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Biological Flora of Central Europe Himantoglossum Adriaticum H

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Biological Flora of Central Europe Himantoglossum Adriaticum H 1 The original published PDF available in this website: 2 https://www.sciencedirect.com/science/article/pii/S1433831918301768?via%3Dihub 3 4 Biological flora of Central Europe Himantoglossum adriaticum H. Baumann 5 6 Judit Bódisa*, Éva Biróa, Timea Nagya, Attila Takácsb,c, Gábor Sramkóc,d, Richard M. 7 Batemane, Lilla Giliánf, Zoltán Illyésg, Jácint Tökölyih, Balázs András Lukácsi, Miklós Csábij, 8 Attila Molnár V.d 9 10 a Department of Plant Sciences and Biotechnology, University of Pannonia, Georgikon 11 Faculty, Festetics u. 7., H-8360 Keszthely, Hungary 12 b Department of Ecology, University of Debrecen, Egyetem tér 1., H-4032 Debrecen, Hungary 13 c MTA-DE ‘Lendület’ Evolutionary Phylogenomics Research Group, Egyetem tér 1., H-4032 14 Debrecen, Hungary 15 d Department of Botany, University of Debrecen, Egyetem tér 1., H-4032 Debrecen, Hungary 16 e Royal Botanic Gardens Kew, Richmond, Surrey, TW9 3DS, United Kingdom 17 f Institute of Botany and Ecophysiology, Szent István University, Páter Károly u. 1., H-2100 18 Gödöllő, Hungary 19 g Mindszenty Youth House, Várberki u. 13., H-8900 Zalaegerszeg-Botfa, Hungary 20 h MTA-DE Behavioural Ecology Research Group, Egyetem tér 1., H-4032 Debrecen, 21 Hungary 22 i Department of Tisza River Research, MTA Center for Ecological Research, DRI, Bem tér 23 18/C, H-4026 Debrecen, Hungary 24 j Kerék u. 4., H-1035 Budapest, Hungary 25 26 *Corresponding author. Tel.: +36305473156. 27 E-mail address: [email protected] (J. Bódis). 28 29 Abstract 30 Himantoglossum adriaticum H. Baumann is a long-lived perennial orchid with an adriato- 31 mediterranean distribution. The species-level separation of this species from the more 32 geographically widespread H. hircinum has only recently been confirmed via a combination 33 of molecular and morphometric techniques, which are further developed here. To provide a 34 comprehensive overview of its autecology we integrated previously published information 35 with extensive unpublished data derived mainly from populations in the Keszthely Hills of 36 Hungary. In this paper we assess the distribution, habitat preferences, life history and seed 37 germination (ex situ and in situ) of H. adriaticum, with special emphasis on its reproductive 38 biology. 39 40 Keywords: 41 Orchidaceae; dormancy; genetic and morphological variation, life cycle; pollination; 42 reproductive biology; 43 44 45 46 47 Contents 48 Introduction ................................................................................................................................ 2 49 Morphology and taxonomy ........................................................................................................ 3 50 Nomenclature and taxonomy ................................................................................................. 3 51 Morphology ............................................................................................................................ 3 52 Is H. adriaticum a genuine species distinct from H. hircinum? ............................................. 4 53 Evolutionary origin of H. adriaticum ................................................................................. 6 54 Distribution and habitat requirements ........................................................................................ 6 55 Geographical and altitudinal distribution ............................................................................... 6 56 Substratum .............................................................................................................................. 7 57 Habitats and associated plant communities ............................................................................ 7 58 Life cycle, phenology and growth .............................................................................................. 8 59 Phenology and growth ............................................................................................................ 8 60 Life cycle and dormancy ........................................................................................................ 9 61 Seed production and dispersal .............................................................................................. 10 62 Seed germination (ex situ and in situ) and seedling morphology ........................................ 10 63 Mycorrhizae ......................................................................................................................... 11 64 Spatial distribution of plants within populations .................................................................. 11 65 Responses to abiotic and biotic factors .................................................................................... 11 66 Response to climate factors .................................................................................................. 12 67 Response to competition and management .......................................................................... 12 68 Herbivores and pathogens .................................................................................................... 13 69 Floral biology ........................................................................................................................... 13 70 Pollination ............................................................................................................................ 13 71 Fruit set ................................................................................................................................. 14 72 Factors affecting fruit set ...................................................................................................... 14 73 Biotic factors .................................................................................................................... 14 74 Abiotic factors .................................................................................................................. 14 75 Physiological and biochemical information ............................................................................. 14 76 Physiological data ................................................................................................................ 14 77 Biochemical data .................................................................................................................. 15 78 Genetic data .............................................................................................................................. 15 79 Chromosome number ........................................................................................................... 15 80 Conservation ......................................................................................................................... 15 81 Acknowledgements .................................................................................................................. 16 82 References ................................................................................................................................ 16 83 84 Introduction 85 86 The genus Himantoglossum W.D.J. Koch includes some of the most conspicuous orchids 87 native to central Europe. Its large and showy flowers are characterized by a greatly elongated 88 central labellar lobe that emerges from the bud in circinnate form but, once extended, 89 transforms into a sinistral spiral (Bateman et al., 2013). The species-level separation of H. 90 adriaticum H. Baumann from the more geographically widespread H. hircinum has only 91 recently been cemented via a combination of molecular and morphometric techniques applied 92 across Eurasia (Sramkó et al., 2014; Bateman et al., 2017). Here, we have assembled an 93 international team to bring together diverse data, both published and unpublished, with the 94 aim of generating a data-rich review of this increasingly well-understood species. 95 96 To provide a comprehensive overview of topics such as morphology and taxonomy, 97 distribution and habitat requirements, life history, phenology, growth patterns and floral 98 biology, we used previously published information as well as unpublished data. Observations 99 of the species’ ecology were conducted across nearly the whole of its distribution area, but the 100 majority of our previously unpublished data were collected in Hungary, mainly in the 101 Keszthely Hills close to Keszthely town. Data were collected from sites along a minor road 102 (approximately 1.8 km in length); from 1992 until 2007, 154 tagged plants were surveyed 103 individually, and from 1999 until 2014, 0.5 × 0.5 m wire-grid plots were established in areas 104 of high juvenile density to capture adult as well as seedling and juvenile data. 105 The growth stage of an individual was recorded as seedling (seedling1: single-leaved 106 individuals with leaf width equal to or less than 0.5 cm; seedling2: single-leaved individuals 107 with leaf width of 0.6–1.0 cm), juvenile (single-leaved individuals with leaf width of 1.1–1.5 108 cm or two-leaved individuals), sterile adult (two-leaved individuals with largest leaf width 109 equal to or greater than 1.6 cm or with three or more leaves), flowering adult (individuals that 110 produced an inflorescence), dormant (individuals that disappeared in one year but re-appeared 111 in a subsequent year) or dead (individuals confirmed dead or typically invisible for three or 112 more years). 113 Morphological observations included number of leaves, plant height (cm), and length and 114 width of the largest leaf (cm). For
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