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Seed micromorphology of Orchidaceae in the Gulf of Guinea (West Tropical Africa)

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Abstract

Qualitative micromorphological characters of seeds were analysed using Scanning Electron Microscopy for 95 species of 34 genera, belonging to the subfamilies Vanilloideae, Orchidoideae and Epidendroideae (Orchidaceae) from the Gulf of Guinea countries. Three genera (Dinklageella, Platylepis and Rhipidoglossum) and 69 species were studied for the first time. Vanilla shows a pattern of seed coat distinct to the rest of the studied taxa. Orchidoideae and Epidendroideae show variation in several features: seed shape, orientation of testa cells, morphology of anticlinal walls, cell corners, shape and ornamentation of the periclinal walls and presence of waxes. Two main patterns have been observed: one with parallel medial cells along the longitudinal axis, thin and straight anticlinal walls, with no raised and no thickened cell corners, wide periclinal walls and lack of waxes, for the terrestrial orchids of the subfamily Orchidoideae, the tribes Tropidieae, Nervilieae, Collabieae, Cymbidieae and the subtribe Malaxidinae (Epidendroideae); the other one showed twisted medial cells, thickened anticlinal zone with prominent ridges, raised and thickened cell corners, narrow-to-not visible periclinal walls and presence of waxes, in epiphytic orchids of Vandeae, Podochileae and Dendrobiinae (Epidendroideae). In the epiphytic genera Ansellia and Graphorkis (Cymbidieae) and in Liparis epiphytica (Malaxideae), thickened anticlinal walls and presence of waxes also occurred but parallel medial cells along the longitudinal axis and wide periclinal walls have been observed. This study confirms the diagnostic value of the qualitative features of the seed coat in Orchidaceae and suggests a probable relation with adaptations to habitat for seed dispersal.

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Acknowledgements

This study was supported by the Ministry of Economy and Competitiveness of Spain through the research project CGL2012-32934. The authors express their gratitude to the curators and collection managers of the herbaria K, MA and WAG. We are much indebted to Yolanda León, the technical staff at the SEM laboratory at the Royal Botanic Gardens, Madrid. We thank Francisco Cabezas, Maximiliano Fero and Mauricio Velayos who participated in Equatorial Guinea’s botanical expeditions. We appreciate Dr Mauricio Velayos and Dr Javier Fuertes for their constructive comments.

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Authors and Affiliations

  1. Departamento de Biología, Universidad Autónoma de Madrid, 28049, Madrid, Spain

    Roberto Gamarra, Emma Ortúñez & Ángela Merencio

  2. Departamento de Sistemas y Recursos Naturales, Universidad Politécnica, 28040, Madrid, Spain

    Pablo Galan Cela

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  1. Roberto Gamarra
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  2. Emma Ortúñez
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Correspondence to Roberto Gamarra.

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Appendix

Appendix

List of studied material with voucher (*species studied for the first, **genera studied for the first):

Ancistrorhynchus clandestinus* (Lindl.) Schltr. Equatorial Guinea: Bata–Sendje, 19 Apr 1993, M. Carvalho 5286 (MA 598003). Angraecopsis tridens* (Lindl.) Schltr. Equatorial Guinea: Musola, s.d., E. Guinea 1765 (MA 784991). Ansellia africana Lindl. Uganda: Bunyuro, 28 Jan 1945, W.J. Eggeling 7051 et al. (K). Bolusiella iridifolia* (Rolfe) Schltr. Tanzania: Morogoro, 21 Oct 2004, D. Konstantine 3571 et al. (MA 802531). Bolusiella talbotii (Rendle) Summerh. Equatorial Guinea: Añisok–Mongola, 27 Oct 1999, R. Pérez Viso 1481 (MA 785018). Brachycorythis buchananii* (Schltr.) Rolfe Zimbabwe: Melsetter Distr., Chimanimani, 2 Feb 1957 J.B. Phipps 429 (K). Brachycorythis macrantha* (Lindl.) Summerh. Gabon: Haute-Ngounyé, Dondo, 31 Mar 1926, G. Le Testu (K). Ivory Coast: Abidjan, Danané, 18 Oct 1974 F. Pérez Vera 702 (K). Brachycorythis pleistophylla* Rchb.f. Zimbabwe: Mufulia, 3 Jan 1956, D.B. Fanshawe 2684 (K). Brachycorythis pubescens* Harv. Ivory Coast: Sémien–Kanebly, 6 Jun 1966, L. Aké Assi 9071 (K). Malawi: Rumphi Distr., Rumphi, 22 Feb 1975, J. Pawek 13817 (K). Uganda: Sebei, Mt Elgon, 26 Jul 1924, J.D. Snowden 928 (K). Brachycorythis sceptrum* Schltr. Senegal: s. loc, s.d., F. Doumbia s.n. (K). Bulbophyllum falcatum* (Lindl.) Rchb.f. Equatorial Guinea: Annobon, 14 Jul 1959, F.A. Melville 31 (MA-293923). R. Pérez Viso 1679 (MA 709345). Bulbophyllum imbricatum* Lindl. Equatorial Guinea: Annobon, s.d., F.A. Melville 200 (MA 293924). Bulbophyllum josephi* (Kuntze) Summerh. Equatorial Guinea: Moka Malabo, 17 Nov 2012, F. Cabezas 1424 et al. (MA 866215). Bulbophyllum pumilum (Sw.) Lindl. Nigeria: Tora Creek, 20 Jan 1983, J.M. Baker 40 (K). Bulbophyllum saltatorium* Lindl. Equatorial Guinea: Monte Alen, 8 Oct 1999, R. Pérez Viso 1517 (MA 784987). Calanthe sylvatica* (Thouars) Lindl. Congo (Kinshasa): Karisimbi, 17 Aug 1937, Louis 5251 (K). Equatorial Guinea: Annobon, 17 Jul 1959, F.A. Melville 161 (MA 293929). Bioko Sur: Malabo–Bioko, 18 Nov 2012, P. Barberá 362 et al. (MA 866581). Calyptrochilum christyanum* (Rchb.f.) Summerh. Guinea-Bissau: Piche, 6 Dec 1961, J.Alves Pereira 2186 (MA 287501). Calyptrochilum emarginatum (Afzel. ex Sw.) Schltr. Equatorial Guinea: Ebebibyin–Mikonesong, 3 Feb 1969, W.W. Sanford 5733 (K). Cheirostylis divina* (Guinea). Summerh. Sierra Leone: Picket Hill, 18 Nov 1951, T.S. Jones 209 (K 541291). Cheirostylis lepida (Rchb.f.) Rolfe Equatorial Guinea: Caldera San Carlos, 11 Jan 1967, W.W. Sanford 4324 (K). Corymborkis corymbis* Thouars Nigeria: Ondo, Akure, 31 Oct 1949, E. Ujor s.n. (K). Cynorkis anacamptoides Kraenzl. Equatorial Guinea: Moka, Las Costeras, 28 Jan 1947, E. Guinea 2125 (K, MA 785020). Moka, 9 Sept 1959, T.C. Wrigley 582 (K). Musola, 20 Jan 1947, E. Guinea 1700 (MA 785003). Cynorkis debilis* (Hook.f.) Summerh. Equatorial Guinea: Moka, 19 Sept 1959, T. Wrigley 616 (K). Diaphananthe bidens* (Afzel. ex Sw.) Schltr. Equatorial Guinea: Bata, Asonga, 22 Mar 1992, M. Carvalho 5062 (MA 610097). Malabo–Rebola, 21 Aug 1988, M. Carvalho 3541 (MA 683196). Diaphananthe sarcorhynchoides* J.B.Hall. Equatorial Guinea: Bata–Bome, 24 Apr 1994, M. Carvalho 5516 (MA 599463). Ivory Coast: J. de Koning 4723 (MA 857399). Dinklageella liberica** Mansf. Equatorial Guinea: Bata–Pembe, 2 Oct 1991, M. Carvalho 4848 (MA 597732). Disperis reichenbachiana* Welw. ex Rchb.f. Angola: Moxico, 12 Jan 1938, E. Milne-Redhead 4084 (K). Disperis thomensis* Summerh. Equatorial Guinea: Bioko, 1986, J. Fernández Casas 4496 et al. (K). Epipogium roseum (D.Don) Lindl. Equatorial Guinea: Annobon, 1 Aug 1959, F. Melville 213 (BR, K). Ghana: Puso Puso ravine, Asiatewa, 22 feb 1960, J.K. Morton 3866 (K). Eulophia angolensis* (Rchb.f.) Summerh. Cameroon: Mbakakou–Tibati road, 30 Jun 1972, A.J.M. Leeuwenberg 10057 (WAG 1135545). Eulophia bouliawongo* (Rchb.f.) J.Raynal Gabon: river Gaboon, Jul 1861, G. Mann 1039 (K). Eulophia caricifolia* (Rchb.f.) Summerh. Equatorial Guinea: Bata–Bome, 1 Jun 1991, M. Carvalho 4705 (MA 597837). Eulophia cucullata* (Afzel. ex Sw.) Steud. Gabon: Estuaire, Pongara, 17 Dec 2012, T.H.J. Damen 542 et al. (WAG 1924526). Eulophia gracilis* Lindl. Equatorial Guinea: Bata–Bome, 6 Dec 1993, M. Carvalho 5430 (MA 598441). Eulophia horsfallii* (Bateman) Summerh. Tanzania: Mafia Distr., 24 Mar 1933, G.B. Wallace 809 (K). Genyorchis apetala* (Lindl.) J.J. Verm. Equatorial Guinea: Bata–Bome, 2 May 1993, M. Carvalho 5298 (MA598113). Graphorkis lurida (Afzel. ex Sw.) Kuntze Equatorial Guinea: Santa Isabel [Malabo], 23 Feb 1969 W.W. Sanford 6025 (K). Habenaria attenuata* Hook.f. Ethiopia: Solulta, 11 Sept 1974, J. Ash 2599 (K). Habenaria bracteosa Hochst. ex A. Rich. Tanzania: Kilimandscharo, 22 Feb 1934, H.J. Schlieben 4835 (MA 385951). Habenaria chirensis Rchb.f. Ethiopia: Bar Dar Airport, 14 Sept 2004, I. Friis 11578 et al. (K). Habenaria filicornis Lindl. Eritrea: Amasen, Dongollo presso, 12 Mar 1902, A. Pappi s.n. (USNH 196981). Habenaria holubii* Rolfe Zambia: Lusaka, III-1967, R. Odgers 255 (K). Habenaria huillensis* Rchb.f. Angola: Cola (Rio Qué), 24 Apr 1954, G. Boss (M). Habenaria ichneumonea (Sw.) Lindl. Zimbabwe: Chakwenga Headwaters, 25 Aug 1963, E.A. Robinson 5637 (K). Habenaria keayi* Summerh. Nigeria: Plateau Province, Naraguta Forest Reserve, VII-1962, D.E.A. King 30 (K). Habenaria macrandra Lindl. Ivory Coast: Mont Momi, 29 Sept 1966, Ake Assri 9146 (K). Habenaria malacophylla Rchb.f. Nigeria: Taraba State, Mambilla Plateau, 6 Oct 1993, Spamin 8 (K). Habenaria procera* (Afzel. ex Sw.) Lindl. Equatorial Guinea: Asoc, VII-1945, E. Guinea 424 (MA 785009). Kukumankok, s.d., E. Guinea 358 (MA 784981). Habenaria rautaneniana* Kraenzl. Tanzania: Ufifa Distr., Milipa, lake Rukwa, 8 Feb 1938, A.P. Michelore 1482 (K). Habenaria schimperiana* Hochst. ex A.Rich. Zambia: Lusaka west, 25 Feb 2000, M.G. Bingham and C.R. Peters 12132 (K). Holothrix aphylla* (Forssk.) Rchb.f. Cameroon: Bamenda, 14 Feb 1971, P. J. Bauer 161 (K). Ethiopia: 27 Dec 1952, J.B. Gillett 14774 (K). Liparis deistelii* Schltr. Equatorial Guinea: Moka, 21 Sept 1959, T.C. Wrigley 652A (K). Liparis epiphytica* Schltr. Uganda: Kigom, VI-1962, C.L.O. Leabey 14 (K). Liparis nervosa (Thunb.) Lindl. Liberia: Monrovia, 31 Aug 1947, J.T. Baldwin Jr. 9203 (K). Equatorial Guinea: Monte Alén, 19 Mar 1999, R. Pérez Viso 825 (MA 702115). Malabo–Luba–Riaba, 8 Nov 1986, M. Carvalho 2665 (MA 492356). Liparis platyglossa* Schltr. Sao Tome and Principe: pico Papagaio, 6 Oct 1997, T. Stévart 221 (MA). Malaxis maclaudii* (Finet) Summerh. Sierra Leone: Loma Mountains, 16 Nov 1965, J.K. Morton SL2552 (K). Malaxis weberbaueriana* (Kraenzl.) Summerh. Cameroon: Plateau of the Adamaoua, 15 Oct 1960, F.J. Breteler 504 (K). Manniella gustavii Rchb.f. Liberia: Bobée Mountain, 9 Jan 1948, W.J. Harley (K). Nervilia bicarinata* (Blume) Schltr. Equatorial Guinea: Annobon, 23 Nov 2010, M. Velayos 11602 et al. (MA-831090). Nervilia crociformis (Zoll. & Moritzi) Seidenf. Tanzania: Kyimbila, 29 Dec 1980, G.P. Leedal 6264 (K). Oeceoclades maculata (Lindl.) Lindl. Equatorial Guinea: Malabo–Baney, 25 Sept 1988, M. Carvalho 3620 (MA 685159). Orestias foliosa Summerh. Congo (Kinshasa): Kisumba, I-1931, Quarré 2375b (K). Platycoryne crocea* Rolfe Kenya: Shimba Hills National Reserve, 2 Jul 2001, W.R.Q. Luke & M. Pakia 7460a (K). Malawi: Rumphi Distr., Nchena Nchena, 28 Jan 1975, J. Pawek 9005 (K). Platycoryne mediocris* Summerh. Malawi: Livingstone scarp, 8 Mar 1972, D. Wedwood 661 (K). J. Pawek 13978 (K-000453701). Platycoryne paludosa* (Lindl.) Rolfe. Sierra Leone: Tower Hill, Freetown, 18 Jul 1967, Morton and Cole SL 4370 (K). F. C. Deighton 3990 (K). Platylepis glandulosa** (Lindl.) Rchb.f. Equatorial Guinea: Annobon, 24 Jul 1959, F.A. Melville 184 (K). Polystachya adansoniae* Rchb.f. Cameroon: Buea, 23 Apr 1948, H. Gregory 294 (K 106752). Equatorial Guinea: Annobon, 14 Jul 1959, F.A. Melville 154 (BR, K). Liberia: Tchien, 16 May 1970, J. de Koning 517 (MA 855185). Polystachya affinis Lindl. Sierra Leone: Senehun (Kamajei), 14 Apr 1960, T.S. Bakshi 128 (K). Polystachya albescens Ridl. Equatorial Guinea: Basilé, 26 Dec 1966, W.W. Sanford 4008 (K 541418). Cameroon: Ndop Plain, 19 May 1962, M. Brunt 450 (K 106651). Polystachya alpina* Lindl. Equatorial Guinea: pico Santa Isabel [Basilé], 20 Feb 1969, W.W. Sanford 5952 (K 541437). Polystachya bifida* Lindl. Equatorial Guinea: pico Santa Isabel [Basilé], 21 Feb 1969, W.W. Sanford 5965 (K). Polystachya calluniflora* Kraenzl. Equatorial Guinea: pico Santa Isabel [Basilé], 21 Feb 1969, W.W. Sanford 5973 (K). Polystachya caloglossa* Rchb.f. Cameroon: Douala Edea Reserve, III-1978, D.W. Thomas 472 (K). Polystachya carnosa* P.J.Cribb & Podz. Equatorial Guinea: Basilé [Malabo], XII-1966, W.W. Sanford 4026 (K). Polystachya concreta* (Jacq.) Garay & H.R.Sweet Equatorial Guinea: Annobon, pico Lago, 23 Nov 2010, M. Velayos 11588 et al. (MA 828136). Malabo–Aeroporto, 18 Dec 1986, M. Carvalho 2818 (MA 492359). Polystachya coriscensis* Rchb.f. Cameroon: Douala, Edea Forest, 27 Jun 1978, D.W. Thomas 210 (K). Polystachya cultriformis (Thouars) Spreng. Equatorial Guinea: Bioko, 1986, J. Fernández Casas et al. 2649 (K). Polystachya dolichophylla* Schltr. Togo: Tasibara, IV-1953, D. Westwood 66 (K). Polystachya elegans* Rchb.f. Equatorial Guinea: Bioko, II-1989, J. Fernández Casas 11918 et al. (K). Polystachya fractiflexa* Summerh. Ghana: Ankasa Forest Reserve, 5 Jan 1969, Halla Enti s.n. (K). Polystachya fusiformis (Thouars) Lindl. Cameroon: Dscahng Distr., Babajou, 25 Nov 1968, W.W. Sanford 5668 (K 106729). Polystachya galeata (Sw.) Rchb.f. Congo (Kinshasa): Penghe, 7 Feb 1914, Bequaert 2363 (K). Polystachya laxiflora* Lindl. Equatorial Guinea: Bioko, 1986, M. Carvalho 2700 et al. (K). Polystachya odorata* Lindl. Equatorial Guinea: Bioko, 1986, J. Fernández Casas 4459 et al. (K). Polystachya polychaete* Kraenzl. Equatorial Guinea: Moka, Las Costeras, s.d., E. Guinea 2130 (MA 294054). Bioko, 1986, J. Fernández Casas 2749 et al. (K). Polystachya puberula* Lindl. Sierra Leone: Loma Mountains, Kinto, 15 Sept 1945, P. Jaeger 1778 (K). Polystachya ramulosa* Lindl. Gabon: Haute-Ngounyé, 11 May 1929, G. Le Testu 5454 (K). Polystachya rhodoptera* Rchb.f. Cameroon: Douala, Edea Forest, 27 Jun 1978, D.W. Thomas 207 (K). Polystachya superposita* Rchb.f. Equatorial Guinea: Moka, Las Costeras, 28 Jan 1947, E. Guinea 2137 (MA 785034). Rhipidoglossum densiflorum** Summerh. Equatorial Guinea: Monte Alén, Enkumekien, 8 Oct 1999, R. Pérez Viso 1513 (MA 784997). Solenangis scandens* (Schltr.) Schltr. Equatorial Guinea: Nfuiñ, 3 Dec 2010, F. Cabezas 1315 et al. (MA 831762). Stolzia leedalii* P.J.Cribb Tanzania: Iringa Distr., Mufindi, 12 Feb 1974, P. Cribb 11450 et al. (K 390281). Vanilla imperialis Kraenzl. Uganda: Bunyoro, s.d., W.J. Eggeling 2062 (K). 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Gamarra, R., Ortúñez, E., Galan Cela, P. et al. Seed micromorphology of Orchidaceae in the Gulf of Guinea (West Tropical Africa). Plant Syst Evol 304, 665–677 (2018). https://doi.org/10.1007/s00606-018-1497-0

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