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Oldenlandia Umbellata L

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Oldenlandia Umbellata L 429 HETEROSTYLISM IN OLDENLANDfA UMBELLATA L. BY BIR BAHADUR Deflartment of J3otany, Osmania University, Andhra Pradesh, India Remarkably little is known about heterostylism in tropical plants, and I know of no studies of natural populations comparable with those of Schoch-Bodmer (1938) on Lylhrum salicaria L., Haldane.(1938) on Primula acaulis and Crosby (I949) on Primula vulgaris. Prof. J. B. S. Haldane suggested that I might make such a study on one of the Rublaceae, a family in which heterostyly is widespread , as Darwin (1892) pointed out in his classical hook, The differe~ztforms offlowers on pIants of the same species. The genus Oldenlandia is represented by 200 species (Rendle, 1959) out of which 21 species display dimorphic heterostylism. Bremeka~p (1952) has monographed the -Aft-lean species of.this genus, in which he has shown the distribution of heterostyly and homostyly. Bremekamp (1956, 1958) reported some more species which are also heterostyledl Oldenlandia umbellata L. shows dlmorphic heterostylism, i.e., two types of flowers occur on separate pIants, conventionally termed as "Thrum eyed", with short style and long anthers, and "Pin eyed", with long style and short anthers (see Plate t5). OIdenlandia umbellata is common in the neighbourhood and accessible to me. It is a conspicuous white flowered annual, and a successful weed. At Adigmet (Osmania University Campus) the popu]atiOn was dense over a large area. At I-Iimayat Sugar and Miraulam it was distributed over w~ste land and road sides, sometimes growing in sand or in rock crevices. At 1Narsapur it was not common in lawns, but occun'ed along-dry water courses, confirming Ridley's (t930) statements as to the dispersal of its seeds by water. Table 1 is a list of all species of 1Lubiaceae kn.own to me in which heterostylism has been reported. These number 155 out of a total of about 5,500 species in the family. Table I, Recorded cases of Heteros~lism i,~ the family .R'ubiaceae S. Species Tribe Floral References No. situationJ" 1. *Cinchonamicrantha. Cinchoneae. D Darwin (t892), East (1940) 2. Sipania sp. Rondeletieae. D Verdcourt (1958) 3. Pentaslanceolata. I-tedvotldeae. T or D Verdcourt (1953d),Baker (1958), - Bir Bahadur (Pin form only) P. pl~rseglovei. ,, D "~lerdcourt(1953c1), Baker (1958) P. ct.eisostoma. ,, D Vcrdcourt (1953d} P. parviflora. ,, D "Verdcourt (1953d), IIiern (1878) P. arvensis. ,, D tliern (1878) P. zanzibarica. ,, D Verdcourt (1953d) p. l~ub~'t~ra. ,, D ,, p. decao,ana. ,, D ,, P. pseudom.agnifica. ,, D quoted by Verdeourt (1953d) P. jn~rlmrea. ,, D I-Iiern (1878) '*East (1940): 4.0 ~pecies are heterosilylic, ID Dimorphic, T Trirnorphic. 430 I-Ieterostylism in Oldenlandia umbellata Table 1. Recordedcases of Heteros~ylism in the family Rubiaceae--((~,ontd,) S. SpEcies Tribe Floral Rc.fe• _.No. - situation .?entas nobilis. Hedyotideae. D Verdcourt (1953d) P. longiflora. ,, T or D 2":'. cocchwa. ,, D quoted by Baker (1958) 4.. 2:'arapentas balllscombei. ,, D Verdcourt (1953c) 2. setigera. ,, T or D 2 ~. gabonica. ,, D Bremekamp (1.952) 5. ?~plnopentas tat~folia. ,, D Verdcourt (i953c) 7-. ul,~gurica. , D :'.'_-. cameronica. ,, D Bremekamp (I952) ',5. J~,:ntodonpentander. ,, D P. laurentioides. ,, D 7. Dibrachi~zos~yhzs kaessnerL ,, D ~. Otomeria gMneensis. ,, D Verdcourt (I953a) O. lanceolata. ,, D ;J O. maNensis. ,, D O. oculata. ,, D 9. t:hamaebenlas greenwayl. ,, D Brcmekamp (1952) O. 5;.'.cosl~erm~ sp. ,, D t 1. ,57ephanococcus crepinianus. ,, D 12. ..~])moslad~ya s~. ,, D sJ !3. I-{edythyrsus sb. ,, D J :t'. A gathisanthemum bojeri. ,, D A. angolense. ,, D A. chtorophyllum ,, D A. quadricostatum. ,, D ~J !5. .~ionitis s~. , D i 6. Amphiasma luzuMdes. ,, I) A. redheadii. ,, D A. robijnsii. ,, D A. bengueUense. ,, D A. merenkyanum. ,, D :i. di~a~'icatum, ,, D ! 7. Lelya osteo~arpa. ,,, D [8. 2-~&iocraterium verlicillare. ]l, DOI Bremekamp (1939) t9. 2Dirichletia slZ r D Hiern (1878) 20. 2~e~,2anisia sp. o, D 2 I. 57&robombyx kinabdunlsis. ,, D Bremekamp (1947a) 22. Danais sp. ,, D Verdcourt (I958) 23. 2~oustonia caerule& ,, D Darwin (1892), Stevens (I912), ]Robinson and Fernald (1908), Lewis (I961a) 24. ~4 ~otis nummzdarla. ,, D Bond (1953) -d. richardiana. ,, D 2~. Schismatodada grichopharynx. ,, D Verdcourt (1958) 26. O ldenlandia cryptoca~pa. ,, D Bremekamp (1952) Ct. tzervosa. ~ D O. chevaliieri. ,, D O. ocellata.. D O. aretioides. ,, D O. pulvh~ata. ,, D O. monanthos. ,, D O. hockii. ,, D O. geophila. ,, D O. frieslorum. ,, D O. greenwa)~i. ,, D ~z O. johnstoML ,, D O. "ru]~icola. ,, D O. rattscosa. , D O. tenella. ,, D O. sipaneoides. ,, D O. ta~davdina. ,, D BIa Ba~AI~Ua 43t TabLe 1. Recorded cases of HeterostSism i~ the family R'ubiaceae--(Contd.) S. Species Tribe Florat References No. situation Oldenlandia taborensis. Hedyotideae D Bremekamp (1952) O. duemmeri. ,, D ,, O. scopulorum. ,, D ,, 0 wiedemanii. ,, D ,, O. icthyoderma. ,, D ,, O. herbacea var. holstli. ,, D :, O. pringlai. ,, D ,, O. sax~fragoides. ,, D ,, O. rosMala. ,, D , O. microaa~x. ,, D ,, . O. q~'inis. ,, D Bremekamp (I952), Baker (1958) O.flosculosa. ,, D Bremekamp (1952) O. patula. ,, ]3 ,, O. umbellata. ,, D Bremekamp (1952), Bit Bahadur, .Lewis (1961a) O. somala. ,, D Bremekamp (1952) O. eludens. ,, D ,, O. marginata. ,, D Bremekamp (1958) O. oxycoccoides. ,, D ,, O. vertidllata. ,, D Bremekamp (1952) O. filifies. ,, D Bremekamp (t956) O. aapensis. ,, D Post (1932) O. sd~imfieri. ,, D Crowfoot (1928) 27. Hedyotis sp. ,, D Darwin (1892) H. n~gricans ,, D Lewis (I 961 b) H. humifusa ,, D ,, I-Z. purpurea. ,, D ,, 28. Bo~tvardia Mantha. D Darwin (I892) 29. Knoxia sp. Knoxleae D Darwin (1892), East (1940) tO. mollis. ,, D Fyson (1932) K. platycarpa. ,, D Bond (1953) 30. Cmckshankia gladatis. Cruekshankleae D Skottsberg (1915) C. lo,menodon. ,, D Verdcourt (1958) 3 i. 2-vlussaenda e~ythrophylla, i'vlussaendeae. D Baker (1958) A,ir. degans. ,, D Baker ( 1958 ), Hall~ .( 1961 ) M. temdflora. ., D Petit (1954) Mussaenda sp. ,, D Bremekamp (1947b) M. chippi ,, D Hail6. (196I) eo~throphylla ,, D ,, 32. Pseudomussaenda stenocarpa. ,: D Petit (1954); Hall6 (1961) l~.flava. ,, D Petit (1954) 33. Gouldia s#, ,, D ~, 34. Adenm'acme lo~lgi.folia. ,, T or D Darwin (]'892), :Hooker (I882) 35. Aphaenandra unlftora. ,, D Bremekamp (1937) Aphaenandra sp. , D t-l"all~ ( 1961 ) 36. Coccocypselum ~p. ,, D Darwin (1892) 37. Pauric{iantha sp. ,, D Verdcourt (1958) P. hirtella. ,, D I-Ial16 (196I) 38. Sablcea venosa. ,, D ,, S. hierniana. ,, D ,, 39. Stipularia africana. ,, D ,, 40. Ecpoma apocynaceum. ,, D ,, 41. Temnopter),x sericea. ,, D , 42. Pentaloucha sp. , D ,, 4,3.' ,gacosperma paniculala ,, D ,, 44. Anthospermum herbaceum. A~lthospermeae D Verdcourt (1958) 45. Coprosma sf~. ,, D Darwin (1892) 4"6. Mitchellarepens. ,, D Darwin (1892), i%obinson and Fernald (1908) 47. Paederia sp. Paederieae D ~Verdcourt (1958) 48. Tricdysia sp. Ixoreae D Hiern (1878) ,2t 432 Heterostylism in OMenlandia umbellata Tabte I. Recordedcases of Heterostylism in the family Rubiaceae~(CcJztd.) s. Species Tribe Floral References No. si tuatioa 49. Craterispermum lauKmm~. C,raterispermeae D Verdcourt (1958) 50. Rudgeajasminoides. Coussareae D Baker (1956) J~. coriaeea. (R. erlantha.) ,, D Baker (1956) "Darwin's (1892) Faramea spY' 51. Psychotria vogeliana. Psychotrleae D Baker (1958) l~o,cholrla ~b. ,~ D Darwin (1892) P, benlhamiana. ,, D Baker (1958) P. warnecM. ,, D P. malyana. ,, D Ernst. (t932) P. elongata. ,, D Fyson (1932) 52. Uragoga nimbmm. ,, D SehneI1 (1953) quoted by Baker (1958) U. peduncularis. ,, D Baker (1958) 53. Lasianlhzts c@itMazus. ,, D Bir Bahachu" (Pin form only) 54. l~ap?uria s/~. ,, D Bremekamp (1961) 55. Grumilea sp. ,, D 56. Geophila sp. D Baker (1958) (Pin ibrm only) 57. Borrerie sp. Spcrmacoceae D Darwin (1892) 58. Diodiasp. ,, D Darwin (1892), Baker (1958) 59. Spermacoce sp. ,, D Darwin (t892) 60. Selonocera sp. ,, T Willis (193I) 61. Suteria ssO. :, D Darwin (I892) 62. Lipostome sp. ,, D 63. Schwendenera sp. ,, D East (1940) 64. Perama sp. D 65. Nertera sp. '~ D Darwin (1862) 66. 3/Ianettla sp. ~ ? D ,, During September and December i960, the collections of plants were made in the localities mentioned in Table 2. As the species is a strict-annual, there was never any doubt as to What constituted a single ;plant, as there ds in perennials such as Primula acauIis and Lythrum salicaria. I carried two bags, into one of which I put long styled plants into the other short styled plants. The counts are given in Table 2. There is an excess of "Pin" or long styIed plants, which is highly significant (2~=44.7). But the different i;opulations are extremely heterogeneous (X~----72.37) and it is quite possible that "Thrum" or short styled plants might predominate in a larger total sample, as they did in two of my six. This inequality or anisopIethy and heterogeneity is very different t)om Haldane's (1938) finding in Primula acaulis. Mention may be made of Faberg~ (1959)who recently counted populations of Oxalis, k trimorphic genus, tie counted the populations of Oxalis grandis and Oxalis suksdo~fii, in which he found anisoplethy as against isoplethy (the equiIibrium condition according to certain theories). His counts are given below (Table 3). To investigate the mode of inheritance of style length, iong and short styled plants Were crossed and selfed in the combinations shown in Table 4. The cross P • is more fertile than T • P as deduced from the greater seed set per capsule, Selfing pin and thrum produced very few- or no seeds. Occasionatly long styled plants set .2-5 seeds. Thus the incompatibility in thrum is absolute, and in the case of pin is relaxed a litt!e. BIR BA~AD~JR 433 Table 2. Counls of'Natural populations of Oldentandia umbellam Locality Pin Thrum Total % of % of eyed eyed Pins Thrums I. Adlgmet .... 337 150 487 69.20 30.80 2. s .... 146 112 258 56.59 43.4I 3~ Narsapur .... 1i 1 133 244.
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