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1 This article was downloaded by: [ ] On: 26 April 2015, At: 06:56 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Grana Publication details, including instructions for authors and subscription information: Pollen morphology of the lianescent species of the genus Croton (Euphorbiaceae) Lea Maria Medeiros Carreira a, Ricardo De Souza Secco a & Ortrud Monika Barth b a Department of Botany, Museum Parense Emilio Goeldi, C.P. 399, Belem, Para b Instituto Oswaldo Cruz, C.P. 926, 20001, Rio de Janeiro, Brazil Published online: 01 Sep To cite this article: Lea Maria Medeiros Carreira, Ricardo De Souza Secco & Ortrud Monika Barth (1996) Pollen morphology of the lianescent species of the genus Croton (Euphorbiaceae), Grana, 35:2, 74-78, DOI: / To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the Content ) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at
2 Grana: 35: 74-78, 1996 Pollen morphology of the lianescent species of the genus Croton (Euphorbiaceae) LEA MARIA MEDEIROS CARREIRA, RICARDO DE SOUZA SECCO and ORTRUD MONIKA BARTH Downloaded by [ ] at 06:56 26 April 2015 Carreira, L. hl. hl., Secco, R. de S. 8: Barth, 0. hl Pollen morphology of the lianescent species of the genus Crotoiz (Euphorbiaceae). Grana 35: ISSN The pollen morphology of lianescent species of the genus Crotoii L. was analysed with regard to the size, number of apertures, shape and ornamentation of the exine. The pollen grains are apolar, atreme and spheroidal. They vary from medium to tery large in size nith croton- and reticulate pattern with supramural elements (clavae). The species are di\ided into two groups: ( 1) exine ornamentation with psilate sexine elements (C grazielae) and (2) exine ornamentation with striate elements (remaining species). The latter group can be subdivided according to the formation or not of a croton-pattern by the sexine elements and by its ornamentation. Lea hlaria Aledeiros Carreira arid Ricardo de Soim Secco. Deparrrrierir of Botaiij, Aliiseiiiii Paraise Eiiiilio Goeldi, C. P. 399, Beleii~, Para; Ortnid Aloriika Barrh. bisritiiro Ositaldo Crii:, C. P. 926, Rio de Jaiiciro, Brad. (Afaniiscript accepted 20 hlorcli 1996) Crototi L. is one of the largest genera of the Euphorbiaceae, in terms of number of species (about 1000, according to Webster & Burch 1968) and requires a detailed up-to-date taxonomic review, since it has only been comprehensively studied by Mueller (1866, 1873). The majority of its species occur in the Antilles and South America. Numerous authors, e.g., Croizat (1940) and Hutchinson (1969) have recorded the type of habit of the species of Crotori as a tree, shrub or herb, but there is no mention in their works of its lianescent habit. Lanjouw (1931) cited Crotori pirllei as the first species of the genus to feature the lianescent habit. Subsequently he described the variety glabrior to be a tree (Lanjouw 1939 a) and suggested that C. pullei must also have been a tree; this correction was then published (Lanjouw 1939 b). While carrying out field work in the Municipality of Peixe-Boi (Para), Secco & Rosa (1992) collected specimens of Crotori pullei, which confirm the lianescent habit of the species. In the MG and IAN herbaria the lianescent habit was only found for the species C. pirllei var. glabrior. Secco (l992), in an effort to contribute to the clarification of the complex taxonomy of the genus, provided an up-todate discussion on the lianas of the genus, including C. pullei and C. pirllei var. glabrior and described the new amazonian species C. jararisaisis, C. grnzielae and C. peririietralerisis, which are h as. While carrying out field works in eastern Amazonia (Paragominas, Para) Secco & Rosa (1992) described C. ascertderis, another species of lianescent habit. Recently, Webster ( 1993) established a provisional revision of the sections of the genus Croton. With regard to pollen morphology of the genus Crotoit, Erdtman (1952) analysed the structure of the exine in detail, describing the structure of the croton-pattern referring to Crotori arboreits. Punt (1962) studied the pollen grains of C pirllei; Salgado-Labouriau ( 1973), analysed the pollen grains of C. agrophilirs, C aiitisjphilitictrs, C. paiiliiiiis, C. parafinis, C. polilinriirs and C. sc1erocalj.s var. interinediirs and Thanikaimoni et al. (1984) discussed the morphology of the pollen grains of C. niatoirrersis, C. erytliroxjloides, C. te-wrnis and C. gratissiimrs. Punt (1987) stated that in relation to the classification system of Webster (1975), the genus Crotori is included in the Crotonoideae because of the specific feature of the Crotori-pattern of the pollen sexine. Nowicke (1994), in a palynological study of the Crotonoideae subfamily, analysed the pollen grains of C. argjratiis, C. balsnriiifer, C. cnliforriicra, C. draco, C. floribimidiis, C. lochnocarpus, C. niatoirrerisis, C. iiiirtisiaiiirs, C. pjrnniidalis and C. texerzsis, without making any reference to the lianescent species of the genus. Since the pollen morphology of the lianescent species of Crotori has not yet been sufliciently investigated, we present a study in order to complement the taxonomic information provided by Lanjouw (1939 a, b) and Secco (1992). MATERIALS AND METHODS Adult flower buds were remowd from specimens of the hiuseu Paraense Emilio Goedi (hfg), CPATU/EhfBRAPA (IAN) and Utrecht (U). The pollen grains nere processed according to the acetolysis method of Erdtman (1952) for light and scanning electron microscopy. The terminology follows that of the glossaries of Barth &: hlelhem (1988) and Punt et al. (1991). The size classes are in accordance with Erdtman (1952). RESULTS Pollen descriptions Crotori sp. (Fig. 1 A-D). - Pollen grains large, diameter 54k2.1 (48-68) pm, inaperturate, spheroidal, with a dense croton-pattern. In SEM the sexine (3.1 pm thick) consists of mqre or less elliptical, striate elements (clavae), with an apical spinule. Grano 35 ( 1996) Scandinavian Uni\ersity Press. ISSN
3 Pollen of Crotori Iiatias 75 Downloaded by [ ] at 06:56 26 April 2015 Crotori aseerideris R. Secco & Rosa (Fig. 1 E-H). - Pollen grains large to very large, diameter 98k1.9 (82-112) pm, inaperturate, spheroidal, with croton-pattern consisting of irregularly and sparsely distributed elements. In SEM the sexine (3.5 pm thick) consisis of distinctly striate and subtriangular elements, (clavae); an apical spinule is not evident. Small, psilate, clavate processes are found in between the larger elements. Crototi grazielae R. Secco (Fig. 1 I-L). - Pollen grains medium to large, diameter 50f 1.2 (45-55) pm, inaperturate, spheroidal, with croton-pattern. In SEM the sexine (3.5 iim thick) consists of rounded, psilate elements (clavae) without an apical spinule. Small, psilate processes occur in between the clavae. Crotori jararisaisis R. Secco (Fig. 2 A-D). - Pollen grains of medium size, diameter 48 ko.9 (45-51) pm, inaperturate, spheroidal, with croton-pattern. In SEM the sexine (2.6 pm thick) consists of rounded, striate elements (clavae) without an apical spinule. Small, psilate elements are found. Crotoii prrllei Lanj. (Fig. 2 E-H). - Pollen grains very large, diameter 123k 1.6 ( ) pm, inaperturate, spheroidal, with a coarsely reticulate pattern, and sparsely spaced sexine elements on top of the muri. The lumina are pentagonal. In SEM the sexine (6.0 pm thick) consists of more or less elliptical, striate elements (clavae), situated at the meeting points of the muri. Numerous small, psilate clavae are found in the lumina. Croton pilllei var. glabrior Lanj. (Fig. 2 I-L). - Pollen grains very large, diameter 134k3.6 ( ) pm, inaperturate, spheroidal, with a coarse reticulate pattern, and sexine elements on top of the muri. The lumina are pentagonal. In SEM the sexine (6.5 pm thick) consists of more or less elliptical, striate elements located on the meeting points of the muri. Numerous small, irregularly distributed psilate clavae are found in the lumina. Pollen key 1. Pollen grains with croton-pattern 1.1. with psilate sexine elements 1.2. with striate sexine elements C. grazielae sexine elements with an apical spinule Crorori sp sexine elements without an apical spinule elements ca. 2 pm in diameter with shallow striae C. jararisarsis elements ca. 4 pm in diameter with deep striae C. aseariferrs 2. Pollen grains reticulate, sexine elements sparsely spaced at the meeting points of the muri, numerous small clavae in the lumina. c. p1rllei C. pirllei var. glabrior DISCUSSION AND CONCLUSIONS The pollen grains of the species examined show a wide variation regarding exine ornamentation. Typical crotonpattern does not occur in C. ptillei and C. pilllei var. glabrior, which have a broad, reticulate pattern with widely separated sexine elements and pentagonal lumina. Nowicke (1994), analysing the pollen morphology of the Crotonoideae, emphasizes the pollen grains of C. nintoirretuis, which have supramural, widely spaced sexine elements, similar to clavae, on the contact area of the segments of the muri. The sexine ornamentation of C. pilllei and C. pzrllei var. glabrior described here, agree perfectly with that of C arntoure~isis (Nowicke 1994). According to Secco (1992), the characteristic leaf and floral morphology suggests that Crotori grazielae should be placed near Crotori pirlki. However, Crotori grazielae has pollen grains which are very different from those of Crotori pirllei. In the former species the pollen grains have psilate and juxtaposed sexine elements, while in the latter they are widely spaced and striate. The morphology and disposition of the sexine elements separates Crotori grazielre (a species endemic to Serra da Neblina-Amazonas) from Croton pirllei (Suriname and Para/Brazil). Webster (1993) included the species C. ascerideirs and C. pirllei in the subsection Palariostigim Mart. ex. Baill., section Cyclostigrim Griseb. He pointed out that in a future study they might be placed in a separate subsection. Considering the morphological characteristics of the lianescent species of Crototr (Secco 1992) and the pollen morphology reported in the present study, it is clear that Webster (1993) Fig. 1. A-D. Croton sp. (A) grain at high focus. LM x 880. (B) the same at low focus. Lhl x 880. (C) optical cross-section. Lhl x 880. (D) densely spaced striate clavae with an apical spinule. SEhl x E-H. Crotori ascazdetzs. (E) grain at high focus. LM x 680. (F) the same at low focus. Lhi x 650. (G) optical cross-section. LM x 680. (H) distribution and shape of striate clavae and smaller elements. SEhi x I-L. Croroti gro:ie/oe. (I) grain at high focus. Lhl x 680. (J) the same at low focus. Lhl x 680. (K) optical cross-section. Lhl x 680. (L) distribution and shape of clacae. SEhl x Fig. 2. A-D. Crototi jowrisetuis. (A) grain at high focus. LM x 600. (B) the same at low focus. Lhl x 600. (C) optical cross-section. Lhl x 600. (D) distribution and shape of clavae. SEhl x E-H. Crorotz pullei. (E) grain at high focus. Lhl x 525. (F) the same at low focus. LM x 525. (G) optical crosssection. Lhl x 525. (H) distribution and shape of clavae. SEhl x I L. Croroti pidlei var. glubrior. (I) grain at high focus. Lhl x 525. (J) the same at low focus. Lhl x 525. (K) optical cross-section. Lhl x 525. (L) distribution and shape of clavae. SEhl x Gruna 35 (1996)
4 Downloaded by [ ] at 06:56 26 April 2015 Fig. I. Grana 35 (1996)
5 Downloaded by [ ] at 06:56 26 April 2015 Pollen of Crotori limas 17 Fig. 2. Grana 35 ( 1996)
6 Downloaded by [ ] at 06:56 26 April L. A t hi. Cnrreircr et nl. was correct in suggesting that the lianescent species of Crotort should be placed in a separate subsection. The key to separate lianescent species of Crototi presented by Secco ( 1992) is supported by the results of pollen morphology and confirms that the remaining species are diflerent from Crotori piillei Lanj. ACKNOWLEDGEMENTS We express our appreciation to the Instituto Oswaldo Cruz, FIOCRUZ, Rio de Janeiro, for use of the Zeiss DhlS-930 scanning electron microscope and to Dr. P. Whelan for the translation of the text. All the authors express their thanks to the CNPq for research grants, the second author in particular for grant number '86/ZO/FV to study Croton. SPECIMENS EXAMINED Crorori sp., N.T. Silva 1667, IAN Crorori ascendens R. Secco & N. Rosa, Rosa & Uhl5212, MG Crorori gra:ielae R. Secco, Silva & Brado 60834, hlg Crorori jarariseitsis R. Secco, Lleras et al. P17086, MG Crororr pitllei Lanj., Stahel 76, Utrecht Crorori pitllei Lanj., P. Cavalcante 2565 MG Crorort ptcllei var. glabrior Lanj., Rombouts 654, IAN REFERENCES Barth, 0. M. & Melhem, T. S Glossario ilustrado de Palinologia - UNICAMP, Campinas. Croizat, L Euphorbiaceae. - In: A collection of flowering plants from Mount Roraima and adjacent Venezuela, British Guiana and Brazil. (ed. A. C. Smith, I. 0. Williams, D. C. Stanley, L. Croizat et al.). - Bull. Torrey Bot. Club 67: Erdtman, G Pollen morphology and plant taxonomy Angiosperms. - Almqvist & Wiksell, Stockholm. Erdtman, G A Handbook of Palynology. - Hafner, New York. Hutchinson, J Tribalism in the family Euphorbiaceae. - Am. J. Bot Lanjouw, J The Euphorbiaceae of Surinam. J. H. de Bussy, Amsterdam. Lanjouw, J. 1939a. New or noteworthy Euphorbiaceae from Suriname. - Rec. Trav. Bot. Ned Lanjouw, J b. Euphorbiaceae. In: Flora of Suriname. Vof. 2 p. 1. Additions and corrections (ed. A. Pulle). pp K. Ver. Kolon. Inst., Amsterdam. hlueller, J Euphorbiaceae. In: Prodromus systematis naturalis regni vegetabilis. Vol (ed. A. P. de Candolle), pp Treuttel 8: Wiirtz, Paris. Mueller, J Euphorbiaceae. - In: Flora Brasiliensis. Vol (ed. C. F. P. von hlartius), pp S. Endlicher (E. Fenzel), Munchen - Men - Leipzig. Nonicke, J. W A palynolopical study of Crotonoideae (Euphorbiaceae). Ann. hl. Bot. Card. 81: Punt, W Pollen morphology of the Euphorbiaceae with special reference to taxonomy. - Wentia: Punt, W A survey of pollen morphology in Euphorbiaceae with special reference to Phjllari~ltiis. - Bot. J. Linn. SOC. 9.1: Punt, W. Blackmore, S., Nilsson, S. & Le Thomas, A, (eds) Glossary of pollen and spore terminology. LPP Contributions Series no. 1, LPP Found., Utrecht. Salgado-Labouriau, hl. L Contribuiciio a Palinologia dos Cerrados. - Acad. Bras. Cienc., Rio de Janeiro. Secco, R. de S Notas sobre as lianas do genero Crotort L. (Euphorbiaceae). - Bol. hfus. Par. E. Goeldi, Ser. Bot. 8: Secco, R. de S. & Rosa, N. A Crotort asrerideits (Euphorbiaceae), a new liana from Eastern Amazonia. Novon 2: Thanikaimoni, G., Caratini, C., Nilsson, S. 8: Grafstrom, E Omniaperturate Euphorbiaceae pollen with striate spines. - Bull. Jard. Nat. Belg. 54: Webster, G. L Conspectus of a new classification of the Euphorbiaceae. Taxon 24: Webster, G. L A provisional synopsis of the section of the genus Crotori (Euphorbiaceae). - Taxon 42: Webster, G. L. & Burch, D Euphorbiaceae. ( In: Flora of Panama. Part 6., ed. R. E. Woodson, Jr.) -Ann. M. Bot. Card 54: Grana 35 ( 1996)
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