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CONSIDER the LILIES by Dean G

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The onion Allium platycaule (now in Alliaceae) from Soldier Creek, Modoc County. Photograph by J. Vale; its use courtesy of the Jepson Herbarium, UC. CONSIDER THE LILIES by Dean G. Kelch Consider the lilies of the field, flower, it wasn’t a member of the are a host of other “imposters” (see ho w they gro w; they to il no t, true lily genus Lilium. Table 1). neither do they spin. This particular response reflects These common names refer to Matthew VI: 28. a general tendency toward vague- plants that aren’t necessarily closely ness on the part of botanists. The related to each other. Not all of here is some controversy as term lily has been used in a very them are placed in the lily family, to just what wildflowers are loose fashion to describe a broad Liliaceae s.l. (sensu lato, in the broad Treferred to in the biblical array of plants. Consider the great sense). The most you can say is that passage above. Some have suggested diversity falling under the rubric they are all monocots. However, if that it is the autumn daffodil, lily. Besides the true lilies (Lilium) you count the water lily (Nymphae),a Sternbergia lute.a Most scholars are that include the tiger lily, the Ori- you can’t even say that! agreed, however, that whatever the ental lilies, and the Asian lilies, there The monocots (short for mono- VOLUME 30:2, APRIL 2002 FREMONTIA 23 Flowers of most monocots are TABLE 1. PLANTS THAT HAVE THE three-parted, an ancient character in flowering plants that is otherwise COMMON NAME OF LILY THAT ARE NOT found in only a few ancient lineages TRUE LILIES ( LILIUM ) of non-monocots. In many of the “lilies” s.l., the three sepals and three Daylily (Hemero calli)s petals are similar in appearance and Toadlily (Tricyrtu)s are referred to collectively as tepals. Corn lily (Veratrum) The vast majority of monocots are Desert lily (Hesperocall)is herbaceous. The few woody groups Trout lily (Erythronium) (such as palms and yuccas) have odd Lily-of-the-valley (Co nvallaria) wood that is unlike the dense, often Lily turf (Liriope) ringed wood of non-monocot trees Rain lily (Zephyranthe)s and shrubs. Voodoo lily (Amorphophallu)s As you can see from the above Sego lily (Calochortu)s characteristics, most monocots are Calla lily (Zantedeschi)a relatively simpler in structure than Lily-of-the-Nile (Agapanthus) other flowering plants. Therefore, Scarborough lily (Vallo ta) the various natural groups of mono- Plantain lily (Hosta) cots are less different from each Ginger lily (Hedychium) other than many groups of non- Glory lily (Gloriosa) monocots. For example, a strap- Checker lily (Fritillaria) shaped leaf does not lend itself to as Fortnight lily (Dietes) much shape variation as a net-veined Foxtail lily (Eremurus) leaf (think of the great leaf variety Spear-lily (Doryanthe)s in such non-monocot families as the Palm lily (Curculigo) carrot family, Umbelliferae, and the rose family, Rosaceae). This explains cotyledons) are one of the largest monophyletic groups of flowering plants. The remainder of the flow- ering plants are generally referred to as the dicots (dicotyledons), but this latter group is not monophyl- etic, as the monocots seem to have arisen from within the “dicots” early in the history of flowering plant evolution. The monocots include such diverse groups as arums, palms, gingers, grasses, irises, orchids, and, of course, lilies s.l. The members of these groups all have a single seed leaf (cotyledon), while most flower- ing plants have two seed leaves (think of a bean or alfalfa sprout). The cotyledon often isn’t around to look at for very long, but there are other characteristics that are common in monocots. Most monocots have linear or strap- shaped leaves with parallel veins. These leaves generally lack a well- defined petiole (stalk), and the base Sand lily, Leucocrinum montanum (now in Anthericaceae), from Lassen County. forms a sheath around the stem. Photograph by B. Ornduff; its use courtesy of the Jepson Herbarium, UC. 24 FREMONTIA VOLUME 30:2, APRIL 2002 TABLE 2. GENERA INCLUDED IN THE LILIACEAE SENSU LATO IN THE JEPSON MANUAL AND THEIR ASSIGNMENTS TO LILIACEAE SEGREGATE FAMILIES DISCUSSED IN THE TEXT Assignment of genera to families is based on the references provided. Agavaceae Lilium (lily) Agave Sc o lio pu(fos etid adder’s tongue) Camassia (camas) Strepto pus(t wisted-stalk) Chlo ro galum( soap plant, amole) Melanthiaceae Hastingsia Stenanthium Hesperocalli(sd esert lily) Veratrum (corn lily, false hellebore) Hesperoyucc(aour lord’s candle) Xero phyllum(b ear-grass, Yucca( Spanish bayonet) Indian basket-grass) Alliaceae Zigadenus(d eath camas) Allium (onion, garlic) Nartheciaceae Ipheio n(st ar flower) Narthecium( bog asphodel) Nothoscordum(fa lse garlic) Smilacaceae Anthericaceae sensu stricto Smilax (green briar) Leucocrinum(s and lily) Tecophilaeaceae Asparagaceae Odontostomum Asparagus Themidaceae Asphodelaceae Andro stephium Alo e Bloomeria( goldenstar) Aspho delu(sa sphodel) Bro diaea Convallariaceae Dichelostemma(b lue dicks, snake lily) Maianthemum( false lily-of-the-valley) Muilla Nolina(b eargrass) Triteleia( Ithuriel’s spear, pretty face) Smilac ina (false Solomon’s seal) Tofieldiaceae Hyacinthaceae Tofieldia( bog asphodel) Muscari (grape hyacinth) Trilliaceae (possibly within the Melanthiaceae) Liliaceae Trillium (wakerobin, trillium) Calochortu(sm ariposa lily, globe lily) Uvulariaceae Erythronium(fa wn lily) Clintonia (queen’s cup; potential placement) Fritillaria (fritillary) Disporum( twin bells) why the lily family (Liliaceae) has way. In the late 19th century, it approach was not useful in discern- long been a “catch-all” group that was common to recognize all “lil- ing the natural groups within the included most monocots with some- ies” bearing flowers with inferior Liliaceae s.l. Luckily, the work of what showy, radially symmetrical ovaries as the amaryllis family many botanists resulted in the accu- flowers and no obvious specializa- (Amaryllidaceae). While propo- mulation of a lot of information tions. (This contrasts with the very nents of this approach could boast about the microscopic and chemical specialized flowers of orchid family of its simplicity, it resulted in such characters of monocots. A group members, for example, which have diverse plants as daffodils and led by the Danish botanist Rolf the fused male and female flower agaves being lumped together, while Dahlgren decided to synthesize all parts forming a column.) yuccas (close relatives of agaves) of this information and revise the The lily problem was no secret were left in the Liliaceae. The tra- classification of monocots. This to botanists. There never was a ditional Liliaceae s.l. was like a gi- work was published in 1985 as The doubt that the lily family contained gantic house of cards: remove one Families o f Mo no co tyledo(Dnsahlgren, groups of quite disparate plants. The piece and the whole structure was et al. 1985). In this book, plants problem consisted in how to break liable to come crashing down. formerly in the Liliaceae s.l. were up the family in a practical, natural Obviously, the single character placed in 40 different families in VOLUME 30:2, APRIL 2002 FREMONTIA 25 Beargrass, Nolina parryi (now in Convallariaceae), from the Kingston Mountains. Photograph by C.S. Webber; its use courtesy of the Jepson Herbarium, UC. three different orders! Botanists and cluding the members of the mono- system of Dahlgren et al., such as horticulturists are, by and large, cotyledons. (For more information the listing of cultivated taxa in Kelch rather conservative, and so it has on this approach, see the sidebars (2000), will have to be modified in taken a while for these changes in on pages 4–7 and page 15.) the future. taxonomy to be accepted. The preliminary published re- While it may seem as if our en- During the 1990s, acceptance sults of this research confirm many tire classification of plants is sliding of the Dahlgren classification has of the conclusions reached in The into the abyss, things aren’t as bad as been hastened by the explosion of Families o f Monoco tyledoThens. re- they appear. The molecular data, by the academic discipline molecular sults also show that the story may and large, has confirmed much of systematics. By comparing the DNA be more complicated than we had our understanding of plant relation- sequences for a particular gene or hoped. This should come as no ships. Many traditional plant groups genetic marker for a large number surprise, as increased knowledge seem to be monophyletic. Of the of species, biologists found large leads us to a more sophisticated un- seriously unnatural groupings, per- numbers of new characters to use in derstanding of things. This is how haps the families of monocots rep- elucidating the evolutionary rela- science advances. Nevertheless, resent the most extreme case. How- tionships of living organisms, in- publications that closely follow the ever, there were earlier indications 26 FREMONTIA VOLUME 30:2, APRIL 2002 that the traditional taxonomy of The inclusion in the agave Hyacinthaceae and close to Hosta monocots was seriously flawed from group, based on evidence from com- based on serological data. the work of Dahlgren et al. Other parison of sequences of the chloro- Placing Hastingsia, Chlorogalum, groupings that have proven unnatu- plast gene rb cL, of some New World and Camassia in the Agavaceae ren- ral include the figwort family genera that Dahlgren had placed ders that family difficult to identify (Scrophulariaceae s.l.; see the article in the Hyacinthaceae (Camassia, based on macroscopic characters. It by Olmstead on page 13), and the Hastingsia , and Chlo ro galum) was is possible that further sampling will dogwood family (Cornaceae). something that no one had pre- identify two related lineages: one a Liliaceae s.l.
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  • Complete Chloroplast Genomes Shed Light on Phylogenetic

    Complete Chloroplast Genomes Shed Light on Phylogenetic

    www.nature.com/scientificreports OPEN Complete chloroplast genomes shed light on phylogenetic relationships, divergence time, and biogeography of Allioideae (Amaryllidaceae) Ju Namgung1,4, Hoang Dang Khoa Do1,2,4, Changkyun Kim1, Hyeok Jae Choi3 & Joo‑Hwan Kim1* Allioideae includes economically important bulb crops such as garlic, onion, leeks, and some ornamental plants in Amaryllidaceae. Here, we reported the complete chloroplast genome (cpDNA) sequences of 17 species of Allioideae, fve of Amaryllidoideae, and one of Agapanthoideae. These cpDNA sequences represent 80 protein‑coding, 30 tRNA, and four rRNA genes, and range from 151,808 to 159,998 bp in length. Loss and pseudogenization of multiple genes (i.e., rps2, infA, and rpl22) appear to have occurred multiple times during the evolution of Alloideae. Additionally, eight mutation hotspots, including rps15-ycf1, rps16-trnQ-UUG, petG-trnW-CCA , psbA upstream, rpl32- trnL-UAG , ycf1, rpl22, matK, and ndhF, were identifed in the studied Allium species. Additionally, we present the frst phylogenomic analysis among the four tribes of Allioideae based on 74 cpDNA coding regions of 21 species of Allioideae, fve species of Amaryllidoideae, one species of Agapanthoideae, and fve species representing selected members of Asparagales. Our molecular phylogenomic results strongly support the monophyly of Allioideae, which is sister to Amaryllioideae. Within Allioideae, Tulbaghieae was sister to Gilliesieae‑Leucocoryneae whereas Allieae was sister to the clade of Tulbaghieae‑ Gilliesieae‑Leucocoryneae. Molecular dating analyses revealed the crown age of Allioideae in the Eocene (40.1 mya) followed by diferentiation of Allieae in the early Miocene (21.3 mya). The split of Gilliesieae from Leucocoryneae was estimated at 16.5 mya.