some Ledebouria and Scilla (and help needed with ID)

[Marlon Machado]

Digging on the internet, I found more references on the phylogenetic relationships of Ledebouria etc. and the Hyacinthaceae. Unfortunately I could only get the pdf versions of the last three articles:

Hankey A. 2003. Distinguishing between Ledebouria, Drimiopsis and Resnova. PlantLife (S. Afr.) no. 29. 38-39.

Stedje B. 1998. Phylogenetic relationships and generic delimitation of sub-Saharan Scilla (Hyacinthaceae) and allied African genera as inferred from morphological and DNA sequence data. Plant Systematics and Evolution 211(1-2): 1-11.

Abstract:
The genus Scilla has recently been split into two genera, Scilla s. str. and Ledebouria, based on morphological characters of South African species. This splitting, as well as the phylogenetic relationships between these genera and the related genera Drimia and Drimiopsis, is here discussed in the light of cladistic analyses of morphological and DNA sequence data of species from tropical Africa, the Mediterranean and India. Two chloroplast DNA regions are analysed: the intergenic spacer between the tRNA genes L(UAA) and F(GAA) and the L(UAA) intron.

Tindamanyire N. 2007. Studies of generic and specific delimitation of the genera Drimiopsis and Ledebouria (Hyacinthaceae). M.Sc. thesis. Oslo University, Norway.

Abstract:
The monocot genera Ledebouria and Drimiopsis belong to the family Hyacinthaceae in the order Asparagales. The aim of this study was to investigate the generic delimitation between the two genera using DNA sequencing, and to investigate the specific delimitation within Ledebouria using both morphological and molecular characters.
For the molecular analysis the Drimiopsis and Ledebouria specimens were sequenced for three chloroplast regions; trnL intron, trnL-F intergenic spacer and trnS-G intergenic spacer. In the strict consensus tree of the combined analysis of the three regions, the two genera each form a monophyletic group (though without any bootstrap or jackknife support), supporting the recognition of two separate genera.

For the morphological investigation 22 Ledebouria specimens were investigated using univariate and multivariate analyses of altogether 19 qualitative and quantitative characters. The analyses of the morphological data provided not enough information for the proposal of a final specific delimitation within Ledebouria, but together with the molecular phylogeny they suggested that six tentative taxa may be identified within the genus Ledebouria; L. kirkii, two groups of broad leaved taxa (L. cordifolia and one unnamed taxon), and three groups within the L. revoluta complex. The characters leaf width, leaf length, perianth segment length, leaf threads, spotting pattern of the leaves and ovary shape seem to be the most important morphological characters for delimiting these taxa. However, more investigations (morphological and molecular) including more specimens from all geographical regions is needed to confirm this result.

Lebatha P., Buys M.H., Stedje B. 2006. Ledebouria, Resnova and Drimiopsis: a tale of three genera. Taxon 55(3):643-652.

Abstract:
The monophyletic status of Ledebouria, Resnova and Drimiopsis (Hyacinthaceae) was investigated using cladistic analyses of morphological characters as well as by combined trnL-F and morphology dataset. Based on morphology alone, numerous synapomorphies support Ledebouria, Resnova and Drimiopsis as three separate clades. In contrast, trnL-F data in a reduced taxon sample suggest that Resnova and Drimiopsis are imbedded within Ledebouria, while in combination with morphology again suggest them to be separate. The analysis of morphological data shows that the majority of morphological characters listed by Manning & al. (2004) to support Ledebouria s.l. are homoplasious or symplesiomorphies. A revised taxonomy of Ledebouria, Resnova and Drimiopsis is premature.

Müller-Doblies U., Müller-Doblies D. 2008. A partial revision of the tribe Massonieae (Hyacinthaceae) 1. Survey, including three novelties from Namibia: A new genus, a second species in the monotypic Whiteheadia, and a new combination in Massonia. Feddes Repertorium 108(1-2):49-96.

Abstract:
The tribe Massonieae with about 150 species in 15 genera is subendemic to southern Africa (Namibia, South Africa, Lesotho, and Swaziland) radiating with two genera into tropical East Africa and one of them to S Asia. Two of these 15 genera, Periboea and Resnova, are formally resuscitated in the present paper. Four genera are known from Namibia, to which Namophila urotepala gen. et sp. novae is added.
The hitherto green-flowered genus Whiteheadia is no longer monotypic: the white-flowered W. etesionamibensis with distinctly longer perigon-segments and with many other minor differences is described from several localities in Diamond Sperrgebiet no. 1 and surrounding areas eastwards to the Hunsberge.

The only Massonia known from Namibia (MERXMÜLLER & ROESSLER 1973) is not M. echinata but a species of its own which deserves a new combination: M. sessiliflora; it also occurs in the Cape, namely on the southern bank of the Orange River.

Massonieae as a whole are subdivided into three subtribes. A key to the subtribes, keys to the genera within the subtribes, and in several genera with important changes also keys to the species are provided. Fourteen species of southern Africa are resucitated from synonymy: in Resnova (4, incl. one comb. nova), Drimiopsis (1), Massonia (6, incl. the above species from Namibia), Periboea (1, a new combination), and Polyxena (2); as two accepted species proved to be younger synonyms (Drimiopsis maxima and Neobakeria heterandra) the increase in species by reinstatements is reduced to 12. Seven new species are proposed in addition to the two Namibian ones: in Periboea (P. oliveri), in Polyxena (P. calcicola). and in Drimiopsis (D. comptonii, D. davidsoniae, D. pusilla, D. reilleyana, and D. stolonissima). For tropical East Africa three new combinations (altogether six) are proposed, Ledebouria hildebrandtii. L confertiflora, and Drimiopsis fischeri which are transferred from other genera.

Pfosser M., Wetschnig W., Ungar S., Prenner G. 2003. Phylogenetic relationships among genera of Massonieae (Hyacinthaceae) inferred from plastid DNA and seed morphology. Journal of Plant Research 116(2): 115-132.

Abstract:
The tribe Massonieae Baker (Hyacinthaceae-Hyacinthoideae) presently consists of about 19 genera and 230 species distributed from Africa (south of the Sahara) to Madagascar and India. Based on atpB and trnL-F DNA sequences the tribe is monophyletic only when the genus Pseudoprospero is excluded from Massonieae. In most trnL-F trees, this genus occupies a basal position within subfamily Hyacinthoideae and is sister to the rest of the subfamily. Molecular data suggest that the remaining genera of Massonieae do not share common ancestry with the Eurasian/North-African tribe Hyacintheae Dumort. (Scilla, Hyacinthus and allies), and thus a narrow concept of the essentially Eurasian genus Scilla is supported. Members of well-supported clades in Massonieae usually show similarities in seed characteristics as determined by scanning electron microscopy. Phylogenetic position and seed morphology indicate that Massonia angustifolia and M. zeyheri do not belong to the genus Massonia but fall into a clade together with Daubenya, Androsiphon and Amphisiphon. The genus Whiteheadia appears paraphyletic in the 50% majority rule trnL-F tree and occupies a basal position next to Massonia. However, in the strict consensus tree neither monophyly nor polyphyly can be excluded for this genus. Seed appendages are documented for members of the genera Ledebouria and Lachenalia. Within the genera of Massonieae there is a tendency towards bending of the seed axis. This phenomenon is most obvious within the genus Lachenalia. Delimitation of genera based on seed morphology largely agrees with the results of molecular studies. Correlation between number, size and color of seeds, geographical distribution and phylogenetic position of the genera are discussed.

Wetschnig W., Pfosser M. 2003. The Scilla plumbea Puzzle-Present Status of the Genus Scilla Sensu Lato in Southern Africa and Description of Spetaea lachenaliiflora, a New Genus and Species of Massonieae (Hyacinthaceae). Taxon 52(1):75-91.

Abstract:
We have studied a taxon from South Africa that has been hidden for more than 150 years within Scilla plumbea (Hyacinthaceae). However, phylogenetic reconstruction based on trnL-F data, morphological evidence (including seed characters), and karyology suggest independent status. DNA data place it close to the Daubenya alliance but not close to any particular genus of Massonieae previously included within Scilla s.l. Unique characters of seeds indicate that this taxon does not belong to the Daubenya clade. Analysis of the iconotype of Scilla plumbea Lindl., on the other hand, reveals morphological details suggesting membership in the Scilla natalensis clade (= genus Merwilla in the recent classification of Speta). Merwilla, however, occupies a basal position among Massonieae and is not related to the Daubenya clade. Based on all data, we describe this taxon as a new genus and species Spetaea lachenaliiflora Wetschnig & Pfosser.

Pfosser M., Speta F. 1999. Phylogenetics of Hyacinthaceae Based on Plastid DNA Sequences. Annals of the Missouri Botanical Garden 86(4):852-875.

Abstract:
Hyacinthaceae presently consist of approximately 70 genera and 1000 species. To investigate the monophyly of the family and the generic relationships, we sequenced the trnL intron and the trnL-trnF intergenic spacer region of chloroplast DNA for 105 taxa in Hyacinthaceae and 18 species of related families. By testing different outgroup compositions, we provide evidence for the monophyly of the family if the North American genera Camassia and Chlorogalum, which are more closely related to Agave and Hosta, are excluded from Hyacinthaceae sensu Dahlgren. Several generic implications can be deduced from the analysis, the most prominent one of which is the polyphyletic origin of the Linnaean genera Scilla, Ornithogalum, and Hyacinthus. Especially members of the genera Scilla and Hyacinthus are extensively intermixed with each other. According to the DNA sequence data, the only true Scilla species are found in the Mediterranean region and appear as a monophyletic clade. A tetrapartition of the family into (1) the monotypic subfamily Oziroeoideae Speta, accommodating the South American Hyacinthaceae; (2) the subfamily Urgineoideae Speta, housing relatives of the squills; (3) the subfamily Ornithogaloideae Speta, including the tribes Ornithogaleae Rouy and Dipeadieae Rouy; and (4) the largest and most advanced subfamily, Hyacinthoideae Link, consisting of the tribe Massonieae Baker (including species from Africa south of the Sahara and from India) and the Mediterranean/Asian tribe Hyacintheae Dumort., is proposed. Previously included in Hyacinthaceae sensu Dahlgren, the North American genera Chlorogalum and Camassia show affinities to Agavaceae and Funkiaceae and appear as a distinct clade together with Anthericaceae. Furthermore, the occurrence of taxa from southern Africa at basal positions in all subfamilies points to the origin of evolution of the Hyacinthaceae in this region.

Stedje B. 2001. Generic Delimitation of Hyacinthaceae, with Special Emphasis on Sub-Saharan Genera. Systematics and Geography of Plants 71(2):449-454.

Abstract:
Different opinions on the generic delimitation of Hyacinthaceae are reviewed with emphasis on sub-Saharan taxa. Morphological characters used for identification of genera as well as results from DNA-sequencing data are discussed in relation to different generic delimitations proposed by various authors. Recommendations on the generic delimitation of the family are put forward. The use of a wide generic concept is suggested with emphasis on the use of qualitative characters as diagnostic characters for genera.