrated in early herbals: Otto Brun- num, foetidum, H.L. Bat.”). E. foetidum was illustrated fels(1488–1534)referstoitinhisherbalas“Mannstrew” first by Hans Sloane (1660–1753) (Sloane 1707: pl. (Brunfels 1532: 281), which is still a German vernacu- 156) based on a herbarium sheet from “Jamaica and lar name for Eryngium campestre today. There is, how- Barbados”. Robert Morison (1620–1683) presented ever, a discrepancy in Brunfels: his illustration is clear- early illustrations of North American species: the reis- ly a Cirsium species.Sample Leonhard Fuchs (1501–1566) sued volumespages of his Plantarum Historiae Universalis presented a correct wood-cut of Eryngium campestre Oxoniensis (Morison 1715) include on pl. 37, besides L. (Fuchs 1543: Cap. CXII, CLXVI, Fig. 1) and indi- Hernandez’ “Ocopiaztli” (no. 22), one further illustra- cated some of the medicinal uses, including its use as a tion of E. yuccifolium Michx. (no. 21). diuretic and a remedy for bites of poisonous animals. In his Species Plantarum Carl von Linné (1707– Mathias de L’Obel (1538–1616) published the first il- 1778) mentioned eight Eryngium species (Linnaeus lustration of E. alpinum (L’Obel 1576: 490, actually a 1753: 232), two of which (E. aquaticum L., E. foetidum plate taken from Dodoens; see Baumann 1998: 122) L.) originate from the Americas. Many descriptions of and some nice illustrations of Spanish Eryngium spe- new species followed the introduction of the binominal cies, including E. pusillum and E. tenue, both probably nomenclature. A very important treatment is found in the first illustrations of these rare western Mediterra- volume 4/2 of the Encyclopédie méthodique of Jean nean plants. They were later published again in L’Obel Baptiste Antoine Pierre de Monnet de Lamarck (1744– History of Illustration and Research in Eryngium 11
um material collected by Luis Née during the Malaspi- na Expedition. A first summarizing monograph of Eryngium was presented by François Delaroche (1780–1813). In his most important study he revised both Eryngium L. and Alepidea L. (DELAROCHE 1808). His monograph in- cludes excellent illustrations, treats altogether 50 spe- cies, and gives 19 new names, of which only two are synonyms of earlier names. 14 of the 17 new species originate from the New World. Delaroche’s herbarium was incorporated into the Geneva herbarium, where most of the types are located today. In the following decades, many species were de- scribed following the exploration of hitherto unknown parts of the world. In the Old World, the most important author of names in the genus Eryngium was Pierre Ed- mond Boissier (1810–1885), who validated 16 new names, occasionally together with Heinrich Karl Haussknecht (1838–1903). Most new species came from the Near and Middle East, the region covered by his Flora Orientalis. Augustin Pyramus de Candolle (DeCandolle 1830) validated in his Prodromus nine new names: four are still accepted species names, one has been recombined, and four are synonyms. In the New World, the discovery and description of species boomed as a result of the voyages of such col- lectors as Friedrich Sellow (1789–1831) and Auguste François Marie Glaziou (1828–1906), both more trav- ellers than scientists, who themselves described rela- tively few or no species. Glaziou collected the types of nine species, Sellow the types of no fewer than 22 spe- Fig. 1: Eryngium campestre from Leonhard Fuchs (1543: cies. Many species from the latter were published by CLXVIII). Courtesy of Württembergische Landesbibliothek Ludolf Karl Adalbert von Chamisso (1781–1838) in Stuttgart. Chamisso (1833). Together with Dietrich Franz Leon- hard von Schlechtendahl (1794–1866), Chamisso de- scribed further species from the Romanzoff expedition (Chamisso & Schlechtendahl 1826) and from the col- 1829) (Lamarck 1798: 751ff.). 25 species of “panicaut” lectors Christian Julius Wilhelm Schiede (1798–1836) are mentioned, all of whichSample in fact belong to the genus and Ferdinand pages Deppe (1794–1861) (Schlechtendahl & Eryngium. This treatment was written by Lamarck Chamisso 1830). Turczaninow (1847) added three himself and signed by him at the end of the last para- names. Joseph Decaisne (1807–1882) validated six graph. new names and thoroughly discussed the differences The first to realize the great diversity of the New between the Old World and the New World species. 11 World Eryngiums was Antonio José Cavanilles (1745– species were described by William Jackson Hooker 1804). He described in several publications at least (1785–1865), mostly together with George Arnott seven species (Cavanilles 1800 a, b, 1801a, b), six of Walker (1799–1868). Names of five further New World which come from the New World; some of them are the taxa were published by August Heinrich Rudolph first species of the subgenus Monocotyloidea (e.g., Grisebach (Grisebach 1854, 1874, 1879), based mostly E. longifolium Cav., E. monocephalum Cav.) to become on material of Paul Günter Lorentz (1835–1881). The known. The descriptions are based mostly on herbari- Californian botanist Willis Linn Jepson (1867–1946) 40 General Part: Taxonomy, Biogeography and Evolution
E. alpinum L.
Sample pages
E. carlinae Delaroche E. humile Cav. E. tenue Lam.
Fig. 4: The petal types of Eryngium shown in lateral view. E. humile and E. tenue from Froebe et al. (1981). Scale = 1 mm. The Characters and Their Taxonomic Values 45
Fig. 5: E. amethystinum (subg. Eryngium), cross-section of the ripe fruit with seed coat (sc), endocarp (enc), mesocarp (mc), bundles (b), crystals (c), oil ducts (od), and tubercles (t) on the epidermis. Scale bar = 0.1 mm.
Fig. 6: E. bithynicum (subg. Eryngium), cross-section of ripe fruit with oil ducts (od), bundle (b) and tannin inclusions (ti). Scale bar = 0.1 mm. Sample pages
Fig. 7: E. amethystinum, cross-section of the ripe fruit with dorsal/lateral bundles and oil ducts respectively, the lignified ventral band (lvb), tannin inclusions (ti) and the wings (w); other abbreviations see before. Scale bar = 0.5 mm. 346 Systematic treatment of the Eurasian and North African Species
Köyceğiz, Sandras Da., P. nigra ormani acıklikarı, 1500 m, pentin ana kaya Makiuk, 150–200 m, 15.7.1991, A. Güner, 18.7.1968, A. Pamucuoğlu (HUB); Prov. Muğla: Marmaris, H. Duman, H. Sagban, Z. Aytaç 9615 (GAZI, HUB); Muğla 30 m, Pinetum brutiae, on serpentine, 24.3.1956, P.H. Davis Vil.: Köyceğiz – Terasakan, 11.7.1960, H. Peşmen 949, M. & O. Polunin: Flora of Turkey 1956 (BM); C2 Muğla: be- Ayder (G); Muğla, between Köycegiz and Fethiye, 21.5.1967, hind Marmaris, 50–100 m, Rocky serpentine slope. T. Baytop (E); C2 Muğla (Caria): Muğla – Fethiye, Macchie 14.4.1965, Davis 41414 (E, K); C 2 Muğla: Marmaris, Mar- in Pinetum brutiae 47 km südöstlich Muğla, 70 m, 20.6.1954, maris milli Parki, Balar daği güneybatsi, Serpentin arakaya, A. Huber-Morath, Flora Anatolica 12197 (G); C 2 Muğla: 500 m, 29.6.1997, H. Sagban 1851 (HUB); C2 Muğla: Fethiye’den Köycegiz’z 23 km Pinus brutia ormani, serpen- Reşadye Yarimadasi (R. peninsula) between Marmaris and tin arazi, ca. 200 m, 10.4.1981, A. Güner 3423, B. Yildiz Datça, 10 km E of Reşadye, alt. 150 m, Steep slopes with (HUB); Muğla, Fethiye: Göcek köyü, Tuglu-tepe, Kizilcam very sparse vegetation, conglomerate,Sample 22.7.1997, Peter Hein, ağaclamma pages salare, 29.4.1958, F. Yoltirik (E). Vil. Denizli: Plants from Turkey 4288 (B); C 2 Muğla: Marmisten C2 Denizli: N Karabayir (an der Str. Fethyie – Çameli) 1040 Datça‘ya 20 km, yangin alani, serpentin arazi, 80–150 m, m, steile Schutthänge (Kalkmergel), W-exp., 28.5.2005, R. 9.4.1981, A. Güner 3360 - B. Yildiz (HUB); Ophiolithgestein Ulrich 5/7 (STU). Vil. Antalya: 300 m N-W Antalya, Ant.- Datça-Halbinsel westl. Marmaris, W-Anatolien, 120 m, Kork., 25 km, ausgetrocknetes Bachbett, H bis 200 m, 21.4.1992, R. Ulrich (M); Turkey, Prov. Muğla, Datça Mar- 27.7.1979, H. Kehl S (B); Antalya: Tabak Deresi (NW Anta- maris, 6 km from Bsiçk [Emick], 110 m, Serpentine outcrop, lya), 320 m, 18.4., I. Bozakman, K. Fitz 1970 184 (W); Lyci- basal scree, 7.6.1963, Dudley D. 35449 (E, G, K, W); C2, en, Vilayet Antalya: Bucht von Tekirova, Fehas gegen Tahta- Vilayet Mugla. Marmaris – Büyükkaraağaç, Weidefläche li Dag, 150 m, 31.5.1950, Mehpare Başarman, A. Huber- nach ehem. Pinus brutia-Wald, 10 m, 8.5.1986, E. Dorn (M); Morath, Flora Anatolica 9785 (G). Prov. Mugla: distr. Marmaris, Ordugah. Cultivated dry rocky Unknown localities: C1 Muğla : d. Marmaris: Yarimadasi, slopes, 13.7.1960, Khan, Prance & Ratcliffe 25 (ANK, E, 300 m, Rocky serpentine northern slopes, 18.4.1965, Davis K); C 2: Muğla: Köyceğiz. Beyobasi Süpürgeliktepe, Ser- 41284 (K); Nif Dagh, Luschan, Pflanzen aus Pisidien (WU). Eryngium subg. Eryngium 347
Sample pages Fig. 68: Eryngium thorifolium Boiss. Drawing by Gillian Meadows in Davis, Harper & Hedge (1971: 24). a: habit, b: fruit, c: bract. Courtesy of Ian Hedge, Royal Botanic Garden Edinburgh.
Floristic records: Akman et al. (1979a: Tab.12): 10 km versant méridional du Col (Muğla). Ulrich (pers. comm..) après Köycegiz (route de Muğla), région de Fethiye (30 km 26.11.2004: Termessos, NW Antalya. route de Muğla); entre Fethiye et Kemer, A 10 km à l’E de Ula entre Köycegiz et Muğla. Akman et al. (1979b: Tab. 26): Ecology: E. thorifolium grows mostly in open Pi- Massif du Sandras Dağ, en divers points du versant sud sur nus brutia and Pinus nigra subsp. pallasiana forests in la piste de Koyceğiz (Muğla); Route de Fethiye à Çameli, partially shady and relatively dry places, sometimes Eryngium subg. Eryngium 349
Serpentine is toxic for many plants because of the The locality given in Boissier’s protologue “ad rad- release of Al and Ni protons during withering. Plants ices montis Ararat” (“at the roots of Mt. Ararat”) is ex- growing on this rock must be adapted to these heavy tremely confusing, as Mount Ararat is isolated and far metals. Furthermore, serpentine rocks are fairly nutri- outside the main distribution range of the species. All ent poor. Unlike many other serpentine species, E. specimens with the number 3570 of Aucher-Éloy bear thorifolium does not hyperaccumulate nickel, which is “Khoï” as the collecting site. This refers to the city of one of the most toxic heavy metals. It contains only 6 Khoy in Iran, well south of Ararat and close to the re- mg/kg Ni, which is particularly low for a serpentine mainder of the populations of E. thyrsoideum. plant (Reeves, pers. comm.). Although E. thorifolium is Derivation of the name: The name of this species able to grow under these conditions, it is not dependent refers to the thyrsoid synflorescence. on serpentine and may also grow on other, mostly non- Plant perennial, hemicryptophyte; green, 30–100 calcareous rock types. cm tall. Basal leaves long petiolate with broad, flat- E. thorifolium prefers a Mediterranean to Mediter- tened petioles and sheathed bases, trifoliate with undi- ranean-montane climate from sea level to 1650 m vided or irregularly divided, often trifoliate segments, (for the highest population see Akman et al. 1979b: leaves with large marginal spines, with a rounded to Tab. 26). cordate base, palmately to reticulately, rarely pinnately Akman et al. (1979a: Tab. 12; 1979b: Tab. 26) re- veined, blades 8–15 cm long, 12–20 cm broad, seg- corded E. thorifolium from two different but similar ments up to 50 mm broad. Stem with an unbranched pine-forests types: one is called the Pinus brutia–Cyti- rhizome up to 10 mm in diameter and a conspicuous sopsis dorycniifolia subsp. reesii association, which tuber, erect, with a fibrous tuft of old leaf sheaths atthe grows at altitudes of 100 to 500 m. The other one are base, whitish, up to 12 mm in diameter, single, un- the Pinus nigra subsp. pallasiana forests mostly on branched except for the short branches of the synflores- serpentine and above 1200 m, where apart from E. cence. Cauline leaves similar to the basal leaves in the thorifolium other endemics occur, including Teucrium lowest part of the stem, in the middle trifoliate, with sandrasicum, Alyssum dicolor, Asperula brevifolia, and parallel-veined petioles, pinnately to reticulately Sideritis pisidica. The relevés with E. thorifolium are veined, sheaths broad and entire, spiny, upper cauline compiled in Table 29. leaves and bracts of the synflorescence branches sim- Modified indicator values for Eryngium thorifo- ple, spiny, sessile with a broad, reticulately veined base. lium: Capitula very numerous, arranged in a condensed, of- Light: 4 Temperature: 6 Moisture: 3 Acidity: v ten extremely long thyrse, umbels globular to ovoid. Nitrogen: 3. Involucral leaves seven to eight, linear, slightly broad- Illustrations: Drawing by Gillian Meadows in ened towards the base, one-veined with a thickened Davis, Harper & Hedge (1971: 24) – Fig. 68; photo by midrib and mostly one pair of marginal spines, pun- A. Wörz – Plate 9, fig. 1. gent, about 1.5 to 2 x as long as umbels. Bracts entire, Populations and threat level: E. thorifolium is not linear with a thickened midrib, broadened at the base, very common within its small distribution range. The pungent. Sepals ovoid, obtuse, mucronate, with a white populations are scattered, most of them with no more margin. Petals linear, glabrous to hispid, white or blu- than 100 plants. The species may grow in secondary ish, wings short, about 1/4 of the petal length, lobulum habitats, such as on roadsidesSample where slopes have been inflexum pages as long as the petal, fused at its whole length, freshly exposed. Overall, the populations seem to be apex emarginate. Fruits relatively small, obovate, with stable at present. Therefore, despite its rareness, the unequal, long-acuminate, relatively large marginal ap- species seems not to be acutely endangered. It is con- pendages and very small appendages on the back. sidered rare in the IUCN Red Data List (Walter & Gil- Variability: The type specimens Aucher 3570 bear lett 1998: 589). basal leaves which are trifoliate with undivided, obo- Chemical components and ethnobotany: – vate segments similar but not identical to those of E. pyramidale, as Aucher 3570 have larger, pungent mar- 47. Eryngium thyrsoideum Boiss. in Ann. Sci. Nat. ginal teeth and more prominent veins, with bases that Bot. 3e Sér. 1: 121 (1844). – Lectotype (designated are more rounded and not prominently decurrent on the here): [Iran] “Eryngium thyrsoideum Boiss./Khoï/ petiole. Their cauline leaves are usually undivided, Aucher-Eloy-Herbier d’Orient No. 3570” (Lecto-: K). ovate, and sessile. Hybrids of Eryngium 441
13 Hybrids of Eryngium
This chapter contains the full treatment of the hybrids volucral leaves six to seven per capitulum, lanceolate, of Eryngium arranged alphabetically. with one central vein and pinnate secondary venation, General pungent, with some marginal spines, 1.3 to 3 x as long as umbel. Bracts entire, pungent. Sepals lanceolate, Hybrids are rare in the Apiaceae and also rare in acuminate. Petals linear, white, shortly ciliate, wings Eryngium subg. Eryngium. They are probably much about 1/4 of the length of the petal, lobulum inflexum more common in the American subgenera, notably in nearly reaching the base, with an obtuse, ciliate, sec- subg. Monocotyloidea, where intermediate specimens ondarily inflexed lobe, fused most of its length. Fruit and a fluctuation of characters are frequent. Most- hy unknown. brids recorded here are found growing between the par- ent species (inter parentes) and not far removed from Variability: Intermediate between the parents, clos- them, or are garden taxa known exclusively from culti- er to E. campestre, but variable in the shape of the um- vation. Some of them are imperfectly known, and her- bels, the leaves, and many other characters. Sennen barium material is scarce. (1927: 374) distinguishes var. bourgatiforme and var. campestriforme according to their habitus without pro- Eryngium x chevalieri Sennen in Bull. Soc. Bot. viding a valid description. France 49: 375 (1902). – Type material: [France] Chromosome number: 2n = 15 (Perdigó Arisó & “Cerdagne française à Llo (…), vallon d’un affluent de Llauradó Miravall 1984: 190 f.), intermediate between la Sègre rive droite.” The lectotypification of Perdigó E. campestre (2n = 14) and E. bourgatii (2n = 16). Arisó & Llauradó Miravall (1984: 192) is not valid, be- Hybrid status and biology: E. x chevalieri is a cause the specimen was collected in 1916 well after the fixed hybrid occurring in the contact zone of the parent publication of the protologue (see Reduron 2007: species (Baudière et al. 1978). The pollen is normal, 1201). A possible lectotype specimen would be: but most of the anthers are abandoned and the fruits are [France] “Environs de Llo, Pelouses, 1902 3/8, Fre. sterile (Molinas i de Ferrer & Perdigo i Ariso 1981: Sennen (BC, MPU, P)”, provided its collection really 182). predates the publication of the protologue. E. bourgatii x E. campestre Distribution: Europe: Pyrenees, in the contact Derivation of the name: The name honours J. zone of E. campestre and E. bourgatii, more or less in- Chevalier, inspector of the railways of western France ter parentes. and a friend (“mon excellent ami”) of Sennen. Specimens seen: France: Cerdagne: Environs de Llo, Pelouses, 3.8.1902, Plant perennial, hemicryptophyte; green to bluish Fre. Sennen (BC, MPU, P); Cerdagne: Valle de Llo à Castel- in the synflorescence. Basal leaves long-petiolate, lvidre, 1600 m, E loco, inter parentes, 22.8.1916, F. Sennen sheathed, palmate to trifoliate with trisect to trifid seg- – Plantes d’Espagne 2678 (BM, MA), lectotype selected by ments, or larger middle segment pinnate, occasionally Perdigó & Llauradó (1984: 192); Vallée de Vallcebolère, segments irregularly pinnate or incised, decurrent on Nord du chemin, 1990 m, 14.7.1920, Fre. Sennen, Plantes de rhachis (NOT on petiole!), ovate to rounded in outline, Cerdaigne (MPU). pinnae covering each other, bases of pinnae sometimes Spain: Catalogne: Pyrenées à Montgrony, prairies de fused, producing an intralaminar peltation, leaves pin- Mayans, 1500 m, Inter Parentes, 25.8.1913, Sennen, Plantes Sampled’Espagne pages1690 (JE, MA, W). nately to reticulately veined, spiny, similar to E. camp- estre. Stem with a short rhizome, more or less erect, Floristic records: Sennen (1917: 132): prairies de Mayens. Sennen (1927: 374): “Val de Llo, pentes herbeuses, inter pa- less branched and spreading than in E. campestre. rentes, près de Mas Jaubert, entre Osséja et Valcebollère, Cauline leaves long-petiolate, tripartite with trifid to Serra de Montgrony à Mayans”. tripartite segments, rounded in outline, spiny, palmate- Illustrations: Molinas I de Ferrer & Perdigo i Ariso ly veined, prophyllae trisect to tricuspidate, with large (1981), plate of Gagnepain in MPU, see for example marginal spines. Capitula globose to ovoid, intermedi- Reduron (2007: 1200). ate between E. campestre and E. bourgatii, also in number, arranged in a dichasium with fewer branches Eryngium x embergeri Sennen ex Sennen & Mauricio, than in E. campestre, with a conspicuous additional Cat. fl. Rif orient.: 46 (1933), nomen nudum. whorl of linear spines below the involucral leaves. In- E. dichotomum x E. tricuspidatum 474 Systematic treatment of the Eurasian and North African Species
18 General Index
(E)-caryophyllene, 260 bees, 101, (E)-nerodilol, 423 bergamotene, 257, 435 (phyllocladane, 176 bergapten, 213, 326 10-hentriacontanone, 213 betulin acid, 55 10-hentriacontanone, 304 Betulo-Adenostylietea, 105, 380 12-0-tetradecanoylphorbol acetate, 55 bicyclogemacrene, 137, 435 1-ketose, 55, 137, 176, 213, 257, 304, 326 bog, 380 1-tridecanole, 257 Boissier, Edmond, 11 2,3,5-trimethylbenzaldehyde, 423 bootstrap, 19, 81 2,4,6-trimethybenzaldehyde, 423 Bornmüller, Johannes, 159 9-hentriacontanone, 440 Brachypodio-Chrysopogonetea, 73, 122 acetlenes, 51, 326 bracteoles, 38 Achilleetum santolinae mareoticum, 226 bracts, 38 Actinolema, 46, 49, 55, 74, 75, 77, 79, 86, 88 Braun-Blanquet-method, 14 Adenostylion, 105 broadleaf forest, 73 Adenstylo-Doronicetum, 111 Bromeliaceae, 35 aegelonol benzoate, 213 Brometalia erecti, 122 aesculentin, 137 Brometalia, 198 agasyllin, 213 Brometum fibrosi, 336 alcanes, 51, 326, 440 Bromion erecti, 132 aldehydes, 440 Bromo-Plantaginetum, 132 Alepidea, 39, 46, 49, 77, 79, 81 Brunfels, Otto, 10 allogamous, 438 bumblebees, 101, 244, 376 allopolyploids, 56 bundles, 44 alpine meadows, 112 Bupleurum, 35, 51, 89 Alyssum masmenaeum, 72 butterflies, 101, 294 Ammophiletea arundinaceae, 298 C17 polyacetylenic alcohol, 112 andesite, 267 caffeic acid, 55, 213 Angelica, 46 Cakiletea maritimae, 297, 298 anthocyan, 76 Campanulo-Laserpitietum latifoliae, 111 Anthoxantho-Agrostidetum, 213 Candolle, Augustin Pyramus de, 11 aphrodisiac, 186, 214, 226, 304 carboxylic acis, 51 Apioideae, 44, 46, 55, 56, 86 Cardaegean relict element, 72 Araliaceae, 34, 36, 56, 87 Caricetalia fuscae, 382 archaeoendemic, 244, 259, 343 Caricetea curvulae, 382 Arcto-Tertiary flora, 304, 378 Carici centaureetum rupestris, 132 armeno-kurdic distributionSample pattern, 350 Caricion pages ferrugineae, 105, 111 Aruncus, 76 Carlina, 214 asparagus, 298 carpophore, 44, 47 Astragalo-Brometea, 289 caryogram, 446 Astrantia, 49, 55, 74, 75, 77, 86, 88 caryophyllene, 55, 137, 213, 313, 435 autopolyploids, 56 Cavanilles, Antonio José, 11, 143 Azorelloideae, 34 Ceanothus, 67 Banister, John, 10 cedar forests, 170 barrigenol esters, 137, 186, 310 Centaureo odessanae-Elymetum gigantei, 298 barrigenol, 213, 257, 326 cerambycidae, 320 barringtogenol esters, 326 Chamisso, Ludolf Karl Adalbert von, 11 Bauhin, Caspar, 10 chasmophyte, 72, 140, 143, 343, 344 General Index 475
Chelsea Physic Garden, 176 DNA-content, 67 Chenopodiaceae, 90 docosanal, 440 Chevalier, J. 441 dolomite, 389 chlorogenic acid, 55, 137, 176, 235, 257, 304, 326 driftline, 297, 298 chromosome base number, 56 dulcitol, 55, 226 chrysanthenyl acetate, 423 dunes, 73, 297, 298. 409 Chrysopogonetum gryllii, 213 Dürer, Albrecht, 10 Chrysopogoni-Airietum capillaris, 132 Durieu de Maisonneuve, M.C., 242 Chrysopogoni-Euphorbietum nicaeensis, 132 Echieto-Silenion villosae, 403 Chryspopgoni-Saturejon, 132 Elymo-Ammophiletum, 297 Cicendion filiformis, 439 emetic, 214 Cichorio-Eryngietum plani, 325 endocarp, 44, 47 Cirsietalia flavispinae, 306, 380 endopolyploidie Cirsio-Brachypodio pinnati 212 endosperm, 44 Cirsio-Scrophularietum, 140 Epiolobio-Meliloletum albae, 212 Cisto-Lavanduletea, 409, 414 episperm, 44 citrate, 304 Eryngio-Brometum fibrosi, 336 citric acid, 51, 213, 326 Eryngio-Bromion, 122, 134, 137 cladistic analysis, 19 Eryngio-Centaureetun rhaponticae, 111 collenchyma, 37 Eryngion variifoliae, 380 coma, 37 eryngiumgenine, 55, 326 Constance, Lincoln, 13 Erysimo crepidifolii-Festucetum valesiacae, 212 continental drift, 78 essential oils, 55, 423 Coronillo ramosissimae-Juniperetum phoenicae, 403 Eurasian-continental floristic element, 320 cotyledon, 51, 355 Euxinian floristic element, 253 Coulter, John Merle, 12 exocarp, 44 coumarin, 213, 226, 326 expectorant, 214 crispane, 435 falcarindiol, 51, 176, 213, 440 cryo-mediterranean zone, 259 falcarinol, 51, 213 257, 341, 440 curcumacrene, 440 falcarinone, 51, 326 Cynancho-Artemisietum santonicae, 323, 325 falconines, 326 Cynaro-Agropyretum pungentis, 213 farnesene, 55, 213, 342, 435 Cynosurion, 213 farnesol, 423 Dalmatian pine forest, 122 fatty acids, 55 Danthonio-Scorzoneretum villosae, 132 Ferstuco pseudoviniae-Poetum bulbosae, 132 Daphneeto-Festucetea, 73 ferulol, 257 dayas, 74, 417 Festucetum pseudovinae-valesiacae, 132 Decaisne, Joseph, 11 Festucion pungentis, 105, 111 Delaroche, François, 11 Festucion valesiacae, 212 SampleFestuco valesiacae-Stipetumpages capillaries, 212 deltoin, 226, 404 Deppe, Ferdinand, 11 Festuco-Brometea, 73, 242, 409 desert, 229 Festuco-Koelerietum splendentis, 132 diabetes, 226 fissures, 344 diaphoretic, 214 flavonoids, 55, 257, 440 dioecism, 32 flavonol, 55, 213, 304 Dioscurides, 10 floodplain, 409, 429, 434, 439 diptera, 294, 320 fossil records, 88 diterpene, 112, 176, 257, 304, 326, 341, 440 freezing microtome, 16 diterpenoids, 213 fructose, 304 diuretic, 214 Fuchs, Leonhard, 10, 11 476 Systematic treatment of the Eurasian and North African Species
furanocoumarins, 55 Hydrocotyloideae, 34, 44, 46, 56, 87 gabbro, 250 hydroxylized triterpene alcohols, 257 Galio-Potentilletum bifurcatae, 325 hymenoptera, 32, 294, 394, 438 garrigue, 241, 270, 361, 368, 369 Hypochaeris, 67, 90 geitonogamy, 32 hypocotyl, 51 Gentianella, 90 hypsophylls, 38 germacrene D, 337 ichneumonidae, 320 germacrene, 55, 137, 213, 257, 341, 440, 435 Ilex aquifolium, 179 germination, 101 inclusions, 47 glaciation, 86, 99 intralaminar peltation, 34, 150 Glaziou, Auguste François Marie, 11 Inulo oculi-christi-Stipetum pulcherrimae, 212 glucose, 304 Inulo-Caricetum praecocis, 325 glycolic acid, 304, 326 Inulo-Celmisietum arcturi, 344 glycolic acid, 51, 213 involucral leaves, 37, 38 Gondwana continent, 87, 89 irano-turanian element, 150, 156, 176, 181, 350, 384 grandivittin, 213 Isauriae, 281 granite, 245 Isoetalia, 429 granulite, 439 Isoeto-Nanojuncetea, 423, 434 grazing, 112 isoimperatorin, 213 Grisebach, August Heinrich Rudolph, 11 Jepson, Willis Linn, 11 Gros, M. 269 Junco-Isoetum velatae, 424, 430 habitat modification, 369, 419 Juncus, 35 Hacquetia, 74, 75, 77, 86, 87 Jussieu, 217 haemorrhoids, 186 kaempferol, 55, 112, 137, 176, 213, 242, 257, 261, 304, Haussknecht,. Heinrich Karl, 11 326, 341, 369, 404, 440 heavy metals, 349 kaempferol-0-3,7-dirhamnoside, 326 Hedera helix, 51 karyotypes, 56 Hedysario-Helianthemetum rubellii, 366 Kotschy, C.G.T, 286 Heldreich, T. v., 273 lactones, 137 Helianthemetea annuae, 409, 414 Lactucetum orientalis, 180 Helichryso-Armerietum dalmaticae, 132 lacunae, 35 hemicryptophytes, 33 Lagoecia, 77 Hemsley, William Botting, 12 Lamarck, Jean Baptiste, 10, 235, 261, 353 hentriacontane, 52, 176, 213, 304, 435 Laurasia, 89 heptacosan, 112, 326 leaf primordial, 355 Heracleum, 46 lemans, 324 Hermann, Paul, 10 leoss hill, 292 hermaphroditic, 193 lepidoptera, 394 Hernandez, Francesco, 10 Lichtensteinia, 46, 77 Samplelife forms,pages 33 heterobathmy, 39 heterophylly, 36 lignanes, 304 hexacosanol, 213 limestone, 72, 140, 344 holly, 394 linalool, 55, 260 Hooker, William Jackson, 11 Linné, Carl von, 10 humulene, 137 linolic acid, 55, 112, 213, 304 Huter, R., 276 L‘Obel, Mathias de, 10 hybrid section, 446 lobulus inflexus, 39 hybridization, 88, 339 Lomatium, 67 hybrids, 441 longibornene, 341 Hydrocotyle, 44 long-range dispersal, 72, 78, 86, 90 General Index 477
Lorentz, Paul Günter, 11 ornamental, 341 luteoline, 55, 440 oromediterranean, 74, 361, 380 lycaenidae, 394 orophyte, 170 Mackinlayoideae, 34 oxalic acid, 51, 213, 304, 326 Madro-Tertiary flora, 90 oxypeucedanin hydrate, 213 malic acid, 51,, 213, 304, 326 oxypeucedanin, 213 malonic acid, 51, 213, 304, 326 palaeoendemic, 159 mannitol, 55, 112, 213, 226 paraphyly, 86 marble, 389 Paronychio echinulatae-Plantaginetum serrariae, 369 marl, 389 parsimony tree, 19 marmesin, 226, 404 pentacosan, 112, 440 Mathias, Mildred Ester, 13 peridotite, 250 Mediterranean Sea Holly, 176 Petagnaea, 77, 86, 87 mericarps, 44 petroselicic acid, 55, 112, 186, 213, 226, 257, 304, mesocarp, 44 326 Messinia salinity crisis, 87 peucedanin, 213 microsatellite markers, 99, 101 Peucedanum, 46, Miocene, 87, 89, 90 phenylpropanoids, 137 Miss Wilmott’s Ghost, 257 Philippi, Rudolf Amandus, 12 Molinio-Arrhenatheretea, 256, 306 phlorogluciol glycoside, 226 monocarpic, 219, 244, 253, 339 phosphate gravels, 399 monoterpene glycosides, 213 phrygana, 198, 267 monoterpene, 226, 440 phyllocladene isomer, 423 monoterpenoids, 176, 257, 304, 326, 435 phyllocladene, 55, 260 Morison, Robert, 10 phyllomes, 38 muurolene, 257, 341 phytadiene, 213 Myrrhis, 46 phytol, 213 Myrto communis-Querceto suberis, 366 pinene, 137, 435 n-alcanes, 137, 304, 326 Pistacio lentisci-Quercetum suberis, 366 Navarretia, 74 Pleistocene, 72, 86, 96, 99, 115, 139, 304, 311, 378 nectarium, 39 Pleurotus eryngii, 214 Née, Luis, 11 Pliocene, 88, 96, 343, 378 neoendemic, 72, 281, 416 Plukenet, 10 neophytadiene, 213, 257, 341 Poion alpini, 276 nickel, 72, 349 Polemanniopsis, 47, 77 n-octanol, 326 pollen, 44, 88, 438 n-propyl-sesquiterpene, 226 pollinator, 32 ocimene, 423 polyacetylenes, 213 octacosanol, 213 Polygonetum idaei, 213 SamplePolygono-bistortae-Eryngietum pages alpini, 111 oil ducts, 44, 46 oleanolic acid, 55, 292 Polygono-Trisetion, 105 oleic acid, 55, 112, 213, 257, 304, 326 polyins, 51 Oligocene, 87 polytomy, 81, 86, 87, 88 oligosaccharides, 55, 213, 304 Porta, P., 276 Olivier, G.A. 443, 444 pozzines, 380 Ononidi-Brometum condensati, 132 Prangetea ulopterae, 155 Ononido-Rosmarinetea, 213 pre-Pleistocene steppe flora, 343 Oreomyrrhis, 90 Preslio-Eryngietum corniculati, 424, 430 organogenesis, 355 primordial leaves, 36 ornamental plant, 112, 137, 176 prophyllae, 36 478 Systematic treatment of the Eurasian and North African Species
proterandric, 101, 294 Schiede, Christian Julius Wilhelm, 11 proterandry, 32 Schlechtendahl, Dietrich Franz Leonhard von, 11 psammophyte, 409 sclerenchymatic ring, 37 pseudanthia, 38 Scorzonerion villosae, 132 pseudocorolla, 38 screes, 72 Pseudocarum, 77 Secalinetea, 73 pseudomaquis, 313 secondary rosettes, 32 pseudovivipary, 32 sectional classification, 91 Pulsatillo-Pinetum nigrae, 389 sedaneolide, 137 pyranocoumarines, 55 seedlings, 51 quartzite, 245 segetal community, 226 quercetin, 55, 112, 137, 257, 326, 440 self-incompatibility, 32, 101 quercitol, 226 Sellow, Friedrich, 11 Quercus suber forest, 361, 369, 409 Senecio delphinifoliae-Fedietum cornucopiae, 235 radiation, 86, 89, 96 sepals, 38 Rafinesque, Constantine Samuel, 13, serpentine, 72, 122, 180, 229, 316, 336, 337, 348, 349, Ranunculus thora, 345 376 ratchet search, 19 serpentinophyte, 336 receptaculum, 37 seseli acetylene, 176 refuge, 99 Seslerietea, -alia, 105 rendzina, 170 Seslerio-Festucion pall., 212 reproductive biology, 32 sesquicineole, 313 reticulate evolution, 88 sesquiphellandene, 257 rhizomes, 33 sesquiterpenes, 55, 213, 257, 313, 326, 337, 341, 440 Rhynchophorus, 155 sesquiterpenoides, 55, 112, 176, 304 Rigo, G., 276 Shakespeare, William, 214 Rocher, E., 445 šibljak, 313 rock crevices, 143 Sloane, Hans, 10 rock debris, 348 solifluction, 155 rock fissures, 72, Sparganium, 35 rock habitats, 72, 74 spathulenol, 337 Rose, Joseph Nelson, 12 Sphagnum, 380 Rosmarinetea, 213 squalene, 137 rosmarinic acid, 55, 112, 137, 176, 213, 235, 257, 304, static populations, 72 326 Steganotaenia, 46, 47, 77 ruderal habitats, 73 Stellarietea mediae, 73, 375 Rudereto-Secalinetea, 415 stigmasterol, 55 Rumici acetosellae-Artemisietum austriacae, 325 Stipeto-Morinion, 134, 137 Rumici-TragoponogetumSample orientale, 325 stolon, pages 32, 438 sabinene, 313 stomata, 37 saccarides, 55 Streptanthus glandulosus, 67, 72 saccharose,137, 176, 213, 304 stylopodium, 32, 39, 44 Salicetum waldsteinianae, 111 sudorific, 242 sand dunes, 198 tall herb communities, 72, 75, 380 Sanicula, 55, 75, 76, 77, 79, 86, 87, 88 Tamarici-Hippophaetum, 187, 213 sapogenines, 55, 137 tannin, 47, 49, 304 saponins, 55, 137, 176, 186, 213, 257, 304, 326 temporary wetlands, 73, 74 Saturejo-Edrianthetum, 132 terpene aldehyd esters, 55, 112, 137, 186, 257, 326, 337 Saturejo-Ischaemetum, 132 terpenoids, 304 Scheuchzerio-Caricetea, 382 Tethys Ocean, 87 General Index 479
Theophrastus, 10 vernal pool, 73, 74, 422, 434 thermo-mediterranean climate, 361 Villars, D., 338 therophyte, 405 Visiani, R., 445 thymol, 435 vittae, commissural, 44 tonic, 186 vittae, intrajugal, 46 Tournefort, Jospeh Pitton de, 10, 261 vittae, vallecular, 46, 47 trichomes, 34 Vulpio-Lotion, 132 Trifolio-Lolietum, 213 Walker, George Arnott, 11 Trigonelleto-Althaeetum ludwigii, 403 Wheeler Haines, R., 270 trimethybenzaldehyd, 137 Wiegand, E. 387 Trinio glaucae-Caricetum humilis, 212 Winclada, 19 trisaccharide, 326 winter-rain climate, 90 triterpene saponoid, 292 Wolff, Karl Friedrich August Hermann, 12 triterpenes, 137 woodiness, 33, 79 triterpenes, 310, 326 xanthotoxin, 326 triterpenoids, 304 Xerobrometum, 242 tubercles, 47 Xerobromion, 198 tuberculosis, 298 xero-mesogaeic element, 72, 139, 343 tubers, 33 yangambin, 304 tuff, 348 Zable, H., 446 tumbleweed, 32, 193 α-pinene, 423 typification, 14 β-elemene, 337 unestered sapogenins, 304 β-eudesmol, 423 Urban, Ignatz, 12 β-sitosterol, 226 valencene, 55, 260, 440 β-sitosterole, 55
Sample pages 480 Systematic treatment of the Eurasian and North African Species
19 Geographic Index
Aegaean Islands, 139, 140 Croatia, 99, 101, 104, 118, 122, 132, 197, 220, 442, Aegaean, 72 443, 445, 446 Afghanistan, 73, 150, 155, 176, 178, 181, 183, 185, Cyprus, 193, 197, 222, 226, 263, 265, 294, 297, 298 215, 290, 292 Czech Republic, 322 Albania, 119, 220, 311, 312, 335 Dalmatia, 114, 122, 132, 442, 446 Algeria, 193, 231, 232, 235, 297, 354, 355, 357, 358, Denmark, 294 359, 361, 366, 371, 371, 394, 398, 404, 431, 434 Dinarids, 122 Algerian Atlas, 361, 366, 368 Egypt, 198, 224, 226, 355, 358 Alps, 105, 115 England, 193 Amanus, 156 Espadanel mine, 88 Anatolia, 155 Estonia, 294 Andalucia, 147, 148, 162 Finland, 294 Andes, 67, 86 France, 101, 164, 196 , 212, 219, 294, 298, 337, 339, Antarctica, 88 434, 435, 438, 440, 441, 445 Anti-Atlas, 140, 141, 143 French Alps, 99, 104, 112 Anti-Taurus, 213 Galicia, 436, 439, 440 Apennines, 115 Georgia, 181, 193, 256, 297, 323 Ararat, Mount, 349 Germany, 75, 112, 193, 196, 213, 256, 294, 295, 320, Argentina, 56, 78, 87 326 Armenia, 152, 193, 198, 384 Gibraltar, 236, 294 Atlas, 96, 145, 381, 416 Great Britain, 295 Australia, 74, 78, 79, 86, 87, 89, 90, 294, 297 Greece, 119, 122, 136, 137, 139, 140, 193, 197, 220, Austria, 103, 112, 197, 212, 320, 324, 326 225, 248, 263, 273, 296, 298, 303, 310, 311, 313, 343, Azerbaijan, 152, 182, 193, 256 355, 387, 388 Balkan Peninsula, 73, 96, 99, 114, 115, 133, 335, 343, High Atlas, 378, 380, 399, 404 387 Hungary, 320, 322 Balkans, 56, 105, 311, 376 Iberian Peninsula, 56, 89, 394, 404, 415, 419, 423, 426, Baltic Sea, 294 428 Belgium, 193, 295, 298 Iran, 73, 150, 153, 155, 159, 160, 176, 178, 181, 183, Black Sea, 253, 257, 294, 303 185, 186, 187, 219, 332, 349, 350, 384 Bohemia, 212 Iraq, 150, 152, 224, 263, 266, 270, 272, 350 Bolivia, 56 Ireland, 294 Bosnia and Herzegovina, 104, 112, 118, 197, 220, 311 Israel, 224, 226, 248, 249, 266, 267, 294 Bosnia, 101 Istria, 132 Brazil, 56, 66, 78, 87, 90 Italy, 56, 103, 114, 116, 197, 219, 231, 235, 295, 296, British Isles, 298, 294 335, 339, 358, 359, 340, 371, 371, 421, 434, 435 Brittany, 303, 436, 438,Sample 439 Jordan, pages 224, 226, 266 Bulgaria, 193, 197, 219, 220, 296, 311, 313 Juan Frenandez-Islands, 79, 88, 89 Calabria, 371 Jura, 105 California, 73, 74, 78, 90, 422 Karatau Mountains, 283, 284, 290 Carinthia, 99 Kashmir, 181, 183, 323 Caucasus, 181, 185, 231, 253, 256 Kazakhstan, 183, 283, 285, 289, 290, 292, 323 Central Europe, 257, 306 Kyrgyzstan, 289, 290 Chile, 56, 74, 86, 89, 90 Lebanon, 150, 151, 224, 229, 248, 249, 250, 263, 265, China, 75, 320, 323 267, 274, 275, 297 Corsica, 431, 434, 435, 436 Libya, 193, 197, 431, 434 Crete, 56, 72, 139, 193, 217, 219, 261, 267, 343, 376, Luxemborg, 193 384 Macedonia, 119, 220, 311, 313, 335, 385, 387, 389 Geographic Index 481
Mallorca, 303 Serbia, 118, 311, 313, 314, 322, 334, 335, 337 Mercantour, 99 Sibiria, 75 Mesopotamia, 155 Sicily, 115, 353, 354, 355, 369, 371, 431 Mexico, 56, 66, 67, 87 Sierra Nevada, 257, 259, 260, 261 Mongolia, 323 Slovakia, 212, 213, 322 Montenegro, 101, 119, 220, 296, 311, 312 Slovenia, 115, 117, 132, 219, 296 Morocco , 67, , 73, 74, 89, 162, 140, 141, 142, 143, Spain, 88, 145, 146, 147, 162, 164, 166, 168, 170, 171, 145, 147, 148, 169, 170, 175, 189, 192, 193, 198, 231, 193, 213, 236, 241, 242, 244, 246, 259, 260, 261, 270, 232, 241, 242, 259, 277, 294, 297, 298, 304, 306, 355, 278, 279, 296, 354, 355, 356, 361, 394, 405, 407, 409, 356, 359, 361, 366, 367, 371, 371, 373, 374, 378, 380, 415, 419, 423, 424, 426, 429, 431, 436, 438, 440, 441 381, 382, 391, 393, 394, 397, 399, 400, 403, 404, 405, Swiss Jura, 101 409, 416, 419, 421, 423, 443 Switzerland, 101, 112, 193, 256 Moyen Atlas, 304, 306 Syria, 150, 151, 198, 223, 229, 230, 263, 265, 267, 274, Netherlands, 193, 196, 295 275, 431, 434 New Zealand, 90 Tajikistan, 183, 289, 290, 292 Norway, 294 Taurus, 156, 263, 267, 274, 281, 282, 287, 316, 327, Olympus, 122, 136 329, 376 Oregon, 78 Transcaucasia, 253, 257 Pakistan, 150, 183, 215, 217 Transcaucasus, 181, 185, 186, 187, 213 Palestine, 224, 263 Tunisia, 193, 231, 233, 234, 235, 263, 267, 297, 298, Peloponnes, 115 355, 358, 359, 361, 366, 369, 371, 373, 399, 374, 375, Pindus Mountains, 122, 136, 219 394, 404, 431, 433, 435 Poland, 197, 295, 320 Turkey, 67, 73, 93, 114, 150, 151, 155, 156, 159, 160, Portugal, 88, 193, 236, 240, 242, 244, 245, 295, 405, 181, 192, 193, 198, 213, 222, 227, 248, 250, 253, 263, 419, 421, 424, 426, 428, 439, 440 265, 267, 273, 274, 279, 281, 287, 294, 297, 298, 303, Pyrenees, 162, 441 316, 323, 326, 327, 329, 330, 343, 345, 348, 376, 384 Quarnero-Islands, 132 Turkmenistan, 150, 155, 176, 183 Rif Montains, 189, 259, 260, 443 Ucrainia, 193, 323 Romania, 197, 322, 442 Ural, 320 Russia, 75, 181, 193, 198, 231, 256, 320, 323 Uruguay, 56, 66, 78, 87 Sahara, 74, 143, 399 USA, 56 San Marino, 117 Uzbekistan, 181, 183, 290, 292, 308, 310 Sardinia, 355, 358, 419, 421, 431, 434 Vermion, 122 Saudi-Arabia, 263, 267 Sample pages 482 Systematic treatment of the Eurasian and North African Species
Plate 1
Fig. 1: E. alpinum, Nenzinger Himmel, Vorarlberg, Austria. Fig. 3: E. antiatlanticum, Col de Kerdous, Morocco.
Sample pages
Fig. 2: E. amethystinum var. tenuifolium, Pisidia, Greece. Fig. 4: E. aquifolium, Grazalema, Spain. Peter Leins Claudia Erbar Flower and Fruit Morphology, Ontogeny, Phylogeny, Function, Ecology
2010. X, 440 pp., 258 figures, 3 tables, 17 x 24 cm, hardcover. ISBN 978-3-510-65261-7 € 69.00 pages
"Flower and Fruit describes the fascinating role that flowers and fruits play in the evolution of plants. It represents an updated translation of the second edition of the successful standard textbook Blüte und Frucht (2nd completely revised edition, 2008) by the same authors. The origin of flowers and their important role in the life cycle of plants are described accompanied by an explanation of basic terms. The genetic approach to floral organ determination is introduced. On the basis of ontogeny, a morphological analysis of the manifold flower structures follows, laying the foundation for exploring further phylogenetic and diverse functional and ecological questions: the hermaphroditism problem, pollination optimization, pollen tube competition, adaptationsSample to the pollinators, seed dispersal patterns, modes of diaspore dispersal and adaptations of plants and their diaspores to the dispersing agents. A very useful fruit classification is included in the book. A comprehensive appendix containing more than 400 floral formulas serves as reference for placing the mentioned taxa in a classification of flowering plants. This book addresses plant scientists, students, lecturers and teachers as well as readers interested in botany, plant reproduction, biodiversity and evolution.
Schweizerbart Science Publishers · Stuttgart 2010 Johannesstr. 3 A, 70176 Stuttgart, Germany Tel. +49(0)711/3514560 Fax + 49(0)711/35145699 [email protected] www.schweizerbart.de Part 3 Bryophytes and seedless Vascular Plants Syllabus of Plant Families Wolfgang Frey (Editor) 2009. X, 419 pp., 72 figs, 1 tab. hardcover ISBN 978-3-443-01063-8 € 89,- pages
Part 3 provides a thorough treatment of the world-wide morphological and molecular diversity of a part of lower plants [Marchantiophyta, Bryophyta, Anthocerotophyta, Polysporangiomorpha, Protracheophytes, Rhyniophytina, SampleLycophytina, Trimerophytina, Moniliformopses (Cla- doxylopsida, Psilotopsida, Equisetopsida, Marattiopsida, Polypodiopsida)], and Radiatopses (Progymnospermop- sida). The advent of DNA sequencing and advances in phylogenetic analysis has raised new interest in the rela- tionships of liverworts, mosses, hornworts, ferns, and fern allies as extant representatives of early land plant evolu- tion. Following the tradition of Engler with the morphologi- cal-anatomical data and incorporating latest results from molecular phylogenetics and phylogenomics, an up-to- date overview of families and genera has been created that will serve as reference for a long time. Borntraeger BScience Publishers • Stuttgart Johannesstr. 3a, 70176 Stuttgart, Germany. Tel. +49 (711) 351456-0 Fax. +49 (711) 351456-99 [email protected] www.borntraeger-cramer.de Bibliotheca Botanica, Volume 159, 2011 A. Wörz, Revision of Eryngium L. (Apiaceae-Saniculoideae)
This well illustrated monograph exhaustively treats The monograph is divided into two parts: and revises the Eurasian and African species of The general part presents the general morphol- the genus Eryngium [E:Eryngo, D:Mannstreu, F: ogy and results of a cladistic analysis of Eryngium Panicaut]. Eryngium is an annual or perennial (using morphology, fruit anatomy, ecological data) cosmopolitan. The genus has the largest number and compares these with genetic data. The re- of species (230) within the family Apiaceae. sults validate the classification of the genus into Numerous characteristics of Eryngium are con- five subgenera (subgg. Eryngium, Monocotyloi- sidered: general morphological, chemotaxonomic dea, Semiaquatica, Foetida, and Lessonia) and and genetic characteristics, chromosome counts lead to the description of a small new subgenus, as well as ecological properties, distribution, fossil E. subg. Ilicifolia. record and reproductive biology. Every species, The special part of this revision treats 61 Eurasi- every individual is part of its habitat: biological an and North African species in detail, providing and non-biological parameters form the basis of descriptions,pages distribution maps, illustrations and an organisms existence by defining its evolution- keys for identification. Herbarium studies form the ary challenges. basis of this work. Phytosociological data serve as indicators for ecological requirements. An ex- tensive list of more than 600 references, supple- mented by lists of valid and illegal names, rounds up this volume.
Sample
www.schweizerbart.de ISBN 978-3-510-480430-2 ISSN 0067-7892
eschweizerbart_XXX Bibliotheca Botanica Original Contributions to Botany,Botany founded in 1886 ISSN 0067-7892
Bibliotheca Botanica, Volume 159
Arno Wörz: Revision of Eryngium L. (Apiaceae-Saniculoideae): General part and Palaearctic species 2011. 498 pages, 84 gures, 41 tables, 12 plates, 31 x 23 cm. ISBN 978-3-510-48030-2, bound 189.– € www.schweizerbart.com/9783510480302
This new and profusely illustrated part of its habitat: biological and Lessonia) and lead to the descrip- monograph exhaustively treats non-biological parameters form the tion of a small new subgenus, E. and revises the Eurasian and Af- basis of an organism’s existence subg. Ilicifolia. rican species of the genus Eryn- by de ning its evolutionary chal- gium [E:Eryngo, D:Mannstreu, F: lenges. The special part of this revision Panicaut]. Eryngium is an annual The monograph is divided into two treats 61 Eurasian and North Afri- and perennial cosmopolitan. The parts: can species in detail, providing de- genus has the largest number of scriptions, distribution maps, illus- species (230) within the family The general part presents the trations and keys for identi cation. Apiaceae. general morphology and results Extensive herbarium studies form Numerous characteristics of Eryn- of a cladistic analysis of Eryngium the basis of this work. Phytosocio- gium are considered: general mor- (using morphology, fruit anatomy, logical data serve as indicators for phological, chemotaxonomic and ecological data) and compares ecological requirements. genetic characteristics, chromo- these with genetic data. The re- An extensive list of more than 600 some counts as well as ecologi- sults validate the classi cation references, supplemented by lists cal properties, distribution, fossil of the genus into ve subgenera of valid and illegal names com- record and reproductive biology. (subgg. Eryngium, Monocotyloi- pletes this volume. Every species, every individual is dea, Semiaquatica, Foetida, and
Schweizerbart Science Publishers Johannesstr. 3A, 70176 Stuttgart, Germany., Tel. +49 (0)711 351456-0, Fax +49 (0)711 351456-99, [email protected], www.schweizerbart.de Bibliotheca Botanica 159
Preface ...... 5 7. Eryngium: Classi cation and Evolution ...... 77 Abstract ...... 6 7.1 The Position of Eryngium within the Saniculoideae . . 77 I General part: Taxonomy, Biogeography and Evolution . 9 7.2 The Subgeneric Classi cation ...... 77 1. Introduction ...... 9 7.3 Cladistic Analysis ...... 79 2. History of Illustration and Research in Eryngium . . . . .10 7.3.1 General ...... 79 3. Material and Methods ...... 14 7.3.2 Cladistic analysis of Subg. Eryngium ...... 81 3.1 Nomenclature and Herbarium Studies ...... 14 7.3.3 The clade of Subg. Ilicifolia ...... 81 3.2 Phytosociological Methods ...... 14 7.3.4 The cladistic analysis of subgg. Foetida, Monoco- 3.3 Distribution Maps ...... 14 tyloidea, and Semiaquatica ...... 81 3.4 Chromosome Numbers...... 15 7.4 Comparison with Molecular Results ...... 86 3.5 Fruit Anatomical and Petal Morphological Studies . . .15 7.5 Fossil Records ...... 88 3.6 Cladistic and Systematic Methods ...... 19 7.6 Biogeography...... 88 4. Reproductive biology ...... 32 8. Sectional classi cation and keys ...... 91 5. The Characters and their Taxonomic Values ...... 33 8.1 Division of Eryngium into subgenera ...... 91 5.1 Morphological and Anatomical Characters ...... 33 8.2 Division of Eryngium subg. Eryngium into sections . . 91 5.1.1 The Habits and Life Forms ...... 33 8.3 Division of the sections of Eryngium subg. Eryngium 5.1.2 Rhizomes and Tubers ...... 33 into species ...... 92 5.1.3 Aerial Stems ...... 33 8.4 Description of the anomalous Eryngium sect. Aquifolia 5.1.4 Basal Leaves ...... 33 and the division of this section into species ...... 96 5.1.5 Cauline Leaves ...... 36 8.5 Description of Eryngium subg. Ilicifolia and the division 5.1.6 Leaf Sheaths ...... 37 of this subgenus into species...... 97 5.1.7 Stomata Types...... 37 8.6 Division of Eryngium subg. Semiaquatica into species 97 5.1.8 Collenchyma...... 37 8.7 Division of the European and North African species of 5.1.9 Capitula, In orescences, and Syn orescences . .37 Eryngium subg. Semiaquatica ...... 98 5.1.10 Involucral leaves...... 37 II Systematic treatment of the Eurasian and North 5.1.11 Bracts ...... 38 African Species ...... 99 5.1.12 Sepals ...... 38 9. Eryngium subg. Eryngium ...... 99 5.1.13 Petals ...... 39 10. Eryngium subg. Ilicifolia ...... 391 5.1.14 Nectarium ...... 39 11. Eryngium tenue ...... 405 5.1.15 Pollen ...... 44 12. Eryngium subg. Semiaquatica...... 441 5.1.16 Mericarps ...... 44 13. Hybrids of Eryngium ...... 441 5.1.17 Seedlings ...... 51 14. Nomina illegitima, nomina nuda ...... 447 5.2 Chemical Characteristics ...... 51 14.1. Nomina illegitima ...... 447 5.3 Cytological Characteristics ...... 56 14.2. Nomina nuda ...... 447 5.3.1 Results and General Remarks ...... 56 15. Acknowledgements ...... 448 5.3.2 Discussion and Evolutionary Trends ...... 56 16. References ...... 450 5.4 Ecological Characteristics ...... 67 17. Index nominum ...... 471 5.4.1 General ...... 67 5.4.2 The Habitat Types ...... 67 5.4.3 Discussion of the ecological characteristics . . . . 74 6. Nomenclature and Description of the genus Eryngium L. 76
Order form
I (we) order from Schweizerbart Science Publishers (Nägele u. Obermiller), Johannesstr. 3A, 70176 Stuttgart, Germany Tel. +49 (0) 711/351456-0, Fax +49 (0) 711/351456-99 [email protected], www.schweizerbart.de
____ Ex. A. Wörz, Revision of Eryngium L. (Apiaceae-Saniculoideae): General part and Palaearctic species Bibliotheca Botanica 159. 2011. ISBN 978-3-510-48030-2, 189.– €. Name: Address:
Date: Signature:
To our customers in the USA and Canada: We invoice in US$. US$-cheques are accepted. Airmail delivery at an extra charge. Our books are available in the US through Balogh International, 1911 N. Duncan Rd., Champaign, IL 61822, USA. Phone (217) 355 9331, Fax (217) 335 9413. [email protected] Prices subject to change without notice 11.2011/10.000