ArticlePDF Available

Accommodating Nomocharis in Lilium (Liliaceae)

Phytotaxa 277 (2): 205–210
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
Accommodating Nomocharis in Lilium (Liliaceae)
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:
*Author for corresponding
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
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-
=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
=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.
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.
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.
Balfour, B. (1918) The Genus Nomocharis. Transactions of the Botanical Society of Edinburgh 27: 273–300.
Brummitt, R.K. (2014) Taxonomy versus cladonomy in the dicot families. Annals of the Missouri Botanical Garden 100: 89–99.
Ehrendorfer, F. & Barfuss, M.H.J. (2014) Paraphyly and polyphyly in the worldwide tribe Rubieae (Rubiaceae): challenges for generic
delimitation. Annals of the Missouri Botanical Garden 100: 79–88.
Evans, W.E. (1925) A revision on the genus Nomocharis. Notes from the Royal Botanic Garden, Edinburgh 15: 25–26.
Franchet, A.R. (1889) Nomocharis. Journal de Botanzque 3: 113–114.
Franchet, A.R. (1892) Les Lis de la Chine et du Thibet. Journal de Botanique 6: 305–321.
Franchet, A.R. (1898) Platarum sinensium ecloge secunda. Journal de Botanzque 12 : 177–196.
Gao, Y.D., Harris, A., Zhou, S.D. & He, X.J. (2013a) Evolutionary events in Lilium (including Nomocharis, Liliaceae) are temporally
correlated with orogenies of the Q–T plateau and the Hengduan Mountains. Molecular Phylogenetics and Evolution 68: 443–460.
210 Phytotaxa 277 (2) © 2016 Magnolia Press
Gao, Y.D., Harris, A.J. & He, X.J. (2015) Morphological and ecological divergence of Lilium and Nomocharis within the Hengduan
Mountains and Qinghai–Tibetan Plateau may result from habitat specialization and hybridization. BMC Evolutionary Biology 15:
Gao, Y.D., Hohenegger, M., Harris, A., Zhou, S.D., He, X.J. & Wan, J. (2012) A new species in the genus Nomocharis Franchet (Liliaceae):
evidence that brings the genus Nomocharis into Lilium. Plant Systmatics and Evolution 298: 69–85.
Gao, Y.D., Zhou, S.D. & He, X.J. (2013b) Lilium yapingense (Liliaceae), a new species from Yunnan, China, and its systematic significance
relative to Nomocharis. Annales Botanici Fennici 50: 187–194.
George, A.S. (2014) The case against the transfer of Dryandra to Banksia (Proteaceae). Annals of the Missouri Botanical Garden 100:
Hayashi, K. & Kawano, S. (2000) Molecular systematics of Lilium and allied genera (Liliaceae): phylogenetic relationships among Lilium
and related genera based on the rbcL and matK gene sequence data. Plant Species Biology 15: 73–93.
Léveillé, H. (1913) Decades plantarum novarum. Repertorium specierum novarum regni vegetabilis 12: 281–288.
Harrow, R.L. (1930) Nomocharis. In: Cox, E.H.M. (Ed.) The New Flora and Silva. Dulau & Co. Ltd., London, pp. 75–78.
Hörandl, E. (2014) Nothing in taxonomy makes sense except in the light of evolution: examples from the classification of Ranunculus.
Annals of the Missouri Botanical Garden 100: 14–31.
Liang, S.Y. (1980) Flora Reipublicae Popularis Sinicae, Vol 14, Anagiospermae, Monocotyledoneae Liliaceae (I). Science Press, Beijing,
pp. 116–157.
Liang, S.Y. (1984) Studies on the genus Nomocharis (Liliaceae). Bulletin of Botanical Research 4: 163–178.
Linnaeus, C. (1753) Species Plantarum. Laurentius Salvius, Sweden, 302 pp.
Nishikawa, T., Okazaki, K., Arakawa, K. & Nagamine, T. (2001) Phylogenetic analysis of section Sinomartagon in genus Lilium using
sequences of the internal transcribed spacer region in nuclear ribosomal DNA. Breeding Science 51: 39–46.
Nishikawa, T., Okazaki, K., Uchino, T., Arakawa, K. & Nagamine, T. (1999) A Molecular Phylogeny of Lilium in the Internal Transcribed
Spacer Region. Journal of Molecular Evolution 49: 238–249.
Peruzzi, L., Leitch, I.J. & Caparelli, K.F. (2009) Chromosome diversity and evolution in Liliaceae. Annals of Botany 103: 459–475.
Ronsted, N., Law, S., Thornton, H., Fay, M.F. & Chase, M.W. (2005) Molecular phylogenetic evidence for the monophyly of Fritillaria and
Lilium (Liliaceae; Liliales) and the infrageneric classification of Fritillaria. Molecular Phylogenetics and Evolutuion 35: 509–527.
Sealy, J.R. (1950) Nomocharis and Lilium. Kew Bulletin 5: 273–297.
Sealy, J.R. (1983) A revision of the genus Nomocharis Franchet. Botanical Journal of the Linnean Society 87: 285–323.
Stuessy, T.F. & Hörandl, E. (2014) Evolutionary systematics and paraphyly: Introduction. Annals of the Missouri Botanical Garden 100:
Stuessy, T.F., König, C. & Sepúlveda, P.L. (2014) Paraphyly and endemic genera of oceanic islands: implications for conservation. Annals
of the Missouri Botanical Garden 100: 50–78.
Wilson, E.H. (1925) The lilies of Eastern Asia: a monograph. Dulau & Co. Ltd., London, 111 pp.
... introduction All true lilies -roughly 125 species of Lilium, now including the former genus Nomocharis (Gao and Gao 2016) -are restricted to the Northern Hemisphere, with centers of diversity in East Asia (where more than 50% of all species are native to China), southern Europe, the Caucasus, and eastern and western North America. Individual species are mostly restricted to limited areas within continents. ...
... This analysis produced a single, fully resolved tree with 95% to 100% statistical support for two-thirds of all inferred relationships, including 69 lily species and five outgroups spanning the three genera most closely related to the true lilies (Fritillaria, Cardiocrinum, Notholirion). Nomocharis species are now classified as part of Lilium (Gao and Gao 2016). Each pair of adjacent "twigs" in the tree represents a pair of closest relatives -so-called sister species -which diverged from each other and their inferred ancestor where the twigs split from the preceding eVoLuTioN, from the previous page eVoLuTioN, next page u branch. ...
... The genus Lily belongs to the family Liliaceae includes more than 110 species throughout the world, out of which 50-60 species are being cultivated in Asia, 24 species are being exploited in North America, and 12 species are exploiting in Europe (Dhiman et al., 2018). In the context of Asia, the East Asian region especially Japan, Korea, and China are thought to be the native of the many wild lily species (Mc Rae, 1998;Gao & Gao, 2016). The southwest and north parts of China are taken as natural habitats of more than 55 species of wild lily, and it is believed to be the center of the diversity of Lilium species (De Jong, 1974). ...
Full-text available
This experiment was carried out to evaluate the Lilium leichtlinii var maximowiczii germplasm collected from the different natural habitats from all over Korea. In total 30 accessions were studied for nine traits viz. plant height, leaf length, leaf width, the numbers of flowers, flower diameter, length of outer tepal, width of outer tepal, the number of leaf burn, and days to flowering in randomized block design with three replications. The ANOVA revealed highly significant variability prevailing among the investigated genotypes for almost all studied traits (except leaf width). The higher estimated value of GCV, PCV, heritability (H 2), and genetic advance as percent of the mean was obtained for the number of flowers and leaf burn. The moderate to high GCV and PCV coupled with higher heritability estimates (H 2) and GAM were found for plant height and flower diameter. Progeny selection would be effective as the prevalence of additive gene effect for these traits. Besides, leaf width, leaf length, length of outer tepal, the width of outer tepal, and days to flowering traits were possessed moderate to low GCV and PCV value coupled with the moderate value of heritability estimate with the low level of GAM proved to be the prevalence of non-additive genetic effect thereby indicating the necessity of alternative breeding approach for these traits improvement and breeding scheme. For the former group of traits breeding hybridization and selection would be an effective method, and primarily mean performance of these traits would be very handy for the decision of selection.
... We used the huge ITS phylogeny constructed by Gao et al. (2015), which includes 293 accessions from 103 of ca. 130 species of Lilium (including the genus Nomocharis; Gao and Gao, 2016) across the world. Ancestral morphological states were reconstructed across the ITS consensus tree, resulting from the Bayesian analysis and using the character matrices in Appendix S1. ...
Full-text available
This study aimed to characterize the morphological variations in the vegetative and floral traits of 73 wild Lilium amabile plants from six habitats in Korea. It was observed that L. amabile is distributed nationwide at any altitude from 300 m (Mt Mangdaeam) to 1550 m (Mt Halla). The majority of the natural habitats of L. amabile were found on mountain slopes, and some were found in rugged mountain regions. The down-facing flowers of this species not only had many blotches but also dense trichomes, and the flowering time was found to be from mid-June to mid-July. ANOVA revealed significant variations in vegetative and floral traits among the six habitats, indicating that the environment has substantial influences on the various growth parameters of L. amabile, such as plant height; number of leaves, bracts, papillae, and flowers; leaf angle; and lengths of the anther, longest blotch, and nectary of the petiole. In addition, the vegetative and floral traits were found closely correlated with each other under the direct impact of the environment. These findings will facilitate to find the appropriate environmental conditions for the conservation and development of L. amabile population as future lily-breeding materials.
Full-text available
Biological classification aims at establishing ordering systems for organisms. Principles of classification, however, differ in their criteria and in their information content. Cladistic classification emphasizes information on descent, but the strict application of logical inclusiveness leads in practice to disregard of modification and to a lack of information on evolutionarily relevant features. Phenetics provides information on similarity regardless of descent. Evolutionary classification maximizes information on evolution by combining information on descent and modification, but it relaxes the requirement of inclusiveness. In practice, this means accepting holo- and paraphyletic taxa, but rejecting polyphyletic groups. Review of a recently published case study of the species-rich and cosmopolitan genus Ranunculus demonstrates how evolutionary classification can be performed in practice. A hypothesis of descent was reconstructed by phylogenetic analysis of DNA sequence markers plus morphological and karyological characters. Based on this backbone phylogeny, information content on morphology, karyology, and ecology was used as a criterion for delimitation of infrageneric taxa. This concept resulted in the subdivision of a more basal, paraphyletic Ranunculus subg. Auricomus and the holophyletic Ranunculus subg. Ranunculus. On a sectional level, 14 holophyletic and two paraphyletic sections plus one monotypic section were classified. Holophyletic sections mostly reflect extinction gaps, while paraphyletic groups appear in clades that have reticulate evolution and/or ecological shifts. Classification of paraphyletic and monotypic sections preserves information on morphology, ecology, and evolutionary processes. This pluralistic approach is justified as it best reflects the diversity of the genus. The principle of broadening criteria maximizes information on descent and modification. Evolutionary classification facilitates practicability and stability of taxonomic work, as the broadening of criteria restricts the number of equally valid options for classification. For users, preserving information content on phenotypes aids practicability, because the connection to traditional literature and to modern information systems is optimally maintained.
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.
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.
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.
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.
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.