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Trillium flexipes Raf., Autik. Bot. 133. 1840. Type:— USA. Kentucky: s.d., Rafinesque s.n., (P 1776245 digital image!, lectotype, designated here). Rafinesque (1840) described Trillium flexipes from plants in “West Kentucky and Tennessee ” with sessile leaves, 3-nerved, the peduncle “inclinato flexuoso,” lanceolate-acuminate sepals and white obovate-acute petals. No collector was provided in his description, but it was presumably one of his. A specimen at P (P01776245) from the Durand Herbarium has a label that provides only “ Kentucky, Rafinesque. ” Given that this is the only likely type material that we have located, we have designated this as the lectotype. Heterotypic synonyms: Trillium album Small, in part, see under T. rugelii. Trillium cernuum var. declinatum f. walpolei Farw., Rep. Michigan Acad. Sci. 21: 363. 1920. Type:— USA. Michigan: Washtenaw County, Ypsilanti, copses, flowers purple, 19 May 1918, Farwell 4849 (BLH0000359 digital image!, lectotype designated here; isolectotypes: BLH0000360 digital image!, GH00030152 digital image!, MICH1192832 digital image!, MICH1192834 digital image!). Note:— Patrick annotated the BLH specimen as lectotype, but he never published this. We formalize this here and designate BLH0000359 as lectotype. Trillium cernuum var. declinatum f. billingtonii Farw., Rep. (Annual) Michigan Acad. Sci. 21: 363. 1920. Type:— USA. Michigan: Washtenaw County, Ypsilanti, copses, flowers brown, 19 May 1918, Farwell 4850 (MICH1192831, lectotype designated here, digital image!; isolectotypes, BLH0000361 digital image!, BLH 00030151 digital image!, BLH0000362 digital image!, GH00030149 digital image!). Note:— McVaugh annotated the MI specimen as lectotype. Patrick later also annotated this specimen as the lectotype, but neither officially published this designation. Trillium erectum var. blandum Jennison, Rhodora 40: 486. 1938. Type:— USA. North Carolina: Swain Co., Opp. Ela, in rich shaded woods loam, elevation 1850 ft., 9 May 1937, Jennison 2185 (holotype: TENN; isotype: GSMNP01232). A paratype is at US. Trillium flexipes f. walpolei (Farwell) Fernald, Rhodora 46: 17. 1944. Basionym: Trillium cernuum var. declinatum f. walpolei Farw. Fernald (1944) considered this to be a red-flowered form of T. flexipes and the anther and pollen color support this, though the status of these red-flowered T. flexipes -like plants remains unclear, but they could be hybrids with sympatric T. erectum. Trillium flexipes is widely distributed over a large part of the Midwest from South Dakota eastward to Pennsylvania and south into northern Arkansas and Alabama. Over this area, it is variable in floral position (erect vs. reflexed or pendent) and fragrance (Case & Case 1997, AF pers. obs.). Within some parts of this distribution, it is uniform in morphology, e.g., in northern Alabama (Case & Case 1997, AF pers. obs.) where the plants have large upright flowers, overlapping large petals and an intense fragrance. In Missouri where AF has observed many plants the flowers are variably reflexed, declining, or horizontal and held just above the leaves to erect, but the flowers are smaller with narrowly ovate-lanceolate petals and without the distinctive powerful fragrance of some southern forms. Even within a geographic region, fragrance varies over the flowering period, and during a single day the fragrance varies from morning to afternoon (AF pers. obs.). For instance, Alabamian T. flexipes open with a distinctive mild tea rose-like fragrance discernible at close range in the early part of the day, but a southern magnolia-like fragrance is noticeable at a meter or more away by midday. Missouri forms have a mild, but pleasant floral fragrance., Published as part of Floden, Aaron J. & Knapp, Wesley M., 2023, Typification of the North American species of Trillium subg. Trillium (Melanthiaceae: Parideae), pp. 193-200 in Phytotaxa 599 (3) on page 196, DOI: 10.11646/phytotaxa.599.3.6, http://zenodo.org/record/8012591, {"references":["Rafinesque, C. S. (1840) Autikon botanikon. By the author, Philadelphia, 200 pp.","Case Jr., F. W. & Case, R. B. (1997) Trilliums. Timber Press, Portland, 285 pp."]}

Trillium vaseyi Harbison, Biltmore Bot. Stud. 1: 24. 1901. Type:— USA. North Carolina: M. N. Caro, 1878, Vasey s.n. (US00091979 digital image!, lectotype, designated here: isolectotypes: F0046074 F digital image!, GH00030177 digital image!, NY00319910 digital image!, NY00319911 digital image!, P01776194 digital image!, PH00028410 digital image!). Homotypic synonym: Trillium erectum var. vaseyi (Harbison) Ahles, J. Elisha Mitchell Sci. Soc. 80: 172. 1964. Heterotypic synonym: Trillium vaseyi f. album House, Muhlenbergia 6: 73. 1910. Type:— USA. North Carolina: Haywood Co., No. Pigeon Gap Trail, 4000 ft. alt., 9 May 1909, House 4146 (holotype: US 00091980 digital image!). Notes:— Harbison (1901) named Trillium vaseyi for G. R. Vasey, who collected this species in the North Carolina mountains in 1878. There are many specimens collected by Vasey in several herbaria. The Biltmore Herbarium was transferred to US, and we select the US collection (00091979) as the lectotype. One of the duplicates at NY (00319911) has a collection number of “ No 477 ” handwritten on the preprinted labels similar to that of most other duplicates. We consider this to be an isolectotype despite the collection number on the label. Annotations on NCU00000631 were noted by Patrick, and he considered this a syntype because Harbison had labeled it “co-type.” No other collection was mentioned in the protologue aside from Vasey’s gathering. Trillium vaseyi and T. simile co-occur in some regions, and in these mixed populations they hybridize if their flowering periods overlap (Case & Case 1997). Although they do share pendent flowers and long anthers, the anther and sometimes pollen color of T. vaseyi is yellow or purple to lavender, whereas that of T. simile is always yellow. Moreover, observation in the field over two decades shows that T. vaseyi typically flowers two to three weeks later than T. simile when they are sympatric (AF pers. obs). Forms of T. vaseyi with white petals still have a reddish ovary and purple anthers (Peattie 1927)., Published as part of Floden, Aaron J. & Knapp, Wesley M., 2023, Typification of the North American species of Trillium subg. Trillium (Melanthiaceae: Parideae), pp. 193-200 in Phytotaxa 599 (3) on page 198, DOI: 10.11646/phytotaxa.599.3.6, http://zenodo.org/record/8012591, {"references":["Harbison, T. G. (1901) New or little-known species of Trillium. Biltmore Botanical Studies 1: 19 - 24.","Case Jr., F. W. & Case, R. B. (1997) Trilliums. Timber Press, Portland, 285 pp.","Peattie, D. C. (1927) Trillium in North and South Carolina. Journal of the Elisha Mitchell Scientific Society 42: 193 - 206."]}

Trillium rugelii Rendle, J. Bot. 39: 331–332, pl. 426, f. B. 1901. Type:— USA. North Carolina: on the mountains of the Broad River, Mar 1841, Rugel s.n. (BM 327335 & BM 001009870 [one sheet] digital image!, lectotype, designated here; isolectotypes: E00346019 digital image!). Heterotypic synonyms: Trillium affine Rendle, J. Bot. 39: 334. 1901. Type:—USA. Georgia: s.d., Rugel s.n. (holotype: BM 000547280 digital image!) Trillium album (Michx.) Small, Fl. S.E. US. 278. 1903. Basionym: Trillium rhomboideum var. album Michx., Fl-Bor. - Amer. 1: 215. 1803. Type:— P 00730404 digital image!, lectotype, designated here) (see also T. erectum). Notes:— Trillium rugelii was described in 1901 by Rendle from a Rugel collection at BM. In the protologue, he stated “Specimens in Herb. Mus. Brit.” There is one sheet at BM that has two stems, one of which was selected as a lectotype, as noted on the annotation label by Patrick (c. 1981), but this typification was never formally published. The stem on the upper left was selected as the lectotype by Patrick and the stem on the right was selected as an isolectotype despite the presence of a single collection label. The ICN rules (ICN 8.2) state that this should be treated as a single gathering. Thus, we select the sheet at BM as the lectotype (BM327335 & BM001009870). The combination T. album was made by Small (1903). The basionym was Michaux’s T. rhomboideum var. album , which was based on specimens that are in fact T. rugelii. Here, we typify the name T. rhomboideum var. album and designate the specimen at P (00730404) as the lectotype of T. rhomboideum var. album. Notes:— Case (1997) reported differences in fragrance between the forms in the Southern Appalachians vs. plants from the middle and northern parts of Alabama. He described them as strongly fragrant of old garden roses but did not mention that of the mountain plants. Observations by AF of T. rugelii from Tennessee south to Alabama and east to South Carolina have shown that floral fragrance is variable and often similar to some forms of foamflower (Tiarella), fragrant violets, or Cyclamen. The intensity of the fragrance varies over the distribution, but the floral fragrance is often stronger in more southern populations., Published as part of Floden, Aaron J. & Knapp, Wesley M., 2023, Typification of the North American species of Trillium subg. Trillium (Melanthiaceae: Parideae), pp. 193-200 in Phytotaxa 599 (3) on pages 196-197, DOI: 10.11646/phytotaxa.599.3.6, http://zenodo.org/record/8012591, {"references":["Small, J. K. (1903) Flora of the southeastern United States. New York, published by the author, 1370 pp.","Case Jr., F. W. & Case, R. B. (1997) Trilliums. Timber Press, Portland, 285 pp."]}

Trillium hibbersonii (T.M.C.Taylor & Szczaw.) D.O’Neill & S.B.Farmer, Phytotaxa 436: 193. 2020. Basionym: Trillium ovatum Pursh forma hibbersonii T.M.C. Taylor & Szczaw., Syesis 7: 250. 1974. Type:— CANADA. British Columbia: near Boat Basin, Hesquiat Harbour, west coast of Vancouver Island, elev. 2000 ft., 1938, Hibberson s.n. (holotype: UBC!). Homotypic synonyms: Trillium ovatum Pursh var. hibbersonii (T.M.C.Taylor & Szczaw.) G.W.Douglas & P.Pojar, Canad. Field-Naturalist 115: 343. 2001. Trillium hibbersonii Wiley, nom. nud., Rare wildflowers of North America. 1968., Published as part of Floden, Aaron J. & Knapp, Wesley M., 2023, Typification of the North American species of Trillium subg. Trillium (Melanthiaceae: Parideae), pp. 193-200 in Phytotaxa 599 (3) on page 196, DOI: 10.11646/phytotaxa.599.3.6, http://zenodo.org/record/8012591

Trillium simile Gleason, Bull. Torrey Bot. Club 33: 391. 1906. Type:— USA. North Carolina: moist woodlands near Tryon, 28 April 1904, Gleason 14933-a (holotype: NY00051079 digital image!). Homotypic synonyms: Trillium vaseyi var. simile (Gleason) Barksdale, J. Elisah Mitchell Sci. Soc. 54: 285. 1938. Trillium erectum var. vaseyi f. simile (Gleason) Ahles, J. Elisah Mitchell Sci. Soc. 80: 172. 1964. Notes:— The concept of T. simile has been greatly expanded and the morphological characters misapplied since its description. Gleason (1906) compared T. simile to T. rugelii in his description and diagnosis of this new species, stating it differed by its longer stamens, yellow anthers (vs. purple), and proportionately longer filaments. Gleason was familiar with T. rugelii and its typical habit of declining to reflexed flowers held beneath the leaves, so his comparison of this novelty suggests that the type gathering of T. simile had flowers held beneath the leaves. In his description, he even stated that the peduncles were “declined or cernuous” (Gleason 1906). Barksdale (1938) placed T. simile at varietal status under T. vaseyi, also confirming the original concept of Gleason (1906) that the flowers were pendent. Barksdale (1938) was confident that T. simile was merely a color variant of T. vaseyi occurring in mixed populations with the white form being rare but also in isolated uniformly white-flowered populations. Peattie (1927) argued that T. simile has erect peduncles, but sometimes declined flowers based on the type gathering and his field experience. Although the type at NY does have the flowers in an upright position, it is not apparent whether this is due to a pressing artifact or the plants truly having erect peduncles. The current concept is that of a large plant, with erect peduncles, large creamy-white petals that usually are overlapping at their bases, a dark ovary, and yellow pollen (Case & Case 1997, Case 1997). Forms like this occur throughout the Southern Appalachians (Case 1997). Some specimens have been observed in herbaria that share this morphology from Ohio, West Virginia, and Virginia (AF pers. obs.). Barksdale (1938) said that Small’s (1933) treatment could apply to nearly any white-flowered erect Trillium with a dark ovary, and it appears that the current concept does not match the type based on the protologue in which Gleason (1906) compared his new species to T. rugelii. Floral fragrance in Trillium has been used as a field character when comparing related species. Peattie (1927), Small (1933), Case (1997), and others have mentioned the sweet or delicious floral scent of T. simile, which the authors have observed on various erect and reflexed forms in the Southern Appalachians that fit both concepts of T. simile. A form that is common in and around the Great Smoky Mountain National Park (North Carolina and Tennessee) at lower elevations on rich mesic slopes has been considered the typical form with large creamy-white, erect flowers (Case & Case 1997), but this form has a fragrance identical to that of the higher elevation reddish forms of T. erectum, i.e., wet dog or egg whites. Forms with erect peduncles have been observed with a sweet applelike fragrance, and declining forms observed in western North Carolina and northern Georgia have been observed with a distinctive spicy clove-like or a tea rose-like fragrance. Like T. flexipes, it may be that the fragrance changes temporally over the flowering period to attract potential pollinators or to even deter herbivory, and that the volatiles producing the fragrance vary during the day due to temperature. In addition to the fragrance difference, T. simile differs from T. erectum in its much longer filaments and anthers that surpass the ovaries, but there has never been a wide-range study that surveys this character and its taxonomic value. Molecular analyses of T. subg. Trillium may resolve this complex of white-flowered species., Published as part of Floden, Aaron J. & Knapp, Wesley M., 2023, Typification of the North American species of Trillium subg. Trillium (Melanthiaceae: Parideae), pp. 193-200 in Phytotaxa 599 (3) on page 197, DOI: 10.11646/phytotaxa.599.3.6, http://zenodo.org/record/8012591, {"references":["Gleason, H. A. (1906) The pedunculate species of Trillium. Bulletin of the Torrey Botanical Club 33: 387 - 396. https: // doi. org / 10.2307 / 2478819","Barksdale, L. (1938) The pedicellate species of Trillium found in the southern Appalachians. Journal of the Elisha Mitchell Scientific Society 54: 271 - 296.","Peattie, D. C. (1927) Trillium in North and South Carolina. Journal of the Elisha Mitchell Scientific Society 42: 193 - 206.","Case Jr., F. W. & Case, R. B. (1997) Trilliums. Timber Press, Portland, 285 pp."]}

Trillium sulcatum Patrick, Brittonia 36: 27–35, f. 1–5. 1984. Type:— USA. Tennessee: Grundy Co., rich bouldery woods at base of N-facing slope near Deer Lick Falls in Layne Cove, elev. ca. 420 m, western escarpment Cumberland Plateau, 2 km due N of Monteagle, 21 Apr 1980, Patrick, Perkins & Horn 1122 (holotype: TENN!; isotypes: BH, DUKE10000089 digital image!, GH00030175 digital image!, MO-104974!, NCU00000629 digital image!, NY00319900 digital image!, UNA, US 00406395 digital image!). Heterotypic synonyms: Trillium erectum var. sulcatum Barksdale, J. Elisha Mitchell Sci. Soc. 54: 280–281. 1938. Type:— USA. North Carolina: Surry Co., in deep woods near Roaring Gap, 27 Apr 1937, Lyons s.n. (holotype: NCU 00000625 digital image!; isotypes, NCU 00000621 digital image!, NCU 00000622 digital image!, NCU 00000623 digital image!, NCU 00000624 digital image!, NCU 00000626 digital image!). Note:— This name was published without a Latin diagnosis and is thus invalid. The date of collection is given as “1934” on plants.jstor. org, but the specimen labels read 1937. Trillium sulcatum f. albolutescens T.S.Patrick, Brittonia 36: 30. 1984. Type:— USA. North Carolina: Ashe Co.; rich shady western slopes of “The Peak,” in western part of Ashe County, 16 May 1950, Harper 4167 (holotype: MO-104973!; isotypes: GH00030176 digital image!, NY00319901 digital image!, US 00406185 digital image!). Notes:— This species is similar to T. erectum but occurs mostly occurs to the west, largely on the Cumberland Plateau in Kentucky, Tennessee and adjacent Alabama. The two are sympatric in a small portion of their distributions in eastern Kentucky, north-eastern Tennessee, south-western Virginia, and north-western North Carolina along the Blue Ridge escarpment of the Southern Appalachians. In some cases, the two species are hard to separate based and may be better regarded as a single variable species with a broad range of variation and similarities in floral fragrance and habitat preferences. However, in cultivation, lower-elevation T. sulcatum is a better garden plant and more vigorous. In the Ridge and Valley of Tennessee, white-flowered forms in uniform populations have been misidentified as T. flexipes, but these forms have dark ovaries and the wet-dog floral fragrance of the typical forms (AF pers. obs.)., Published as part of Floden, Aaron J. & Knapp, Wesley M., 2023, Typification of the North American species of Trillium subg. Trillium (Melanthiaceae: Parideae), pp. 193-200 in Phytotaxa 599 (3) on page 198, DOI: 10.11646/phytotaxa.599.3.6, http://zenodo.org/record/8012591

Trillium cernuum L., Sp. Pl. 1: 339–340. 1753 Type:— USA. In Carolina, [no collector given], 469.1 (LINN!, lectotype designated by Gleason, Bull. Torrey Bot. Club 33: 390, 1906) Heterotypic synonyms: Trillium brevipetalum Raf., Med. Fl. 2: 100. 1830. Type:—Near the lakes, Ontario and Erie. Type not located. Trillium glaucum Raf., Med. Fl. 2: 102. 1830. Type:— In Pennsylvania, near Philadelphia, Maryland, Virginia, “& c.”.” Type not located. Trillium hamosum Raf., Med. Fl. 2: 101. 1830. Type:—In the Pocono Mountains, Pennsylvania, discovered by Steinhauer. Type not located. Trillium medium Raf., Med. Fl. 2: 102. 1830. Type:—from New England south to Virginia. Type not located. Trillium declinatum (A.Gray) Gleason, Bull. Torrey Bot. Club 33: 389. 1906. nom. illeg. Basionym: Trillium erectum var. declinatum A.Gray, Manual [Gray], ed 5: 523. 1867. Type:— USA. 1863, Robbins s.n. (holotype,: GH 00030145 digital image!; isotypes, GH00030146 digital image!, MO-104984!). Notes:— Patrick annotated MO 104984 as Trillium cernuum, against which Gray (1867) compared his var. declinatum in the protologue. The name at species level is preceded by T. declinatum Raf., which is T. catesbaei, and Fernald (1932) provided the new name T. gleasonii. Patrick also annotated NY00319876(!) as lectotype but never published this. This collection, NY00319876, was annotated by Patrick as T. flexipes. This may have been one of the specimens that Gray (1867) used as part of his description of var. declinatum , but he cited only collections by Robbins from Lake Superior with no mention of additional specimens in his description of var. declinatum. Fernald (1932) later provided a new name, T. gleasonii, because T. declinatum Raf. preceded T. declinatum (A.Gray) Gleason. We here assign this concept to T. cernuum. Trillium cernuum var. macranthum Eames & Wiegand, Rhodora 25: 191. 1923. Type:— USA. New York: Tompkins, Valley of Fall Creek, Round Marshes (Gracie’s Swamp) and vicinity, Dryden, low woods, north end of marshes, 31 May 1914, Eames & MacDaniels 2012 (holotype: GH00030148!; isotypes: CM0361 digital image!, F0046068 F digital image!, MIN1002612 digital image!, S-G-7311 digital image!). Note:— Eames & Wiegand (1923) discussed variation in T. cernuum. Trillium cernuum f. viride Cayoutte & Cayoutte, Naturaliste Canad. 111: 326. 1984. Type:— CANADA: Quebec: Saguenay –Lac-Saint-Jean (Quebec), Saguenay, Ilets-Jermie, 48°53’N- 68°48’W, 12 Jul 1973, Cayouette 73-418 (holotype: not located; isotypes: DAO000466224 digital image!, CM0491 digital image!, GH00030151 digital image!, MICH1192833 digital image!, NY00319873 digital image!, US00288583 digital image!). Trillium cernuum f. marginatum Cayouette & Cayouette, Naturaliste Canad. 111: 326. 1984. Type:— CANADA. Quebec: Comté de Gaspé-quest, Mont-Saint-Pierre, 49°13’ N-65°49’0. Erablière à bouléau jaune, en flanc de côteau, exposition sud-est. 13 Jun 1973, Cayouette & Quattara 73-145 (holotype ” QFA; isotypes: CM0360 digital image!, DAO000466223 digital image!, GH 00030150 digital image!, MICH1192832 digital image!, NY00319872 digital image!, US00288582 digital image!). Trillium cernuum f. tangerae Wherry, Bartonia 23: 49. 1945. Type:— USA. Pennsylvania: Adams County, 2 miles west-southwest of Iron Springs, moist rocky woods, 5 May 1945, Wherry s.n. (holotype: PH 00028390, lectotype designated here, digital image!; isolectotype: PH 00028389 digital image!). Note:— Patrick annotated PH 00028390 as lectotype. We formalize this designation here. Trillium cernuum var. terrae -novae Boivin, Rhodora 55: 101–102. 1953. Type:— CANADA. Newfoundland: west coast near Stephensville, west side of E. Harmon Air Force Base, in wet black soil under Alder bushes, 9 June 1949, Bassett 293 (holotype: DAO 000466288 digital image!)., Published as part of Floden, Aaron J. & Knapp, Wesley M., 2023, Typification of the North American species of Trillium subg. Trillium (Melanthiaceae: Parideae), pp. 193-200 in Phytotaxa 599 (3) on pages 194-195, DOI: 10.11646/phytotaxa.599.3.6, http://zenodo.org/record/8012591, {"references":["Gray, A. (1867) A manual of botany of the northern United States (ed. 5). Putnam, New York, 703 pp. https: // doi. org / 10.5962 / bhl. title. 65524"]}

The mitochondrial DNA has been used widely in molecular baraminology studies. Besides the mitochondrion, the chloroplasts of plants harbor an entire organelle genome, which can also be harnessed for molecular baraminology studies, just like the mitochondrial DNA. The ten-fold size of the chloroplast genome allows a much larger sequence space to be analyzed, but it also brings its own set of analytical challenges to the table. In this study, the chloroplast genome of 163 species from the plant order Liliales were examined, with the genus Stemona (order Pandanales) as an outgroup. A multiple alignment was created, and a sequence similarity matrix was derived from it, which was then clustered into several putative holobaramins. Of these, six groups had more than five members. These putative baramins are: Tulipa+Amana, Disporum, Fritillaria+Lilium+Nomocharis+Notholirion, Daiswa+Paris+Trillium, Smilax, and Veratrum. Compared to the results of a previous morphological analysis, there are some discordances. The previous analysis separates Trillium from Daiswa and Paris, and unites Alstroemeria with Disporum. Ultimately, genetics should decide baraminic classification, albeit the chloroplast genome still makes up only a small fragment of the entire plant genome. Following this study, it is hoped that chloroplast genome analysis may enrich the toolkit of molecular baraminology.

At present, the evaluation methods for pharmaceutical properties of Chinese medicinal films have many problems, such as poor objectivity for the indexes and no quantitative and standardized evaluation methods. This study established a new method using three important physical property parameters, i.e., flow index, weight loss rate, and elongation rate, which were closely related to the pharmaceutical properties of films. On this basis, the above parameters were taken as indicators to optimize the film formulation of Trillium tschonoskii total saponins and verify the feasibility and suitability of the established method and parameters in formulation optimization. A self-made flow distance detection device and a viscometer were used to measure and characterize the fluidity, where the flow index refers to the ratio of the flow distance per unit time to the viscosity. The weight loss rate was measured by the 3 M transpore~(TM) surgical tape. The film-forming property was characterized by the weight loss rate of the sample within a certain period of time. An electronic tension machine was employed to measure the elongation rate after drying, which was used to characterize the ductility of the film. The results showed that the established method for the determination of flow index, weight loss rate, and elongation rate was stable and reliable. The optimal film formulation of T. tschonoskii total saponins could be obtained by optimization with those indicators. As demonstrated, the above evaluation indicators(flow index, weight loss rate, and elongation rate) can guide the optimization and design of formulation, and the new evaluation method constructed based on this shows a good application prospect in formulation optimization and formulation quality evaluation of medicinal films.

Steroidal sapogenins (SS) are structural analogues of steroidal drugs, which are frequently used for the treatment of several diseases including reproductive, malignancies, neurological, and inflam...

Key to the subgenera of Trillium 1 Flowers usually pedicellate; leaf uniformly green, usually not mottled............................................................................................ 2 - Flowers sessile; leaf green usually with darker shades of green and silver mottling scattered over the surface4. T. subgen. Sessilia 2 Stigmas uniform (not tapered), united in a common style or at least basally connate; ovary at anthesis somewhat obscured by corolla and stamens and relatively less exserted, 3- or 6-angled, typically white, greenish white or green; petal with margins undulate, basally overlapping, pink or white aging to various pink shades or remaining white (T. nivale); adaxial petal venation not engraved; flower funnelform or at least with basally erect petals shallowly encircling ovary; leaf ovate, elliptic-ovate, ovaterhombic, ovate-lanceolate or oblong.................................................................................................................................................. 3 - Stigmas thicker basally and tapered above, separate; ovary at anthesis exposed, exserted or only slightly obscured, typically strongly 6-angled or ridged, generally dark red, to creamy white or white with red markings; petal with margins entire, separate or only slightly overlapping, if white, aging to brown; adaxial petal venation typically engraved; flower open, or widely agape, not funnelform; leaf shape rhombic, broadly elliptic, rhombic-ovate or lanceolate-elliptic (T. hibbersonii)..........1. T. subgen. Trillium 3 Stigmas connate; leaf elliptic-ovate, ovate-lanceolate or oblong.................................................................. 3. T. subgen. Delostylis - Stigmas only basally connate; leaf ovate, elliptic-ovate or ovate-rhombic................................................ 2. T. subgen. Callipetalon, Published as part of Lampley, Jayne A., Gereau, Roy, Floden, Aaron & Schilling, Edward E., 2022, A revised subgeneric classification of Trillium (Parideae, Melanthiaceae), pp. 278-286 in Phytotaxa 552 (5) on page 280, DOI: 10.11646/phytotaxa.552.5.1, http://zenodo.org/record/6796567

Premise One of the best-documented ecological responses to climate warming involves temporal shifts of phenological events. However, we lack an understanding of how phenological responses to climate change vary among populations of the same species. Such variability has the potential to affect flowering synchrony among populations and hence the potential for gene flow. Methods To test whether an earlier start of the growing season affects the potential for gene flow among populations, we quantified the distributions of flowering times of two spring-flowering plants (Trillium erectum and Erythronium americanum) over 6 years along an elevational gradient. We developed a novel model-based metric of potential gene flow between pairs of populations to quantify the potential for pollen-mediated gene flow based on flowering phenology. Results Earlier onset of spring led to greater separation of peak flowering dates across the elevational gradient for both species investigated, but was only associated with a reduction in potential gene flow in T. erectum, not E. americanum. Conclusions Our study suggests that climate change could decrease gene flow via phenological separation among populations along climatic gradients. We also provide a novel method for quantifying potential pollen-mediated gene flow using data on flowering phenology, based on a quantitative, more biologically interpretable model than other available metrics.

PREMISE Environmental heterogeneity influences plant distributions and diversity at several spatial scales. In temperate forests, fine-scale environmental variation may promote local coexistence among herbaceous species by allowing plants to spatially partition microsites within forest stands. Here we argue that shallow soils, low soil water-holding capacity and fertility, and reduced light near tree boles should favor short, shallow-rooted, evergreen species like Anemone acutiloba with low moisture, nutrient, and light requirements. Farther from trees, richer, deeper soils should favor taller, deeper-rooted herbs with greater moisture and nutrient demands, such as Sanguinaria canadensis and Trillium flexipes. METHODS We tested these hypotheses by mapping the fine-scale distributions of Anemone, Sanguinaria, and Trillium individuals within a 50 × 50 m plot, comparing local species' distributions with respect to soil depth and proximity to neighboring trees, and characterizing intraspecific and interspecific spatial associations. RESULTS Local plant distributions were consistent with our predictions based on leaf height, physiology, and phenology. Anemone was found in microsites on shallower soils and closer to trees than either Sanguinaria or Trillium. In all three species, individual plants were spatially aggregated within 2 m, but spatially segregated from individuals of the other species beyond 2 m. CONCLUSIONS Differential plant responses to fine-scale environmental heterogeneity and observed spatial associations suggest that local species-environment associations could facilitate coexistence. These findings illustrate how fine-scale environmental heterogeneity coupled with phenological and physiological differences likely contribute to spatial niche partitioning among spring-flowering forest herbs and maintain high local plant diversity within temperate forests.

Models of ecological responses to climate warming predict species’ migration towards higher latitudes or elevations. However, models often neglect nonclimatic factors, such as herbivory, which could slow down or prevent geographic range expansion. A previous study in Mont Mégantic National Park (Quebec) found that in one year (2016), browsing by white-tailed deer on Trillium erectum L. was substantially higher at high elevations than low elevations. Under the hypothesis that deer herbivory could limit the upper elevational range expansion of T. erectum, here we ask (i) whether herbivory increased with elevation during two additional years (2017 and 2018), (ii) whether the rate of deer visitation increases with elevation, and (iii) whether the effect of herbivory on relative growth rates increases with elevation. Contrary to the earlier study, we did not find a significant trend of herbivory with elevation, although there was a weak positive relationship between deer visitation and elevation. We found a strong negative impact of browsing on relative growth rates, but the magnitude of this effect did not vary with elevation. Our results thus do not support the hypothesis that herbivory limits the range of T. erectum at high elevation, but herbivory could have a negative impact on populations in general if the browsing rate remains high.

INTRODUCTION Trillium govanianum (Nag Chhatri and Teen Patra) is traditionally used for curing joint pains, wounds, and sexual disorders. Steroidal saponins are the main active components of this species. However, only a small amount of information is available about steroidal saponins of this plant. OBJECTIVE To develop an ultra-high-performance liquid chromatography-quadrupole time of flight tandem mass spectrometry (UHPLC-QTOF-MS/MS) and ultra-high-performance liquid chromatography-evaporative light scattering detector (UHPLC-ELSD) methods for the qualitative and quantitative determination of steroidal saponins in T. govanianum. METHOD The dried rhizomes of T. govanianum (100 mg) were extracted with ethanol-water (80:20, 10 mL) by ultrasonic treatment for 30 min at 40°C. The prepared sample was analysed by UHPLC-QTOF-MS/MS and UHPLC-ELSD for the qualitative and quantitative determination of steroidal saponins. RESULT A total of 24 saponins were identified using UHPLC-QTOF-MS/MS; seven of them were characterised by comparing with standards. Furthermore, five saponins [govanoside B (2), protodioscin (6), pennogenin tetraglycosides (11), borassoside E (21) and borassoside D (24)] were quantified using UHPLC-ELSD method in different extracts and fractions of T. govanianum. The method showed good linearity (R2 ≥ 0.993), limit of detection (0.92-4.09 μg/mL), limit of quantification (3.1-13.5 μg/mL), precision [intra-day relative standard deviations (RSDs) < 4.3% and inter-day RSDs < 5.5%], and accuracy (84.0-110.3%). This is the first report on the quantification of 2, 6, 11, 21 and 24 in T. govanianum. CONCLUSION The present study provides an efficient analytical method for the identification and quantification of steroidal saponins and will be helpful for the quality evaluation of T. govanianum.

In Himachal Pradesh tribal migratory shepherds carry rich repository of traditional knowledge of wild medicinal plants and its uses, in this respect, an ethnobotanical survey was carried out in Tungasigarh and its surrounding area of Thunag Subdivision district Mandi Himachal Pradesh from 2018 to 2019. The required information on ethnomedicines used by tribal migratory shepherds was collected through personal field visits, interview method and by using a pretested questionnaire. Total 64 medicinal plant species were reported viz. Aconitum heterophyllum Allium ursinum, Allium humile, Trillium govaninum, Bergenia ciliata, Berberis lyceum, Cannabis sativa, Dioscrea deltoidea, Rhododendron arboratum, Pistacia integerrima, Zanthoxylem armatum was recorded. Total of 64 species were documented herb species were dominant (48) followed by shrub (9), tree (7). This study shows that shepherds in tribal areas are highly dependent on ethnobotanical medicines, which evolved over generations of experience, for the healthcare. This survey can help as baseline data on ethnomedicinal plants used in Thunag sub division of Mandi district and could be helpful in conservation of traditional knowledge as well as medicinal plants.

Introduction: Breast cancer cases, mastectomy and following reconstruction procedures are growing in numbers. Despite being lifesaving, mastectomies have a destructive psychological impact on the patients. On the other hand, breast reconstruction improves psychological damages within the same population. Various techniques for nipple reconstruction were described in literature. Trillium flap is an innovative technique to reconstruct neo-nipple with several advantages that make it superior to other popular flaps. Objectives: To come up with an innovative design for reconstructing a neo-nipple post mastectomy, that is superior to other popular flaps. Results: The Trillium flap design has less visible and easily camouflaged scars, is geometry-based, specific, well-detailed and flexible to produce a tailored nipple with any desired height and diameter and ensures the flaps good vascularity and the neo-nipple projection sustainability. Conclusion: Trillium flap is an innovative technique to reconstruct neo-nipple with several advantages that make it superior to other popular flaps. The results shown in the study are for experimental procedures done on human tissue samples of excised flaps from abdominoplasties and brachioplasties. Further application on actual cases is needed with monitoring of neo-nipple projection sustainability on the long term.

The antidiarrheal effect of methanolic extract of

Hurricane Hugo was a category five storm in September of 1989 that significantly impacted natural areas along the Carolina coastal plain through wind damage and storm surge flooding. Francis Beidler Forest, an Audubon wildlife sanctuary in Four Holes Swamp, suffered severe damage to its forest canopy. In response to concerns that the rare spring ephemeral Trillium pusillum var. pusillum may be negatively impacted by the loss of the mixed hardwood canopy, we established permanent plots in the spring of 1990 with single leaf, triple leaf, and flowering individuals recorded by Ecology faculty and students at The Citadel. Disruption to the forest canopy would significantly alter forest floor microhabitat conditions negatively affecting T. pusillum var. pusillum population demography. There was no expected negative effect of Hurricane Hugo, and the loss of canopy cover on the population. Evidence suggests that the decrease in canopy cover and increased light was associated with increased flowering. The coefficient of variation, as a measure of cohort variability among years, increased from flowering, to triple leaf, to single leaf across the 29 years of population monitoring. There was a significant positive association between the number of named storms in the previous two and three years and the number of single leaf plants. There was no evidence that the population is decreasing, even though the number of flowering individuals has decreased. The results of this long-term demography suggest that even severe natural disturbances, like hurricanes and tropical storms, may have a positive effect on Trillium population dynamics.

Ethnopharmacological relevance Trillium tschonoskii Maxim. is one of traditional Chinese medical herbs that has been utilized to treat brain damages and cephalalgia. The neuroprotective effect of total saponins from Trillium tschonoskii rhizome (TSTT) has been demonstrated efficacy in rats following ischemia. However, the axonal remodeling effect of TSTT and the detailed mechanisms after ischemic stroke have not been investigated. Aim of the study We aimed to estimate therapeutic role of TSTT in axonal remodeling using magnetic resonance imaging (MRI) technique, and explored possible mechanisms underlying this process followed by histological assays in ischemic rats. Methods Male Sprague-Dawley (SD) rats underwent permanently focal cerebral ischemia induced by occluding right permanent middle cerebral artery. TSTT was intragastrically administrated 6 h after surgery and once daily for consecutive 15 days. Neurological function was assessed by the motor deficit score and beam walking test. T2 relaxation mapping and diffusion tensor imaging (DTI) were applied for detecting cerebral tissues damages and microstructural integrity of axons. Luxol fast blue (LFB) and transmission electron microscope (TEM) were performed to evaluate histopathology in myelinated axons. Double immunofluorescent staining was conducted to assess oligodendrogenesis. Furthermore, the protein expressions regarding to axonal remodeling related signaling pathways were detected by Western blot assays. Results TSTT treatment (65, 33 mg/kg) markedly improved motor function after ischemic stroke. T2 mapping MRI demonstrated that TSTT decreased lesion volumes, and DTI further confirmed that TSTT preserved axonal microstructure of the sensorimotor cortex and internal capsule. Meanwhile, diffusion tensor tractography (DTT) showed that TSTT elevated correspondent density and length of fiber in the internal capsule. These MRI measurements were confirmed by histological examinations. Notably, TSTT significantly increased Ki67/NG2, Ki67/CNPase double-labeled cells along the boundary zone of ischemic cortex and striatum. Meanwhile, TSTT treatment up-regulated the phosphorylation level of Ser 9 in GSK-3β, and down-regulated phosphorylated β-catenin and CRMP-2 expression. Conclusion Taken together, our findings indicated that TSTT (65, 33 mg/kg) enhanced post-stroke functional recovery, amplified endogenous oligodendrogenesis and promoted axonal regeneration. The beneficial role of TSTT might be correlated with GSK-3/β-catenin/CRMP-2 modulating axonal reorganization after ischemic stroke.

Trillium Linnaeus (1753: 339) species have a reputation for being difficult to distinguish from one another, both on morphological (e.g. Freeman 1985, Schilling et al. 2019) and initial studies with standard plastid markers (Osaloo et al. 1999). Thus, the species that is sister to the rest of the Parideae, Pseudotrillium rivale (Watson 1885: 378) Farmer (2002: 687), was only recognized to be distinctive after DNA sequence data became available (Farmer & Schilling 2002). In the somewhat similar case discussed here, an entity that was considered to be only a form or variety of one species, T. ovatum Pursh (1803: 216), has turned out to be so genetically distinct that it is not even placed in the same clade as T. ovatum (Farmer 2006, Lampley et al. unpubl.).

Background Robust phylogenies for species with giant genomes and closely related taxa can build evolutionary frameworks for investigating the origin and evolution of these genomic gigantisms. Paris japonica (Melanthiaceae) has the largest genome that has been confirmed in eukaryotes to date; however, its phylogenetic position remains unresolved. As a result, the evolutionary history of the genomic gigantisms in P. japonica remains poorly understood. Results We used next-generation sequencing to generate complete plastomes of P. japonica, P. verticillata, Trillium govanianum, Ypsilandra thibetica and Y. yunnanensis. Together with published plastomes, the infra-familial relationships in Melanthiaceae and infra-generic phylogeny in Paris were investigated, and their divergence times were calculated. The results indicated that the expansion of the ancestral genome of extant Paris and Trillium occurred approximately from 59.16 Mya to 38.21 Mya. The sister relationship between P. japonica and the section Euthyra was recovered, and they diverged around the transition of the Oligocene/Miocene (20 Mya), when the Japan Islands were separated from the continent of Asia. Conclusions The genome size expansion in the most recent common ancestor for Paris and Trillium was most possibly a gradual process that lasted for approximately 20 million years. The divergence of P. japonica (section Kinugasa) and other taxa with thick rhizome may have been triggered by the isolation of the Japan Islands from the continent of Asia. This long-term separation, since the Oligocene/Miocene boundary, would have played an important role in the formation and evolution of the genomic gigantism in P. japonica. Moreover, our results support the taxonomic treatment of Paris as a genus rather than dividing it into three genera, but do not support the recognition of T. govanianum as the separate genus Trillidium. Electronic supplementary material The online version of this article (10.1186/s12870-019-1879-7) contains supplementary material, which is available to authorized users.

Trillium has fascinated the botanical world with its peculiar biology and high medicinal value. The genus appears in North America and Asia, and is widely used for traditional medicine on both continents. Medicinal value has been validated recently through the isolation and ex situ testing of constituent compounds. This study reviews biology, medicinal value, ecological threats and knowledge gaps, and offers suggestions for conservation and sustainable utilization. Findings indicate that the biology of the genus is well explored and the reproductive constraints are well recognized. Steroidal compounds and pharmaceutical potential of the genus are reported, but controlled clinical trials on humans are yet to be conducted. High medicinal value and the peculiar biology have made the genus susceptible to several threats.

Ethnopharmacological relevance The development of a multidrug-resistant (MDR) phenotype is a main obstacle to the successful treatment of breast cancer. Saponins of several herbs are considered as promising candidates for drug resistance treatment. We extracted Paris saponin VII (PS VII) from Trillium tschonoskii Maxim. and investigated whether it could sensitize chemoresistant breast cancer cells MCF-7/ADR to the cytotoxic effects of adriamycin. Materials and methods MCF-7/ADR cells were exposed to 0.5 μM PSVII plus different concentrations of adriamycin (0–100 μM). Then, MTT assay and adriamycin accumulation assay were used to assess cell proliferation and intracellular adriamycin retention. P glycoprotein levels and intracellular rhodamine 123 (Rh-123) accumulations were investigated to measure the expression and activity of P-glycoprotein. A xenograft model of nude mouse was utilized to observe the effect of PSVII in vivo. Results Treatment with PSVII influenced cell viability of MCF-7/ADR cells, as well as sensitized MCF-7/ADR cells to the cytotoxic effects of adriamycin. Moreover, PSVII significantly downregulated MDR1 expression in MCF-7/ADR cells. Intravenous administration of PSVII significantly enhanced anticancer efficacy of adriamycin to MCF-7/ADR xenograft model in nude mice. Conclusion These findings suggested a possible application of PSVII in combination with chemotherapy and/or as neo-adjuvant therapy in the treatment of MDR breast cancer.

—Analysis of molecular phylogenetic data was used to reveal the existence of a novel species of Trillium subgen. Sessilium. Trillium delicatum sp. nov. differs from the two other low-growing species of the subgenus, T. decumbens and T. reliquum, in features such as straight stems and dung-scented flowers. It occurs in wet floodplain forests in the Oconee and Ocmulgee river drainages in central Georgia, and has a distinct floral scent compared to the wide spectrum in Trillium. The molecular phylogenetic results based on four markers (nuclear ribosomal ITS and plastid trnH-psbA, trnL-trnF, and rpl32-trnL spacers) included samples of all species of the subgenus, and documented the distinctiveness of T. delicatum as a distinct taxon and also provided resolution of species relationships that suggested that the subgenus originated in southeastern North America with two migrations to western North America. The molecular results also suggested the need for closer examination of T. cuneatum and T. lancifolium, and also to assess whether T. decipiens and T. underwoodii are distinct species. Based on the few known populations and susceptibility to damage by hogs, Trillium delicatum is a rare species requiring protection.

Phenological overlap with pollinators is crucial for reproductive success in insect-pollinated plants. In this study, we examined whether pollinator visitation successfully occurred during an entire flowering season in two populations of the insect-pollinated spring ephemeral Trillium camschatcense in the Tokachi region of Hokkaido, northern Japan. We bagged flowers and excluded pollinator visitation during either the first or the last half of the entire flowering season to compare pollination success between the two periods. The two populations have experienced differing levels of climate warming in the last 60 years, which impacted pollinator visitation. In the population experiencing temperature rise more rapidly, fertilization rate and seed set decreased sharply when bagged during the first half period, indicating that pollinator visitation is skewed to the early part of the flowering season. The temporal skewness of pollination success would be an early warning signal of the impacts of climate warming on the reproductive success of T. camschatcense.

Trillium govanianum rhizomes are traditionally consumed as a raw powder and decoction for the treatment of health complications. Hence, the present study aimed to investigate whether aqueous and alcoholic extracts of T. govanianum rhizomes under hot and cold extraction conditions have similar or dissimilar chemical, nutrient, and antioxidant profiles. The total phenolics, flavonoids, carbohydrates, proteins, fats, and energy values were estimated in all the conditionally prepared samples. The total phenolics (21.23±1.4 mg GAE/g extract), flavonoids (70.57±3.24 mg RE/g extract) were found higher in hot ethanolic extract (TGHEt), while cold water extract (TGGC) showed higher nutrients including amino acids (10.545±0.219 mg/g) and nucleosides (1.803±0.018 mg/g). The nutrient energy value (2.60 and 2.49 Kcal/g extract) was higher in cold and hot ethanolic extracts. Further, TGHEt scavenged the DPPH. (IC50 ; 870±22 μg/mL) and ABTS.+ (IC50 ; 80±1.49 μg/mL) effectively and proved its highest antioxidant activity compared to other samples. In LC/MS/MS-based metabolite profiling, twenty-six metabolites (fatty acids, steroidal saponins, triterpene saponins, ecdysteroid hormones) were confirmed with mass fragmentation and literature, while one hundred nine metabolites were identified using the METLIN database. The principal component analysis showed clustering of hot condition extracts while cold extracts were differentially located in quadrants. The heatmaps exhibited the associations and differences between metabolite composition, solvents, and extraction conditions. The identified metabolites speculatively predicted the biosynthesis pathway of T. govanianum. Findings also illustrated that T. govanianum is a source of bioactive nutritional components and saponins. The current metabolite profiling of T. govanianum will help in its agricultural and biotechnological interventions for higher quality produce.

The genus Trillium belongs to the family Melanthiaceae (alt. Trilliaceae) and comprises around 50 accepted species. Steroidal saponins are the principal phytomolecules in Trillium. A total of around 83 different steroidal saponins with three structurally different sapogenin skeletons have been reported from the Trillium genus. Steroidal saponins from Trillium have exhibited in vivo neuroprotective properties, and sapogenin portions are structurally similar to neurosteroids. Steroidal sapogenins, such as diosgenin, have been utilized as a raw material for the synthesis of steroidal hormones and bio-active steroidal derivatives in industries. Sapogenins offer the opportunity to design and develop natural neuroactive steroidal derivatives. This book chapter provides an overview of sapogenins and saponins characterized from the genus Trillium and an outline of their spectroscopic assignments. Besides this, in silico study has been carried out to understand the molecular interactions of steroidal sapogenins with the GABAA receptor. The amalgam of computational biology, medicinal synthetic chemistry, and clinical approach can be used for the development of reliable, novel, and potential neuromodulatory agents from Trillium or other steroidal saponins enriched natural sources.

Trillium apetalon (4x) and T. tschonoskii (4x) hybridize commonly where both species grow sympatrically, leading to the formation of tetraploid T. miyabeanum in Hokkaido, Japan. The present study aimed to determine which isolation factor is responsible for the frequency and asymmetry of hybrid T. miyabeanum formation in a sympatric population of T. apetalon and T. tschonoskii. We examined the contributions and strengths of four reproductive isolation barriers of T. miyabeanum formation: flowering phenology, breeding system, genetic isolation, and hybrid inviability. In addition, we also investigated the effect of flowering phenology on reproductive success (i.e., seed production and outcrossing rates) and outputs (i.e., ovule production) for T. apetalon and T. tschonoskii. We calculated the absolute contribution of each isolation barrier to the total reproductive isolation and found that flowering phenology and differences in breeding systems between the two parental species were more effective when T. apetalon was the maternal parent. Furthermore, hybrids with T. apetalon as the maternal parent had lower viability than those of the reciprocal cross and did not reach the flowering stage. Particularly, absolute contribution of premating isolation, especially by flowering phenology and breeding system, was higher than that of other isolation factors for both crossing directions. For the formation of T. miyabeanum, we concluded that asymmetry of hybridization between T. apetalon and T. tschonoskii would be caused by strong premating isolations. The asymmetry of the isolating barriers may promote T. tschonoskii as the maternal parent of T. miyabeanum.

Paris polyphylla var. alba is a medicinal plant commonly used in the southwest of China. This study characterized the complete chloroplast (cp) genome sequence of P. polyphylla var. alba to investigate its phylogenetic relationship in Melanthiaceae. The cp genome of P. polyphylla var. alba is 165,079 base pairs (bp) in length with 36.96% G + C content. The cp genome is divided into (a) large single copy (LSC) (84,393 bp), (b) small single copy (SSC) (16,066 bp), and (c) two inverted regions (32,310 bp). The cp genome contains 135 genes, including 89 protein-coding genes, 38 tRNA genes, and 8 rRNA genes. Phylogenetic analysis indicated that P. polyphylla var. alba is closest to P. polyphylla var. emeiensis, and Paris had a close relationship with Trillium in Melanthiaceae.