Lilies of the world: Lilium species database

Lilium majoense, picture by Riz Reyes

The Genus Lilium

The genus Lilium belongs to the family Liliaceae and comprises approximately 100+ recognized species. There are obvious and not so obvious divisions among the species into subspecies (ssp.) or varieties (var.), that when included brings the number closer to 200.

Almost all native lily species can be found in the northern hemisphere in the temperate zones between roughly 30° - 60° latitude. Though today, due to human activity, lily species have been introduced to nonnative areas, and in some instances have become feral or naturalized to those areas, such as England and Scotland. Even Africa now has a lily species. Lilium zairii was discovered in Zaire in the Central Africa, though it now appears it is just a feral form of L. formosanum that escaped. This goes to show the adaptability of the genus to survive in new habitats.

The genus can roughly be divided as follows, in North America, there are roughly 30 species, mostly on the west and east coast, with some into the northeastern plains of the US and Canada. Roughly 10 species can be found in Europe proper, and 70 species can be found in Middle and East Asia (Eurasia). The Japanese archipelago has about 30 species it shares with its neighbors Korea and Russia, of which 10 or so are found only on the Japanese islands, and in many cases only in very specific areas. Indochina (Vietnam, Cambodia, and Laos) is home to some of the most exotic looking of the genus, and arguably some of the most complex in terms of taxonomy.


Distribution of the genus lilium globally

Though DNA analysis is showing that there are many more species than previously thought, there is considerable debate over if these genetic differences in fact represent new species or are representative of a larger complex or group of plants that occupy a geographically similar area or region and have simply taken on different phenotypical characteristics due to factors such as geographical isolation, genetic drift, and variation in phenotypes due to local environmental, or micro-habitat, or geographical factors.

What is a species?

Since the LSF is dedicated to the genus Lilium, it is necessary to discuss the long debate of, what is a species? A species is the most specific (species) rank or classification of an organism in the hierarchy of biological classification. A species simply defined is a related group of organisms that can interbreed and produce viable offspring. That is a pretty broad definition. For example, despite our broad differences in appearance, all humans belong to a single species Homo sapiens, which means, “wise upright man” (though some would debate if that's an accurate reflection of the species). Humans can obviously interbreed to produce viable offspring all over the world, so all humans would be considered to belong to a single species.

This can be extended to lilies as well. Not all Lilium species can interbreed though. We can cross some species that would not cross naturally with the aid of science and a laboratory and thus create “Frankenlily” (hybrids). This is an important distinction to make; only individuals that can interbreed successfully without the aid of science or special, artificial non-natural circumstances, are considered a species. “Frakenspecies” are not species. (I'm sure someone will argue this point.) Most lily species will interbreed within a specific larger regional area or complex of geographically related species or individuals, but not outside that complex. For example, most Western American lilies will hybridize with other Western American lilies, but won’t hybridize (at least without help) with Eastern American, European, or Asian lily species.

To further complicate this debate is that there is more than one definition of a species, just like there is more than one definition of a “mile” (American mile, metric mile, Olympic mile, etc.). It is important then to make sure that when we are discussing “species” we are using the same definition so to avoid confusion.

Species can also be discussed using the following criteria.

  • Taxonomical species: A group of individuals that share enough similar phenotypical characteristics to suggest they are genetically related. A good example would be the Lilium pardalinum complex.
  • Genetic species: A genetic species (as defined Baker & Bradley) is a set of genetically isolated interbreeding populations. Often determined by using DNA and other genetic analysis. A good example would be the various isolated populations of Lilium occidentale.
  • Evolutionarily significant species: a “wildlife/type species” is a population of individual specimens/organisms considered distinct for the purpose of conservation or research. A good example would be any proposed species such as Lilium crystalense.
  • Phylogenic or evolutionary species: A phylogenic species is a group of individuals in a evolutionarily divergent lineage that has maintained its genetic integrity though time and thus is unique and recognizable over time and space. A good example would be Lilium regale.
  • Ecological or “niche” species: ecological species is a group of individuals adapted to particular or unique set of factors or resources (geographical, environmental, etc.) called a “niche.” According to this concept, populations form the discrete phenological clusters that we recognize as species because the ecological and evolutionary processes controlling how resources are divided up tend to produce those clusters. Often called micro populations. A good example would be Lilium eupetes.

When we discuss a “species” we need to be sure we are all using the correct definition that is appropriate to the discussion at hand.

Subspecies & variants/varieties

This is where things can get really complicated really fast. The most specific hierarchical rank is that of a species as we have already discussed. But, as always, there are exceptions. If a species has similar taxonomical characteristics with another obviously related species, enough to suggest say, they're not siblings but perhaps cousins, then it may be described as a subspecies or variant (ssp. or var.), which are both the same thing.

Subspecies and varieties are common in large related complexes of individuals that show a wide range of phenological diversity and inhabit different, yet similar, habitats. Several good examples are the L. pardalinum complex and the L. nepalense complex. These are large genetically related populations that show a great deal of phenological variation, yet share a similar geographical region and habitat, yet are distinct enough, at least in terms of phenotype, to suggest they are possibly genetically related. True subspecies or variants are probably few. A big factor to consider when determining whether a population of individuals in a population is a subspecies is whether or not it has become geographically isolated, and the normal flow of genetic information from other populations has become inhibited or cut off altogether, allowing factors like genetic drift, microenvironment, inbreeding, etc. to lead to a unique phenotype, or possible unique genotype that is distinct, but yet still recognizable as the original species. The population could be thought of as being in the process of becoming a separate species.

In theory, there is no end to subtyping. This is called “splitting.” The number or taxonomical differences that one can claim to find in an individual population or related populations is endless. This does not mean that every plant or population that has a slight tweak here or there is a subtype or variant. I think of the example of Pacific Northwest salmon on the mighty Columbia river. There are endless tributaries on the Columbia. And salmon return to the tributary they hatched in to spawn and die. Does this mean that each tributary and its resident salmon are distinct subtypes? Or are they simply all a single species that call the Columbia river and its tributaries home? It's easy to see how this can lead down a dark endless rabbit hole.

Now, going on, there is the Alaskan salmon as well. You might argue and be correct to say that Columbia river salmon and Alaskan salmon are related and might be subspecies (related species) of a larger complex referred to as Pacific salmon. You can see where this argument can go on forever without any real agreement on anything expect it's a slippery slope once you start making minute distinctions in populations and calling them “subspecies.” The same holds true for lily species. It has been a common practice to split or subtype lily species, sometimes to a ludicrous extreme, making it seem like there are more species when actually they all belong to one species that simply has many diverse phenotypes.

Sections

The genus Lilium is further divided into seven taxonomical divisions of the genus called sections. This follows the classical division laid out by Comber. The sections help to further classify the genus by region such as Europe, Asia, etc.

There are seven sections of the genus Lilium:

  1. Martagon - (European section)
  2. Pseudolirium (North American section)
  3. Liriotypus (Eurasian section) (Mediterranean, Persia)
  4. Archelirion (oriental section) (Japan, China)
  5. Sinomartagon (Asiatic section) (China)
  6. Leucolirion (trumpet section) (Indochina)
  7. Daurolirion (Miscellaneous)

A section is a taxonomic hierarchical rank below the rank of genus, but above the rank of species. Sections may in turn be divided into subsections. Sections are typically used to help organize very large genera, which may have hundreds of species.

Example:

  • Family (Liliaceae)
    • Genus (lilium)
      • Section (e.g. Martagon, Pseudolirium, Etc.)
        • Species (e.g. martagon)
          • Subspecies or variety (e.g. album)

Conclusion

The Lilium Species Foundation is concerned with the preservation of the genus Lilium and related plants. The distribution of lilies around the world is largely within the Northern hemisphere in the temperate regions. When we discuss a species we need to be sure to define which term we are using so we can all be on the same page. When we discuss subspecies we need to keep in mind to ask where did it come from? Was it isolated for a reasonable amount to time (think centuries to millennia)? When growing a species, keep in mind that it might not be as labeled. So keep good notes and get as much information as possible before sowing to ensure you have the best possible chance of success. Take note of where it came from: climate, seasons, elevation, soil type, and so on. These are all characteristics that help determine a species as much as can it interbreed. You can’t have too much information. Don’t just order some seed and sow it and hope for the best. Especially with rarer species. If you prepare with passion, you'll have much better success than just winging it.

Lily species by section

(Source, Lilies: a guide for growers and collectors, by Edward Austin McRae)

Note: The hierarchy we are using is that as proposed by Comber (1949). There are other groupings as well, most notably by Baranova (1988). She further subdivided the genus into eleven sections and classified the European lilies into four different sections. She placed L. bulbiferum in Lilium. sect. Pseudolirium same as E. H. Wilson (1925) with other Asian and American erect-flowering lilies and L. candidum in the unispecific section Lilium. L. martagon is placed in L. sect. Martagon together with other lilies having a verticillate leaf arrangement. The remaining European lilies with Turk’s cap flowers and scattered leaf arrangements were placed in L. sect. Eurolirium Baranova (1988). (McRae 1998)

Lilium Genus section as proposed by Comber (1949)

Section 1 martagon, Eurasia (Europe & Asia): The name martagon is derived from Turkish which denotes a type of turban worn by Sultan Muhammad I. A reference to the shape of the flower which resembles a type of turban. (McRae 1998)

  • Lilium martagon (Linnaeus 1753)
    • Lilium martagon album (Weston 1772)
    • Lilium martagon albiflorum (Vukotinovi)
    • Lilium martagon cattaniae (syn. ssp. dalmaticum) (Visiani 1865)
    • Lilium martagon caucasicum (Mischenko 1928)
    • Lilium martagon daugava (syn. ssp. koknese) (Malta 1934)
    • Lilium martagon hirsutum (Weston 1772)
    • Lilium martagon pilosiusculum (Freyn 1890)
  • Lilium distichum (Nakai 1915)
  • Lilium hansonii (D.T. Moore 1871)
  • Lilium medeoloides (A. Gray 1859)
  • Lilium tsingtauense (Gilg 1904)

Section 2 Pseudolirium, North America (USA & Canada): pseudolirium contains all the erect outfacing flowered lily species (21 species roughly) found in North America. It is comprised of two words. The first pseudo or false. The second Lírio means 'flower' or 'lily' meaning 'false flower/lily'. This section is composed of four subsections.

  • Section 2, subsection A (S2a)

    • Lilium bolanderi (S. Watson 1885)
    • Lilium columbianum (Hanson ex. Baker 1847)
      • Lilium columbianum ingramii (Anderson 1942)
    • Lilium kelloggii (Prudy 1901)
    • Lilium humboldtii (Duchartre 1870)
      • Lilium humboldtii ocellatum (Elwes 1877)
    • Lilium rubescens (Watson 1879)
    • Lilium washingtonianum (Kellogg 1859)
      • Lilium washingtonianum purpurescens (Stearn 1948)
      • Lilium washingtonianum minus (Prudy 1919)

  • Section 2, subsection B (S2b)

    • Lilium maritimum (Kellogg 1875)
    • Lilium kelleyanum (Lemmon 1903)
    • Lilium occidentale (Purdy 1897)
    • Lilium pardalinum (Kellogg 1859)
      • Lilium pardalinum fragrans (Prudy)
      • Lilium pardalinum giganteum (syn. Lilium harrisianum) (Bean & Vollmer)
      • Lilium pardalinum shastense
      • Lilium pardalinum vollmerii (Eastwood 1948)
      • Lilium pardalinum wigginsii (Bean & Vollmer 1955)
    • Lilium parryi (Watson 1878)
      • Lilium parryi kessleri (Davidson 1924)
    • Lilium parvum (Kellogg 1862)
      • Lilium parvum crocatum (Stearn 1947)
      • Lilium parvum 'Hallidayi'
    • Lilium crystalense (B. Small)
    • Lilium nevadaense

  • Section 2, subsection C (S2c)

    • Lilium canadense (Linnaeus 1753)
      • Lilium canadense coccineum (Pursh 1814)
      • Lilium canadense editorum (Fernald 1943)
      • Lilium canadense flavum (Pursh 1814)
      • Lilium canadense immaculatum (Jenkinson 1932)
    • Lilium grayi (Watson 1879)
    • Lilium iridollae (M.G. Henry 1947)
    • Lilium michauxii (Poiret 1813)
    • Lilium michiganense (Farwell 1915)
    • Lilium philadelphicum (Linnaeus 1762)
      • Lilium philadelphicum andinum (Ker-Gawler 1822)
    • Lilium pyrophilum (M.W. Skinner & Sorrie)
    • Lilium superbum (Linnaeus 1762)

  • Section 2, subsection D (S2d)

L. crystalense and L. nevadaense are not currently recognized species. They are however often listed with the North American lily species. We are including them in the hopes that further research will settle the debate.

Section 3 Liriotypus, Mediterranean (Caucuses, Persia): liriotypus consists of two words, Lírio meaning 'flower' or 'lily', and typus meaning 'a mystical figure or symbol', 'something that represents the classical'. Most probably referring to Lilium candidum which is seen as the classic lily type in terms of flower form.

According to Comber (1949), Lilium section Liriotypus has a total number of 20 species and includes all European, Turkish and Caucasian species except for L. martagon, which has the widest distribution range of all Lilium species and belongs to L. sect. Martagon. Since no lilies are distributed between Asia Minor/Caucasus and eastern Afghanistan (Stern 1938), the section thus contains all but one Lilium species occurring west of this gap. With the exception of L. candidum with widely trumpet-shaped flowers and L. bulbiferum with erect bowl-shaped flowers, all members of L. sect. Liriotypus have scattered leaves and Turk’s cap flowers (McRae 1998).

  • Lilium akkusianum
  • Lilium albanicum (syn. L. pyrenaicum albanicum)
  • Lilium bosniacum (syn. Lilium carniolicum bosniacum, L. pyrenaicum bosniacum)
  • Lilium bulbiferum (Linnaeus 1758)
  • Lilium candidum (Linnaeus 1753)
    • Lilium candidum cernuum (Weston 1772)
    • Lilium candidum plenum (Weston 1772)
    • Lilium candidum salonikae (Stoker 1935)
  • Lilium carniolicum
  • Lilium chalcedonicum ((Linnaeus 1753)
  • Lilium ciliatum (P. H. Davis 1965)
  • Lilium heldreichii
  • Lilium jankae (syn. Lilium pyrenaicum jankae) (Hayek 1932)
  • Lilium kesselringianum (Mischenko 1914)
  • Lilium ledebourii (Baker) Boissier 1882
  • Lilium monadelphum (Bieberstein 1808)
    • Lilium monadelphum armenum (Eremin 1966)
  • Lilium rhodopeum
  • Lilium polyphyllum (D. Don 1840)
  • Lilium pomponium (Linnaeus 1753)
  • Lilium ponticum
    • Lilium ponticum artvinense
  • Lilium pyrenaicum (Gouan 1773)
  • Lilium szovitsianum

Section 4 Archelirion (oriental selection): achelirion is composed of arche which means 'over and encompassing', and Lírio meaning 'flower' or 'lily,' referring to 'open flower', and includes most of the lilies native to Japan, as well as some Chinese species.

  • Lilium auratum (Lindley 1862)
    • Lilium auratum pictum (Carriere 1867)
    • Lilium auratum platyphyllum (Baker 1880)
    • Lilium auratum praecox (Baker 1880)
    • Lilium auratum rubrovittatum (Duchartre 1870)
    • Lilium auratum virginale (Duchartre 1870)
  • Lilium brownii
    • Lilium brownii australe (Staph 1921)
    • Lilium brownii viridulum (syn. L. brownii colchesteri) (Baker 1885)
  • Lilium japonicum (Thunberg ex Houttuyn 1780)
  • Lilium nobilissimum (T. Makino 1914)
  • Lilium rubellum (Baker 1898)
  • Lilium speciosum (Thunberg 1794)
    • Lilium speciosum album (Baker 1873)
    • Lilium speciosum rubrum (Baker 1873)
    • Lilium speciosum clivorum
    • Lilium speciosum alba-novum (Mallet 1925)
    • Lilium speciosum gloriosoidies (Baker 1880)
    • Lilium speciosum kraetzeri (Baker 1874)

L. nobilissimum is sometimes recognized as a subspecies of L. alexandrae (e.g., L. alexandrae nobilissimum).

Section 5 sinomartagon (Asiatic section) China, Bhutan, Indochina: sinomartagon is composed of two words. Sin- meaning Chinese, from Latin Sinæ the Chinese, from Ptolemaic Greek Sinai, from Arabic Sin China. All forms are probably from Chinese Ch'in, the name of the fourth dynasty of China. And martagon which is already described in section 1, referring to the shape of a turban. This section contains three subsections.

  • Section 5, subsection A (S5a)
    • Lilium davidii (Elwes 1877)
      • Lilium davidii macranthum (Raffill 1938)
      • Lilium davidii unicolor (syn. L. davidii biodii, L. davidii willmottiae unicolor) (Cotton 1938)
      • Lilium davidii willmottiae (Raffill 1938)
    • Lilium duchartrei (Franchet 1887)
    • Lilium henryi (Baker 1888)
      • Lilium henryi citrinum (Wallace 1936)
    • Lilium lancifolium (Thunberg 1794)
      • Lilium lanciflolium flore-pleno (Regel 1890)
      • Lilium lancifolium fortunei (Standish 1866)
      • Lilium lanciflolium splendens (Leichtlin 1870)
    • Lilium lankongense (Franchet 1892)
    • Lilium leichtlinii (Hooker 1867)
      • Lilium leichtlinii maximowiczii (Regel) Baker 1871)
    • Lilium papilliferum (Franchet 1892)
    • Lilium rosthornii
  • Section 5, subsection B (S5b)
    • Lilium amabile (Palibin 1901)
      • Lilium amabile luteum (Constable 1939)
      • Lilium amabile unicolor (Comber 1935)
    • Lilium callosum (Siebold & Zuccarini 1839)
      • **Lilium** callosum flaviflorum* (Makino 1913)
    • Lilium cernuum (Komarov 1901)
      • Lilium cernuum candidum (Nakai 1917)
    • Lilium concolor (Salisbury 1806)
      • Lilium concolor coridion* (Baker 1871)
      • Lilium concolor partheneion (Baker 1871)
      • Lilium concolor pulchellum (Fischer) Regel 1876
      • Lilium concolor strictum (Hooker 1872)
    • Lilium pumilum (syn. Lilium tenuifloium) (de Candolle 1812)
      • Lilium pumilum flore-albo (E. H. Wilson 1925)
    • Lilium fargesii (Franchet 1891)
    • Lilium xanthellum (Want & Tang 1980)
  • Section 5, subsection C (S5c)
    • Lilium amoenum (Wilson 1920)
    • Lilium arboricola (W. T. Stearn 1954)
    • Lilium bakerianum (Collett & Hemsley 1890)
    • Lilium euxanthum
    • Lilium henrici (Franchet 1898)
    • Lilium mackliniae (Sealy 1949)
    • Lilium majoense
    • Lilium lophophorum
    • Lilium nanum (Klotzsch & Garcke 1862)
    • Lilium nepalense 'complex' (D. Don 1821)
    • Lilium oxypetalum (Baker 1874)
    • Lilium paradoxicum (Stearn 1956)
    • Lilium poilanei (Gagnepain 1934)
    • Lilium primulinum (Baker 1892)
      • Lilium primulinum burmanicum (Stearn 1948)
      • Lilium primulinum ochraceum
    • Lilium sempervivoideum (Léveillé 1915)
    • Lilium sherriffiae (Stearn 1950)
    • Lilium souliei (Franchet 1898)
    • Lilium stewartianum (Balfour & W. W. Smith 1922)
    • Lilium taliense (Franchet 1892)
    • Lilium wardii (F. Stern 1932)

The designation of these species have been debated for some time. It has been argued they should be placed under the L. nepalense complex of lilies. It is widely accepted that this section of species in Indochina, Bhutan, Myanmar (Burma), etc. are a taxonomical disaster. Until further research can accurately separate them into their proper sections we will leave them for now in the section they have been placed with the acknowledgment they might be moved or reclassified later.

  • Section5, subsection C?
    • Lilium brevistylum
    • Lilium lijiangense
    • Lilium anhuiense
    • Lilium habaense
    • Lilium huidongense
    • Lilium jinfushanense
    • Lilium matangense
    • Lilium medogense
    • Lilium pinifolium
    • Lilium pyi
    • Lilium saccatum
    • Lilium tiaschanicum
    • Lilium floridum

The species under the this heading have not been formally placed under any section but it's assumed based on their location they are best suited for Sec. 5c. We will place them in their appropriate section as more information becomes available.

Section 6 Leucolirion (trumpet section) (China): section Leucolirion is divided into two subsections. Leucolirion is composed of the words, leuco which means 'white' or 'light colored', and Lírio which means 'flower' or 'lily'.

  • Section 6, subsection A (S6a)
    • Lilium leucantheum (Baker 1901)
      • Lilium leucanthum centifolium (Stearn 1935)
      • Lilium leucanthum chloraster (Wilson 1925)
    • Lilium regale (Wilson 1912)
    • Lilium sargentiae (Wilson 1912)
    • Lilium sulphureum (Baker 1892)
    • Lilium wenshanense
  • Section 6, subsection B (S6b)
    • Lilium anhuiense
    • Lilium formosanum (Wallace 1891)
    • Lilium longiflorum (Thunberg 1794)
    • Lilium neilgherrense (Wight 1858)
    • Lilium philippinense (Baker 1873)
    • Lilium wallichianum (Schultes & Schultes 1830)
    • Lilium puerense

Lilium neilgherrense is sometimes placed as a subspecies of L. wallichianum (e.g., L. wallichianum neilgherrense)

Section 7 Daurolirion (miscellaneous): This section is composed of only two species. Daurolirion is composed of the words Dauria, a province in Siberia, and the word Lírio which means 'flower' or lily'.

  • Lilium dauricum (syn. L. pennsylvanicum) (Ker-Gawler 1809)
  • Lilium maculatum (syn. L. wilsoni) (Thunberg 1794)