Palynotaxonomic studies in Tabernaemontana L. sensu lato (Apocynaceae)

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1 Grana ISSN: (Print) (Online) Journal homepage: Palynotaxonomic studies in Tabernaemontana L. sensu lato (Apocynaceae) Madeleine Van Campo, Siwert Nilsson & Anthony J. M. Leeuwenberg To cite this article: Madeleine Van Campo, Siwert Nilsson & Anthony J. M. Leeuwenberg (1979) Palynotaxonomic studies in Tabernaemontana L. sensu lato (Apocynaceae), Grana, 18:1, 5-14, DOI: / To link to this article: Published online: 01 Sep Submit your article to this journal Article views: 207 View related articles Citing articles: 10 View citing articles Full Terms & Conditions of access and use can be found at

2 Grana 18: 5-14, 1979 Palynotaxonomic studies in Tabernaemontana L. sensu lato (Apocynaceae) MADELEINE VAN CAMPO, SIWERT NlLSSON and ANTHONY J. M. LEEUWENBERG Van Campo, hl., Nilsson, S. & Leeuwenberg. A. J. hl.: Palynotaxonomic studies in Tnberrraemontana L. sensu lato (Apocynaceae).-Grana 18: 5-14, Uppsala 7 February ISSN The pollen grains of some genera related to Tabernaemontana (Apocynaceae) were examined by means of light and electron microscopy. Some genera ought to be included in Tabernaemontana with reference to the general pollen morphology as well as to the fine sculpture of the inner exine surface. Quelques genres d'apocynaceae proches de Tabernaemonrana ont tt6 examints en Mph et hleb. On suggere qu'un certain nombre de genres allits soient inch dans le genre Tabernaenionrana la suite de I'examen du pollen et notamment de la face interne de I'exinc. Key words: Tnbertiaetriontntrn (Apocynaceae), paly notaxonomy Madeleine Van Campo. Laboraroire de Palynologie.CNRS-EPHE, UniversitP des Sciences er Tecliniqttes, Montpellier, France. Si\cerf Nilsson, Swedish Mirseitm of Natrtral Iiistov, Palyiological Laboratory, S Srockholm 50, Sweden. Anthony J. hf. Leeuwenberg, Laboratorium wor P1antensystema:iek en GeogrnJie, Gen. Foulkesweg 37, Wageningen, Nerherlands. The genus Tnbertinetiiotitntza L. (Apocynaceae) has been subjected to various taxonomic treatments with reference to grouping and delimitation. In order to contribute further evidence to the taxonomic interpretations, Tnbenzaeniotztatza and some related genera were selected for a preliminary pollen morphological study. Special attention was moreover paid to the type species of the genus names as synonyms of Taberticiemontnnn by Pichon (1938a, b) and Leeuwenberg (1976). In connection with revisional work on the Apocynaceae, Pichon (1938cr, b) included some pollen morphological data which were further referred to by Erdtman (1952). Pollen morphology of some randomly chosen apocynaceous genera with special reference to fine sculpture of the inner and outer exine surface was recently described by Nilsson & Van Campo (1979). MATERIAL AND METHODS Pollen material from dried specimens was obtained from the following herbaria: BR, K, L, P, U and WAG. The material was acetolysed. For light microscopy the pollen samples were mounted in glycerine jelly and examined with a Leitz Ortholux microscope (objective x 100; N.A. 1.30). Where possible, ten measurements of pollcn were usually made; but fewer measurements were made of other pollen features. The photomicrographs were taken at a magnification of x For scanning electron microscopy (SEM) the pollen grains were coated with gold and examined with a Jeol JSM-35 electron microscope. Some of the grains were divided in halves for later examination of their inner exine surface. For transmission electron microscopy (TEM) the pollen grains were cut with an LKB-Ultrotome; the sections were post-stained with uranyl acetate and lead citrate. POLLEN DESCRIPTION Pollen grains 3 to 4- (rarely S-)colporate, radially symmetrical. isopolar, suboblate to prolate, 22-70x23-55 pm. The shape in equatorial view (plane shape) is usually oval with curved to straight Erdrmoii dleniorial Voliiiiie II Groirfl 18

3 6 Al. Viin Ctrrirpo el ill. sides. In some cases the exine bulges at the equator. Amb circular to triangular. Colpi 8-25 pm long. narrow (slit-like) with rounded or pointed ends, or wider (up to 5 pm) and oval to lanceolate in shape. Sometimes they are equatorially constricted. The endoapenures are situated in a continuous band-like equatorial zone varying in width between 2 to 12 pm, or in shorter, discontinuous areas. The endoapertural, equatorial zone (and areas) consists of thinner exine covered with granular elements and is delimited by thickened nexine margins (costae equatoriales). Exine I to 3 prn thick. Sexine usually thicker than nexine, smooth with perforations or slits, to finely reticulate. On the inner exine surface there are granules or wart-like thickenings which become coarser towards the poles. The exine is stratified into a tectum, a granular stratum and a foot layer (Fig. 1 H). INTER- AND INTRAGENEKIC VARIATION in the material examined a certain pollen morphological variation was observed. Apart from the great size range also shape differences were noted. The following taxa have relatively small grains (polar axis ca. 30 pm, or less): Tnbernnemontmin citrifolia (Fig. IA-G), T. discolor, T. elegntis, T. 1iystri.r (Fig. 5 A-C). T. riiarrritintin and T. sp. (J midncastrirni snccharotirtn (Fig. 4 G-H). T. flovicnns (Fig. 3 A) and Stermiadertin grotidiflorn (genus maintained by Woodson, 1928) have pollen grains exceeding 50 pm. In 7. snriariho small and large grains were mixed on the same slide. Tabeniaenrontorrn discolor, T. elegans, T. lieterophjlln, T. Iirrrnblotii, T. 1iystri.r (Fig. 5 A-C), T. peridrrlijloro. T. retiisn, T. sessilijolin (Fig. 6 E-G), and T. wntricosa (Fig. 6 A-D) have prevailingly spheroidal pollen grains while T. albflorn, T. coriacea. T. riitirrlosn (Fig. 3 B-E), T. srrbmollis. T. rrridrrlata (Fig. 2), T. sp. (Phrissocarpirs rigidirs) and Steniniodenia grandiflorn are usually prolate. The other species were trdnsitional as to size and shape. As indicated above the equatorial zone varies in width outline and continuity. It is evenly wide and consistently continuous in the following species, and range between (2)4-6(12) pm: T. albdlorcr, T. nirrnntioca, T. citrifolin (Fig. 1 A-G), T. corincetr, T. discolor, T. jlaricnns p.p. (Fig. 3 A), T. riniitlosn (Fig. 3 B-E), T. snrin~iho, T. sirbniollis. T. rrrrdirltrto (Fig. 2). T. sp. (Plirissoccrrprrs rixidrrs) 2nd Steninindeliin grancirflora. In the other species the equatorial zone is variable. The exine is perforate to finely reticulate in T. coronoria p.p. (Fig. 4 A-F) and T. cliclrotorrm p-p. (Fig. 5 D-F). otherwise smooth. DISCUSSION Pichon (l9.%7: 233) gave a fairly detailed description of the continuous equatorial zone (ceinture) of rig. 1. A-G 7ciberricie,tiorittrritr cirrijolin. (A) Pollen grain in equatorial view tvith a short colpus, a continuous equatorial band-like zone with thickened margins. and fine. indistinct granules inside the grain. (B) The same in lower focus. (C) The same in optical cross-section. Note the markedly thickened nexine at the equatorial zone. (D) A tetracolporate pollen grain in polar view. high focus. (E) The same in optical cross-section. A-E: Lhl x I OOO. (F) Detail of the outer smooth and perforated exine. SEhL Xca (G) A grain in slightly oblique equatorial view. The exine is markedly smooth and bulges at the equator due to the thickened margins of the equatorial zone. There are numerous minute perforations visible particularly towards the poles. The colpus membrane is granular. SEhl x ca (H) Tuberrioemorirann sirhniollir. Section of the exine divided into a smooth tectum (1). a gnnular layer and an unevenly thick, irregular footlayer V, which forms the granular to vcrrucose inner exinc surface. TEhl x ca Fig. 2. Tnbertinenioritnnn rmdrrlnrn. (A) 3-colporate pollen grajn showing one colpus and-the continuous cndoapertural equatorial zone in high focus. (B) The same in lower focus. (C) The same in optical cross-section. (D) A gnin showing part of the coarsely granular equatorial zone between two colpi. (E) The same in lower focus. A- E are all in equatorial view. Note the markedly thickened nexine margins that border the equatorial zone and the distinctly granular inside of the grain. (R A pollen grain in polar view in relatively low focus. Note the granular inner exine pattern. (G) The same in optical cross-section. AG: Lhlx 1 OOO. (H) A grain showing a short colpus with gnnular membrane, and a smooth, perforated exine surface. SEhi X ca (I) Detail of the partly distorted and ruptured exine surface between two colpi. A finely radiating surface pattern is evident near the ruptures. SEhl X ca Fig. 3. (A) Tnbernneniontana fluricans. 4-colporate pollen grain in equatorial view with two colpi visible and the continuous equatorial zone. There are fine, densely spaced granules inside the exine. Lhl X 1 OOO. B-E T. rinirrloscr. 3-C. Two tetracolporate pollen grains in equatorial view. SEhl X ca (D) One half of a cut pollen grain showing the granular to vermcose (towards the poles) inner exine surface and the equatorial zone. SEhl x ca (E) Detail of the inner exine and the granular equatorial zone. SEM X ca

4 Fig. I

5 Fig. 2

6 Fig. 3 Palytiotnxotiottiic stiidies iti TnDertinet?iotitrrtiu 9

7 10 hf. Vori Cnrtipo c f ti/. Fig. 4. A-F ~n6~r~nett1orif~i~~~1 coromrio. (A) A perforate (F) A tetracolporate grain in polar view. optical crosslo finely reticulate pollen grain in equatorial view, rela- section. A-F: Lhlx I OOO. G-11 f~nrlncnsfrrrni SUCtively high focus. (B) The same in lower focus. (C) The cliurafiori. (G) a granular to verrucose grain slightly obsame in optical cross-section. (D) A 3-colporate pollen lique in equatorial view. SEhl x (H) Half of a pollen grain in equatorial view, high focus. showing one colpus grain showing two discontinuous thin. elongate areas, subtended by a thin, elongate, lanceolate area (discon- almost fused to form ;in equatorial zone; a granular to tinuous equatorial zone). (E) The same in lower focus. verrucose inner exine surface. SEhl x 2300.

8 Palytiortrroiioiiiic sriidies iti T~iberiiaeiiiontaiin 1 1 Grann I8

9 Fig. 6 Grimti 18

10 PciI~riofn.rorioriiic studies iri Tciberricierrioritnrici I3 pollen in Tabernaemontaneae. He pointed out that the American Tabernaemontanene have pollen grains with a continuous and equally \vide endoopertural zone, while those of the Old World have uneven or interrupted zones. He concluded, ho\vever, that this distinction was not absolutely consistent on a specific level. In another work (Pichon 1938b) Poginntlin and Rejoira were described as having pollen grains without endoapertural zone (ceinture) while the other genera, e.g. Encitnrriin, Hazirri~ci, hfrrritafora, I'nridcicnstrirni, Stenittinderria and Tciberrinerrioritarin all had such a zone, if not always distinct. Erdtrnan (1952: 50) noted that the inner exine surface of the Apocynaceae was often uneven. Zonorate griins were met with in Tabernaemontaneae. As regards apertures, size and shape, he pointed out that those characteristics were as a rule consistent in taxa supplied by Pichon from hladagascar. Sculptural elements on the inner exine surface in Apocynaceae have been described and depicted in various connections by, e.g., Van Campo (1971). Schill & Leuenberger (1972) and Nilsson & Van Campo (1979). In the taxa studied here such ele- Fig. 5. A-C T~bef/J~Jet/io/Jf~f/la hysrrix. (A) A pollen grain in equatorial view showing one colpus subtended by an elongate, lanceolate area. (B) Another pollen grain in the same view, in optical cross-section. (C) A grain in polar view, in optical cross-section. A-C: Lhl X 1 OOO. D-I: T. diclrorom. (D) Half of a cut pollen grain showing two thin, elongate, non-fused areas, a finely to coarsely granular (venucose) inner surface. and a portion of the outer smooth, perforate exine. SEhl x ca (E) A pollen grain slightly oblique in equatorial view. SEhl x ca (F) Part of the smooth exine surface with perforations of varying size and shape, either round or slit-like. SEhl X ca Fig. 6. A-D ~ubefrr~e~~iorrrunu wtrlricosu. (A) A grain showing a narrow colpus subtended by a thin oval area (B) The same in lower focus. Note the fine granules inside the grain (C) A grain in polar view. high focus with numerous granules inside. (D) The same in optical cross-section. A-D: Lhi x 1OOO. E-G T. sessilifoliu. (E) A tetracolporate pollen grain in equatorial view with a rough, uneven, and densely perforated exine surface. SEhl x ca (F) A tetracolporate grain in polar view. The polar and colpal areas are smoother than the mesocolpia. SEhl x ca (G) Part of a colpus and adjacent margin. The colpus is of irregular shape with a jagged margin. a coarsely granular membrane delimiting the endoaperture. The granules on the f smooth colpus margin seem to be extraneous though they may derive from the colpus membrane. The areas around the perforations appear slightly elevated. SEhi x ca ments were consistently found. The exine fine structure of Neriirrri olenritler ins described by Van Campo & Lugardon (19731, and of Aiinccrrriptn siibrriollis (= Tnborriaerriortnrin sribrriollis) by Nilsson & Van Campo (1979). In the present stage it is difficult to evaluate the observed inter- and intraspecific variations and to dmw conclusions of decisive taxonomic significance. In agreement with Pichon (19380, b) \ye confirm that T. dichofor~icr (Pcigiciritlin clicliotorrr(r) lacks "ceinture". As regards Kejoirci, however, our observations differ from those of Pichon. but he never examined T. tir~rtiiiriacn (Kejoiiri) included in our study. We also disagree regarding T. coroiinria (Eriwfciriiici), 7. elegnris (Lepfophnryrigia), r. refirsa (Paridaca), T. sessilifolici (hfiiritufara), T. \witricosn (Sarcoplinryigici) and T. sp. (Paridncasfrrrni sncclicircifirrri). In all other cases we can fully support I'ichon's results with reference to the equatorial zone. From a pollen morphological standpoint we provisionally suggest that T. cilbiflora, T. nirrcirificicci, T. coriciceci, T. discolor, T. Jnvicnris, 7. ri~tirilosn, T. striicirilio. T. srrbniollis, T. irriclirlcifci. T. sp. (I'lirissocnrpris rigidirs) and Stcnininderiio grow diyorn be regarded as more closely related to T. citrifolici (type species of Tnberrineriioritoricir; cf. Leeuwenberg 1976) than to the other species examined. In order to express a more definite view of possible relationships within the Taberrincrriorimiin-complex far more material would, however, be required. ACKNOWLEDGE hl EN TS We are grateful to the Directors of the herbaria in Hrussels. Kew, Leiden, Paris. Utrecht and Wageningen for supplying material for this study. For expert technical assistance we are thankful to hlr J. hl. Fillols, hlontpellier and hliss Elisabeth Grafstrom. Stockholm. Linguistic revision was made by hlr Alan Dixon, Stockholm. SPECIhlENS INVESTIGATED * illustrations ~ubcrriuerfionlnncr dbi/loru (hliq.) Puk (type species Of Tubernu hlarkgraf. not hliers). - Surinam, Versteeg 2s g (U). T. nirruniiucu Gaud. (type species of Rejoita Gaud.). - New Guinea. Kostermans 2917 (L); v. Royen (L). T. cirrifoliu L. (lectotype species of Tubrrnacnmnfann L.). - Guadeloupe, Proctor (K); Sastre 2737 (P). Santo Domingo, v. Tuerckheim 2818 (U).* Erdtrirtrii,\fet~ioritil Yoliriiw I /

11 14 rzt. V[ttl cntirpo El trl. T. coriacea Link ex K. & S. (lectotype species of Atiacampta hliers). - Brazil, Spruce 675 (1'); Trail1 514 (K). 7. coronnria (Jacq.) Willd. (type species ofeniiruuiiir (A. DC.) Stapf). - India. hlasters s.n. (L). Indonesia. LBrzing I1973 (L). Thailand, Zimmerman 16 (U).* T. dicliotonio Roxb. (type species of Pagiantlia hlarkgnf). - India, Wallich 1871 (K). Formosa, Chang 3159 (L).* T. discolor Sw. (lectotype species of Tubernu hliers, not hlarkgraf). -Cuba, Rugel 263 (L). T. elegnns Stapf. (type species of Leptopharpngia (Stapf) Boiteau). - hloqmbique, Le Testu 484 (P). T.Javicnns R. & S. (lectotype species ofannrtia hliers). -Brazil, Harley C.S (U).* T. lieteropliyllu Vahl (type species of Stenosolen hlarkgraf). - Brazil. Krukoff 1643 (U). Surinam. Lindeman & Teunissen L.B.B (U). T. hunrblotii (Baill.) Pichon (type species of Oclironeriiini Baill.). - hladagiascar, Humblot 435 (P). T. hpstrix Steud. (type species of Peschiera A. DC.). - Brazil, Pabst 7138 (K); Saint-Hilaire B 19 (P).* Indonesia, W. Java, Koerniasih 37 (L). 7. titurtririnna Poir. (type species of Oistunthera hlarkgnf. later in Pmidnca with hlarkgraf). - hlauritius, Lorence h158 (WAG). T. modesta Bak. var. nietlirienii (Stapf & L. Green) Pichon (T. niodesta is type species of Hozirririi Pichon). - hfadagascar, Sew. Forest. 49(P). T. pendiiliflora K. Schum. (type species of Camerunia (Pichon) Boiteau). -Cameroon, \V. de Wilde & B. de Wilde-Duyfjes 1%7 (WAG). T. rerrisa (Lam.) Pichon (type species of Pandacii Noronha ex Du Petit Thouars). - hladagascar, Geay 6775 (P). T. rimilosn Woods. - Venezuela, hlaguire. Wurdack & Keith Keith (K).* T. sananho Ruiz & Pavon (lectotype species of hferizudeniu). - Colombia, Plowman, Lockwood, Kennedy & Schultes 2371 (K). T. sessili/olin Bak. (type species ofafuntofura Pichon). - Madagascar, Bosser 8516 (P);* Humbert 6436 (P). T. srrbniollis hlart. (Anucanipfa with hliers). - Brazil, Prance & Pennington 1985 (K).* T. undiilata Vahl (type species ofbonnfoitsia A. DC.). - Fr. Guyana. Sastre & hloretti 3948 (P). Surinam, Boswezen 5470 (U);* Roberts L.B.B. I1219 (U). T. ventricosa Hochst. ex. A. DC. (type species of Snrcophary@a (Stapf) Boiteau). - Tanzania, Semsei 1917 (BR).* Zaire. Corbisier 105 (BR). 7. sp. (Pnndaciisrrioii socclinrotirrri Pichon). - hladagascar. Cours 2587 (P).* T. sp. (Phrissocarpirs rigidits. later in Anocnnrpta with hiarkgraf). -Brazil, Sandeman 2164 (K). Related species Stemniudenia graridiyora (L.) hliers. - Surinam, Lanjouw & Lindeman 1371 (U). REFERENCES Erdtman, G Pollen morphology and plant taxonomy. Angiosperms. - Almqvist & Wiksell. Stockholm. Index Herbariorum 1Y74. Part 1. Herbaria of the world. - Oosthoek, Scheltema & Holkema, Utrecht. Leeuwenberg, A. J. hl The Apocynaceae of Africa. 1: Tabernaemontana L. 1. Introductory remarks to a revision of the species represented in Africa. - Adansonia ser : Nilsson. S. & Van Campo, hl Pollen morphological' studies in the Apocynaceae with special reference to tine structure. - Proc. 4th Int. Conf. Palynol., Lucknow 19?6/77 (in press). Pichon. hl. 1948n. Classification des Apocynackes: VI. Genre "Tabernaemontana". - Not. Syst. 13: Pichon, hl Classification des Apocynacies: IX. Kauvolfiies, Alstoniies, Allamandies et Tabern& montanoidkes. - hlh. hlus. Natl. Hist. Nat. 27: Schill, R. & Leuenberger, B Pollenmorphologie und Cytotaxonomie der Gattung Pachypodium Lindl. - Rot. Jahrb. 92: Van Campo. hl Precisions nouvelles sur les structures comparfes des pollens de Gymnospennes et dangiospermes. - C.K. Acad. Sc. Pans Van Campo, hl. & Lugardon, B Structure grenue infratectale de I'ectexine des pollens de quelques Gymnospermes et Angiospermes. - Pollen Spores 15: Woodson, R. E Studies in the Apocynaceae. 11. A revision of the genus Stemmadenia. -Ann. hlo. Bot. Gard. IS: Grrrriii IR

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