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Accommodating Nomocharis in Lilium (Liliaceae)

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Yundong Gao at Chinese Academy of Sciences
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Phytotaxa 277 (2): 205–210
http://www.mapress.com/j/pt/
Copyright © 2016 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Li-Bing Zhang: 12 Sept. 2016; published: 27 Sept. 2016
http://dx.doi.org/10.11646/phytotaxa.277.2.8
205
Accommodating Nomocharis in Lilium (Liliaceae)
YUN-DONG GAO & XIN-FEN GAO*
CAS Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization & Ecological Restoration and Biodiversity Con-
servation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences; email: xfgao@cib.ac.cn
*Author for corresponding
Introduction
Controversy regarding the status of the genus Nomocharis Franchet (1889: 113) has been undergoing since its recognition
by Franchet (1889). Recent molecular studies (Nishikawa et al. 1999, Hayashi & Kawano 2000, Nishikawa et al. 2001,
Ronsted et al. 2005, Peruzzi et al. 2009) have resolved Nomocharis as being nested within Lilium Linaeus (1753: 302).
Results of our own previous studies (Gao et al. 2012, Gao et al. 2013a, Gao et al. 2013b), with expanded sampling of
species of Nomocharis have been congruent with those of previous studies by others. Thus recognition of Nomocharis
would render Lilium paraphyletic. We prefer to recognize a monophyletic Lilium here, although paraphyletic groups
are sometimes advocated in literature (e.g., Brummitt, 2014; Ehrendorfer & Barfuss, 2014; George, 2014; Hörandl,
2014; Stuessy & Hörandl, 2014; Stuessy et al., 2014). Most recently, we proposed that the morphological divergence
between Nomocharis and Lilium was the result of habitat specialization (Gao et al. 2015). The extensive introgression
caused by hybridization within Lilium and Nomocharis (Gao et al. 2013a, 2015) supports a single-genus concept.
Taxonomic treatment and nomenclatural account
We here transfer the eight species of Nomocharis to Lilium. All eight relevant species are distributed in alpine zone of
southwestern China, northeastern Upper Burma and northeastern India (i.e., the Hengduan Mountains and neighboring
regions) and can be distinguished from each other with the following key (cf. (Sealy 1983, Liang 1984).
Key to species
1a. Leaves predominantly in whorls of 3-9 leaves; nectaries at base of inner tepals a series of low flanges of tissue standing erect and
arranged flabellately on either side of a short median channel.
1b. Leaves alternate; filaments nearly subulate, tapering from flat, widened base to filiform apex; inner tepals entire at margin.
2a. Perigone shallowly cupular to cupular, neither spotted nor blotched .........................................................................1. L. basilissum
2b. Perigone white or pale pink, blotched with purple.
3a. Leaves elliptic to lanceolate rarely ovate; perigone blotched all over, or spotted for short distance at the base .................................
............................................................................................................................................................................. 2. L. pardanthinum
3b. Leaves narrowly elliptic-lanceolate or linear to lanceolate; inner tepals with margins entire to minutely erose.
4a. Perigone spreading out flat, white or pale pink, blotched all over with purple ..................................................... 3. L. meleagrinum
4b. Perigone shallowly cupular at first, opening out widely, white to pale pink or rose with spots or small blotches of crimson or purple
for a short distance at the base .......................................................................................................................................... 4. L. farreri
5a. Perigone pale yellow and not swellings or flanges on either side at the base of inner tepals ..............................5. L. gongshanense
5b. Perigone not pale yellow and nectaries of the inner tepals either flanges of tissue arranged flabellately or swellings or ridges on
either side of the short median channel.
6a. Nectaries of inner tepals a series of flanges of tissue arranged flabellately on either side of a short median channel ........................
.................................................................................................................................................................................. 6. L. synapticum
6b. Nectaries of inner tepals swellings on either side of the basal median channel.
7a. Perigone opening out flat, blotched in the lower half or all over; style longer than (rarely equal to) the ovary .......... 7. L. apertum
7b. Perigone cupular or shallowly cupular, finely spotted at the base and lightly so over the lower half; style shorter than the ovary ....
.................................................................................................................................................................................. 8. L. saluenense
GAO & GAO
206 • Phytotaxa 277 (2) © 2016 Magnolia Press
1. Lilium basilissum (W.E.Evans) Y.D.Gao, comb. nov. (Fig. 1a–c) ≡Nomocharis basilissa W. E. Evans (1925: 25).
Type:—MYANMAR. Upper Burma, Chawchi Pass, in dwarf cane brakes, alt. 3000–3100 m, 21 Jul 1920, Farrer 1738
(holotype E-00394034!).
This species occurs in China (northwestern Yunnan) and north Myanmar.
FIGURE 1. Field pictures of western China Lilium (formerlly Nomocharis): a–c, L. basilissum; d–f, L. farreri; g–i, L. gongshanense; j–l,
L. meleagrinum.
2. Lilium pardanthinum (Franchet) Y.D.Gao, comb. nov. (Fig. 2a–c) ≡Nomocharis pardanthina Franchet (1889 :
133). Type:—CHINA. Yunnan, Dali (Tali), Cangshan Mountians (Tsang-chan), in glades of Bamboo, alt. 3000–4000
m, 2 Jun 1883, Delavay 257 (holotype P-00730859!; isotypes P-00730860!, A-00030008!).
TREATING NOMOCHARIS IN LILIUM Phytotaxa 277 (2) © 2016 Magnolia Press • 207
=Nomocharis mairei Léveillé (1913: 287). Type:—CHINA. Yunnan, Plateau de Ta-Hai, 3200 m, Maire, E.E. s.n. (holotype E-
00381777!).
=Nomocharis leucantha Balf.f. (1918: 276). N. mairei Balf. f. f. leucantha (Balf. f.) Evans (1925: 29). Type:—CHINA. Yunnan, on the
eastern flank of the Tali range, alt. 3350–3650 m, Forrest G. 3845 (lectotype E-00381779!, isolectotype K–000900808!; designated
here!).
=Nomocharis mairei Balf.f. f. candida Evans (1925: 29). Type:—not designated.
=Nomocharis pardanthina Franchet f. punctulata Sealy (1978: 295). Type:—CHINA. Yunnan, Lijiang (Lichiang), Forrest G. 5816
(holotype K-000900813!).
This is a very complicated species exhibiting many forms that have, at various times, been considered as different taxa
(Sealy 1950, 1983, Liang 1984). It is endemic to China (SW Sichuan, NW Yunnan).
FIGURE 2. Field pictures of western China Lilium: a–c, L. pardanthinum; d–f, L. saluenense.
3. Lilium meleagrinum (Franchet) Y.D.Gao, comb. nov. (Fig. 1j–i) ≡Nomocharis meleagrina Franchet (1898: 196).
Type:—CHINA. Yunnan, Sila, 1–15 Jun 1895. Soulié 1032 (holotype P–00730856!).
=Nomocharis biluoensis S.Y.Liang (1984: 169). Type:—CHINA. Yunnan, Weixi, 13 Jul 1981. Hengduan Mt. Expedition of Institute of
Botany 1485 (holotype PE–00036078!).
This species occurs in SW Sichuan, SE Xizang, NW Yunnan of China.
4. Lilium farreri (Harrow ex W.E.Evans) Y.D.Gao, comb. nov. (Fig. 1d–f) ≡Nomocharis farreri (W. E. Evans) Harrow
(1930: 76). N. pardanthina var. farreri (Cox) Evans (1925: 20). Type:—MYANMAR. Upper Burma, Hpimaw (Pian-
Ma) Pass, in light glades of Bamboo, alt. 3000 m, 13 Jun 1919, Farrer 1031 (holotype E-00394032!).
= Nomocharis pardanthina sec. Farrer in Gardeners’ Chronicle series 3 (1919: 29). Type:—not designated.
This species occurs in China (northwestern Yunnan) and north Myanmar.
GAO & GAO
208 • Phytotaxa 277 (2) © 2016 Magnolia Press
FIGURE 3. Field pictures of Lilium apertum in western Yunnan: a–c, population from Zhongdian, Yunnan showed spot variation; c–e,
population of Fugong, Yunnan showed variations in tepal color; f–h, habits of L. apertum under different habitats; i–j, anatomical pictures
showed two types of L. apertum from Zhongdian and Fugong, as well as a comparison of outer and inner tepals.
TREATING NOMOCHARIS IN LILIUM Phytotaxa 277 (2) © 2016 Magnolia Press • 209
5. Lilium gongshanense (Y.D.Gao & X.J.He) Y.D.Gao, comb. nov. (Fig. 1g–i) ≡Nomocharis gongshanensis Y.D.Gao
& X.J.He (2012: 69). Type:—CHINA. Yunnan, Gongshan, alt. 3200m, Gaoligongshan Range, sunny grassy and bushy
slopes on limestone soils, 7 Jul 2009, Gao Y.D. G09003 (holotype SZ!).
It is endemic to northwestern Yunnan (Gongshan), China.
6. Lilium synapticum (Sealy) Y.D.Gao, comb. nov. ≡ Nomocharis synaptica Sealy (1950: 296). Type:—INDIA.
Assam, Delei Valley, alt. 3000 m, 7 Jul 1928, Kingdon Ward 8399 (holotype K-000900820!).
It is endemic to northeastern Assam, India.
7. Lilium apertum Franchet (1898: 220). (Fig. 3) ≡ Nomocharis aperta (Franchet) E.Wilson (1925: 13). Lilium
oxypetalum (Royle) Baker sec. Franchet (1892: 320). Type:—CHINA. Yunnan, Dali, Cangshan Mountians (Tsang-
chan) alt. 3000m, 18 Jun 1889, Delavay 4178 (holotype P-00730870!).
=Nomocharis forrestii Balf.f. (1918: 293). Nomocharis aperta var. forrestii (Balf. f.) W. Smith et W. E. Evans (1925: 96). Type:—CHINA.
Yunnan, Mountains in the N.E. of the Yangtze bend, alt. 3960 m, 15 Jul 1913, Forrest G. 10620 (holotype E-00381776!; isotypes
BM-000629649!, K-000900817!).
This species occurs in SW China (SW Sichuan, SE Xizang and NW Yunnan) and north Myanmar.
8. Lilium saluenense (Balf. f.) S.Y.Liang (1980: 154) (Fig. 2d–f). ≡Nomocharis saluenensis Balf. f. (1918: 293). L.
apertum var. thibeticum Franchet (1898: 221). Type:—CHINA, Yunnan, Sila, 25 Jul 1895, Soulié 1031 (holotype P-
00730868!, isotypes K-000900818!, P-00730869!).
=Nomocharis tricolor Balf.f. (1918: 296). Type:—CHINA, SE Xizang (Tibet), alpine meadow, alt. 4300 m, 19 Jul 1913, Kingdon Ward
801 (holotype BM).
This species occurs in SW China (SW Sichuan, SE Xizang and NW Yunnan) and north Myanmar.
Acknowledgments
The author appreciate the assistance of the staff in the herbaria of BM, CDBI, E, HUH, K, KUN, P, PE, and SZ in the
study of specimens. This research was supported by the National Natural Science Foundation of China (Grant No.
31500163) and Plant resources sharing platform project of Sichuan Province.
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Paraphyly and Polyphyly in the Worldwide Tribe Rubieae (Rubiaceae): Challenges for Generic Delimitation
Article
  • Nov 2014
On the basis of multidisciplinary studies on the tribe Rubieae, we contribute to the current discussion on paraphyly and supraspecific taxa that do not contain all descendant species of an ancestral clade. Rubieae belong to the large, predominantly tropical and woody family Rubiaceae and include possibly ≤ 1000 mostly temperate and herbaceous species with worldwide distribution. Our studies span distinctive groups throughout the tribe, consist of a maximum parsimony analysis of plastid atpB-rbcL and rpL32-trnL DNA sequences, and are summarized in a condensed strict consensus tree. A corresponding two-dimensional scheme illustrates alternative hypotheses for phylogenetic relationships among all major Rubieae clades identified. The small relictual genus Kelloggia Torr. in Benth. & Hook. f., formerly excluded from the Rubieae, is supported as a remnant of the ancestors of the tribe. Didymaea Hook. f. and Rubia L. represent early phylogenetic side lines. All other Rubieae form a large monophyletic crown group with the traditional genera Asperula L. being polyphyletic and Galium L. paraphyletic. Changes in the circumscription of these and other genera are thus inevitable. The necessity of accepting paraphyletic taxa as well as the positive and negative aspects of taxonomic splitting versus lumping within Rubieae are discussed. Additionally, lectotypes are designated for one section of Asperula---Asperula sect. Dioicae Airy Shaw & Turrill, typified by A. conferta Hook. f.—and for three sections of Galium---Galium sect. Leiogalium (DC.) Ledeb., typified by G. sylvaticum L.; Galium sect. Lophogalium K. Schum., typified by G. multiflorum Kellogg; and Galium sect. Depauperata Pobed., typified by G. songaricum Schrenk ex Fisch. & C. A. Mey.
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Taxonomy Versus Cladonomy in the Dicot Families
Article
  • Nov 2014
The theoretical basis for cladistic classification into monophyletic (holophyletic) ranked taxa is fatally flawed and is promoting bad taxonomy. Biological classification that takes account of evolutionary history may be based on two main factors—lines of descent and extent of divergence represented by morphological and other characters. In taxonomy a balance must be found between lines of descent and characters, and insistence on one at the expense of the other will give unacceptable results. Much confusion has arisen in systematics from the failure to appreciate that taxonomy, which groups organisms into ranked taxa (families, genera, etc.), is essentially different from grouping them into clades. These two processes are based on conflicting hierarchies and have different methodologies and functions. For several decades, however, the cladistics movement has adopted lines of descent rather than characters as the sole basis of taxonomy, insisting that only complete clades should be recognized as taxa. But as soon as one imposes ranks on a phylogeny, one must create paraphyletic taxa. These are natural products of evolution, which should be recognized in taxonomy. When ranks are adopted without acceptance of paraphyletic taxa, taxonomic free fall sets in, and every clade sinks into those taxa to which its original ancestor is referable. The clash of hierarchies results in absurdity, extending to sinking the entire plant kingdom into one family and one genus, but this has been strangely overlooked by the cladistic side. Adoption of ranked taxa is incompatible with recognition of only complete clades. Merely because one taxon falls phylogenetically within the clade of another taxon at the same rank does not necessarily mean that it must be included in it taxonomically. New characters will have arisen during evolution, which should be taken into account. A monophyletic (= holophyletic) system recognizing only complete clades is logically possible only if ranks are abandoned, as in the PhyloCode. It may be referred to as a “cladification” and the process producing it as “cladonomy,” which are quite different concepts from a “classification” and “taxonomy.” In a classification we have a hierarchical series of taxa at different ranks, while in a cladification we merely have a hierarchy of clades nesting within successively bigger clades. Cladistic taxonomy is particularly nonsensical in paleobotany, where our Linnaean taxonomic system and our code of nomenclature apply just as they do for extant plants. Cladograms are not classifications, and they need critical taxonomic assessment. The great majority of users of taxonomy are interested in characters and not cladistic theory. A general purpose classification is needed, requiring acceptance of paraphyletic taxa that are defined by characters as well as lines of descent. Examples in the dicot flowering plant families are given in which cladistic principles have imposed excessive insistence on lines of descent at the expense of evolution of characters, producing what many regard as bad taxonomy.
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The Case Against the Transfer of Dryandra to Banksia (Proteaceae)
Article
  • Nov 2014
The transfer of Dryandra R. Br. to Banksia L. f. was based on the use of holophyly (monophyly s. str.) as an essential criterion for recognition of taxa. The transfer was significant in scope and focuses on two iconic genera of plants in Western Australia. It has been accepted by some and rejected by others. It is one of many examples in a debate that pits recent genetic analysis against centuries of field and herbarium studies, and cladists against classical taxonomists. I argue that: (1) there are sound morphological characters distinguishing Dryandra from Banksia and they should be maintained as genera; (2) paraphyly should be accepted in biological classification; (3) scientifically, and for a morphologically complex genus of 137 specific and infraspecific taxa, the use of 11 taxa for the molecular analysis of Dryandra was insufficient; (4) some morphological data, mapped onto the cladogram a posteriori, were incorrect; (5) molecular cladistic approaches should complement rather than override pre-existing and extensive classifications based on phenotypic traits; (6) the acceptance of the transfer for the Australian Plant Census was premature according to guidelines published by Australian herbaria.
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Paraphyly and Endemic Genera of Oceanic Islands: Implications for Conservation
Article
  • Nov 2014
Genera of flowering plants that are endemic to oceanic islands are often of great biological interest. These groups represent adaptive complexes that confer distinction to the islands or archipelagos in which they are found, and this often results in a focus on their conservation. In recent decades, numerous molecular phylogenetic (and other evolutionary) studies have been done on island genera, hence providing much valuable new information on relationships and evolution of island groups. Genera restricted to oceanic islands derive evolutionarily from parental stocks usually in continental regions. These parental genera are often themselves evolutionarily successful, being particularly adept at dispersal, adaptation, and speciation. These immigrants to isolated oceanic islands derive from common ancestors of large and diverse parents or directly from within the lineages themselves. If in the latter case the island derivatives are treated at the generic level, then the parental genus becomes paraphyletic in a cladistic sense. In this circumstance there are three alternatives to classification of the island group: (1) treat both the island complex as a distinct holophyletic genus and the progenitor as a coordinate, but paraphyletic, genus; (2) submerge the island complex into the parental genus, perhaps at the subgeneric or sectional level, creating a larger holophyletic genus; or (3) divide the parental genus and island complex into a series of smaller genera in such a manner that all become holophyletic. A synthesis of recent investigations on 100 endemic island genera and relatives was completed in the Bonin Islands, Canary Islands, Galápagos Islands, Hawaiian Islands, Madeiran Islands, Robinson Crusoe Islands, and St. Helena. The results show that 64 genera are still accepted and remain uninvestigated or are seen as holophyletic in phylogenetic analyses. Seven have already been submerged based on non-cladistic results, and 29 are viewed as being nested within larger parental genera. Of this latter group, 15 of the genera are still being recognized at this time; six have been recommended as belonging within their parental genera; and eight have been formally transferred into the progenitor genera with combinations made. If further actions were to be taken based on strict holophyly, following the second alternative mentioned above, then these 29 genera would disappear as endemics in their islands or archipelagos. This would result in an overall average drop of 31.9% endemic genera in oceanic islands worldwide (based on the sample analyzed). With the third alternative, new generic concepts for the island and progenitor taxa would need to be worked out. Instead of recognizing genera on the basis of simple holophyly, genera should be based on cohesiveness, distinctness, and monophyly s.l. (i.e., including paraphyly and holophyly). A statistic is provided as a means for making these assessments quantitatively. The importance of unique and/or divergent character change for classification of island lineages is also stressed.
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Evolutionary Systematics and Paraphyly: Introduction
Article
  • Nov 2014
Phylogenetic systematics, especially involving molecular data, has had a remarkable impact on systematic biology. Numerous tree-building computer programs exist for the reconstruction of phylogenies, and many packages are available for analysis of population genetic data for estimating genetic divergence within and among populations. These advances have come about through the joining of statistical algorithms, computer programs, and DNA base-pair sequence and fragment data. Deeper genomic data are on the horizon for use with similar questions, and the next several years will witness many spectacular genetic advances. While great progress is being made on analytical approaches with molecular data in systematics, the use of the results of these analyses in biological classification has solidified into a dogmatic view, which has impeded further progress. Emphasis still remains on using only synapomorphies, even single characters, for delimitation of groups, on insisting that sister groups should have the same rank, and admitting only holophyletic (= monophyletic s. str.) groups. Evolutionary divergence within lineages and reticulate evolution are often ignored. As a result of these processes, paraphyletic groups, i.e., monophyletic groups that do not contain all descendants from a common ancestor, are often rejected. Evolutionary systematics takes the processes of descent and modification into consideration for reconstructing phylogenetic relationships involving many dimensions. This symposium presents various approaches for recognizing cladogenetic, anagenetic, and reticulate evolution in different organisms, which help reveal micro- and macro-evolutionary processes. Controversy still exists regarding how taxonomists should incorporate the diversity of evolutionary patterns and processes into biological classification. Case studies demonstrate that purely phylogenetic (cladistic) concepts of classification are unsatisfactory in cases of non-hierarchical relationships. Contributions also deal with the controversial question of recognition of paraphyletic groups in classification.
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