Crisan Ioana et al. WILD SPP. FROM ROMANIAN MEADOWS AND THEIR IMPORTANCE FOR ORNAMENTAL BREEDING

CRIȘAN Ioana*, VIDICAN Roxana*,**, STOIAN V.*, STOIE A.*

*Department of Plant Culture, Faculty of Agriculture, University of Agricultural Sciences and Veterinary Medicine Cluj-Napoca **Corresponding author e-mail: [email protected]

Abstract Many iris that had contributions for obtaining current cultivated garden irises can be found in Romanian flora. It is known that had a great contribution in obtaining dark shade of today and had a contribution in obtaining first cultivars of Amoena type flowers admired in many current hybrids. genotypes from were documented contributors to the obtaining of fertile dwarf garden irises. Also, recently in Romanian villages were identified nameless tetraploid old garden irises similar with ‘Amas’ that can present importance for breeding disease resistant ornamental irises. All these suggest the importance as source of genetic material, the irises from our flora could still hold for breeding programs, highlighting the need of increased awareness and raised conservation efforts for native irises from meadows, even more since Red lists indicate a current decreasing trend for many wild irises in including Romania.

Keywords: garden iris, genotype, wild traits, tetraploids, flower type,

INTRODUCTION

History of the Family hybrids as well (White et al., 1997; was traced back to Colasante and Vosa, 2000), Australasia during Cretaceous while presenting many unresolved the Iris diverged from its taxonomic and nomenclatural African relatives about 45 million challenges. These include problems years ago (Goldblatt et al., 2008). of synonymy, populational Today the genus Iris is divided in 6 polymorphism, and origin subgenera, 12 sections and 260 (Colasante and Vosa, 2000). For species (Wilson and Morrison, 2011) example, not only the well-known reaching the greatest diversity in species Iris germanica it is believed (Goldblatt et al., 2008). to have appeared by crossing Irises have been known as between species (Lim, 2016), but ornamental and medicinal in others as well, like Iris pumila Romania for a long time (Butură, (White et al., 1997). Also, Tutin et 1979; Ardelean et Mohan, 2008). al. (1980) mentioned a hybrid The genus Iris is rich not only in between Iris aphylla and Iris pumila species but natural and artificial allegedly found in Central Romania.

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Crisan Ioana et al. Besides native species spuria as vulnerable (Allen et al. mentioned in this paper, in 2014; IUCN, 2017). Romanian literature are described The aim of this paper is to several others especially in older bring into attention the importance works like the one of Săvulescu et that wild irises have for plant Nyarady (1966), that currently are breeding in the context of decreasing not unanimously recognized as wild iris populations across Europe, separate species, fact acknowledged highlighting the need of increased by Romanian authors in last years awareness and raised conservation (Robu, 2005). While some can be efforts for protecting our native found in international literature as species. synonyms to existing species, others are considered of some ORNAMENTAL IRISES RELATED known species (Tutin et al., 1980; TO ROMANIAN FLORA White et al., 1997; Cullen et al., 2011). In this sense, Iris brandze First Iris breeders from past Prodan that is spread across several centuries, conducted hybridizations counties in Eastern Romania: Buzău, intuitively. They made crosses, Vrancea, Galați, Vaslui, Iași, harvested seed, raised seedlings, Botoșani (Sârbu et al. 2013) is made selections, all without considered actually Iris sintenissi awareness about cellular biology. Janka ssp. brandze. Similarly, Iris From the ornamental perspective, the halophila Pall. is considered to be conversion of garden hybrids from ssp. halophila Pall. diploids to tetraploids was the most (White et al., 1997; Tutin et al., significant event in ornamental iris 1980). Also, Iris mangaliae Prodan history (Norris, 2012). is actually referring to Because the current Linn (White et al., 1997) and Iris ornamental Iris assortment is very arenaria Waldst. et Kit. often wide (Beresford-Kroeger, 2004), the described as subspecies of Iris horticultural classification used humilis in our literature (Sârbu et al. worldwide for irises, is designated 2013) was merged simply in Iris by the abbreviated English name for humilis as of 2011 (Biltz et al., each category (ISA Handbook, AIS 2011). Handbook, British Iris Society, The Red List of European Iridarium of Komarov Botanical medicinal plants, is listing 7 Iris Institute). To these main ornamental species, from which 5 are found also categories, can be added a in Romania: Iris aphylla, Iris nomenclature of different crosses pseudacorus, Iris pumila, Iris between species and series within the sibirica, Iris spuria. Among these, genus as the one described by IUCN is listing Iris aphylla and Iris Komarnicki (2012) for the Apogon sibirica as near threatened and Iris irises.

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Crisan Ioana et al. Knowing the characteristics due to slender habitus but was little ascribed to each horticultural class is used. The successful MDB cultivars important for breeding of ornamental originated from crosses between Iris irises, constituting criteria for pumila and modern tetraploid TB evaluation of cultivars (ISA Handbook, AIS Handbook). followed by generations of selection Further are presented the for miniature proportions that lead to main horticultural classes of irises the creation of hundreds MDB related to species found in our wild registered cultivars. This paved the flora and major sources consulted for way to obtaining also SDB and IB describing the classes were those of later on. Species Group of the British Iris Standard Dwarf Bearded Society, The American Iris Society (SDB) and Iris Society of Australia, all bloom after MDB but before indicated at the end of the the IB. They have between 21-40 cm classification together with other in height. The flowers have a popular sources used. The diameter of 7-8 cm and the flower horticultural classes of irises related stem presents 3 flower buds. They to species found in our wild flora are: have a wide color range and were A. gon irises Po obtained by crosses between TB and Miniature Dwarf Bearded MDB, thus presenting the (MDB) are the shortest, with stems characteristics of both. Their history often unbranched and having begins in 1940 USA with breeders between 15-20 cm. They are the first Cook and Douglas, who managed to to flower among the bearded types in obtain fully fertile hybrids from the early spring. Their origin is in the crosses between Iris pumila and TB, meadows of , Eastern and soon other hybridizers followed. Europe (including Romania) and Cook launched the first SDB . The existence of this class cultivars. Later, breeders used owns the most to introduction of Iris existing SDB for further crosses. pumila in mainstream breeding Intermediate Bearded (IB) reach a during 1930-1950. Notably Bob height of 41-65 cm and have a Schreiner from USA used imported diameter between 8-12 cm. They seeds from Romania in late 1930 to present minimum 2 branches per obtain the miniature cultivars stem and have 4 to 5 flowers per ‘Carpathia’ and ‘Sulina’ that were stem. They were obtained by crosses the ones further used by other between SDB and TB. breeders. For some early breeders, Border Bearded (BB) are also attracted interest the shorter versions of TB. They

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Crisan Ioana et al. reach 64 cm in height and flowers Most modern TBs, BBs, and MTBs have 12.5 x 9 cm. have 48 while MDBs Miniature Tall Bearded and SDBs can have variable (MTB) have the strictest numbers, yet many classification. These diploids are produce fertile offspring by crossing distinguished by the delicacy of the on one another. SDBs x TBs crosses flowers that have to form on slender give IBs that are usually infertile and flexible stems. Are often called hybrids (Spoon, 2011). ‘table irises’. The branched flower B. Apogon irises stems reach 41-70 cm in height and Spuria (SP) have heights present 6-9 flowers each having a between 60 to over 100 cm and wide diameter of 9 cm but not larger than color range from white, blue to 15 cm. brown and often with yellow signals. Tall Bearded (TB) exceed The species belong to the series 70 cm in height. The flower stem is Spuriae from subgenus Limniris, branched and presents several buds rooted in the horticultural literature (7 or more for new cultivars). through Iris spuria and its Reunites the tall bearded garden subspecies. irises either hybrids or species: Iris Siberians (SIB) have blue, germanica, , Iris violet and white flowers. They can variegata, Iris trojana etc. Many TB reach 60-100 cm in height. Includes available at the beginning of XXth 10 species of the subgenus Limniris century were diploids, making them divided in 2 groups: first with 28 more difficult to cross with other chromosomes (sibiricae) is found species (often resulting sterile also in Romania. Tetraploids of this hybrids), much to the frustration of group are not found in the wild, but early hybridizers like of Schreiner many of today cultivars are and cytologist Marc Simonet. tetraploid. However, their crossing is Eventually, successful crosses with more difficult, and they usually set dwarf species lead to the creation of less seed than the diploids. There SDB and other horticultural Iris were also less breeding programs for categories (White et al., 1997; Norris them (White et al., 1997; Norris 2012; AIS Handbook, ISA 2012; AIS Handbook, ISA Handbook, The Blog of AIS, British Handbook, The Blog of AIS, British Iris Society, Iridarium of Komarov Iris Society, Iridarium of Komarov Botanical Institute). Botanical Institute).

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Crisan Ioana et al. IRISES OF ROMANIAN FLORA All species prefer meadows. Some are restricted to open habitats Many Iris species from on hills and plains () Romanian flora are xerophytes (Iris while others can be met in various suaveolens, Iris reichenbachii) or environments. For example, some xero-mesophytes (Iris aphylla, Iris irises can grow in glades and at forest pumila, Iris variegata, Iris edges (, Iris variegata, graminea), with only a few species ), bushy areas (Iris met in swampy meadows, or pallida, Iris variegata, Iris watersides: Iris spuria, , graminea) and grass-covered . rockeries (Iris reichenbachii, Iris Two taxa were long known pumila, , Iris aphylla) for their ability to establish on saline (Sârbu et al., 2013). soil: Iris sintenissi ssp. brandze and In table 1 are named 15 Iris Iris spuria ssp. halophila (Tutin et species from Romanian meadows al. 1980; Sârbu et al. 2013) while a avoiding synonyms for the species. few others also can tolerate to some For the spread or status of the species degree a saline substrate: Iris in the Romanian wild flora, was used germanica, Iris sibirica, Iris the latest consulted source pseudacorus, Iris pumila, Iris mentioned in the last column, for graminea (Grigore, 2008). each species.

Table 1 Inventory of Iris species from Romanian meadows (unanimously accepted names for distinct species) Species Flowering Phytocoenosis Spread or Sources time status Iris aphylla L. V-VI Geranion sanguinei, vulnerable Tutin et al., 1980; (2n= 24, 40, 48) Festucion valesiacae, Ciocârlan 2000; Sârbu et Jurineo- al., 2013; Marinescu et Euphorbinenion Alexiu, 2013 Iris germanica L. V-VI sub-spontanous cultivated Tutin et al., 1980; (2n= 36, 44, 48) Ciocârlan 2000; Sârbu et al., 2013 Iris graminea L. V-VI Brometalia erecti, frequent Tutin et al., 1980; (2n=34) Geranion sanguinei Ciocârlan 2000; Sârbu et al., 2013 Iris humilis Georgi V-VI Quercetea rare Tutin et al., 1980; (2n=22) pubescentis, Ciocârlan 2000; Biltz et Festucetalia al., 2011; Sârbu et al., vaginatae, 2013 Festucetalia valesiacae Iris pallida Lam. V-VI Festuco-Brometea rare Tutin et al., 1980; (2n=24) Ciocârlan 2000; Norris, 2012; Sârbu et al., 2013

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Crisan Ioana et al.

Iris pontica Zapal V Festucetalia vulnerable White et al. 1997; Sârbu (2n=72) valesiacae et al., 2013; Irimia et Mânzu, 2013 Iris pseudacorus L. V-VII Phragmitetalia common Tutin et al., 1980; (2n= 24, 30, 32, 34, Ciocârlan 2000; Sârbu et 40) al. 2013; Iris pumila L. IV-V Festucetalia frequent Tutin et al., 1980; (2n=32) valesiacae, Stipion Ciocârlan 2000; Sârbu et lessingianae, al. 2013; Festucion valesiacae Iris reichenbachii V Festucion valesiacae rare Tutin et al., 1980; Heuff. (2n=24, 48) Ciocârlan 2000; Sârbu et al. 2013 Iris ruthenica Ker V-VI Seslerietalia frequent Tutin et al., 1980; White Gawl. (2n=84) et al. 1997; Ciocârlan 2000; Sârbu et al. 2013 Iris sibirica L. V-VI Molinion, Molinio- sporadic Tutin et al., 1980; (2n=28) Arrhenatheretea Ciocârlan 2000; Sârbu et al. 2013; Iris sintenisii Janka V Quercetea sporadic Tutin et al., 1980; (2n=16, 18) pubescentis Ciocârlan 2000; Sârbu et al. 2013 Iris spuria L. V-VI Molinietalia, sporadic Tutin et al., 1980; (2n=20, 22) Puccinellietalia Ciocârlan 2000; Sârbu et al. 2013; Iris suaveolens IV-V Festucion valesiacae rare Tutin et al., 1980; Boiss. et Reut Ciocârlan 2000; Sârbu et (2n=24) al. 2013 Iris variegata L. V-VI Geranion sanguinei, frequent Tutin et al., 1980; (2n=24) Quercetalia Ciocârlan 2000; Sârbu et pubescentis, al. 2013 Festucetalia valesiacae

The wild Iris species found also in selection. No rebloom is possible Romanian flora although without four doses of recessive contributors to many ornamental iris alleles of the gene locus that is called classes have only yellow, blue and Frigida in Arabidopsis. Rebloomers purple shades as described in need support from high doses of autochthonous works (Ciocârlan dominant genes for vigor and 2000; Sârbu et al., 2013). However, resistance to bacterial soft rot, other there is a dominant inhibitor of diseases and pests (Spoon, 2011). anthocyanins (IA), and one for Topographical information for carotenes (IC) at a separate locus that several important species, reveals a is involved in color pattern found more or less widespread area of today in so many current cultivated distribution across Romanian irises (Spoon, 2011). All wild irises geographical regions. While Iris bloom only once a year (Table 1). spuria can be found in many counties The cultivated rebloomers have lost in Transilvania, Banat, Crișana and wild type, dominant traits by , Iris spuria ssp. halophila

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Crisan Ioana et al. is restricted to counties of Dolj, from species that created the hybrid. Prahova, Ialomița, Vrancea, Galați Modern traits for flower form, like and Iași. Limited to several counties wide and ruffled petals, come from in east and south-east Romania is Iris recessive genes that in order to be sintenisii while Iris sintenissi ssp. expressed, must occur in four doses brandze is further restricted to in tetraploids, and there is an counties of Botoșani, Buzău, increased difficulty to express Vrancea, Galați, Vaslui, Iași. recessive traits in tetraploids Also, Iris suaveolens can be compared to diploids (Spoon, 2011). met in Tulcea and Constanța. Iris It is known that , pallida is limited to Mehedinți, while can increase plant size, flower size, Iris reichenbachii can be found in fragrance, vigor and many other Mehedinți, Caraș-Severin and desired horticultural traits (Norris, Hunedoara (Ciocârlan 2000; Sârbu 2012). But more specialists today et al., 2013). start to ask if the tendency to create In flora of Cluj County, can oversized, heavy and multiple be met several important Iris species: flowers that with rain can hardly Iris pumila and Iris humilis open or even start to rot, together (Ciocârlan 2000; Sârbu et al., 2013), with disease susceptibility is what the vulnerable Iris aphylla ssp. the public is going to seek in the hungarica (Marinescu et Alexiu, future. 2013; Fig. 1), as well as Spoon (2011) citing a another rare and vulnerable species British author, said: “the natural, signaled across several counties in graceful lines of the wild iris flower Transylvania including Cluj (Irimia are being lost”. In this context and Mânzu, 2013). perhaps, a returning to the older genotypes and even wild species in CHALLENGES FOR BREEDING search for suitable genitors for a new generation of resilient garden irises There are five directions to is not far-fetched. consider in current breeding of Regarding old garden irises, ornamental irises: 1) color: flowers Blažek (2016) reports that in (including patterns) and leaves Romanian villages, he found (variegated/non-variegated), 2) tetraploids of Iris aphylla as well as flower morphology and plant tetraploid irises related to ‘Amas’, morphology according to each similar with the bitone or bicolor corresponding ornamental class clones of Iris trojana or Iris (branching stem, height etc.), 3) germanica. He concluded that blooming time (early/late, although old tetraploid irises are not rebloomers), 4) resistance to rare in European gardens they rotting. Dominant genes of Iris remain nameless. This might be of cultivars express wild type traits great interest because according to

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Crisan Ioana et al. Historic Iris Preservation Society type presents white standards and most modern tall bearded irises colored falls and it had as starting descended from ‘Amas’, an early base the species Iris reichenbachii tetraploid collected by Sir Michael and two TB cultivars. Due to dosage Foster in 1885 in (Historic effect, four doses of dominant (IAs) Iris Preservation Society). In this leads to obtaining an Amoena with sense, the “nameless” old genotypes no anthocyanin in the standards identified by Blažek (2016) in while four doses of recessive (ias) Romania could be of great interest gives an anthocyanin Self type for breeding ornamental irises, flower. The Bicolor type flower specially disease resistant ones. presents standards and falls of Although most cultivars sold today different colors. The Bitone type as garden plants are the known flowers present different shades of breaded hybrids some species might the same color on their falls and present interest too. Iris pontica, also standards, Blend type flowers found in Romania, was once present a combination of different considered to have great potential as colors. There are also other types of a rockery plant (White et al., 1997). flowers like Luminata, Neglecta, Today bearded garden irises can Plicata, Glaciata, Zonal and present flowers with various colors Variegata type flowers (Beresford- displayed in a variety of ways, and is Kroeger, 2004; Cundy et Bartlett, one of the main characteristic that 2007; Spoon, 2011; Norris 2012; distinguishes them. Most Historic Iris Preservation Society). pigmentation of the flower is in the The emblematic blue and purple upper and lower epidermal cells. The flowers rich in anthocyanins remain water-soluble violet, blue, maroon, the preferred colors for iris lovers to and cardinal red anthocyanin this day, from fine blue-violet to pigments are in the central fluid nearly black. Also, a very sought vacuoles. The lipid soluble yellow color in Iris is red, which xanthophyll and carotenes as well as preoccupied many breeders (Norris, pink, red, and orange lycopene 2012). In many cases the carotenoids pigments are in the cytoplasm and not the anthocyanins, are (Spoon, 2011). responsible for red and orange hues Besides the one-colored flowers of Iris flowers (Lim, 2016). It is called Self-type, there are several acknowledged that it is a challenge to others. The IA gene by inhibiting obtain a cultivar with intensely red anthocyanin pigments in standards, spectrum. Pink, peach, red, yellow- style arms, and falls is responsible orange, and orange in Iris are for most white, yellow, pink, and produced by the carotenoid pigment, orange in single colored flowers. lycopene. The recessive allele (t) for One dominant dose can remove most lycopene expression in four doses of the anthocyanin. Amoena flower (tttt) produces the “tangerine factor”

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Crisan Ioana et al. (Spoon, 2011). Many widespread iris have reached the highest point in cultivars display orange, white and 1990, have arisen from crosses yellow colors. Pink was the second between dark purple and blue color after yellow that triggered the cultivars, and Iris aphylla (figure 1) interest of breeders in the XXth- is credited with the intensifying century. To less interest were green pigmentation of today black iris flowers. Most black iris cultivars that cultivars (Norris, 2012).

Fig.1. Iris aphylla in Botanical Garden of UASVM Cluj-Napoca (2017)

Variegated foliage admired most effect, as a result only 5 to 10 % of notably in certain cultivars of Iris the seedlings have variegated pallida and Iris pseudacorus (Ondra, foliage. Allan Ensminger worked 2007), is controlled both by many years to obtain irises with cytoplasmic genes of the variegated flowers and foliage with chloroplasts and modifying genes in some degree of success (Spoon, the chromosomes that give a dosage 2011).

CONCLUSIONS

Most irises cultivated today The efforts of hybridizers in are tetraploids and modern traits of past decades are remarkable flowers are caused by recessive especially since to obtain the genes. expressing of recessive traits in

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Crisan Ioana et al. tetraploids is much harder compared special care. It might seem that wild to diploids. However, some recent traits could still be helpful in authors and hybridizers have thrown breeding more resilient garden irises a shadow of doubt on the old trend of today, with more graceful lines too. breeding for exacerbated increased Several autochthonous iris number and size of flowers in new species have a decreasing trend of cultivars at the cost of the elegance their populations, as it was of wild flower, that now tend to meet acknowledged by many authors and also practical difficulties for plants in IUCN, indicating the need for opening their blooms during raining sustained conservation efforts. weather and their proneness to rot. Species that had a Also, the rebloomers having lost the contribution in obtaining some wild type dominant traits are ornamental classes of irises can still susceptible to pests, lack of vigor, be found in Romanian flora and sensitivity to drought and need could be used in plant breeding.

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