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Notes on Alophia (Iridaceae) and a New Species, A. veracruzana, from Vera Cruz, Mexico PDF

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Preview Notes on Alophia (Iridaceae) and a New Species, A. veracruzana, from Vera Cruz, Mexico

Number Volume 79, 4 & Howard Goldblatt 903 1992 Alophia resemblance goes no and but the further, C. kin- in the genus. reportedly has pendent It lilac -violet probably does not have a tonii particularly close flowers with a yellow center, connate filaments, Alophia relationship to or Ainea, pandurate anthers, and deeply forked branch- style mm Little attention has been paid to the broad con- es, the arms of which are 14-15 long. In the nective condition in Iridaceae, and although seems article in which he described A, intermedia, Ra- it intuitively to be apomorphic, the character venna (1964) included A. Alophia^ is dis- trijiora in a two now tributed in of three genera of Maricaeae and species regarded as conspecific with Tigridia several species of CjKpeZ/a, possibly the basal genus meleagris (Molseed, 1970). Examination of the of Tigridieae. Thus, a broad connective probably type of A. intermedia^ not available to us, nec- is is synapomorphous for the clade that includes Mari- essary before status and generic position can be its ceae and Tigridieae (see Goldblatt, 1990) but pie- assessed. siomophic for Ainea and Alophia, although the Alophia veracruzana seems most closely re- pandurate shape in Alophia seems unique. There lated to A. drummondii, which resembles ui the it no longer seems any reason to place Alophia in size and general form of the flower (Figs. lA, 2A), Tigridiinae. The shared polyploid base number Alophia drummondii normally a taller plant, is is almost certainly convergent, and /i/op/ija does not and the flowers are violet to red-purple with the i have marked cream either the united filaments or bisulcate pollen well-diflFerentiated tepal claws to grains that are apomorphic Beyond yellow and densely speckled throughout with dark for Tigridiinae. a possible close relationship with Ainea, the red-purple. The major diff"erence be ween affin- ities of Alophia seem more uncertain than ever. species in the shape of the inner tepals. In A, lies drammondiij the inner tepals are narrower than the outer tepals, and the limbs are sharply bent at Distribution and Review Species base 2 The edges of the lower part of the (Fig. A). Alophia occurs across a wide portion of tem- the inner tepal limbs are closely pinched together perate and tropical America. The most known below where they are whitish to yeUow and arched well and widespread species, A. drummondii (Graham) over the median lower part of the limb. This fold R. Foster (synonyms Eustylis purpurea (Herbert) covers a narrow median zone of stalked glandular, & Engelm. The part of Gray, Ravenna), presumably nectariferous tissue. distal E, />artc/a/a (Herbert) extends from limb dark red-purple and as- the southern United States (Arkansas, the outer tepal is Louisiana, Oklahoma, and Texas) southern Mex- cending. Alophia veracruzana (Fig. lA) lacks a to «co limb (including Vera Cruz, Chiapas, and Tabasco) & (Goldblatt, 1974; Henrich The mner 1992). Goldblatt, species, or one closely related to also occurs in undulate margins, have no nectariferous band. it, Guyana Maas aL 1774, seems (cf. et U). This to have SystematICS slightly smaller flowers, filaments united in - the lower and half, style branches that are shorter - veracruzana ^nd do not A arch over the stamens. second species, Mexico. TYPE: Ae smaller-flowered A, (Loesener) Gold- silvestris Missouri ted Watt (synonyms & Nemastylis hequaertii Standley, IJouard Garden, June 990, Goldhlatt ical 1 rotacea Ravenna), Amer- ^. Central restricted to MEXU; is K, MO), 9070 i.^otypcs, (holotype, ica, where extends from southern Mexico Costa to it L Figure A '^ica. medusa third species, A. (Baker) Goldblatt (basionym Chlamydostylis medusa tiini Baker), ap- o mm c _' , . ,- . 5-0 i 4-5, laminis phcatis, latis, FPa^rreenmiilyv m g-. •' cis atrobrunneis, foliis pr.or^o^r-.li,y known, . i occurs Goias, centrali '__ ^f/;;.°"^4"-"5^ spatha iin„t,e^rri\oLr^e 3-x2o 3-x8iK mm,i,u. llnonncg.ap, florum, rhipidiis orazil. It is particularly distinctive in large flow- its interioribus marulis albis vel luteis tepalis floribus iilacinis ers and cm spathes 2.5-3 (tepals long, inner spathes obscure unguiculatis. exlerioribua obovatis tepalis notatis, mm 16-19 mm *^^)j a conspicuous band of glandular tissue in 17-20 longis, ca, 10 latis, interioribus mm mm staminibus atropnrpurffb, 10 ^he median of the inner tepal limbs, and terete longis, ca. latis, mm angubtc contiguis anlheris 2 longis infra eaves. To filamentis ca. now these the new A. veracruzana can mm ^ 1.1 loiigts* added. This has a restricted distribution in north- mm 2 supra. ca. eastern Mexico, known from Vera currently only 25-30 cm Bulbs nar geophyte high. Perennial Leaves 4-5, One dark brown. Ravenna rowly ovoid, the scales species, Alophia intermedia ± shrath- entirely (1964) except the uf>permosl (known laminate to us only from the description) from abc as may and elamtnate, blad<^s pllr.ae, iiK^-ily t Sinaloa western Mexico, be misplaced ing in ^ 904 Annals of the Garden Missouri Botanical 8 long to slightly longer than the stems, the lower begin to separate at about A.M., and by 9 A.M. 2-3 inserted below the ground, the upper two they have spread sufficiently to allow access to the inserted well above ground, the lower of these near style and stamens; by this time the anthers have dline and short- dehisced, revealing the bright yellow pollen. At mm 5-8 est, narrowly lanceolate (to almost linear), about A.M. the inner tepals reach their open 1 full Stem The at the widest, apices attenuate. terete, gen- position. tepals begin to collapse at about 2:30 erally with one branch produced in the upper third, P.M., and by 3:30 P.M. are closed together. Like drummondii^ A. A. veracruzanaisstM- relative its Infl> 4-5 rhipidia, single and terminal, usually -flowered; compatible. Self-fertilization often occurs In one or spathes green, attenuate, the apices generally dry two flowers of a rhipidium with resultant capsule mm 32-38 mner and seed production. As the stigmas are not in secund, contact with the anthers while the flowers are open, inner spathe, pollination presumably occurs after the flowers col- curved outward at the apex; perianth actinomor- lapse, when the parts of the flowers are pushed phic, pale to dark lilac-purple (rarely whitish), paler together by the inward-falling tepals. Outcrossing toward the center and lightly flecked toward the presumably the normal mode of reproduction for is mner with the species in view of the colorful and fragrant lis undulate margins and each The presumably serves with a pair of large perianth. occasional selfing white to yellow marks near the base of the limbs, to ensure seed production in the absence of insect- ± can sweet ob- mediated Plants of ^. t;eracr«za/ia pollination. , ovate, claws weakly and growing differentiated spreading be raised from seed to flowering in a single mm 17-20 outward, outer tepals long, the limbs season. Seeds sown in the greenhouse in late fall mm spreading, ca. 10 wide, inner tepals 16-19 in St. Louis grew to flowering size in about six limb months. right angles to the claws and with the lower lateral areas and median raised the MORPHOLOGY portion recessed. Sta- Karyology AND POLLEN mm mens dark purple, filaments 2 ca. long, broader veracruzana Chromosome number Alophia in and contiguous below, narrow in the upper half; pair one chromosomes, Four 28. long anthers coherent at their bases, with a broad nar- ^^ conlras submetacentric and the other acrocentric, rowly pandurate connective narrowest in the mid- much« mailer ^^^ mm ^^^'^^^ remaining s mm the 12 die %, weakly ascending, 4 LI ca. long, the e - mm The karyotype accords with that of pairs. wide at the base, nearly 2 at the apex, spo- IV -. members (Goldblatt, raploid of Tigridiinae rangia marginal and nearly latrorse; pollen yellow, & Benton Heywood, 1984). Ovary broadly sulcate. narrowly 4 obovoid, ca. mm ^^^ long, exserted from P°"^" ^^""^ ^ '""^'^ 7 the spathes; style 5.5 S'"^"'" ca. j^r IHblatl mm expanded^oo when ^^"^ as the grains fully ^ long, dividing opposite the middle of the an- * those^^ ^^ Thomas, and resemble in prep.) thers into three thickened branches, the branches ^^ mm drummondii '^^^^!'^'"' (Goldblatt, 1 974). erect, ca. 1 .5 long, each dividing into two long °„,ity of the mm correspond with those J ascending-recurved arms, '^^^"^ ^^"^^ these 3 ca. lone ^^ ^' (7I^"d'«' - of Tigridieae excluding Tigridiinae tubular, stigmatic apically, arching over the an- Cobana—Uo\see ^ and teria, Sessilanthera, Capsules thers. obovoid -oblong, trigonous and grams- have bisulcate 1970; Ravenna, which 1974), truncate obovoid-pyrlform, narrowest mi at the cropylar end, the chalazal end concave, Literature Cited Ughtly brown CaLDERON DE RZEDOWSKI, G. 1987- '^'S'f^^^^ie & 28 Howard (Goldblatt "^'^ 9070). una nueva de iridaceas nezii, especie „ ^ Mexico ^^[^ Hidalgo (Mexico). BoL Soc. Bot. '^^ maa bulbous Goldblatt. 1974. Revision of the P. Flower Phenology and Compatibility 373-385. of North America. Brittonia 27: ^^ Chromosome cytology "^^^ 1982. As . in Tigridieae, ^^^^y^. all the fugacious flowers last Iridace neotropical suprageneric systematics of befo Bot 186-198. 7: t^^^. r . oi fuU -. 1990. Phylogeny and classificatmn during Ann. Missouri Bot Herbert unfold, but until 7 A.M, the inner tepals remain n^- on (Iridaceae-Tigridieae): notes t\lh^ixisi closed c/vli^ Salp^^b nus and synonymy of reduction to " Number 4 & Howard 905 Volume Goldblatt 79, Alophia 1 Cardiostigma, Ilysa, and Catila. Ann. Missouri Bot, Ravenna, P. 1964. Notas sobre Iridaceae. Revista Inst. 504-511. Mun. 51-60. Card. 78: Bot. 2: & Henrich, E. P. Goldblatt. 1992. Iridaceae. In: 974. Cobana, a new genus of Central Amer- 1 J. . Flora Mesoamericana 1. (In press.) ican Iridaceae. Bot. Not. 127: 104 108. & Kenton, A. C. A. Heywood. 1984. Cytological 1979. Ainea, a new genus of Iridaceae from . 467-469. studies in South American Iridaceae. PL Syst. Evol. Mexico. Bot. Not. 132: & 146: 87-104. RuDALL, P. A. Wheeler. 1988. Pollen morphology Kew 693-701. MoLSEED, E. 1968. Fosteria, a new genus of Mexican in Tigridieae. Bull. 43: Iridaceae. Brittonia 20: 232-234. Shinners, L. 1964. Tigridia purpurea (Herbert) Shin- 1970, The genus Tigridia (Iridaceae) in Mex- ners, comb. nov. (Iridaceae). Sida 1: 295. . t and ico Central America. Univ. California Publ. Bot. 54. ! I -I I ^ CHROMOSOME NUMBER IN Snow^ and Neil Peter Goldblatt^ TICODENDRON (FAG ALES, TICODENDRACEAE) Abstract number of Ticodendron incoe:nitum reoorted for the time 26, and the phylogenetic is first number ignificance of this within among Fagus, Castanea, After exhaustive collaboration several Castanopsis^ Cyclobalanopsis, & Qu (Hammel authors Burger, 1991) the monotypic i & family Ticodendraceae was described (Gomez- Hans, 1969; Raven, 1975; Morawetz Samuel, & Gomez x 21 22 Laurito 1991) accommodate Trigonobalanus has or P., to the 1989), while Mesoamerican tree Ticodendron incognitum Go- (Hou, 1971; Soepadmo, 1972). Colombobalanus & mez-Laurito Gomez P. (1989). Evidence from and Formanodendron, segregated from Trigono- & wood anatomy remain un- (Carlquist, 1991), leaf anatomy balanus by Nixon Crepet (1989), & may which (Hickey Taylor, 1991), anatomy and mor- counted Nothofagus, floral the family. in = x 13 phology (Tobe, 1991), pollen (Feuer, 1991), and merit own family (Nixon, 1982), has its & Within sieve element characters (Behnke, 1991) strongly (Armstrc ig Wylie, 1965; Ono, 1977). — We n.^ J /" DOcs/trr-iy/onpm/s.us form suggests its inclusion in Fagales. report for the Betulaceae Carpin• us, Ostrya, and = o x first time the chromosome number of Ticodendron a evidently paleodiploid group with distinct, = & incognitum 2n 26 and Betula,Alnus as discuss the possible (Raven, 1975; Love Love, 1982); 1984, number (Kovanda, phylogenetic significance of this within Fa- (Raven, 1975), and Dusckechia == may have gales. Pashuk, 1987) have x Corylus 1 4; = = 14 equire (records of 2 2fi ac = genus as Materials and Methods 11 the but there are records of in re & monogeneric The L5ve 1982). well Liive, (e.g., Root were tips collected at approximately 9:30 (Morawetz 20 x B^l^^oP^^^ae appear to have from specimen A.M. a single growing in the green- 21 in n , count of Samuel, 989) (the sn^gle 1 J^ houses at the Missouri Botanical Garden (Voucher: f may erroneous^ be W„™™./^ R.w„ 1979) Hammel & 17R^A MH^ Af.J fi.au5^raZma(Goldblatt, Rivera 17834, MO). After pretreat- Our Fagales. Qearly, x 3 infrequent in 1 is ment with aqueous -bromonaphthalene 4 1 for hours, x = suggests count ot 2n 26 from a single plant the were tips fixed in 3:1 ethanol (95%)-gIacial = order the 3 Ticodendraceae, so within for /^ 1 10% acetic acid. After hydrolysis HCl in for 7 no with codendron number only shares this minutes at 70*C, root tips were squashed FLP in fagus, a genus from which distinct. is it orcein (Jackson, 1973). basic x Although Raven proposed (1975) ^^ scenario for for Fagales, a plausible Discussion ^^^^^-''''^'J^, x on based evolution can be suggested - The = din 26 and small, suming x R- nremimed base numbe' '"^ & ^^n- uniform chromosomes relatively of Ticodendron weU (Goldblatt related Hamamelidales as corrobo & ^ n.cii dress, 1977; Morawetz Samuel, l^^^^^*"^* .v. J o^^ indicate the distinctness of taxon from reduction this other members dysploid early of Fagales, families of Fagaceae. = and sut*^*!"^^ 4 7 followed by a doubling to 1 , The Fagales (sensu Cronqust, 1981) have base seems us^^^ reduction and II »« ^^ to 13, 12, = numbers x e and 4 withm 8, 1 1 , 1 2, 1 3, 1 (Raven, 1 975). reasonable explanation for change = 13, Fagaceae x In 12 predominates, numbe occurring The in ibers of the order. base De{u.rt,nent of Biology. Washington • University, St. Louis. Missouri 63130. U.S.A. 63166 B. A. Krukoff Curator of African Botany. Missour. Missouri Botanical Garden, P.O. Box 299, Louis. St. Missouri Bot 907. 1992. _^^j~ -L-^i.n^#^i~\^ > J- Number Snow 4 & Volume 79, Goldblatt 907 Chromosome Number 1992 Ticodendron in & common W. to Ticodendron and Nothofagus, was most Hickey, L. D. Taylor. 1991. The leaf ar- J. = chitecture of Ticodendron and the application of achieved independently from x likely 14, a hy- foliar characters in discerning its relationships. Ann. supported by wood pothesis significant differences in 105-130. Missouri Bot. Card. 78: anatomy (Carlquist, 1991), reproductive mor- Hou, D. 1971. Chromosome numbers of 7a^(>«o6a/a- phology (Tobe, 1991), leaf architecture (Hickey nas Forman (Fagaceae). Acta Bot. Neerl. t;er/fci7/a/a We & 543-558. Taylor, 1991) and other features. note, 20; Chromosomal Haplo- J^^^^^^' ^- evolution in comment however, the by Tobe (1991: 141) that 1^^.^: ^ pappus Evo- gracilis: a centric transposition race. Nothofagus the best candidate comparing in is 243-256. lution 27: Fagaceae with Ticodendron, which at least su- Kovanda, M. 1984. Chromosome numbers selected is in 289-301. perficially similar in a number of important char- angiosperms Preslia 56: (2). & lOPB chromosome Love, A. A. Love. 1982. In: acters. number LXXIV. Taxon 31: 120-126. reports & lUPB chromosome Mehra, Hans. 1969. P. A. S. /«; LrrERATURE Cited XXL number Taxon 310 315, reports 18: & MoRAWETZ. W. M. Samuel. 1989. Karyo- & R. A. Armstrong, M. Wyue. A new A. P. 1965. basic J, 129 154 Pp. patterns the Ilamainelltlae. chromosome number logical in the family Fagaceae (Noth- in & Crane Blackmore Evolution, in P. R. S. (editors), ofagus), Nature 205: 1340-1341. Ilamamclidae, and History of the Systematics, Fossil Behnke, H.-D. 1991. Sieve-element characters of Ti- and 'Lower Hamamelidae'. System- Introduction codendron. Ann. 131-134. 1: Missouri Bot. Card. 78: Volume 40A, Clarendon Association Special Carlquist, Wood atics S. 1991. and bark anatomy of Ti- Oxford. Press, codendron: comments on Ann. Mis- relationships. Nixon, K. 1982. In support of recognition of the family souri Bot. Card. 78: 96-104. Kuprianova. Misc. Ser. Bot, Sci, Nothofagaceae Cronquist, An a. 1981. Integrated System of Clas- Amer. 162: 102. New sification of Flowering 2nd York Plants, edition. & W. Crepet. 1989. Trigonobalanus (Fa- L. New Bot, Card, Bronx, York. taxonomic status and phylogenetic relation- gaceae): rKUER, S. 1991. Pollen morphology and the systematic 828-841. Amer. Bot. 76: ships. J. relationships of Ticodendron incognitum. Ann. Mis- Chromosome some South numl>er Ono, M. 1977. in souri Bot. Card. 78: 143-151. agaceae >TI t. ^^LDblatt, p. 1979. Miscellaneous chromosome counts m Mag. Tokyo 90: 313 316. Angiosperms including new family and generic II, Chromosome numt)ors species Pashuk, K. T. 1987. in records. Ann. 856-861. Missouri Bot. Card. 66: Chernogora (Ukrainian Carpa- & of subalpine belt of P. K. Endress. 1977. Cytology and evo- 1069-1074. Zurn. 72: Bot. thians). 67- lution in Hamameiidaceae. Arnold Arb. 58: J. The angiosporm phylog- 1975. bases of Raven, H. P. 71. 724- Card. 62: Ann. MLssouri Bot. & eny: cytology. ComezLaurito, Gomez 1989. Ticoden- L. D. P. J. dron: new a tree from Central America. Ann. Mis- 265- nora 103 1972. Fagaceae. Pp. in SOEPADMO, E. souri 1148-1151. Bot. Card. 76: & Malesiana ser. 1, 7. 199L new Ticodendraceae: a . anatomy, morphology, Reproductive Tobe, H. 1991. family of flowering plants. Ann. Missouri Bot. Card. Ticodendron, Ann. Missouri Bot. and relationships of 78: 87-88. 135-142. Hammel, & Card. 78: W. B. G. Burger. 1991. Neither oak nor alder, but nearly: the history of Ticodendraceae. Ann. Missouri 89-95. Bot. Card. 78: ^-J^r - -

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