Scilloideae: Formerly circumscribed as Hyacinthaceae, this subfamily harbors a diverse group of geophytes being specially diverse in Mediterranean regions, widely distributed from Europe and North Africa to Central Asia and South America, renowned for their seasonality, bulbous habit, and iconic role in the spring floras of arid and seasonally arid regions.
Scilloideae, a subfamily of Asparagaceae, comprises around 60+ genera and approximately over 1000 species, primarily distributed in Africa and Eurasia. The group is well known for its geophytic life strategy, characterized by the presence of bulbs as underground storage organ. Plants have one or more basal leaves, forming a rosette, and a racemose inflorescence, and a basic floral plan of six more or less undifferentiated tepals. Many members are adapted to seasonally dry climates and exhibit striking floral diversity with only minimal morphological alteration from the basic plan of the flowers. The subfamily includes well-known genera such as Hyacinthus, Lachenalia and Ornithogalum, among several others.
Historically, the taxa now placed in Scilloideae were previously classified within the broadly defined Liliaceae and later in the segregate family Hyacinthaceae, depending on the system used. Molecular phylogenetic studies have demonstrated the close relationship of Hyacinthaceae with a diverse range of groups, now circumscribed in a broadly defined Asparagaceae, but separate from the Liliaceae.
Today, Scilloideae is divided into multiple tribes and subtribes based on molecular and morphological characteristics. The most widely accepted classification recognizes four tribes: Hyacintheae, Ornithogaleae, Oziroëeae, and Urgineeae (Chase et al., 2009). There are only a few characters to distinguish the tribes, the most consistent being the presence of particular groups of compounds which has led to research on the possible medical uses of plants in the subfamily.
Hyacintheae is the largest tribe and is further subdivided in three subtribes (Hyacinthinae, Massoniinae & Pseudoprosperinae), including around 30 genera and several hundred species. It is predominantly distributed in the Mediterranean Basin and southern Africa. Members of this tribe typically have blue, star-like flowers, that in some genera become connate (tubular) and have a wide range of colours. The tribe includes some highly ornamental and well known examples, such as Hyacinthus, Muscari, and Scilla.
Ornithogaleae includes genera such as Ornithogalum, Galtonia, Albuca and Dipcadi, with a distribution extending from Africa to Europe and Asia. Members of this tribe typically have white, star-like flowers and a wide range of ecological adaptations. Some species are used in horticulture, while others have medicinal properties.
Oziroëeae is a small tribe that occurs in southern South America, representing an interesting biogeographical disjunction from other Scilloideae tribes, that is further reflected in its phylogenetic position as sister to the remaining tribes. It includes a single genus, Oziroë, which differs from other members of the subfamily in several morphological traits, such as bulb structure and floral arrangement.
Urgineeae include around 200 species distributed across Africa, Europe, and southwestern Asia. Its classification has been debated, with some treatments recognizing multiple genera (e.g., Austronea, Drimia, Indurgia, Schizobasis, Tenicroa, Thuranthos, Urginea, Urgineopsis, Vera-duthiea), while others favour a broader concept lumping many species into Drimia sensu lato. Recent phylogenetic studies, however, support a more fine-scaled classification (Martínez‐Azorín et al., 2023).
Despite significant advances in the phylogenetic classification of Scilloideae, some issues remain unresolved. The placement of certain genera, particularly those with intermediate morphological characteristics, continues to be debated. Additionally, hybridization and polyploidy complicate genetic studies, leading to difficulties in defining species boundaries. Ongoing research by this group using genomic approaches will help clarify these issues and provide a more stable classification, which will be reflected in our here proposed taxonomic treatment of the group.
Key Contributors (in alphabetical order)
Böhnert, Tim (University of Bonn, Germany)
Crespo, Manuel B. (Universidad de Alicante, Spain)
Crouch, Neil R. (BRAM, South African National Biodiversity Institute, South Africa)
Culham, Alastair (University of Reading, UK)
David, John (University of Reading, UK)
Dold, Anthony P. (Rhodes University, South Africa)
Duncan, Graham (South African National Biodiversity Institute, Kirstenbosch National Botanical Garden, South Africa)
Eker, İsmail (Bolu Abant İzzet Baysal University, Türkiye)
Hankey, Andrew (South African National Biodiversity Institute, Walter Sisulu National Botanical Garden, South Africa)
Lekhak, Manoj M. (Shivaji University, India)
Martínez-Azorín, Mario (Universidad de Alicante, Spain)
Pinter, Michael (University Graz, Austria)
Yildirim, Hasan (Ege University, Türkiye)
Zarre, Shahin (University of Tehran, Iran)
Selected Literature
- Ali̇Pour S, Maji̇Di̇ R, Eker İs (2024) Muscari zagricum (Asparagaceae, Scilloideae), a unique new species from Iran. Phytotaxa 652: 133–141. https://doi.org/10.11646/phytotaxa.652.2.5
- Böhnert T, Lobin W (2017) Leopoldia neumannii sp. nov. (Asparagaceae, Scilloideae): a new species of Muscari sensu lato from Greece. Willdenowia 47: 179–185. https://doi.org/10.3372/wi.47.47210
- Böhnert T, Lobin W (2023) Lectotypification of Muscari heldreichii (Asparagaceae) from Greece. Phytotaxa 584: 130–132. https://doi.org/10.11646/phytotaxa.584.2.6
- Böhnert T, Del Guacchio E (2023) The type and correct authorship of the name Muscari neglectum (Asparagaceae). Taxon 72: 1334–1339. https://doi.org/10.1002/tax.13025
- Böhnert T, Neumann M, Quandt D, Weigend M (2023) Phylogeny based generic reclassification of Muscari sensu lato (Asparagaceae) using plastid and genomic DNA. Taxon 72: 261–277. https://doi.org/10.1002/tax.12864
- Chase MW, Reveal JL, Fay MF (2009) A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161: 132–136. https://doi.org/10.1111/j.1095-8339.2009.00999.x
- Crespo MB, Martínez-Azorín M, Alonso MÁ (2020) The identity of Drimia purpurascens, with a new nomenclatural and taxonomic approach to the “Drimia undata” group (Hyacinthaceae = Asparagaceae subfam. Scilloideae). Plant Systematics and Evolution 306: 67. https://doi.org/10.1007/s00606-020-01689-1
- Crouch NR, Martínez-Azorín M (2015) Drimia edwardsii (Asparagaceae, Scilloideae), a new urgineoid species from the Mkhomazi River Valley of eastern South Africa. Phytotaxa 195: 135. https://doi.org/10.11646/phytotaxa.195.2.2
- Eker İ (2021a) Taxonomic resurrection of Muscari wallii (Asparagaceae, Scilloideae). Phytotaxa 513: 226–242. https://doi.org/10.11646/phytotaxa.513.3.3
- Eker İ (2021b) What is Muscari massayanum and what is not? Second species born of confusion: Muscari erzincanicum (Asparagaceae, Scilloideae), a new species from Turkey. Phytotaxa 487: 41–55. https://doi.org/10.11646/phytotaxa.487.1.3
- Eker İ, Yildirm H (2021) Muscari inundatum (Asparagaceae, Scilloideae), a new species from southern Anatolia. Phytotaxa 484: 181–194. https://doi.org/10.11646/phytotaxa.484.2.3
- Eker İ, Eroğlu H (2024) The last missing species born of a confusing puzzle: Leopoldia nigdeensis (Asparagaceae, Scilloideae), a new species from Türkiye. Phytotaxa 635: 59–74. https://doi.org/10.11646/phytotaxa.635.1.3
- Eker İ, Duman H, Yildirm H (2020) Muscari muglaensis (Asparagaceae, Scilloideae), a new species from southwestern Anatolia. Phytotaxa 475: 267–278. https://doi.org/10.11646/phytotaxa.475.4.4
- Garnett GJL, Könyves K, Bilsborrow J, David J, Culham A (2020) The complete plastome of Hyacinthoides non-scripta (L.) Chouard ex Rothm. (Asparagaceae). Mitochondrial DNA Part B 5: 1003–1004. https://doi.org/10.1080/23802359.2020.1720543
- Martínez‐Azorín M, Crespo MB, Juan A, Fay MF (2011) Molecular phylogenetics of subfamily Ornithogaloideae (Hyacinthaceae) based on nuclear and plastid DNA regions, including a new taxonomic arrangement. Annals of Botany 107: 1–37. https://doi.org/10.1093/aob/mcq207
- Martínez‐Azorín M, Crespo MB, Alonso‐Vargas MÁ, Dold AP, Pinter M, Wetschnig W. (2018) Austronea (Asparagaceae, Scilloideae), a new genus from southern Africa, including the description of seven new species. Phytotaxa 365: 101–129. https://doi.org/10.11646/phytotaxa.365.2.1
- Martínez‐Azorín M, Crespo MB, Pinter M, Slade JM, Wetschnig W. (2019) Iosanthus (Hyacinthaceae subfam. Urgineoideae), a new genus from Southern Africa to include Ornithogalum toxicarium and its removal from Ornithogaloideae. Plant Biosystems 153: 580–588. https://doi.org/10.1080/11263504.2018.1527793
- Martínez-Azorín M, Crespo MB, Alonso-Vargas MÁ (2022) Reinstatement of Squilla Steinh., a priority name against the illegitimate Charybdis Speta (Hyacinthaceae, Urgineoideae). Mediterranean Botany 43: e78272. https://doi.org/10.5209/mbot.78272
- Martínez-Azorín M, Dold AP, Crespo MB, Pinter M, Alonso-Vargas MÁ, Wetschnig W (2019) A taxonomic revision of Geschollia (Asparagaceae, Urgineeae)—from a monotypic genus towards its diversification, including the description of five new species. Phytotaxa 427: 85–114. https://doi.org/10.11646/phytotaxa.427.2.1
- Martínez-Azorín M, Crespo MB, Alonso-Vargas MÁ, Dold AP, Crouch NR, Pfosser M, Mucina L, Pinter M, Wetschnig W (2019) New combinations in the tribe Urgineeae (Asparagaceae subfam. Scilloideae) with comments on contrasting taxonomic treatments. Phytotaxa 397: 291. https://doi.org/10.11646/phytotaxa.397.4.3
- Martínez‐Azorín M, Crespo MB, Alonso‐Vargas MÁ, Pinter M, Crouch NR, Dold AP, Mucina L, Pfosser M, Wetschnig W (2023) Molecular phylogenetics of subfamily Urgineoideae (Hyacinthaceae): Towards a coherent generic circumscription informed by molecular, morphological and distributional data. Journal of Systematics and Evolution 61: 42-63. https://doi.org/10.1111/jse.12905
- Martínez-Azorín M, Crespo MB, Vargas MÁ, Pinter M, Crouch NR, Dold AP, Mucina L, Pfosser M, Wetschnig W (2023) A generic monograph of the Hyacinthaceae subfamily Urgineoideae. Phytotaxa 610: 1–143. https://doi.org/10.11646/phytotaxa.610.1.1
- Pinter M, Brudermann A, Crespo MB, Deutsch G, Martínez-Azorín M, Müller-Doblies U, Müller-Doblies D, Pfosser M, Wetschnig W (2013) Massonia citrina (Hyacinthaceae, Hyacinthoideae)—a new species from the Western Cape Province (South Africa). Phytotaxa 112: 50. https://doi.org/10.11646/phytotaxa.112.2.3
- Tercero-Araque A, Martínez-Azorín M, Salazar-Mendías C (2023) Cathissa villasina ( Hyacinthaceae ), a new endemic species from the southeastern Iberian Peninsula. Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology 157: 516–529. https://doi.org/10.1080/11263504.2023.2165568
- Yildirimli S (2011) Three new species from Turkey. Ot Sistematik Botanik Dergisi 18: 1–13.
- Yildirimli S, Selvi B (2002) A new species, Muscari sivrihisardaghlarensis (Liiaceae) from Central Anatolia, Turkey. Ot Sistematik Botanik Dergisi 9: 7–12.
- Yildirm H (2015) Muscari atillae (Asparagaceae): a new species from Eastern Anatolia, Turkey. Phytotaxa 213: 291. https://doi.org/10.11646/phytotaxa.213.3.9
- Yıldırım H, Akbaş K, Özdöl T (2023) Leopoldia buseana (Asparagaceae, Scilloideae), a new species from southwestern Anatolia. Phytotaxa 625: 51–65. https://doi.org/10.11646/phytotaxa.625.1.3