Montanoa leucantha ssp. arborescens
- Dataset
- Compositae
- Rank
- SUBSPECIES
Classification
- family
- Compositae
- genus
- Montanoa
- species
- Montanoa leucantha
general
Taxonomy
[synonomy is available at the end of this section] [available protologues are attached as images; not all protologues are available at this time]
Tribe Gymnarrheneae Panero & VA Funk (subfamily Gymnarrhenoideae Panero & VA Funk).
Gymnarrhena Desfontaines, Mémoires du Museum d’histoire naturelle. Paris, 4: 1. (1818), T.: G. micrantha Desfontaines. Currently this genus has only one species.
Type: Gymnarrhena micrantha Desfontaines, Memoires du Museum d’histoire naturelle, Paris. iv (1818) 1. t. 1
Synonym
Gymnarrhena balansae Coss. & Durieu ex Coss. & Kralik, Bulletin de la Societe Botanique de France 4: 180. (1857) [currently in synonomy with G. micrantha Desf.]
Annual amphicarpic herbs with a prostrate rosette, no more than a few cm high; no reports of milky sap. Leaves simple, forming a dense rosette, sessile, smooth, narrowly lanceolate to narrowly ovate, apex narrowly acute to attenuate, margins denticulate, base truncate, surfaces smooth and glabrous. Subterranean heads homogamous, female, cleistogamous, surrounded by the leaf bases; florets enclosed in involucral bracts, corolla vestigial; achenes relatively large, laterally flattened, blackish, sparsely hairy, remaining below the soil surface on the dead parent plant; pappus absent, vestigial, or of short, basally flattened, somewhat scale-like bristles. Aerial heads congested in the center of the leaf rosette, heterogamous, disciform; involucral bracts imbricate in several series, chartaceous, whitish, acute; receptacle convex, marginally bristly; with a few rows of chartaceous phyllaries. Functionally staminate florets in small groups, loosely connected on very short pedicels, interspersed among the small pistillate florets, corollas small 3-4 lobed, whitish; stamens 3-4, anthers calcarate, ecaudate, without apical appendage. Pistillate florets solitary, each enclosed in a prominent, stiff, white and green bract; corolla filiform, style arms long, with rounded apices. Achenes of functionally staminate florets vestigial, pappus of a few irregularly lacerate scales or absent; achenes of pistilate florets numerous, tiny, ovoid, ciliate, villous, with long twin hairs, cell walls thin, pappus of long-lanceolate, ciliate, acutely acuminate scales.
A second species G. balansae Cross. & Durieu was based on two collections from near the coast in Tunisia and Algeria and they appear to be somewhat different from the collections from the Middle East and further inland in Algeria. However, a more detailed study will have to be made to determine the validity of this taxon.
Gymnarrhena has been collected from North Africa to the Middle East. It is a winter annual and its flowering depends on the rain but usually takes place in March and April (May). According to Gutterman (1989) the life span of the plants is around 67 days out-of-doors (full sun) where it produces both types of heads and 140 days in the greenhouse (8 hours of light) where it produced only aerial heads.
Remaining Synonomy
Cryptadia Lindley ex Endlicher, Genera Plantarum secundum ordines naturales…. By Stephan L. Endlicher. Suppl. 1: 1381. Feb-Mar 1841. T.: non designatus
Cryptadia euphratensis Lindley in Chesney, Narrative of the Euphrates Expedition carried on by Order of the British Government during the years 1835-1837. (1868) t. ad p. 441.
Frankia Steudel ex Steudel, Nomenclator botanicus: numerans ordine alphabetico nomina atque synonyma, tum generica tum specifica, et a Linnaeo et recentioribus de re botanica scriptoribus plantis phanerogamis imposita /auctore Ernesto Steudel. ed. 2. 1: 646, (1840); nom. inval.
Frankia schimperi Steud. Nomencl. Bot. [Steudel], ed. 2. 1: 646, (1840); nom. inval. in syn.
general
Verbesina atriplicifolia Pers., Syn. PI. 2: 472. 1807. Type: Hort.ltal. (holotype, P?; non-flowering isotype, FI!). Persoon often used Jussieu's herbarium (Stafleu, 1967) which is now at Paris. No possible holotypes were found in the loan material from P and Persoon's descliption is cryptic. It would not be possible to place this species except that lussieu and Desfontaines published a better descIiption (Colla, 1824), synonymy, and a drawing, and attributed the species to Persoon. The drawing is somewhat questionable, however, as the rays are too blunt for Montanoa, and they have well-developed stigmas. Although there is still some doubt, the evidence is sufficient to warrant recognition of this name. Eriocoma atriplicifolia(Pers.) Kuntze, Rev. Gen. PI. 1: 336. 1891.
Clerodendron phlomoides Juss. & Desf. in Lamarck, Hort. Ital. 8: 460. 1808, nom. nnd. pro syn.
Galinsoga discolor Spreng., Nov. Provo 19. 1819. Type: Hort. Ital., G. B. Balbis s.n. (holotype, Pi; isotype, G-DC; photo of holotype, US!; photo of G-DC isotype, GH! US!).
Montanoa dumicola Klatt in Durand & Pittier, Bull. Soc. Roy. Bot. Belgique (Brussels) 31: 200. 1882. Type: COSTA RICA, San Jose, hedgerow on the "llanos de Alajuelila," 1100 m, I Dec 1889, H. Pillier 1454 (holotype, GH! isotypes, BR[2J! G! US!).
Montanoa pauciflora Klatt, Leopoldina 23: 90. 1887. Type: "Middle America ... in Herb. Klatt." (holotype not located). The most likely place for this specimen is GH. However, the types as well as the general collections of Montanoa have been searched to no avail. Coreopsis trilobata Vahl ex Klatt, Leopoldina 23: 90. 1887, nom. nud. pro syn. This is listed as the type of M. pallciflora Klatt in the type description. The types as well as the general collections at GH have been checked for this specimen (in litt., M. Calloso) but it has not been located. The possibility exists that the specimen will be located at a later date; a neotype has not been selected.
Montanoa schotii Robins. & Greenm., Proc. Amer. Acad. Arts 34: SIS. IS99. Type: MEXICO,Yucatan, on the road between Merida and Sisal, 24 Oct IS65, A. Schall 913 (holotype, Pi)."
REPRESENTATIVE SPECIMENS: BELIZE. CAYO: EI Cayo, 12 Feb 1931, Bartlett 11421 (F, GH, MICH,MO, NA, US); Bejuco, 15 Feb 938, Gentle 2169 (F, K, MICH, US); 5 mi N of Blancaneau Lodge,12 mi S of Georgeville, 24 Jan 1974, Liesner &: Dwyer 1630 (OS); 3 mi S of Grano de Oro, 2 Jun 1973, Gentry 7761 (MO, OS); Georgeville to Augustine rd at mile 8, 3 May 1969, Proctor 30268 (BM, LL); 9 mi S of the Western hwy on rd to Augustine, 28 Jan 1970, Wilbllr &: Weaver 11491 (DUKE); Macal River 1 mi past Guacamallo Bridge, rd to Millionmio, Dwyer &: Liesner 12355 (MO); S of San Luis and E of Camp Six, 19 Mar 1967, Dwyer et al. 377A (LL, MO); Cave Branch Section, Humming Bird hwy, 18 Feb 1956, Gentle 9029 (LL). COSTA RICA. SAN JosE: San Pedro, E of San Jose, 25 Jan 1922, Green11lan &: Greell1/lall 5315 (GH, MO); San Jose, 18 Jan 1916, Holway 444 (GH). EL SAL VADOR. AHUACHAPAN: near Ahuachapan, 9-27 Jan 1922, Sa 20018 (GH, NY, US); vicinity of Ahuachapan, 16-25 Jan 1947, Standley & Padilla 2610 (F). LA LIBERTAD: stream near Colon, 21 Jan 1949, Williams & Molina 15222 (F). SANTA ANNA: just SE of pueblo of Coatepeqlle along old free rd to San Salvador from Santa Anna, 30 Nov 1978, FL 2922* (OS); 3 km SE of Santa Anna, 18 Nov 1963, Porter 1285 (DS, GH, MEXU). SONSONATE: Finca Chilata, 26-27 Dec 1921, Sa 19229 (GH, NY, US). GUATEMALA. AMITITLAN: Moran, 22 Dec 1916, Holway 625 (US). CHIMALTENANGO: above Las Calderas, 15 Dec 1938, Sa 60086 (G); Quisache, 5-6 Jan 1939, Sa 62009 (F). CHIQUIMULA: btw Chiqllimula and La Laguna, 27 Oct 1939, Se 30702 (F); hills near Ipala, 23 Oct 1939, Se 30355 (F); 2 km from village of EsquiPlilas, 6 Dec 1969, Molina & Molina 25168 (F, LL). GUATEMALA: Santa Catarina Pinula, just S of Guatemala City, 7 Dec 1977, FRa 2638 (OS); SE of Guatemala City on rd to San Salvador, 7 Dec 1977, FRa 2645* (OS). JALAPA: N of Jalapa on rd to Volcan jumay, 8 Dec 1977, FRa 2653 (OS). JUTIAPA: near EI Molina, 26 Nov 1940, Sa 78459 (Fl. PETEN: Tikal National Park, Bajo del Hormiguero, ca. 12 km on old Remate rd, 28 Jan 1961, Contreras 1887 (LL); Tikal, top of Temple IV, 5 Feb 1959, Lundell 15366 (LL); La Liberlad, 16 Mar 1935, Aguilar 476 (LL, MICH, MO, US); EI Paso, 25 Apr 1932, Lundell 1537 (MICH, US); Sacpuy, 10 km S of Flores, 28 Jan 1971, Ortiz 1555 (BM, F, MICH, US); Melchor de Mencos rd, blw kms 35-36, 5 Feb 1966, Contreras 5476 (ENCB, F, MICH, US); rd btw San Andres and Santa Elena, [0 Jan 1970, Ortiz 548(ENCB, F, MO, NY). PROGRESSO: along railroad btw Barranquillo and Cruz, 30 Jan 1942, Se 43328 (F); btw Zanarate and EI Chato bridge, 28 Nov 1969, Molina & Molina 24920 (MO, NY, US). SANTA ROSA: SE of Barberena, 7 Dec 1977, FRa 2646 (OS); SE of Barberena, 7 Dec 1977, FRa 2647 (OS); near EI Molina, 20 Dec 1936, Sa 60472 (F); E of Cui1apa, 25 Nov 1940, Sa 78257 (F); near Cuilapilla, 23 Nov 1940, Sa 78117 (F); Casillas, Nov 1892, Heyde & Lux 4242 (F, G, K, M, MO, NDG, NY). SACATEPEQUEZ: ca. 19 km N of Escuintla, 23 Jan 1977, King 7184 (MO, US); barranco above Duenas, 21 Jan 1939, Sa 63203 (F); near Tejar, 14 Dec 1938, Sa 59863 (F). HONDURAS. COMAYAGUA: near Comayagua, 12-13 Mar 1947, Siandley & Chacon 5935 (F); upstream from main vado in EI Zamorano, 2 Dec 1978, FL 2932 (OS); near Esquela Agricola Panamericana on rd to Giiinope, 4 Dec 1978, FL 2939 (OS); EI Quebracho, 29 Nov 1946, Sa 368 (F); Poza del Tacuazfn, 13 Dec 1947, Molina 671 (BH, F, GH, MO, US); near EI Zamorano, 26 Nov-9 Jan 1947, Sa 1495 (F); rd btw EI Jicarito and EI Pedregal, 13 Nov 1948, Sa 14478 (F); Rio Yeguare at Giiinope, Nov 1943, Rodriguez 1568 (F).OLANCHO: btw Catacamas and EI Hatillo along Rio de Catacamas, 27 Mar 1949, So 18866 (F); near Juticalpa, 5-16 Mar 1949, So 17409 (MICH); rd to San Francisco La Paz, 20 Nov 1963, Molilla 13309 (F, LL, NY, US). EL PARAISO: NE of Danli, 3 Dec 1978, FL 2934* (OS); ca. 1 mi N of EI Paraiso on rd to Danli, 3 Dec 1978, FL 2937 (OS). SANTA BARBARA: San Pedro Sula, Feb 1888, Thieme 5324 (F, G, GH, K, M, MO, NY, US). MEXICO: CAMPECi-IE: Campeche/Yucatan state line on hwy 261, 11 Nov 1975, Dwyer 334 (CA); 17 mi E of Xpuii1, 29 Nov 1977, FRo 2593* (OS); near Escorsega, 16 Jan 1966, Maluda 37483 (F, MEXU); km 5 on the rd from Escarcega to Candelaria, 27 Dec 1965, Herndndez el al. ES-229 (MEXU); Tuxpena, 22 Dec 1931, LUlldell 1093 (ARIZ, DS, F, GH, MICH, MO, S, UC, US). CHIAPAS: Piedra de Huixt1a, 20 Jan 1949, Maluda 18603 (F, MEXU); rd from Toliman to Niquivil near Ojo de Agua, 14 Dec 1976, Breedlove 42607 (DS); jct of Rio Perlas and Rio Jatate at San Quintin near Laguna Miramar, 15 Mar 1955, Sohns 1589 (MICH, US). QUINTANA Roo: Coba, 2 Mar 1976, Morella 535 (MEXU); Chichankanab, s.d., Gaulller 1479 (F, G, GH, K, MEXU, NY). YUCATAN: VaUadolid, 2 Jul 1932, Sleere 1688 (LL, NA, MICH); Peto, 26-27 Jul 1932, Steere 2291 (MICH); Chichen ltza, 23 Jun 1932, Steere 1488 (MICH, US); Calotmul, s.d., Gaumer 2108 (B, BM, C, F, GH, MO, NY, S); Izamal, s.d., GaulIler 2499 (F); Temax, s.d., Gaumer 2357 (BR, DS, MICH, UC, US). NICARAGUA. ESTELl: 3-7 km NW of Pueblo Nuevo, 24 Nov 1973, Williams & Molina 42393 (F). MADRIZ: mt. above Somoto, 15 Nov 1946, Williams & Molilla 10943 (F, GH,MICH, MO).
.general
Uhdea bipinnatijida Kunth, Ind. Sem. Hort. Berol. 1847: 17. 1847. In this description the reference Verh. Vereins Beford. Gartenbaues Konigl. Preuss. Staaten, 1847, was cited, however a search of this journal revealed that the description was published in 1849. According to K. Koch (1864) this species was discovered by the Prussian consul Uhde at "Matmameros" in Mexico and was brought to the Botanic Garden in Berlin 1845 from where it was widely spread, Uhde S.n. The holotype has not been located, neotype selected by Morley (1980) is as follows: Herb. Schultz Bip. S.n. ex Hort. 25.ii.1864 (P!).
Polymnia grandis Kunth, Ind. Sem. Hort. Berol. 1847: [3. 1847, nom. nud pro syn. Kunth listed this name in synonymy but no description can be found. According to K. Koch (1864) this was a name under which the plant was sent to the Botanic Garden at Berlin and after it bloomed Kunth assigned it to Montalloa.
Montanoa heracleifolia Brongniart ex Groenland, Ann. Rev. Hart. 544. 1857, nom. nud. Based upon the following collection: MEXICO: seeds collected in 1843 and harvested in Hart. Bot. Paris, Dec 1845, A. B. Ohiesbrecht S.II. (P[2l!). Montagnaea heracleifolia Brongniart ex Andre, Ann. Rev. Hort. 369. 1863.j: 38. Type:' MEXICO, the description is probably based on the same material as that for Montanoa heracleifolia Brongniart ex Groenland but this is not so stated in the protologue (illustration is taken as the holotype; tracing of illustration, GH!). Morley (1980) selected a neotype but this is unnecessary because elements of the original material are available (illustration) and the neotype is therefore rejected.
Montanoa elegalls K. Koch, Wochenschr. Vereines Beford. Konigl. Preuss. Staaten 7: 408. 1864. Type: unknown. In the protologue Koch states that there is no information on how the plant came to Europe and that it has never flowered. The description of the leaf type is sufficient for it to be assigned to this taxon. Eriocoma elegalls (Koch) Kuntze, Rev. Gen. PI. 1: 336. 1891.
Montanoa pyramidata Schultz Bip. in K. Koch, Wochenschr. Vereines Beford. Gartenbaues Konigl. Preuss. Staaten 7: 408. [864. Type: MEXICO, Jalisco, Guadalajara, [853, D. Oliva s.n. (lectotype [Morley, 1980], P). Two collections are mentioned in the protologue. One is the lectotype and the other is A. Aschenborn s.n., which was probably at B (destroyed inWWII; in litt., H. W. Lack). Eriocoma pyramidata (Schultz Bip. in K. Koch) Kuntze, Rev. Gen. PI. 1: 336. 1891."
REPRESENTATIVE SPECIMENS: MEXICO. COLIMA: 20-30 mi NE of Colima, 26 Dec 1958, Thompson & Fields 355 (MSC, TEX); rd btw Tonila and Colima, 12-13 Nov 1971, Dieterle 4175 (ENCB, TEX). DURANGO: Tamazula, La Bajada, Nov 1921, Ortega 4451 (US). GUERRERO: 4.4 mi SE of Taxeo, 14 Nov 1978, FRi 2825 (OS); 8 km SW of Taxeo, 22 Dec 1967, Rzedowski 25264 (DS, ENCB, MICH, MSC); 12 km N of Iguala, 2 Dec 1966, Rzedowski 23515 (DS, ENCB, LL, MSC); btw Iguala and Teloloapan, 16 Dec 1963, Porter 1364 (DS, GH, MEXU); Aehotla, s.d., Reko 4940 (GH, US); rd from Carrizal to Chiehihualco, 5.6 mi SE of jet of rd to Fila de Caballo, 15 Nov 1978, F 2836 (OS); rd to Fila de Caballo, 20 mi W of jet with hwy 95, 13 Nov 1977, FG 2320 (OS); Aeahuizotla, 14 Dec 1963, Porter 1324 (GH, MEXU); [4.0-14.5 mi N of Tiera Colorada, 10 Dec 1966, AL 4507 (MICH, OS). JALISCO: 16 mi NW of Ameea, [ Nov 1970, Breedlove 18665 (MICH); 3-5 mi N of La Cuesta on rd to Talpa de Allende, 20-21 Nov 1960, M 21255 (DUKE, ENCB, MICH); 6-10 mi SW of Talpa de Allende, 22 Nov 1952, M 14344 (MICH, SMU); San Sebastian, Arroyo, 15 Jan 1927, Mexia 1487 (BM, DS, F, G, GR, MICH, MO, NA, NY, UC, US); Guadalajara, Jul-Oet 1886, Pallller 492 (BM, G, K, MEXU, MO, NDG, US); 5 mi S of Guadalajara (cultivated, double-rayed), 5 Nov 1977, F 2272* (OS); 8.5 mi N of Joeotepee, 6 Nov 1977, F 2282* (OS); Wend of Lake Chapala, 8 Nov 1959, McValigh & Koelz 367 (DUKE, ENCB, LL, MICH, NY); ca. 15 km SSE of Aeatlan de Juarez, 7- 8 Nov 1959, McValigh & Koelz 309 (ENCB, MICH); 1.8 mi S of jet of hwy 80 and rd from Union de Tula to Ejutla, 6 Nov 1977, F 2290 (OS); 14.7 mi NE of Autlan, 16 Dec 1966, AL 4546 (MICH, OS); 12-15 mi SSE of Autlan, 22-23 Nov 1959, McVaugh & Koelz 966 (ENCB, MICH); 3-6 km S of La Huerta, 16-17 Mar, M 23052 (ENCB, MICH); 16 km N of Ciudad Guzman, 13 Nov 1971, Dieterle 4/8/ (CA, ENCB, MICH, OS); E of Ciudad Guzman, 18 Nov 1968, Boutin & Brandt 2280 (MICH); Tamazula de Gordiano, 24 Dec 1967, Clarke e/ al. 2185-/ (MICH); 8 km W of Jilotlan de los Dolores, 22 Nov 1970, M 24590 (DUKE, ENCB, MICH); 12.5-14.5 mi SW of Mazamitla, 15 Dec 1966, AL 4539 (MICH, OS); Jiquilpan-Colima hwy, 3 mi above La Garita, 2 Dec 1959, McVaugh & Koelz 336 (DUKE, ENCB, LL, MICH). MEXICO: ca. 5 mi N of Ixtapan de la Sal, 10 Dec 1974, Stuessy & Roberts 3699 (OS, UC); near Amatepec, 29 Dec 1953, Maliala 30047 (NY); btw Mal Pais and Santa Barbara, 6 Dec 1953, Matilda 30125 (MEXU, NY, US); Temasca1tepec, 14 Nov 1932, Hinton 2401 (BM, G, K, GH, US); Valle de Bravo, 23 Dec 1952, Matuda 27335 (DS, MEXU, Ny); Ocotepec, Tejupi1co, 10 Dec 1967, Rzedoll'ski 25290 (DS, ENCB, MICH, MSC). ICHOACAN: btw Zamora & Sahuayo, km 159 hwy 15,3 Nov 1977, F 2265 (OS); 3 mi W of Zitacuaro, 20 Nov 1978, F 2866 (OS); ca. 28 mi E of Patzcuaro, 16 Dec 1978, FCa 2968 (OS); ca. 8 mi S of Uruapan, 16 Dec 1978, FCa 2971 (OS); Rancho Viejo, Apatzingan, 14 Oct 1939, Hinton 15332 (NA); 16 mi N of Coalcoman on rd to Tepalcatepec, 17 Dec 1978, FCa 2976 (OS); Sierra Naranjillo, Coalcoman, 26 Nov 1938, Hinton 12684 (MICH, TEX, US); 15-16 km SE of Asserradero Dos Aguas, 25-26 Nov 1970, M 24727 (DUKE, ENCB, MICH); 11.3 mi E of jct of rd from Tepaicatepec to Coalcoman and side rd to Dos Aguas, 17 Dec 1978, FCa 2982 (OS); ca. 12 mi SW of Coalcoman on rd to Villa Victoria, 17 Dec 1978, FCa 2985 (OS); ca. 15 mi SW of Coaicoman on rd to Villa Victoria, 17 Dec 1978, FCa 2986 (OS). MORELOS: near Cuernavaca, 15 Nov 1865, Bourgeaun 1199 (BR, C, G, GH, K, MPU, US); 16.7mi NW of Cuautla on rd to Cuernavaca, 24 Nov 1966, AL 4306 (MICH, OS); ca. 5 mi SW of Morelos/Mexico border on hwy 95, 17 Nov 1978, FRi 2847 (OS). NAYARIT: 11.7 mi NW of Tepic, 18 Dec 1966, AL 4551 (MICH, OS); 5-10 mi from Tepec on rd to Compostales, 27 Nov 1967, Grashoff 181 (MSC); btw Ixtlan del Rio Tetitlan, 22 Nov 1955, Carter & Kellogg 3639 (MEXU, MICH, US); 10 mi SE of Ahuacatian, 17-18 Nov 1959, McVaugh & Koelz (DUKE, ENCB, LL, MICH). OAXACA: btw Coixtlahuaca and Tamazulapam, 12 Nov 1894, Nelson 1948 (US). PUEBLA: 5.6 mi SE of Izucar de Matamoros, 22 Nov 1978, F 2870 (OS). SINALOA: San Ignacio, 28 Jun 1926, Ortega 4998 (GH, K, US); Balboa, Jan 1923, Ortega 4998 (US); Panuco, 28-30 Dec 1970, Smith 195 (ARIZ); 3.5 mi E of Copala on rd to Durango, 7 Dec 1966, Gentry 22279 (NA, US); 32-33 mi NW of jct of hwys 15 and 40, 18 Dec 1966, AL 4554 (MICH, OS).
general
Montanoa arborescens DC., Prodr. 5: 566. 1836, as "Montagnaea." Type: MEXICO, Cordilliere de Guchilaque, Aug 1827,.1. L. Berlandier 1006 (lectotype chosen from four syntypes, G-DC, photo IDC 800. 565:III.4 top!; iso1ectotypes, PI! G! Pi; photo of P isolectotype, P! MO! US i). This syntype was selected because it has the most collection information, the largest number of specimens at various herbaria and because it was the only one specified by number in the protologue. Other syntypes are: L. Aleman S .11., 1831; L. Aamlan S.n.; and Mairet 59.
Montagnaea c1ematidea Walpers, Linnaea 14: 308. 1840. Type: MEXICO, locality unknown, s.d., W. F. Kwinski s.n. (holotype LE, not seen; isotype, KIELl). Karwinski's original herbarium is deposited at LE (Holmgren and Keuken, 1974), but the specimen was not present in the loan from that institution. Eriocoma c1ematidea (Walpers) Kuntze, Rev. Gen. PI. 1: 336. 1891.
Montanoa uncinata Schultz Bip. in K. Koch, WochenschI. Vereines Beford. Gartenbaues Konig!. Preuss. Staaten 7: 406. 1864. Type: MEXICO, CumbIe de Estepa, Sep 1842, F. M. Liebmann 484 [Liebm. PI. Mex. 9056] (holotype, Pi; isotypes, C[4]! G! K! Pi; photo of holotype, OS!; photos of C isotype, P! MO! NY! US!; tracing of C isotype, GH!). Eriocoma uncinata (Schultz Bip. in K. Koch) Kuntze, Rev. Gen. PI. 1: 336. 1891.
Montanoa patens A. Gray, Proc. Amer. Acad. Arts 21: 388. 1886. Type: MEXICO, Chihuahua, mountains above Batopilas, Aug-Nov 1885, E. Palmer 164 (holotype, GH!; isotypes, BM! G[2]! GH! K! LE! NY[2]!; photo of K isotype, OS!).
Polymnia nervata M. E. Jones, Contr. W. Bot. 12: 44. 1908. Type: MEXICO, Chihuahua, Guayanopa Canyon, Sierra Madre Mts., 24 Sep 1903, 3600 ft, M. E. Jones s.n. [ace. #40414] (lectotype chosen from two syntypes with identical information, POM!; isolectotype, POM!).
REPRESENTATIVE SPECIMENS: GUATEMALA. SACATEPEQUEZ: above Duenas, 21 Jan 1939, Sa 63257 (F). SOLOLA: 3-5 km W of Panajachel, 6-7 Dec 1963, Williallls et 01. 25317 (F, NY, US). MEXICO. CHIAPAS: 16 km NW of Rizo de Oro, 3 Nov 1971, Breedlove &: Smith 21746 (DS, ENCB, MICH, MO); 30 mi E of Tuxtla, 26 Nov 1977, FRa 257/* (OS); ca. 45 km E of Tuxtla, 16 Dec 1977, FK 2700 (OS); ca. 5 km E of Tuxtia, 16 Dec 1977, FK 2699 (OS); hwy 1957 mi N of hwy 190,27 Nov 1977, FRa 2574 (OS); jct of hwys 190 and 195,25 Nov 1977, FRo 2561 (OS); 24.6 mi N of jct of hwys 190 and 195,27 Nov 1977, FRa 2576 (OS); 34.2 mi N ofjct of hwys 190 and 195,27 Nov 1977, FRo 2580 (OS); Zinacatan, 29 Nov 1971, Breedlmoe 22913 (DS); 5 km above Soyalo, 2 Nov 1965, Breedlove 14053 (DS, ENCB, F, LL, MICH); 23 mi SE of San Cristobal, 18 Nov 1966, AL 4248 (MICH, OS); 2-3 km SW of Aguacatenango, 16 Dec 1977, FK 2693 (OS); 3-4 km SW of Aguacatenango, 16 Dec 1977, FK 2695 (OS). CHIHUAHUA: San Jose de Pinal, Rio Mayo, 21 Sep 1936, Gentry 2840 (ARIZ, F, GH, MO, S, UC); Barranca del Cobre, 30 Dec 1977, Hartlllan et 01. 5107* (OS, RM); Arroyo Hondo, II Sep 1935, Gentry 1776 (MEXU, MO, NA, UC, US); Maguarichic, 20 Sep 1939, Knobloch 6029 (LL, MSC, US). DURANGO: jct of hwy 40 and rd to San Lucia, 20 Dec 1978, FCa 3001 (OS); 4 mi E of jct of hwy 40 and rd to Santa Lucia, 20 Dec 1978, FCa 3002 (OS); 4 mi E of jct of hwy 40 and rd to Santa Lucia, 20 Dec 1978, FCa 3003 (OS); 1.5 mi E of La Fraguita, 20 Dec 1978, FCa 3004 (OS); ca. 45 km SW from La Ciudad, 21-22 Oct /971, Dieterle 3828 (ENCB, MICH); 17 mi ESE of Durango, 3 Oct 1965, C 10261 (CA, DUKE, ENCB, GH, MEXU, MICH, MSC, NY, TEX, WIS); 7 mi E of La Palmito, 27 Sep 1973, Reveal &: Atwood3618 (US). GUERRERO: 5.8 mi E of Chichihualco, 15 Nov 1978, FRi 2838 (OS); 22 mi from hwy 95 on rd to Filo de Caballo, 13 Nov 1977, FRo 2325b (OS); Canizal-Chichihualco rd, 4.2 mi SE of jct of road to Filo de Caballo, 15 Nov 1978, FRi 2834 (OS); Cruz de Ocote, 3 Dec 1963, Rzedowski 18099 (DS, ENCB, MICH, MSC, WIS); Otatlan, 14 Nov 1939, Hinton 14852 (F, LL, MO, NA, NY); Pueblo Viejo, 8 Nov 1939, Hintoll 14818 (F, GH, LL, MO, NA, US). JALISCO: 5 mi NNE of Talpa de Allende, 14 Oct 1960, M 20189 (DUKE, ENCB, LL, MICH, NY); rd to Tapalpa from hwy to Ciudad Guzman, 15 Nov 1968, Boutin &: Brandt 2142 (CA, MEXU, MICH); ca. 5 mi SW of Tepatitian, 8 Oct 1976, SG 4096* (OS); 47 mi S of Iiquilpan,8 Nov 1977, F 2297 (OS); N shore of Lago de Chapala, [ Oct 1966, AL 3858 (MICH, OS); btw Lagos de Moreno and San Juan de los Lagos, 7 Oct [976, SG 4088* (OS). MEXICO: 2 km S of Temescaltepec, 30 Oct 1977, F 2232* (OS); 2 km S of Temescaltepec, 30 Oct [977, F 2233* (OS); 3 km S of Temescaltepec, 30 Oct 1977, F 2234* (OS); 3 km S of Temescaltepec, 30 Oct [977, F 2236* (OS); rd N from Valle de Bravo to hwy [5, [9 Nov 1978, F 2852 (OS); 3.2 mi N of jct of hwy I and hwy 5, 19 Nov [978, F 2854 (OS); near Jepantia, 10 Dec [978, FV 2954 (OS). MICHOACAN: ca. [I mi E of Coalcoman de Matamoros on rd to Dos Aguas, 17 Dec 1978, FCa 2981 (OS); 2 km W of Zacapu, 3 Nov 1977, F 2264* (OS); ca. [5 mi E of Morelia, [5 Dec 1978, FCa 2966 (OS); 24 mi W of Zitacuaro, I Nov 1977, FH 2251 (OS); 4.1 mi W of Zitacuaro, 20 Nov [978, F 2865* (OS); ca. 7 mi SE of Ciudad Hidalgo, 4 Sep [976, HF 4245* (OS); just E of Zitacuaro, I Nov [977, FH 2244 (OS). MORELOS: 1.2 mi S of Mexico/Morelos state line on Cuautla-Amecameca rd, 25 Nov [966, AL 4310 (MICH, OS); hwy 95D near km marker 74, 16 Nov 1977, FG 2360 (OS); 6.3 mi SW of Tepoztlan, 25 Nov 1966, AL 4307 (MICH, OS); 2 mi N of Cuernavaca, 27 Jan 197[, Freeland &: Spetzmall 74 (MEXU, NA). OAXACA: 27 mi NE of jct of hwys [90 and [25, 23 Nov [978, F 2877 (OS); 14 mi NE of jct of hwys 190 and [25,23 Nov [978, F 2880 (OS); btw Huahuapan de Leon and Yanhuitlan, 18 Oct [976, SG 4250* (OS); 30.7 mi NW of Oaxaca, 23 Nov [978, F 2895 (OS); [4.7 mi SE ofjct ofhwy 190 and 1.2 mi NW of Tamazulapan, Rio Oro, 23 Nov 1978, F 2891 (OS); [2-14.5 mi S of Suchiztepec, 8 Nov [966, AL 4177 (MICH, OS); Las Sedas, 20 Oct 1907, Conzalli 2079 (F, MEXU, MICH); near Ixtian de Juarez, 4 Nov [966,AL4120 (MICH, OS); 39 km N ofjctofhwys 190 and 131, 20 Nov 1977, Fro 2399 (OS); 6.8 mi N of Oaxaca on hwy [75, [7 Dec 1977, FTK 2701 (OS); Sierra de San Felipe, 9 Oct-[6 Nov 1894, Pringle 4929 (BM, BR, CM, F, G, GH, M, NY, S, US); 2 mi N ofjct of hwy 190 and rd to Iaycatlan, 6 Nov [966, AL 4134 (MICH, OS). PUEBLA: ca. 3 mi E of Puebla, 20 Aug, HF 4135* (OS); near San Miguel Papaxtla, Cholula, 3[ Sep 1967, Rzedmvski 24898 (ENCB); I km NW of San Pedro Yaucuitlalpan, 24 Oct 1976, Mexica 159 (ENCB); 20 mi N of Huajuapan de Leon, 24 Oct 1965, Cronquist &: Sousa 10402 (CA, DUKE, ENCB, GH, MEXU, MICH, MSC, NY, TEX, US). SINALOA: 5 mi NW of Los Ornos, Sinaloa and Vela, 6 Nov 1969, Breedlove &: Kawahara 17076 (MICH); Palmito, 14-15 Nov 1959, Gentry &: Arguelles 18173 (TEX, US). SONORA: 12.5 mi W of Yecora, 12 Aug 1978, FS 2775* (OS); Huchuerachi, 4 Dec 1890, Hartlllan 302 (GH, K, NY, UC, US); near La Nopalera, 4 Oct 1939, MillieI' 3613 (GH, LL, MICH, NA, UC); San Bernardo, 8 Nov 1934, Gentry 1146 (A, ARIZ, F, MICH, MO); Sierra de Alamos, 30 Oct 1939, Gentry 4788 (ARIZ, DS, F, GH, MICH, MO, NA, NY, UC, US).
morphology
The male florets are not always grouped in the center of the head as previously described, in fact some groups of male florets are found at the outer edge of the receptacle. It seems more likely that the aerial heads are actually groups of heads some of which are few flowered male heads and others are single flowered female heads so these could represent dioecious heads grouped on a common receptacle. The subterranean heads flower first, their petals protrude just above the soil surface. Later the aerial inflorescences appear. Likewise the fruits of the two types of heads of Gymnarrhena have different developments, the larger subterranean fruits develop first and the aerial fruits are produced later and only in wet years. The aerial fruits are wind dispersed while the subterranean fruits germinate underground. On the aerial heads the scales of the pappus as well as the bracts surrounding the achenes are hydrocastic (open when wet) and the achenes are dispursed by wind (Guterman, 2002).
Gymnarrhena exibits “dimorphic cleistogamy” or “true cleistogamy” in that it has two different flower types: chasmogamos and cleistogamous (Culley & Klooster, 2007). While about 228 genera have some type of cleistogamous flowers, only 168 are dimorphic. It is estimated that dimorphic cleistogamy has evolved less than 40 times and most of the lineages have very few species (Culley & Klooster, 2007).
Anatomy
The only anatomical data found is from Zamski et al (1983) who examined the taproot of Gymnarrhena and found that it starts to contract soon after emergence. Ultimately, this contraction causes the retraction of the main shoot apex from the soil surface to a depth of about 10 mm.
morphology
A delicate appearing plant with slender, often erect branches and heads with a large number (90-120) of extremely small (2 mm) disc florets. When Montanoa atriplicfolia is found as a vine it is characterized by large (1-2 cm diam) branches sprawling across other vegetation or the ground and short, slender, erect, lateral branches bearing small groups of capitula. This type of growth form is found primarily in the Yucatan Peninsula. At higher altitudes and in wetter habitats this species gradually changes to a shrub habit. Both growth forms have relatively small leaves that have the same range of shapes.
morphology
This striking species has the most diagnostic leaf shape of the genus. The margins are deeply pinnatifid or bipinnatifid, the adaxial surface is dark green with white pustular hairs that appear as spots on the surface. Occasionally some of the leaves near the synflorescence are less lobed, but the lower ones on the plant are consistently lobed. The species is further distinguished by the mature pales which are the only ones in the genus that expose part of the achene because of a narrow base. The disc florets (8 mm), and the stigmata (12 mm) are the largest in the genus. The apex of the disc corolla is covered with short stiff hairs that are interlocking and perhaps serve to protect the head from predation while in bud. If so, it does not work well because this species shows more evidence of insect damage on the heads than any other species in the genus (pers. obser.).
diagnostic
Shrubs to small trees 3-7 m tall. Stems terete, grooved, brown, herbaceous parts moderately pubescent with reddish or brownish-white appressed hairs. Leaves variable; petioles 2.0-18.5 cm long, occasionally with auricles at distal end, moderately pubescent, hairs reddish, 0.5-1.0 mm long; blades ovate to pentagonal (Fig. 37A-G), 6.5-18.5 cm long, 3.0-17.5 mm wide, apex long-acuminate, margin entire to irregularly serrate, adaxial surface dark green, sparsely to moderately pubescent, hairs pustular, abaxial surface paler green, moderately glandular, hairs in axils of major veins. Peduncles 0.3-1.0 cm long, densely pubescent, hairs less than 1.0 mm long. Heads erect, less than 1.0 cm diam in flower, ca. 1.0 cm diam in fruit, few to several in dense cymose synflorescences in oppositely and alternately branched compound corymbs. Phyllaries 3-6, more or less biseriate, elliptic or ovate-lanceolate, loosely reflexed, 3-5 mm long, l.0-1.5 mm wide, green, apex obtuse or mucronate, margin ciliate, entire, abaxial surface densely pubescent, hairs 0.5-1.0 mm long, adaxial surface glabrous. Ray florets 7-8; corollas white, ligules elliptic to obovate, 20-35 mm long, 4-6 mm wide, apex truncate to 2-notched, adaxial surface glabrous, abaxial surface sparsely glandular and pubescent, hairs 0.5-1.0 mm long, tube 1.0 mm long, 0.5 mm wide, glabrous. Disc florets (Fig. 37H) 25-35; corollas yellow, tube 1.5 mm long, 0.3-0.5 mm diam, moderately glandular and pubescent, hairs 0.5-1.0 mm long, throat narrowly cylindrical, 4.0-4.5 mm long, 1.0-1.5 mm diam, sparsely glandular and pubescent especially on veins, hairs less than 0.5 mm long, lobes 5, 1.0-1.2 mm long, 0.5 mm wide, apex acute to narrowly acute, densely glandular and pubescent, hairs less than 0.5 mm long; stamens with filaments 5.0-5.5 mm long, 0.2 mm wide, anthers fully exserted from corolla (Fig. 37H), thecae brown to purple, 2.5-3.0 mm long, 0.5 mm wide, apical appendages yellow, usually with purple on tip, narrowly acute to acute, 0.5-0.7 mm long, 0.25-0.3 mm wide, glabrous; styles yellow, 10.0-11.5 mm long, enlarged at base, stigmatic surface yellow, 1.0 mm long, apical appendage yellow with purple, acuminate, 0.5 mm long, 0.25 mm wide. Pales at anthesis ovate-triangular (Fig. 37I), 5-6 mm long, 1.5-3.0 mm wide, light yellow sometimes with purple on distal half, apex long-acuminate, indurate, often purple, margin ciliate, entire, abaxial surface densely glandular near center and pubescent on the veins, hairs 0.5-1.0 mm long, adaxial surface glabrous; pales at fruiting and achenes not available. Chromosome number unknown.
From Funk, V. A. "The Systematics of Montanoa (Asteraceae, Heliantheae)" in Memoirs of the New York Botanical Garden. Vol. 36. The New York Botanical Garden. Bronx, NY. 1982.
diagnostic
Shrubs: Stems brown, glabrous. Leaves poorly known; petioles 2-4 cm long, partially winged at distal end, densely glandular and pubescent, hairs 0.5-1.0 mm long; blades ovate-Ianceolate to trullate (Fig. 48A, B), 5-12 cm long, 1-5 cm wide, apex long-acuminate, margin serrate, adaxial surface moderately pubescent, hairs pustular, more evident on young leaves, abaxial sllliace moderately glandular and pubescent, hairs 0.5-1.0 mm long. Peduncles 1.5-3.5 cm long, densely glandular and pubescent, hairs 0.5-1.0 mm long. Heads erect, 0.8 cm diam in flower, ca. 1.0-1.2 cm diam in fruit, many to numerous in open cymose synflorescences in oppositely branched compound corymbs. Phyllaries 5-6, uniseriate, subequal, ovate to ovate-lanceolate, 3-5 mm long, 1.0 mm wide, dark green, apex acute to acuminate, mucronate, margin ciliate, entire, abaxial surface sparsely pubescent, adaxial surface glabrous. Ray florets 6-7; corollas white with dark veins, ligules obovate, 6-10 mm long, 3-4 mm wide, apex acute to 2-notched with clusters of hairs, adaxial surface glabrous, abaxial surface moderately to densely glandular and pubescent, hairs less than 0.5 mm long, tube less than 0.5 mm long, almost non-existent, with a few scattered hairs. Disc florets (Fig. 48C) 49-53; corollas yellow, tube 0.75-1.0 mm long, 0.5 mm diam, moderately glandular and pubescent, hairs 0.5-0.75 mm long, throat cylindrical, 1.5 mm long, 1.0 mm diam, sparsely glandular and pubescent primarily on the veins, hairs less than 0.5 mm long, lobes 5, 0.5-0.7 mm long, 0.5-0.7 mm wide, apex acute and moderately glandular and pubescent, hairs less than 0.5 mm long; stamens with filaments 1.0 mm long, 0.2 mm wide, anthers not fully exserted from corolla, thecae yellow with brown, 1.0 mm long, 0.5 mm wide, apical appendages yellow, acute, 0.5 mm long, 0.4 mm wide, abaxial surface of apical appendages glabrous; styles yellow, 4.0 mm long, slightly enlarged at base, stigmatic surfces yellow, 0.6 mm long, apical appendages yellow with dark spots, acute, 0.4 mm long, 0.4 mm wide. Pales at anthesis obtrullate, 2.5 mm long, 2.0 mm wide, light yellow with dark bands on midrib and veins, apex longacuminate, dark, indurate, margin sparsely ciliate, entire, abaxial and adaxial surfaces glabrous; pales at fruiting (not quite mature) deciduous, almost obdeltoid, 8-9 mmlong, 4-S mm wide, papery with netted venation, stramineous with dark bands on veins, apex abruptly short apiculate, dark, indurate, margin sparsely ciliate, entire, abaxial and adaxial surfaces glabrous. Achenes seemingly typical but too young for certainty. Chromosome number unknown.
diagnostic
Shrubs or vines to 3 m tall or 15 m long. Stems terete, light brown, herbaceous parts puberulent. Leaves variable; petioles 1.5-11.5 cm long, usually partially winged and auriculate, densely glandular and pubescent, hairsO.5-1.0 mm long, blades ovate to pentagonal (Fig. 60A-K), 4.5-14.5 cm long, l.5-15.0 cm wide, apex acute to acuminate, margin entire to irregularly serrate, sometimes deeply 3-10bed, adaxial surface moderately pubescent, hairs pustular, abaxial smface densely glandular and pubescent, hairs 0.5-1.0 mm long. Peduncles 1.5-7.5 cm long, densely glandular and pubescent, hairs 0.5-1.0 mm long. Heads pendulous at fruiting, 0.6-0.8 cm diam in flower, 1.7-2.7 cm diam in fruit, few to many in open cymose synflorescences in oppositely branched compound corymbs. Phyllaries 5-6, uniseriate, equal, quickly becoming reflexed, ovate-Ianceolate, 4-5 mm long, 1.0-1.25 mm wide, dark green, margin ciliate, entire, abaxial surface densely pubescent, hairs 0.5-1.0 mm long, adaxial surface with a few scattered hairs. Ray florets 8-]5, corollas white, ligules lanceolate to narrowly elliptic, 12-24 mm long, 3-6 mm wide, apex acute to 2-notched, adaxial smface essentially glabrous, abaxial surface moderately to densely glandular, tube 0.5-1.0 mm long, 0.25 mm wide, essentially glabrous. Disc florets (Fig. 60L) 90-120; corollas yellow, tube 0.5 mm long, 0.5 mm diam, sparsely glandular and pubescent, hairs less than 0.5 mm long, throat cylindrical, 1.0 mm long, 1.0 mm diam, essentially glabrous or sparsely glandular and pubescent, hairs less than 0.5 mm long, lobes 5,0.5 mm long, 0.5 mm wide, apex acuminate, moderately glandular and pubescent, hairs thick, less than 0.5 mm long; stamens with filaments 1.0 mm long, 0.10 mm wide, anthers not fully exserted from corolla, thecae brown, 0.75 mm long, 0.25 mm wide, apical appendages yellow, acute-acuminate, 0.5 mm long, 0.15 mm wide, abaxial surfaces of apical appendages glandular; styles (Fig. 60M) yellow, 2.5 mm long, enlarged at base, stigmatic surface brown, 0.75 mm long, apical appendages yellow, acute, 0.25 mm long, 0.25 mm wide. Pales at anthesis obtrullate (Fig. 60N) 2.25 mm long, 2.0 mm wide, light yellow with dark bands in center and along veins, apex long-acuminate, indurate, yellow, margin ciliate, entire, abaxial surface moderately glandular and pubescent near distal half, not on apex, hairs 1.0 mm long, on adaxial surface glabrous; pales at fruiting deciduous, more or less obdeltoid (Fig. 60M) , 10-14 mm long, 3-4 mm wide, papery with netted venation, stramineous or purple, apex sinus with apiculate tip, margin sparsely pubescent, hairs less than 1.0 mm long, adaxial and abaxial surfaces glabrous. Achenes brown-black, 2.5 mm long, 1.5 mm wide, smooth. Chromosome number, n = 19.
diagnostic
Shrubs 2-10 m tall. Stems quadrangular becoming terete, brown to gray, herbaceous parts puberulent. Leaves consistent; petioles 4-20 cm long with two auricles, sometimes with thin blade margin extending down adaxial surface, usually with two small leaf-like structures at proximal end, sparsely pubescent, hairs less than 1.0 mm long; blades ovate to ovate-Ianceolate, 12- 30 cm long, 9-40 cm wide, dark green, apex acute to acuminate, margin regularly to irregularly serrate, pinnatifid to bipinnatifid adaxial surface densely pubescent, hairs pustular, turning white, abaxial surface sparsely to densely glandular and pubescent, hairs less than 1 mm long. Peduncles 2-7 (usually 3) mm long, sometimes purple, densely glandular and pubescent, hairs less than 1.0 mm long. Heads pendulous, 1.8-2.5 cm diam in flower, 3.5-4.0 cm diam in fruit, several to many in open cymose synflorescences in oppositely branched compound corymbs, lateral branches spreading to ascending. Phyllaries 8-9, reflexed in fruit, biseriate, subequal, ovate-Ianceolate to lanceolate, 6.0-8.5 mm long, 1.0-3.0 mm wide, green, apex acute to acuminate, mucronate, margin ciliate, entire, abaxial surface densely pubescent, hairs less than 1.0 mm long, adaxial surface glabrous to sparsely pubescent on distal half, hairs less than 0.5 mm long. Ray florets 10-12; corollas white, ligules ovate-lanceolate, 25-35 mm long, 6-11 mm wide, apex acute to two-notched with clusters of hairs less than 0.5 mmlong, adaxial surface essentially glabrous, abaxial surface sparsely to moderately glandular and pubescent, hairs primarily on veins, less than 0.5 mm long, tube 1.5 mm long, 0.5 mm wide, essentially glabrous. Disc florets 95-125; corollas green turning yellow, tube 1.5 mm long, 0.5-1.0 mm diam, glabrous to sparsely glandular and pubescent, hairs less than 0.5 mm long, throat cylindrical, 4-5 mm long, 1.5-2.0 mm diam, moderately to sparsely glandular and pubescent, hairs less than 0.5 mm long, lobes 5, 1.0-1.25 mm long, 0.5-0.75 mm wide, apex acute and densely pubescent, hairs less than 0.5 mm long; stamens with filaments 5.0-6.0 mmlong, 0.25 mm wide, anthers fully exserted from corolla, thecae yellow, 2.5 mm long, 0.5 mm wide, apical appendages yellow, acuminate, 0.5-0.75 mm long, 0.5 mm wide, abaxial surface of apical appendages glandular; styles 7.0-9.5 mm long, enlarged at base, stigmatic surfaces yellow, 1. 5-2. 0 mm long, apical appendages yellow, acuminate, 0.5-2.0 111m long. Pales at anthesis pentagonal, 2.0-2.5 mm long, 1.5 mm wide, light yellow, apex long acuminate, yellow, indurate, margin ciliate, entire, abaxial surface glandular and pubescent near center, hairs to 1.1 mm long, adaxial surface glabrous; pales at fruiting deciduous, obtrullate 16-21 mm.long, 4-5 mm wide, papery with netted venation, stramineous, light in color, indurate, margin entire, abaxial surface sparsely glandular near center, adaxial surface glabrous. Achenes brownblack, 3.5 mm long, 2.0 mm wide, smooth. Chromosome number, n= 19.
distribution
Trujillo, Venezuela (Fig. 49); habitat unknown;23S0 m.
distribution
Southern Mexico from Sinaloa to Oaxaca; pine-oak forests, along streams or on hill and road sides; 450-2000 m.
Because this plant is very showy it is often cultivated, in fact it is second only to Montanoa hibiscifolia in numbers cultivated. Data from herbarium sheets show that it is cultivated from the following countries: Argentina, Ceylon, Burma,Southern India, United States (Florida, California), and throughout its native range in Mexico. One collection (F 2272) was found in the "double rayed" form growing in a yard near Guadalajara.
distribution
Mountains of western and southern Mexico from Sonora to Chiapas and Guatemala.
ecology
Gymnarrhena is an amphicarpic herb (see images) of the Mediterranean biome of North Africa and the Middle East growing in dry, mostly bare, sandy areas and resembling a grass (see images). Research on the reproductive biology of this unusual plant is summarized by Koller and Roth (1964) and in the description above. There are three aspects to its reproductive biology that are most interesting, seed heteromorphism, the presence of aerial as well as subterranean heads, and the presence of both casmogamous and cleistogamous flowers.
Seed heteromorphism, that is the production of seeds with variable morphologies and ecological strategies, represents an allocation of different fractions of seed output to different ends and most examples are found in four flowering plant families, one of which is the Compositae (Harper, 1977). Seed heteromorphism appears to be largely restricted to relatively short-lived, fugitive species, particularly weeds. Venable and Burquez (1989) say that it might be a form of ‘bet hedging’ in response to environments that vary spatially or temporally. These morphological heteromorphisms are important because they may be associated with ecological strategies which have evolutionary significance, such as dispersal, dormancy, differential competitive performance, within or among year timing of germination, vulnerability to predators, seedling growth and survival or fecundity (see many references cited in Chmielewski, 1999).
The presence of aerial as well as subterranean heads has several different explanations. Koller and Roth (1964) reported that mean weight of Gymnarrhena aerial fruit was only 5-6% of the weight of a subterranean fruit and that 6-day-old subterranean seedlings were 6 times the weight of aerial seedlings. They reported that the survival was considerably greater for these subterranean seedlings and that plants under dry conditions may fail to produce aerial heads. On the other hand, according to Brown & Böer (2005) the species occurs on firm sometimes rocky substrates which are generally hostile to plant grown. The above ground fruits are small and possess a small pappus to aid in their dispersal so they can travel some distance from the parent plant while the 1-2 large underground fruits (nearly 20 times the weight of the aerial ones) lack a pappus and stay close to the parent plant. It is their opinion that the underground fruits ensure that when the mother plant dies later in the year the same favorable location is re-colonized. Gutterman (1989) thinks that the underground location protects the seeds from predation. Zeide (1978) described dual strategies for Gymnarrhena micrantha in the Negev desert of Israel. He determined that the subterranean fruits were produced according to a “pessimistic strategy” where fruit production begins as soon as possible concurrent with continued vegetative growth. The aerial fruits, on the other hand, were produce according to a more optimistic strategy (in order to maximize yield, the vegetative stage precedes a last and full switchover to heavy fruiting). This pessimistic strategy must therefore arise from other considerations such as unpredictable environments as it is at odds with optimality models of resource allocation in annuals (Cheplick & Quinn, 1982). They summarize by saying that there are three arguments that have been supported for Gymnarrhena: 1) the seedlings arising form the larger propagules are more tolerant to stress or competition, 2) these subterranean seedlings have a higher probability of surviving to produce seed, 3) genotypes with an early production of subterranean seed may be the only ones to produce seeds under stress (Evenari, 1963; Koller & Roth, 1964; Zeide, 1978). And another has been suggested (Cheplick and Quinn, 1982): 4) the larger subterranean seed may show a greater return on the energy invested than an aerial seed. Other explanations that have been provided are the protection afforded by burial against catastrophies and the placement of subterranean seed in the same microhabitat occupied by the parent plant, providing protection from the risks of random dispersal (Evenari, 1977; Koller & Roth, 1964). Cheplick & Quinn (1982) concluded that the early production of few larger seeds followed by the later production of many small seeds was presumably the result of the variable and unpredictable length of their growing periods and to their roles a fugitive species.
There are about 50 amphicarpic species worldwide, approximately one in every 5000 species of flowering plants; most of them occur in either frequently disturbed and or stressful habitats. Eight of these are found in Israel which has only 2500 species in its known flora, one in ca. 310 species, a very high ratio compared to flowering plants in general (Kaul et al., 2000; Lev-Yadun, 2000). Given the suggested reasons above, Lev-Yadun (2000) adds that since many of the amphicarpic annuals are found in disturbed areas that are the result of drought, fire and grazing. He thinks this indicates a long history of disturbances in the eastern Mediterranean region that pre-date human impact.
The cleistogamous aspect of the subterranean florets is more easily explained in that such flowers are invariably cleistogamous (Kaul et al., 2000).
Additional information can be found in the recent book by Gutterman (2002).
habitat
Chiapas and the Yucatan to Costa Rica; wide range of habitats from cool forests to dry roadsides; 450-2000 m, more vine-like at lower altitudes and in drier environments.
habitat
Cool wet forests on mountain slopes, in higher altitudes in fields and along forest margins and at lower altitudes along roadsides and riverbanks as well as on steep slopes; 1000-2300 m, usually between 1500-1800 m.
lifecycle
Flowering (and Fruiting) period: October (October-November?).
lifecycle
FLOWERING (AND FRUITING) PERIOD: November-January (December-March).
lifecycle
FLOWERING (AND FRUITING) PERIOD: Late November-December (December- February).
Name
- Homonyms
- Montanoa leucantha ssp. arborescens
Bibliographic References
- Andre Michaux (1803) Flora boreali-americana :sistens caracteres plantarum quas in America septentrionali collegit et detexit Andreas Michaux. http://compositae.myspecies.info/node/146
- Cassini, H. (1830) Zyégée http://compositae.myspecies.info/node/143
- Charles Louis L’Heritier (1789) Sertum Anglicum seu Plantae Rariores quae in Hortus juxta Londinum, Imprimis in Hortus Regio Kewensi Excoluntur http://compositae.myspecies.info/node/148
- Kurt Polycarp Joachim Sprengel (1826) Caroli Linnaei... Systema vegetabilium/Curante Curtio Sprengel http://compositae.myspecies.info/node/144
- Laurentii Salvii.,Linné, Carl von (1753) Caroli Linnaei ... Species plantarum :exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas... http://compositae.myspecies.info/node/151
- Merritt Lyndon Fernald,Bernice G. Schubert (09/1948) Studies of American Types in British Herbaria http://compositae.myspecies.info/node/126
- Walter, Thomas (1788) Flora caroliniana: secunda systema vegetabilium perillustris Linnaei digesta... http://compositae.myspecies.info/node/149
- William Aiton (1789) Hortus Kewensis, or, A catalogue of the plants cultivated in the Royal Botanic Garden at Kew. http://compositae.myspecies.info/node/147