b. Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China;
c. Institute of Botany, Northwest Normal University, CN-730070, Lanzhou, Gansu, China
In Chenopodium there are some glandular species, which were first divided into the genus Botrydium Spach(1836), even though that is an illegitimate name(non Wallroth 1815 alga); two species were included into Botrydium: Botrydium aromatium Spach(=Chenopodium botrys Linn.)and Botrydium scharderi Spach(=Chenopodium foetidum Schrad., 1808, nom. Illeg. Non Lam. 1779). Small(1933)later transferred Ch. botrys(Linn.)Small into Botrydium. At the same time, Spach(1836)described the genus Ambrina Spach, also consisting of two species from South and Central America: Ambrina pinnatisecta Spach(=Chenopodium multifidum Linn. and Roubieva multifida(Linn.)Moq.)and A. ambrosioides Spach(=Chenopodium ambrosioides Linn.). However, Botrydium and Ambrina have not been accepted by many taxonomists(e.g. Boisser, 1879; Bunge, 1862, 1880; Standley, 1916; Ulbrich, 1934; Iljin, 1936; Tutin, 1964; Grubov, 1966, Kung and Tsien, 1979, Wilson, 1984; Kühn, 1993; Zhu et al., 2003). Moquin-Tandon(1840, 1849)included Ambrina in his Chenopodium monograph, but transferred Botrydium aromaticum into Ambrina, and placed A. pinnatisecta Spach(nom. illeg.)into the new genus Roubieva Moq. as R. multifida(Linn.)Moq. In short, the circumscription and relationship of these glandular groups remains unclear.
In light of molecular evidence(Kadereit et al., 2003), subfamily Chenopodioideae includes four groups(tribes), Chenopodieae I, Chenopodieae II, Chenopodieae III and Atripliceae. Chenopodium has been shown to be a non-monophyletic group, with different species scattered among Chenopodieae I, Chenopodieae II, and Chenopodieae III. Ch. botrys and Ch. cristatum belong to the glandular groups located in Chenopodieae III, and are related to Dasphania, Teloxys, Axyris and Krascheninnikovia.
In order to clarify the diversification of glandular groups and ascertain the circumscription of some genera, we conducted a morphological observation using scanning electron microscope(SEM). Furthermore, a classification of the glandular group Botrydium(Chenopodium-Botrydium)is presented on the basis of specimens from twenty-one herbaria.
2. Material and methodsScanning electron microscopy was used to observe the surface of the glandular Chemopodioideae(Botrydium, Cycloloma, Roubieva, Ambrina and Dysphania)in order to determine their distinct morphology.
Specimens were examined from the following herbaria: USA ASC, BRY, CAS, COLO, CS, DAV, GH, IDO, MO, NY, RM, TEX, UC, China PE, NWTC, HNWP, LZU, WUK, CDBI and SZ, and Australia CANB(see Fig. 1).
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| Fig. 1 Distribution of Neobotrydium corniculatum (triangles) and Neobotrydium ornithopodum (circles) in China. The provinces: XZ = Xizang, YN = Yunnan, SC = Sichuan, QH = Qinghai, GS = Gansu, NX = Ningxia, SX = Shanxi, HB = Hebei. |
The present study supports the separation of two Eurasian species, C. foetidum and C. botrys, from Chenopodium. These two species possess a series of distinctive characters. Both species are covered with glandular hairs or gland-grains, or both; the shape of the gland-grains are always yellow granulated globular(Fig. 2A-C). Both species have a strong smell, which is suitable for entomophily pollination. Both species have inflorescences that are primitive dichasia(Cronquist, 1981)and seeds that are usually horizontal.
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| Fig. 2 Trichomes of Neobotrydium and related genera (SEM): a) glandular hairs of Neobotrydium botrys; b) gland-grains of Neobotrydium peruense; c) gland-grains of Neobotrydium corniculatum; d) gland-grains of Dysphania plantaginella; e) gland-grains of Cycloloma atriplicifolium; f) gland-grains of Ambrina ambrosioides; g) water-hairs (furfuraceous) of Chenopodium opulifolium; h) gland-grains of Neobotrydium graveolens. |
Chenopodium L., with more than 200 species and C. album L. as its type, have neither gland-grains nor glandular hairs; their trichomes are a kind of single-cell vesicular white water-hairs(furfuraceous)(Fig. 2G). They have glomerule glands which can support entomophily to anemophilous pollination processes. Therefore, based on the characters above, we suggest Botrydium is a valid taxon that includes 20 species which occur in the temperate regions of Eurasia, North Africa, North America, Central America to northwestern South America and Oceania.
Since Botrydium was impropriated by a thalloid alga genus in 1815 before Spach 1836, we resurrect Spach's Botrydium and name Neobotrydium as our new taxon.
With the exception of Neobotydium, there are four genera with glands in Chenopodiaceae that also have distinctive characters: Ambrina glomerule is with gland-grains ellipsoidal(Fig. 2F), its seeds are horizontal, and has only one species, A. ambrosioides(Linn.)Spach, distributed in Central America and southeastern U.S.A.; Cycloloma Moq., has dichasia, its gland-grains are roundedglobular(Fig. 2E), its perianth develops wing-like appendages, and it has one species, Cycloloma atriplicifolium(Sprengel)J.M. Coulter, distributed in U.S.A. to northern Mexico; Roubieva Moq., glomerule its gland-grains are ellipsoidal, the seed is vertical, the type is Roubieva multifidum(Linn.)Moq., and it is distributed only in South America along with Roubieva bonariensis Hook.f. and around 5-7 further species; Dysphania R. Br., glomerule is with gland-grains rounded-globular(Fig. 2D), its c. 10 species are endemic to Australia.
4. Taxonomic treatmentsDiagnostic key to Neobotrydium and related genera
1a. Plants covered with gland-grains or glandular hairs, or both; usually with a specific smell; no single-celled vesicular white water-hairs(furfuraceous)
1b. Plants covered with single-celled vesicular whitewater-hairs(furfuraceous); no gland-grains or glandular hair, no specific smell --------------------------------------------- Chenopodium
2a. Flowers in a dichasium
3a. Perianth dorsal no horizontal wing-shape appendages, sometimes with vertical keel-shape tubercle ---------------- ---------------------------------------------------Neobotrydium
3b. Perianth dorsal with horizontal wing-shape appendage ------------------------------------------- Cycloloma
2b. Flowers in a glomerule
4a. Gland-grains elliptic
5a. Seed vertical; perianth only lobed at apex --------- Roubieva
5b. Seed horizontal; perianth parted near to base ------ Ambrina
4b. Gland-grain rounded-globular ---------------------Dysphania
Neobotrydium Moldenke, Amer. Midl. Naturalist 35: 330(1946) Botrydium Spach, Hist. Veg. Phan. 5: 298, 1836. nom. illeg.
TYPE SPECIES: Neobotrydium botrys(L.)Moldenke [=Chenopodium botrys Linn. =Bortrydium aromaticum Spach and Botrydium botrys(L.)Small].
Annual herbs, covered with glandular hairs, or yellow granulated globular glands or both; with strong odour. Leaves complanate, alternate, margins lobed or parted, or dentate. Dichotomous cymes. Flowers bisexual, perianth nearly globular, 1-1.5mm diameter, 5 or 4-parted, segments ovate to narrow-ovate, with membranaceous narrow margins, sometimes with longitudinal keels; stamens 5, filaments compressed, anther nearly globular. Utricle depressed globular, pericarp membranaceous, adnate to seeds. Seeds usually horizontal, rarely vertical, testa thincrustaceous, red-brown to black, lustrous; embryo half-annular or hippocrepiform, surrounding farinaceous perisperm. Ca. 20 species distributed in Asia, Europe, North Africa, North and Central America, north-west region of South America and Australia.
1. Neobotrydium corniculatum G.L. Chu et M.L. Zhang, sp. nov. Fig. 3
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| Fig. 3 Neobotrydium corniculatum G.L. Chu et M.L. Zhang. 1 habit; 2, 3, 4, 5 leaves; 6 flower; 7 stamen; 8 pistil; 9. cyme; 10 perianth in fruit; 11 utricle; 12 seed; 13 embryo. |
TYPE: G. L. Chu 15618, date of collection, China, Gansu, Diebu Xian, Wangzang, on foot of slopes(Holotype, NWTC).
Herba annua, 30-50 cm alta, obtegens glandulosa, odora. Caulis erectus, sparsim ramosus a basi ad apicem, viridis vel purpureorubellus, costati; rami oblique ascendentes, tenues. Folia alterna, petiolata; circumferentia laminae ovata, 2-5 cmlonga, 1-3 cmlata, prope pinnatisecta; pinnula ovato-oblonga usque ellipica, inaequalis, margine plerumque pinnatifida vel dentata, abaxialis velata globulosis granulosis luteis glandula, adaxialis velata brevis pilis; petiolus 3-15 mm longus, tenuis. Dichasia axillaria et terminalia, 2-5 cm longa, patentes. Flores bisexuales et femnei; perianthium sub globosum, ca 1 mm diam., 5-partitum; segmenta perianthii, leviter sucuulenta, circa 0.9 mm longa, fructificatione dorsaliter prope apice cum 1 corniculatis appendcibus; stamina 5, filamentis linearibus, cum segmentis fere aequilongi, antheris obovatooblongis; stigma 2, filiformis; stylus obscurus. Uticulus: pericarpio membranceo, ad semen adpressum. Semina compresse globulos, horizontalia, 0.7-0.8 mm diam., testa tenuiter crustacea, nigra; embryo hippocrepicus, perispermo farinaceo.
Proximum Neobotrydium schraderium quae differt foliis pinnatifidis et segmentis perianthii fruticatione dorsalitier absque corniculatis appendicibus.
Annual herb, covered with gland-grains and gland-hairs throughout, with strong odor. Stem erect, 30-50 cm tall, sparsely-branched, green, or purple-reddish, ribbed; branches obliquely-ascending, slender. Leaves petiolate, blade ovate in outline, margin irregularly 2- or 1- pinnatisect, 2-5 cm long, 1-3 cm wide; pinnules ovate-oblong to elliptical, usually unequal, margin dentate or pinnatifid, with yellow granulated gland-grains adaxially and short gland-hairs abaxially; petiole 3-15 mm long. Dichotomous cyme axillary and terminal, 2-5 cm long, slightly spreading, flower bisexual and female, perianth compressedglobular, ca. 1 mm wide, 5-parted to near base, segments linearoblong, slightly succulent, ca 0.9 mm long, glandular on back, near apex of back each develops a corniculate appendage in fruit; stamens 5, with filiform filaments, slightly widen out to base, nearly equal in length to segments. Utricle: pericarp membranaceous, adnate to seed. Seed compressed globular, horizontal, 0.7-0.8 mm in diameter, testa thin-crustaceous, black at maturation; embryo annular, perisperm farinaceous. Flowering and fruiting period: AugusteOctober. Its distribution is depicted in Fig. 4.
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| Fig. 4 Neobotrydium ornithopodum G.L. Chu et M.L. Zhang. 1:habit, 2:stem, 3:cyme, 4: flower, 5:stamen, 6:pistil, 7 flower and fruit, 8:utricle, 9:seed, 10:embryo. |
Representative specimens examined
China. Gansu: Lanzhou, Aganzhen, G.L. Chu 179(NWTC); Zhouqu Xian, Jiaoerqiao, alt. 1800 m, field margins, Z.P. Wei 2391(NWTC, WUK); Yongdeng Xian, Liancheng, in woods, alt. 2999 m P.Q. Zhong 10147(NWTC, WUK); Yongdeng Xian, Liancheng, in woods of Betula, alt. 2200-2600m, P.Q. Zhong 10107(PE); Lintan Xian, Yang-Sha, alt. 2600 m, W.Y. Hsia 8619(WUK). Sichuan: Mao Xian, Meadow slopes, Z. He & Z.L. Zhou 13852(PE, SZ); Songpan Xian, Heiheqiao, alt. 1600 m, valley, roadsides, T.P. Wang 7939(PE, WUK); Nanpin Xian, Wandwa areas, meadows, collector unknown 0996(SZ).
2. Neobotrydium ornithopodum G.L. Chu et M.L. Zhang, sp. nov. Fig. 4
TYPE: China, Gansu, Kangle Xian, Dangchuan xiang, Dahebalinchang, Xinfeng, slope thickets, alt. 1900 m, Y.S. Lian et al., 781196(Holotype! NWTC).
Herba annua, 30-40 cm alta, obtegens glandulosa et glandularia, odora. Caulis erectus, ramosis a basi. Folia peliolata; circumferentia laminae elliplica usque oblonga, 3-5 cm longa, 2-2.5 cm lata, apice acuta vel obtusa, basi cuneata, margine pinnafidus; pinnulae inaequalis, leviter obliquae, integeri vel lobatae; petiolus 1-1.5 cm longus. Dichasia axillaris, folia brevior, rhachidibus et ramulis flexuosus ornithopodus, dense glandulis. Flores bisexuales; perianthium subglobosum, 0.7-1mmdiam, plerumque 5-partium; segmenta non apertus, inaequalis, ovata usque oblonga, obtegens glandulosis et longitudine carinae dorso; stamen 5, filamentis linearibus; stylus obscurus; stigmata 2. Utriculus ovatus, pericarpio membranceo, adpresso semen. Semen horizontale, 0.6-0.7 mm diam., testa tenuiter crustacea, nigera in maturitas; embryo hippocrepicus, perispermo farinaceo.
Proximum Neobotrydium schraderianum sed axillaria dichasiis foliis brevioribus; rhachidibus et ramulis flexuosis; floribus denseis, sementis perianthii non apertis, longitudine carinis dorso differt.
Annual herb, 30-40 cm tall, covered with gland-hairs throughout the plant; with strong odour. Stem usually branched from base, erect or slightly ascending. Leaves with petioles; blades elliptical to oblong in outline, 3-5 cm long, 2-2.5 cm wide, apex acute or obtuse, base cuneate, margin pinnatifid, pinnules unequal, margins entire; petiole 1-1.5 cm long. Dichasial cymes axillary, usually shorter than leaves, with dense non-patent flowers; rhachis and ramuli slightly flexuose like a bird-claw, densely covered with gland-hairs; flower bisexual, perianth nearly globular, 0.6-1 mm diam., usually 5-parted, segments unequal, ovate-oblong, usually longitudinally keeled and with short gland-hairs on back, not opening; style very short, stigma 2, filiform. Utricle with membranaceous pericarp, slightly brownish, adnate to seed. Seed horizontal, depressed, 0.6-0.7 mm diam., around margin obtuse, testa thin-crustaceous, black at maturation, slightly lustrous. Flowering and fruiting period: AugusteOctober. Distribution as in Fig. 4.
Representative specimens examined
China: Gansu, Diebu Xian Dianga South Mountain, alt. 2360 m, roadsides, Z.X. Peng 437(LZU); Yuzhong Xian, Xinlong Shan, river marshes, alt. 2100 m, S.R. Wang 120(LZU); Kangle Xian, Lulian, sunny mountain slopes, alt. 2100 m, Y.L. Liu et al., 76350(NWTC); Xiahe Xian, Daba valley, Amquhu, meadows, alt. 2850 m, Gansu Public Health Bureau 206(NWTC); Xiahe Xian, Qingshui, K.J. Fu 1101, alt. 2400 m(WUK).
Hebei: Yi Xian, Longya Mountain, X.Y. Liu 15907(NWTC); Wei Xian, Zhuolu Xian, Hsiaowutaishan, alt. 1550 m, X.W. Kung 1174(WUK); Cholu Xian, Ling Shan Kou, alt. 1700 m, velley, W.Y. Hsia 2448(WUK). Shanxi: Ningwu Xian, slope meadows, Loess Plateau Exp. Team 3151(WUK). Ningxia: Guyuan Xian, slope meadows, alt. 1700 m, J.X. Yang 5475(WUK). Qinghai: Angqian Xian, dunes of river marshes bank, alt. 3600 m, Y.C. Yang 01538(WUK); Gonghe Xian, Heimahexiang, side of Qinghai Lake, alt. 3200 m, Y.H. Wu 21745(HNWP); Guinan Xian, Senduoxiang, alt. 3390 m, Y.H. Wu 26842, 26860(HNWP), Taxiuxiang, alt. 3300 m, (HNWP); Zaduo Xian, suluxiang, terraces of riverbank, alt. 4000 m, S.W. Liu 403(WUK); Qinghai Lake, alt. 1750 m, K.M. Liou 5923(WUK); Mengyuan Xian, Xianmisi, river marshes, P.Q. Zhong 10062(WUK); Ledu Xian, Caotai forest-centre, Z.H. Wu & Y.S. Chen 018(WUK). Sichuan: Batang Xian, Yahuangqu, alt. 3200-3400 m, Xigaosuo Zangkao Team 2940(HNWP); Maerkang Xian, hill top, alt. 3800 m, X. Li 72350(NWTC, SZ), alt. 3250 m, X. Li 72027(NWTC, SZ), K.J. Fu 1197(NWNU), alt. 2650 m, Z.Y. Zhang 53307(SZ), Sayangxiang, alt. 2960 m, Xigaosuo Guole Team 761(HNWP), Nanshan, Xiguosuo Zangyao Team 943(HNWP); Litang Xian, Wenqui, alt. 4000 m, collector unknown(WUK); Jiulong Xian, Tangu, roadsides, alt. 3200 m, Q.Q. Wang 21120(CDBI), alt. 3580 m, Y.G. Liu 1273(CDBI). Xizang: Dingqin Xian, meadoes, alt. 3850m, Z.J. Zhao 83346(SZ); Bomi Xian, Yigong-Gongqu, alt. 2300 m, Qingzang Expl. Team 408(HNWP); Leiwuqi Xian, Sangduoxiang, alt. 3900 m, Qingzang Expl. Team 2729(HNWP); Linzi Xian, near Rongchang, alt. 2600 m, Qingzang Expl. Team 3498(HNWP); Yadong Xian, near Renqinggang, alt. 2800m, Qingzang Expl. Team 761(HNWP). Yunnan: Deqin Xian, Adunzi, slopes, alt. 3700 m, Q.W. Wang 70221(WUK).
3. Neobotrydium tibeticum(A. R. Li)M.L. Zhang et G.L. Chu, nov. comb.
Chenopodium tibeticum A. R. Li in Fl. Xizang 1: 638. f. 203. 1982. Distribution: Asia.
4. Neobotrydium botrys(L.)Moldenke, Amer. Midl. Naturalist 35: 330(1946)
Botrydium botrys(L.)Small in Man. Southeast. Fl. 466.1933. Botrydium aromaticum Spach, Hist. Veg. Phan. 5: 299. 1836. Ambrina botrys(L.)Moq. Chenop. Enum. 37. 1840. Chenopodium botrys Linn. Sp. Pl. 219. 1753; Reichb. Ic. Germ. 34: t.250. f. 1-9.1909; Standley in N. Am. Fl. 21(1): 26. 1916. Iljin in Fl. URSS 6: 46.1936; Grubov, Pl. Asiae Centr. 2: 19. 1966. Kung et al. in Fl. Reip. Popul. Sin. 25(2): 81. Dysphana botrys(Linn.)Mosyakin & Clemants, in Ukrayins’K. Bot Zhurn., n.s. 59:383. 2002; Fl. North Am. 4:273. 2003; 5:377.2003.
C. E. Europe, Mediterranean region, C. W. Asia. Turkey, Cyprus; Naturalized in North and South America.
Distribution: Eurasia.
5. Neobotrydium foetidum(Schrad.)M.L. Zhang et G.L. Chu, nov. comb.
C. foetidum Schrad. Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 2: 79. 1808; C. foetidum f. pumilum Kurtz in R.E. Fr.; Chenopodium graveolens Willd. f. pumilum(Kurtz ex R.E. Fr.)Aellen Verh. Naturf. Ges. Basel 41: 108. 1931.
Distribution: N. SW. China, Europe, C. America.
6. Neobotrydium schraderianum
Neobotrydium schraderianum(Roem. et Schult.)M.L. Zhang et G.L. Chu, nov. comb.
Chenopodium schraderianum Schult. in Roem. et Schult. Syst. Veg. Ed. 6:260.1820. Dysphania schraderiana(Schult.)Mosyakin & Clemants, Ukrayins’K. Bot. Zhurn.59: 383.2002; Fl. China 5:3 77.2003.
Distribution: N. Somalia, Tanzania, Sandi Arabia, Kenya, Aethiopiae, Uganda, Ethiopia, Burundi to S. Africa.
7. Neobotrydium incisum(Poir.)M.L. Zhang et G.L. Chu, nov. comb.
Ambrina incisa(Poir.)Moq. Chenop. Enum 36. 1840.
Chenopodium incisum Poir. in Lam. Encyc. Suppl. 1: 392. 1810.
Distribution: Southwestern U.S.A.
8. Neobotrydium dissectum(Moq.)M.L. Zhang et G.L. Chu, nov. comb.
Ambrina dissecta Moq. Chenop. Enum. 38. 1840.
Chenopodium bipinnatifidum Moric.ex Moq. in DC. Prodr. 13(2): 76. 1849.
C. dissectum(Moq.)Standley in N. Am. Fl. 21(1): 26. 1916.
Distribution: Coahuila to central Mexico.
9. Neobotrydium ambrosioides(L.)M.L. Zhang et G.L. Chu, nov. comb.
C. ambrosioides L. Sp. Pl. 1: 219.1753; Dysphania ambrosioides(L.)Mosyakin & Clemants Ukrayins'k. Bot. Zhurn. 59(4): 382(2002); Teloxys ambrosioides(L.)W.A.Weber Phytologia 58(7): 477(1985); Blitum ambrosioides(L.)Beck Icon. Fl. Germ. Helv.(H.G.L. Reichenbach)24: 118. 1908; Botrys ambrosioides(L.)Nieuwl. Amer. Midl. Naturalist 3: 275.1914; B. ambrosioides(L.)Beck Icon. Fl. Germ. Helv.(H.G.L. Reichenbach)24: 118. 1908.
Distribution: Central America.
10. Neobotrydium procerum(Hochst. ex Moq.)M.L. Zhang et G.L. Chu, nov. comb.
Chenopodium procerum Hochst. ex Moq. in DC., Prodr. 13(2): 75(1849).
Distribution: Northern Somalia.
11. Neobotrydium pusillum(Hook. f.)M.L. Zhang et G.L. Chu, nov. comb.
Chenopodium pusillum Hook. f. Handbk N. Z. Fl. 231. 1864. Distribution: New Zealand.
12. Neobotrydium graveolens(Lag. et Rodr.)M.L. Zhang et G.L. Chu, nov. comb. C. graveolens Lag. et Rodr. Anal. C. Nat 5: 70. 1802. C. incisum Poir(1810) Encycl.(Lamarck)Suppl. 1. 392. 1810.
Dysphania graveolens(Willd.)Mosy. & Clem. in Ukrayins’K. Bot Zhurn., n. s. 59:383.2002; Fl. North Am. 4:273. 2003. comb. illeg.
Distribution: North America to Mexico and Bolivia.
13. Neobotrydium pumilio(R. Br.)M.L. Zhang et G.L. Chu, nov. comb.
Ambrina pumilio(R. Br.)Moq., Chenop. Monogr. Enum. 42. 1840.
Chenopodium pumilio R. Br. Prodr. 407.1810.
Dysphania pumilio(R. Br.)Mosy. & Clem.in Ukrayins’K. Bot Zhurn., n. s. 59:382. 2002.
Distribution: Southern Australia.
14. Neobotrydium carinatum(R. Br.)M.L. Zhang et G.L. Chu, nov. comb.
Ambrina carinata(R. Br.)Moq. Monog. Chenop. Enum. 38. 1840.
Chenopodium carinatum R. Br., Prodr. 407. 1810.
Dysphania carinata(R. Br.)Mosy. & Clem.Ukrayins’K. Bot Zhurn., n. s. 59:382. 2002.
Distribution: eastern Australia.
15. Neobotrydium melanocarpum(J.M. Black)M.L. Zhang et G.L. Chu, nov. comb.
C. carinatum var. melanocarpum J.M. Black, Trans et Proc. Roy. Soc. S. Australia 46. 566. 1922.
Distribution: Australia.
16. Neobotrydium saxatile(Paul G. Wilson)M.L. Zhang et G.L. Chu, nov. comb.
Chenopodium saxatile Paul G. Wilson Nuytsia 4: 179.1983. Distribution: Australia.
17. Neobotrydium peruense G.L. Chu & M.L. Zhang sp. nov. Fig. 5 TYPE: Peru, Prov. Lambayeque, Entre Chiclayo Procedencia, Lambayeque, Borde de Carretera, alt. 30 m, 18 Feb. 1982, S. Llatas Q841(holotype MO). S. America.
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| Fig. 5 Neobotrydium ornithopodum G.L. Chu et M.L. Zhang. |
Herba annua, 15-20 cm alta, obtegens glandulae, odorata. Caulis erectus or ascendens. Folia complanata, alterna, petiolata; laminae ovato-elliptica, 2-4 cm longa, 3-10 mm lata, margine pinnatilobata; lobuli repandi, obtusi; petiolus 2-7 mm longus. Dichasia axillaria, abbreviata, ca. 1 cm longa, patentia, breviora folio. Flores bisexuales; perianthium sub globosum, ca 1 mm diam., 4-partium; antheris ca. 0.2 mm longa Uticulus, pericarpio membranceo, adpresso semen. Semen horizontale compresse globulosum, ca. 0.5mmdiam., testis tenuiter crustaceis rubri-brunneis lucenstibus; embryo hippocrepicus, perispermo farinaceous.
Proximum Botrydium graveolenti Lag. sed perianthiis plerumque 4-partis; axillaria dichasiis abbreviates, foliis brevioribus et non sterilibus terminalibus ramulis differt.
Annual herbs, 15-20 cm high, covered with gland-grains, odoriferous. Stem erect or ascending. Leaves complanate, alternate, petiolate; blade ovate-elliptical, 2-4 cm long, 3-10 mmwide, with pinnatilobate margin, l lobes obtuse, repand; petiole 2-7 mm long. Dichasium axillary, shorter than leaves, ca. 1 cm long; flowers bisexual, perianth nearly globular, ca. 1 mm in diameter, usually 4- parted, densely covered with gland-grains; anther ca. 0.2 mm long. Utricle: pericarp membranaceous. Seeds horizontal, compressed globular, ca. 0.5 mm diam. testa thinly crustaceous, red-brown at maturity, smooth, glossy; embryo semi-annular, perisperm farinaceous.
Representative specimens examined
Peru: Prov. Pacasmayo, Dpto, La Libertad, 10 Jun. 1983, I. Sanchez V.3089(MO).
Distribution: Peru.
18. Neobotrydium cristatum(F.Muell.)M.L.Zhang & G.L.Chu. comb. nov.
Blitum cristatum F. Muell. Trans. & Proc. Philos Inst. Victoria 2: 73. 1858.
Chenopodium cristatum(F.Muell.)F.Muell., Fragm. 7:11. 1869.
Distribution: Australia.
19. Neobotrydium truncatum(Paul G. Wilson)M.L. Zhang et G.L. Chu, comb. nov.
Chenopodium truncatum Paul G. Wilson Nuytsia 4: 177. 1983.
Distribution: Australia.
20. Neobotrydium burundiense G.L. Chu et M.L. Zhang &, sp. nov.
TYPE: Burundi, Zaire(Belgian Congo), alt. 5500 ft, Idjw, Is. Lake Kivu, 24 Feb. 1939, M. V. Loveridge 587(Holotype MO).
Herba annua, obtegens glandulae, odorata. Caulis erectus, sparsim. ramosus. Folia complanata, alterna, viridis petiolata; laminae oblonge-lanceolata, 3-4 cm longa, 1-1.5 cm lata, margine lobata; lobulis dentatis, obtusis; petiolus 2-7 mm longus. Dichasia axillaria et terminalia, abbreviata, breviora folio. Flores bisexuales; perianthium oblongum, ca 1.2 mm diam., plerumque 4-partium ad basim, segmentis rotundis vel ovatis, inaequalibus, leviter cucullatis ad apicem; antheris ca. 0.3 mm longa Uticulus, pericarpio membranceo, Semen verticalia compresse-ovoideum, ca. 1 mm diam., testis tenuiter crustaceis, denigratis lucentibus; embryo hippocrepicus, perispermo farinaceous.
Proximum Botrydium schraderianum Schult., Neobotrydium schraderianum sed perianthiis plerumque 4-partis; Semene verticali differt.
Annual herb, covered with glands throughout. Stem erect, sparsely branched, green, leaves complanate, petiolate, blade oblong-lanceolate in outline, 3-4 cm long, 1-1.5 cm wide margin lobed, lobes dentate. Dichasium axillary and terminal, shorter than leaves. Flower bisexual, perianth oblong, ca. 1.2 mm in diameter, usually 4-parted to near base, segments rotund to ovate, unequal in size, apex slightly cucullate; anther ca. 0.3 mm long. Utricle: pericarp succulent rotund to ovate, with testa thinly crustaceous, black at maturity; embryo nearly annular, perisperm, farinaceous.
Representative specimens examined
Burundi: Prov. Muramwya, Bugarama, alt. 2080 m, 28 April 1982, M. Reekmans 11051(MO); Prov. Gitega, jachere postculturale, alt. 2000 m, 13 May 1982, M. Reekamans 11227(MO).
Distribution: North Africa.
AcknowledgmentsWe thank curators of the following herbaria for checking their collections: ASC, BRY, CAS, COLO, CS, DAV, GH, IDO, MO, NY, RM, TEX, UC, US, PE, NWTC, HNWP, LZU, WUK, CDBI, SZ; Dr. Howard C. Stutz at Brigham Young University, and Dr. Stewart C. Sanderson at USDA Forest Service for their exceptional assistance, Dr. Paul G. Wilson at Western Australian Herbarium for his valuable taxonomical suggestion to manuscript, and Dr. David M. Williams at Natural History Museum London for critical comments and improving the English manuscript. We also thank Mr. Bo-yi Peng for assistance with SEM and Mr. Jian-lu Bai for drawings of Plates 2-3. This study is financially supported by Biodiversity Conservation Program(ZSSD-012)of Chinese Academy of Sciences, and China National Key Basic Research Program(2014CB954201).
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