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Additional Information for Efsa Copies of literature review This document contains copies of three recent literature reviews. Hyperlinks to each review are provided in the table below Scope of review EU agency to Date Related which literature dossiers review sent A literature search on information relevant for EFSA August C/NL/09/01 the safety of GM carnation to humans, 2014 and including the safety of delphinidin and C/NL/09/02 potential allergenicity⃰. Updated assessment of the probability of gene Bureau GGO, NL October C/NL/13/02 dispersal from cut-flowers of the cultivated 2013 carnation (Dianthus caryophyllus) imported into Europe Updated review of potential toxicity and Bureau GGO, NL October C/NL/13/02 allergenicity of the acetolactate synthase (ALS) 2013 protein ⃰ Copies of the papers cited in this review are provided. Copies of literature review An assessment of the probability of gene dispersal from cut-flowers of the cultivated carnation (Dianthus caryophyllus) imported into Europe CONTENTS Section Page 1. Baseline information; Dianthus biology 1 1.1 Taxonomy 2 1.2 Distribution 3 1.3 Reproductive biology 7 1.4 Inter- and intra-specific hybridisation 8 1.5 Weediness of Dianthus 10 2. Baseline information: carnation 11 2.1. Types of cultivated carnation 11 2.2. Carnation cultivation methods 12 2.3. Carnation utilization in Europe 13 2.4. Carnation import and distribution in Europe 13 3. Probability of gene dispersal from carnation 14 3.1. Introduction 14 3.2. Probability of gene dispersal by vegetative propagation 14 3.3. Probability of gene dispersal by seed set and seed distribution 14 3.4. Probability of gene dispersal by pollen distribution 15 4. Conclusions 16 5. Literature cited 18 1. Baseline information; Dianthus biology Attachment B4 has been expanded from earlier versions included in previous marketing applications to the EU. As part of the regulatory process for the release of transgenic carnation in Australia, the Australian government has also produced an assessment of the biology and ecology of carnation (OGTR, 2006). 1.1 Taxonomy Dianthus genus The Dianthus genus is a member of the Caryophyllacea, or pink, family and contains about 300 species. The genus is native to Europe, Asia, North Africa and the Arctic region, where one species is found (Hickey and King, 1981; Tutin and Walters, 1993). The second edition of Flora Europaea (Tutin, and Walters, 1993) lists 115 species, and 91 sub-species within 32 of these species. Seventy seven of the species listed are endemic to Europe. Copies of literature review Recent studies have suggested that the Dianthus genus has one of the fastest rates of evolution in plants, possibly due to ployploidization (Weiss et al., 2002; Balao et al., 2009; Balao et al., 2011b). Dianthus chromosomes number ranges from 30 (diploid) to 105 (Weiss et al., 2002; Andersson- Kotto and Gairdner, 1931). Dianthus species are largely perennial, though some species, such as D. armeria, are annuals (Jurgens et al., 2002). Dianthus caryophyllus In its unimproved, single flower form, Dianthus caryophyllus is called, in English, the clove pink or Grenadine (Britannica, 1999) or the clove gillyflower (Harvey, 1978). The English translation of the Latin word “caryophyllus” is “cloves” and caryophyllus was a commonly used name in the early taxonomy of the Dianthus genus (Smith, 1794). Clove pink was grown in the middle ages for its clove like perfume (Harvey, 1978) and was named by Linnaeus in Species Plantarum (De Langen et al., 1984). Dianthus caryophyllus may be taken as the type species for Dianthus. The taxonomy of Dianthus caryophyllus has a long history and there are many sub-species and varieties of this species in the scientific literature. Dianthus siculus, D. arrosti, D. gasparrinii, D. virgatus, D. tarentinus and D. longicaulis were sometimes previously treated as subspecies or varieties of D. caryophyllus (Bacchetta et al., 2011). Bracchi and Romani (2010) include Dianthus caryophyllus var. inodorus L. and Dianthus caryophyllus var. orophilus (Jord.) Rouy & Foucaud in a recent checklist of Piacenza, Italy and Giardina et al. (2007) list a synonym of Dianthus gasparrinii as D. caryophyllus subsp. Gasparrinii. A worldwide Dianthus species, D. sylvestris, is known by the synonym D. caryophyllus ssp. sylvestris or var. sylvestris (Flora Piacentina, 2001; http://www.anthos.es). The e-flora of the French botany network (http://www.tela- botanica.org/site:accueil) lists nine varieties and eight sub-species of D. caryophyllus, in addition to the species itself. A compilation of the sub-species and varieties of Dianthus caryophyllus are shown in Table 1. The confusion in nomenclature can lead to mis-identification in the field – a problem recognized by Smith (1794) two centuries ago. Table 1. Sub-species and varieties of Dianthus caryophyllus recorded in the taxonomic literature Dianthus caryophyllus subsp. sylvestris (Wulfen) Dianthus caryophyllus var. inodorus L., Rouy & Foucaud, 1896 ( = Dianthus sylvestris 1753 Wulfen ). Dianthus caryophyllus var. collivagus (Jord.) Cariot & Dianthus caryophyllus var. juratensis St.-Lag., 1889 Gren., 1865 Dianthus caryophyllus var. consimilis (Jord.) Rouy & Dianthus caryophyllus var. orophilus Foucaud, 1896 (Jord.) Rouy & Foucaud, 1896 Dianthus caryophyllus var. guyetanii (Jord.) Rouy & Dianthus caryophyllus var. saxicola Foucaud, 1896 (Jord.) Cariot & St.-Lag., 1889 Dianthus caryophyllus subsp. longicaulis (Ten.) Dianthus caryophyllus subsp. Arcang. godronianus (Jord.) Sennen, 1932 Dianthus caryophyllus var. longicaulis (Ten.) Dianthus caryophyllus subsp. virgineus P.Fourn., 1936 sensu auct. plur. Dianthus caryophyllus subsp. siculus (C.Presl) Dianthus caryophyllus subsp. Arcang., 1894 coronarius (Lam.) Bonnier, 1913 Dianthus caryophyllus var. coronarius L., 1753 Dianthus caryophyllus subsp. coronarius (Lam.) P.Fourn., 1936 Information;http://www.florealpes.com/fiche_oeilletsauvage.php?photonum=4&PHPSESSID=98ka413mmbrqb38otk1c bh8n5egkvi97# (Fleurs des Hautes-Alpes, de montagne, de Provence et d'ailleurs) and http://inpn.mnhn.fr/espece/cd_nom/133832/tab/rep (INPN, Inventaire National du Patrimoine Naturel, France.) Copies of literature review The chromosome number of Dianthus caryophyllus samples collected from the wild has consistently been counted at 30 (Jones and Cooper, 1968; Gadella and Kliphuis, 1970; Hassall, 1978). Carnation Carnations are double-flowered cultivars and in the general trade, botanical and horticultural literature carnation cultivars are considered to belong to the species Dianthus caryophyllus (Smith, 1794). The common name for Dianthus caryophyllus is carnation. However, the exact taxonomic and breeding history of carnation is not precisely known (Hughes, 1991; Harvey, 1978) and it is almost certain that carnation is a hybrid involving two or more Dianthus species, one of which is likely to be Dianthus caryophyllus (Hughes, 1991; Allwood, 1954). It is believed that carnation breeding began in the 1500’s in France (Holley and Baker, 1963). The double-flowered varieties of the carnation were known as an ornamental plant in Europe in the 15th century (Harvey, 1978). The herbarium specimen assessed by Linnaeus included a double-flowered carnation specimen (this may be viewed at http://www.linnean-online.org). More than one hundred years ago carnation breeding was well established in the USA, and today there are half a dozen large breeders in the world. The second edition of the International Dianthus register (1983) lists over 30,000 cultivars. 1.2 Distribution Dianthus genus The centre of biodiversity for Dianthus is southern Europe and the greatest range of Dianthus species are found in the south eastern European countries. Table 2 (adapted from Tutin and Walters, 1993) lists the number of species recorded in countries with the greatest diversity of Dianthus species. In Europe, most Dianthus species are found in the Balkan region and in the Mediterranean countries. In North Europe Dianthus species are far less common, or even absent. For example, only six species are known in the British flora (Clapham et al., 1987), and five in Holland (Tutin and Walters, 1993). The majority of Dianthus species are not widely spread in Europe, and are confined to one or two countries, to specific mountain regions (Strid and Tan, 1997) or to alpine areas (Schwegler, 1979). Table 2. European countries with the greatest diversity of Dianthus species Country Number of Dianthus species Former Yugoslavia 44 Bulgaria 39 Greece 37 Romania 32 Central + S.W. European Russia 31 Spain 26 Italy 24 Albania 21 France 20 There are six Dianthus species that have been found throughout the world. They are described below using information from Tutin and Walters (1993): • D. barbatus. Not a European species exclusively, D. barbatus is native to all Balkan countries and Eastern Europe. It has naturalized elsewhere after escape from cultivation. Copies of literature review • D. armeria. The most widely distributed Dianthus species in Europe, as far north as Southern Sweden. This species has a world-wide distribution. • D. sylvestris. (Bacchetta et al., 2010). This species is highly polymorphic and closely related to D. caryophyllus. D. sylvestris is found in southern Europe and Mediterranean islands. There are six sub-species, three of these which are endemic to Europe, though the species itself is not. Dianthus sylvestris is found “from SE Spain to Greece and Northwards to the Swiss Jura and the Alps”. The typical habitat is rocky places (Polunin, 1980). • D. superbus. This species, which is not endemic to Europe, is found in all areas except much of the west and south of Europe (Tutin and Walters, 1993). There are three sub-species. • D. deltoides. This species is not endemic to, but is distributed in, most of Europe. It is rarer in the south (Tutin and Walters, 1993). Experiments have shown this species capable of establishment in restored meadows (Kalle et al., 2009). • D. carthusianorum. The only one of the six species which is possibly endemic, this species is found in south, central and western countries and has a very variable form. In northern European countries the majority of Dianthus populations, if present, will be represented by one or more of the above six species (e.g.
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