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Studies on tribal systematics of Amaryllidaceae 1. The systematic position of Lapiedra Lag
... The circumscription of Galantheae has varied considerably, even in recent times, with various constituent genera, for example: Galanthus L., Hannonia Braun-Blanq. & Maire, Lapiedra Lag., Leucojum L. ( Traub, 1970;Müller-Doblies and Müller-Doblies, 1978;Meerow and Snijman, 1998). In a molecular phylogenetic study using plastid matK and nrDNA ITS sequences by Lledó et al. (2004), Leucojum was revealed as paraphyletic and the genus Acis Salisb. ...
... Although not a member of tribe Galantheae, the L. martinezii extract expressed one of the highest activities (IC 50 ¼ 9.3 mg/ml) and contained several alkaloids, which could not be identified from the small quantities extracted. This monotypic genus is endemic but common to the maritime cliffs vegetation in South East Spain and Northern Morocco (Müller Doblies and Müller-Doblies, 1978). Ismine and two other phenantridine alkaloids have been isolated from L. martinezii in a previous study ( Suau et al., 1990). ...
... Collectively these four inflorescence types are referred to as monochasia (Eichler, 1879). While, it may have been possible that the lateral cymes in the Tacca chantrieri inflorescence were ancestrally derived from a bostryces which transitioned to a cincinni through a rhapidum or drapanium, as have been described in the Amaryllidaceae genera Clivia and Lapideria (Müller-Doblies and Müller-Doblies, 1978), it is more likely that the ancestor condition was a thyrsoid with uncondensed internodes; a common condition in this order (Remizowa et al., 2010;Nuraliev et al., 2021;Yudina et al., 2022). As such, the this represents a convergent case of an umbellate inflorescence since the lateral cymes are arrange differently than the bostryx derived umbellate taxa. ...
Inflorescence structure is very diverse and homoplasious, yet the developmental basis of their homoplasy is poorly understood. To gain an understanding of the degree of homology that these diverse structures share, we characterize the developmental morphology and anatomy of various umbellate inflorescences across the monocots and analyzed them in an evolutionary context. To characterize branching order, we characterized the developmental morphology of multiple inflorescences with epi-illumination, and vascular anatomy with Laser Ablation Tomography, a novel high-throughput method to reconstruct three-dimensional vasculature. We used these approaches to analyze the umbellate inflorescences in five instances of presumed homoplasy: in three members of the Amaryllidaceae; in three members of the Asparagaceae, including a putatively derived raceme in Dichelostemma congestum; in Butomus umbellatus (Alismataceae), in Tacca chantrieri (Dioscoreaceae), and in umbellate structure in Fritillaria imperialis (Liliaceae). We compare these with racemes found in three members of the subfamily Scilliioideae (Asparagaceae). We find there are three convergent developmental programs that generate umbellate inflorescences in the monocots, bostryx-derived, cincinnus-derived and raceme-derived. Additionally, among the bostryx-derived umbellate inflorescence, there are three instances of parallel evolution found in the Amaryllidaceae, in two members of Brodiaeoideae (Asparagaceae), and Butomus umbellatus, all of which share the same generative developmental program. We discuss the morphological modifications necessary to generate such complex and condensed structures and use these insights to describe a new variant of metatopy, termed horizontal concaulesence. We contextualize our findings within the broader literature of monocot inflorescence development, with a focus on synthesizing descriptive developmental morphological studies.
... Leucojum is morphologically diverse. Generally According to Muïler-Doblies and Doblies (1978), the two genera should be placed in their own subtribe, Galanthinae, which is closely related to Narcissinae (Narcissus L., Sternbergia Waldst. & Kit.). ...
Phylogenetic analyses of the monocotyledonous genera Leucojum and Galanthus (Amaryllidaceae, Asparagales), using plastid (trnL-F and matK) and largely non-coding nuclear ribosomal (ITS) DNA sequences show the two to be closely related to Lapiedra, Narcissus, Vagaria, Pancratium and Sternbergia. We compare the results obtained with a combined parsimony analysis of these nucleotide sequences with that of a matrix of morphological characters. The sampling included all species of Leucojum and most species of Galanthus (representing all series and subseries of the genus) and used as outgroup the above mentioned genera of Amaryllidaceae shown to be close relatives. The plastid, nuclear and morphological data were analysed independently and in combination, showing that the boundaries between the two genera are not appropriate. Galanthus is monophyletic but embedded in Leucojum. On the basis of chromosome numbers and floral characters Leucojum has been previously divided into four subgenera, which have been accepted as genera by some authors. In our phylogenetic analyses (separate as well as combined), Leucojum species are separated in two primary clades corresponding to L. subgenera Ruminia + Acis and L. Leucojum + Aerosperma. The taxonomic implications of this pattern are discussed, and an alternative classification is proposed. Finally, biogeographic relationships of species of both Leucojum and Galanthus are discussed, emphasising the possible origin of the narrowly distributed taxa of Leucojum relative to the widespread species.
In Egypt, taxonomic aspects of the genus Pancratium L. (Amaryllidaceae, Pancratieae) were thoroughly studied to point out the most reliable characters for taxon delimitation. Here, we base our species on both geographical and morphological features. The detailed taxonomic revision of genus Pancratium is based on field studies and the examination of representative specimens together with the authentic types deposited in the major herbaria of Egypt and Kew. Variations in the aerial scape, spathe, perianth, corona-teeth, pistil, stamens, capsules and seeds, as well as those of leaf base and leaf orientation are good markers to separate the different species. As a result, five species are recognized, including P. trianthum as a new record to the flora of Egypt. It is restricted to the south-eastern corner of Egypt (Gebel Elba region) and being unique in possessing solitary spathe and long attenuate leaf base. A specific key and descriptions are given with typifications.
In absence of original material, a neotype is designated for the name Lapiedra martinezii Lag. (Amaryllidaceae). The neotype is a specimen from SE Spain kept in the herbarium of the Royal Botanic Garden of Madrid (MA).
Bulbous (rarely rhizomatous), mostly geophytic, perennials, terrestrial, occasionally aquatic or epiphytic, rich in family-specific alkaloids. Bulbs tunicate. Leaves annual or persistent, sessile and linear or lorate, or petiolate and lanceolate to widely elliptic, distichous or spirally arranged; sometimes basally sheathing and forming an aerial pseudostem, usually glabrous, rarely with trichomes. Inflorescence scapose, pseudoumbellate (reduced helicoid cymes); scape sometimes wholly subterranean and appearing obsolete, terminated by 2 or more spathaceous, obvolute or equitant, usually marcescent bracts that enclose the flowers in bud (bracts rarely absent); inner bracteoles usually present and successively shorter and narrower. Flowers 1–many, perfect, frequently large and showy, sessile or pedicellate, each usually subtended by a bracteole, actinomorphic or zygomorphic, generally protandrous.
In Egypt, taxonomic aspects of the genus Pancratium L. (Amaryllidaceae, Pancratieae) were thoroughly studied to point out the most reliable characters for taxon delimitation. Here, we base our species on both geographical and morphological features. The detailed taxonomic revision of genus Pancratium is based on field studies and the examination of representative specimens together with the authentic types deposited in the major herbaria of Egypt and Kew. Variations in the aerial scape, spathe, perianth, corona-teeth, pistil, stamens, capsules and seeds, as well as those of leaf base and leaf orientation are good markers to separate the different species. As a result, five species are recognized, including P. trianthum as a new record to the flora of Egypt. It is restricted to the south-eastern corner of Egypt (Gebel Elba region) and being unique in possessing solitary spathe and long attenuate leaf base. A specific key and descriptions are given with typifications.
Ungiminorine N-oxide was isolated from Pancratium maritimum, and homolycorine N-oxide and O-methyl lycorenine N-oxide from Lapiedra martinezii. These compounds represent the first examples of naturally occurring N-oxides from the Amaryllidaceae.
The only Namibian species of Spiloxene is described and illustrated by drawings and photos as Spiloxene etesionamibensis U.MÜLL.-DOBLIES, MARK.ACKERMANN, WEIGEND &. D.MÜLL.-DOBLIES. It was hitherto misinterpreted as Spiloxene scullyi (BAKER) GARSIDE in SÖLCH & ROESSLER (1969) in MERXMÜLLER’s Prodromus einer Flora von Südwestafrika, but its closest ally is S. flaccida (NEL) GARSIDE, found in the Western and the Eastern Cape. Spiloxene etesionamibensis, which was found independently by GIESS & MERXMÜLLER and by MÜLLERDOBLIES, is only known from two or three independent collections from a single locality. S. scullyi is no longer known from Namibia.
In addition to the eight Spiloxene species already known from the Northern Cape Spiloxene namaquana U.MÜLL.-DOBLIES, MARK.ACKERMANN, WEIGEND & D.MÜLL.-DOBLIES is described and illustrated by drawings and photos. Its closest ally is Spiloxene trifurcillata (NEL) FOURC. from the Eastern Cape.
For Spiloxene namaquana the morphological analysis of the branching pattern of an annual shoot taken in 1988 in the field is evaluated by a longitudinal and a horizontal diagram with explanatory remarks. The unusual innovation system of extreme anisoiteration is discussed.
On Map 1 the quarter degree squares of the type localities (± only known localities) of both species is given.
Phylogenetic analyses of the monocotyledonous genera Leucojum and Galanthus (Amaryllidaceae, Asparagales), using plastid (trnL-F and matK) and largely non-coding nuclear ribosomal (ITS) DNA sequences show the two to be closely related to Lapiedra, Narcissus, Vagaria, Pancratium and Sternbergia. We compare the results obtained with a combined parsimony analysis of these nucleotide sequences with that of a matrix of morphological characters. The sampling included all species of Leucojum and most species of Galanthus (representing all series and subseries of the genus) and used as outgroup the above mentioned genera of Amaryllidaceae shown to be close relatives. The plastid, nuclear and morphological data were analysed independently and in combination, showing that the boundaries between the two genera are not appropriate. Galanthus is monophyletic but embedded in Leucojum. On the basis of chromosome numbers and floral characters Leucojum has been previously divided into four subgenera, which have been accepted as genera by some authors. In our phylogenetic analyses (separate as well as combined), Leucojum species are separated in two primary clades corresponding to L. subgenera Ruminia + Acis and L. Leucojum + Aerosperma. The taxonomic implications of this pattern are discussed, and an alternative classification is proposed. Finally, biogeographic relationships of species of both Leucojum and Galanthus are discussed, emphasising the possible origin of the narrowly distributed taxa of Leucojum relative to the widespread species.
The monophyletic,Eurasian clade of Amaryllidaceae,was,analyzed,using plastid ndhF and,rDNA ITS se- quences for 33 and 29 taxa, respectively; all genera were represented by at least one species. Both maximum parsimony and Bayesian analysis were,used on each data set and the combined,data. Both sequence matrices,resolve the Central and East Asian tribe Lycorideae as sister to the Mediterranean-centered genera of the clade, and recognize two large subclades within the greater Mediterranean region: Galantheae, consisting of Acis, Galanthus and Leucojum; and Narcisseae (Narcissus and Sternbergia)/Pancratium. However, there are areas of incongruence between the ndhF and ITS trees. When three predominantly monotypic genera, Hannonia, Lapiedra ,a ndVagaria, centered in North Africa, are removed from the alignments, the two sequence matrices produce fully congruent topologies with increased support at many of the nodes, with ILD between partitions rising from,P 5 0.07 to 0.96. We hypothesize,that lineage sorting took place after the divergence,of Galantheae and Narcisseae/Pancratium from a common genepool with Hannonia, Lapiedra ,a ndVagaria retaining a mosaic of the ancestral haplotypes. We also performed,dispersal-vicariance analysis to reconstruct biogeographic,scenarios on several of the generic level phylogenies found with and without these three genera included, as well as on a species-level phylogeny of Galantheae. After the vicariant divergence of the Asian Lycorideae, North Africa and the Iberian Peninsula are the most likely areas of origin for the rest of the clade. The results of the dispersal-vicariance analysis are discussed,in the context of the complex biogeographic,history of the Mediterranean,basin. The Eurasian clade of the Amaryllidaceae contains
Cladistic analyses of plastid DNA sequences rbcL and trnL-F are presented separately and combined for 48 genera of Amaryllidaceae and 29 genera of related asparagalean families. The combined analysis is the most highly resolved of the three and provides good support for the monophyly of Amaryllidaceae and indicates Agapanthaceae as its sister family. Alliaceae are in turn sister to the Amaryllidaceae/Agapanthaceae clade. The origins of the family appear to be western Gondwanaland (Africa), and infrafamilial relationships are resolved along biogeographic lines. Tribe Amaryllideae, primarily South African, is sister to the rest of Amaryllidaceae; this tribe is supported by numerous morphological synapomorphies as well. The remaining two African tribes of the family, Haemantheae and Cyrtantheae, are well supported, but their position relative to the Australasian Calostemmateae and a large clade comprising the Eurasian and American genera, is not yet clear. The Eurasian and American elements of the family are each monophyletic sister clades. Internal resolution of the Eurasian clade only partially supports currently accepted tribal concepts, and few conclusions can be drawn on the relationships of the genera based on these data. A monophyletic Lycorideae (Central and East Asian) is weakly supported. Galanthus and Leucojum (Galantheae pro parte) are supported as sister genera by the bootstrap. The American clade shows a higher degree of internal resolution. Hippeastreae (minus Griffinia and Worsleya) are well supported, and Zephyranthinae are resolved as a distinct subtribe. An Andean clade marked by a chromosome number of 2n = 46 (and derivatives thereof) is resolved with weak support. The plastid DNA phylogenies are discussed in the context of biogeography and character evolution in the family.
10. Zusammenfassung und Schlußfolgerung
Bei Galanthus liegt eine doppelte Täuschung vor: Scheinbar handelt es sich um die Endblüte eines seitenständigen Schaftes, nach dem Gesagten jedoch anscheinend um die unterste Seitenblüte eines endständigen Schaftes. Wenn man die Pseudo‐Strukturen ganz auskosten will, kann man sagen, daß eine pseudoterrninale Blüte auf einem pseudo‐lateralen Schaft sitzt, was zum pseudo‐monopodialen Zwiebelbau führt (vgl. Abb. 2 und 10).
Es erhebt sich abschließend die Frage, ob es überhaupt echt monopodiale Zwiebeln gibt. Wie einleitend gesagt, stamrnen die Beispiele dafür immer aus den Amaryllidoideen, und flir diese sind alle drei denkbaren Meinungen vorgebracht worden:
. die Amaryllidoideen sind monopodiaI gebaut,
. dic Amaryllidoideen sind sympodial gebaut,
. die Amaryllidoideen sind teils monopodial und teils sympodial gebaut,
wobei über die Abgrenzung der beiden Gruppen keine Einigkeit besteht und wobei auch gar kein Versuch gemacht worden ist, die cine Gruppe von der andelen abzuleiten. Ich hoffe, dafs das wenige, was ich iiber die Amaryllidoideen irn allgemeinen gesagt habe, ausreicht, um anzudeuten, daß hier ein recht einheitlicher Bauplan vorliegt. Mit Galanthus fällt nun das beste Beispiel flir den monopodialen Zwiebelbau, und da keine andere Gattung iiber ein anderes oder gar besseres Argument verfügt, sind wohl die monopodialen Zwiebeln aus den Lehrbiichern zu streichen, und der morphologische Abstand zwischen Allioideae und Amaryllidoideae ist ein Snick geringer geworden.
The anatomy of seeds from three species of Pancratium occupying different habitats was examined. Pancratium maritimum grows on the spray belt of the Mediterranean shore, P. sickenbergii in sandy soil in the steppe regions, and P. parviflorum in the Mediterranean batha. Seed development was found to be similar in all three species. All the walls of the seed coat cells derived from the outer integument are encrusted with a highly resistant brown-black material which apparently consists of phenolic quinones. Two or three compressed outer layers of cells of the testa of P. maritimum and P. sickenbergii form a dark protective layer. The rest of the testa in these two species is composed of large, dead, air-filled cells with large intercellular spaces between them In contrast, the seed coat of P. parviflorum consists of smaller cells with small intercellular spaces; the seed possesses an elaiosome containing two types of lipid. The differences in characteristics of the testa among the three species are of adapt...
The tribe Galantheae
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The prevíous discussion about the systematic position of La piedra referred to ahl eight Mediterranean genera of Amaryllidoideae. Although not yet discussed in detail, the following classification is here proposed
- Of The
- Genera
- Amarylltdoideae
OF THE MEDITERRANEAN GENERA
OF AMARYLLTDOIDEAE
The prevíous discussion about the systematic position of La piedra referred to ahl eight Mediterranean genera of Amaryllidoideae. Although not
yet discussed in detail, the following classification is here proposed (Table 2):