Polish Botanical Journal 56(2): 131–153, 2011 EAST AFRICAN BRYOPHYTES XXIX. THE CERATOLEJEUNEA (LEJEUNEACEAE) SPECIES OF THE INDIAN OCEAN ISLANDS TAMÁS PÓCS Abstract. To date, fi ve species of Ceratolejeunea are known from the East African Islands: C. belangeriana and C. calabariensis from a number of localities, C. variabilis from three, and C. papulifl ora and C. umbonata from single stations. This paper newly establishes the synonymy of C. calabariensis with the Neotropical C. cornuta and of C. stictophylla with C. papulifl ora, and reports several additional localities of the latter species. Ceratolejeunea diversicornua, previously known from West Africa only, is reported from Madagascar, and C. andringitrae and C. saroltae are described as new to science. A key to the species of Ceratolejeunea from the Indian Ocean Islands is provided. The taxonomic position of C. boschiana remains uncertain. Key words: Ceratolejeunea, Comoro, Indian Ocean Islands, Madagascar, Mascarenes, Seychelles Tamás Pócs, Botany Department of Eszterházy College, Eger, Pf. 43, H-3301, Hungary; e-mail: [email protected] INTRODUCTION The genus Ceratolejeunea, with ca 40 species, is From continental Africa, Vanden Berghen (1951, distributed exclusively in the tropics. Its species 1973) identifi ed eight species. Five species have occur on various substrates, mostly bark and living been identifi ed from the East African Indian Ocean leaves. The name of the genus refers to the horned Islands (Grolle 1995; Pócs 1995; Pócs & Geissler perianths of its members. The coloring of the genus 2002; Wigginton 2009). Some 15 species are varies from dark brown to olive, with a dull shine known from Asia and the Pacifi c, some of which that is quite rare among Lejeuneoideae and is have been revised by Mizutani (1981). The exact more typical of the Ptychanthoideae subfamily. number of all Ceratolejeunea species is not known, The genus was established by Jack and Stephani as only the Neotropical species have been mono- (1892) from the subgenus Cerato-Lejeunea defi ned graphed recently; a revision of species on other by Spruce (1884–1885). The following charac- continents is much needed. ters distinguish the Ceratolejeunea species from In the course of several collecting excur- other Lejeuneoidae: their special color and shine; sions throughout East African Islands from the their imbricate leaves which frequently present Seychelles and Comoros to different parts of utriculi and generally have various types of ocelli Madagascar and the Mascarenes (Réunion and in the lobe; their small, single-toothed, infl ated Mauritius), and with contributions from my col- lobule; their predominantly bilobed underleaves leagues (from EGR, MO, NAI, EA and TANA), (subgen. Caduciloba R. M. Schust. – though the I have assembled a rich collection of Ceratole- underleaves are entire in the small subgen. Cera- jeunea species. So far only a smaller portion of tolejeunea); by their Lejeunea-type branching; these materials have been revised, the results of and their usually four-horned perianths with Pyc- which are summarized below. nolejeuneoid innovation. In terms of continental On the basis of this initial revision, seven distribution, America is the richest, with 23 species species can be said with certainty to occur in (Dauphin 2003a) to which two more have recently the area: Ceratolejeunea andringitrae Pócs sp. been added, one by Ilkiu-Borges and Alvarenga nov., C. belangeriana (Gottsche) Steph., C. cor- (2008) and the other by Reiner-Drehwald (2011). nuta (Lindenb.) Steph., C. diversicornua Steph., 132 POLISH BOTANICAL JOURNAL 56(2). 2011 C. papulifl ora Steph., C. saroltae Pócs sp. nov. and tinguish C. belangeriana from C. cornuta. More C. umbonata Steph. It is also clear that C. cala- Asian and Pacifi c populations need to be examined bariensis Steph. and C. variabilis (Lindenb.) to determine whether C. belangeriana really merits Schiffn. are synonyms of C. cornuta, and C. stic- species rank. If its perianth length remains the only tophylla Herzog a synonym of C. papulifl ora. Al- difference, it should be distinguished at subspecifi c though Grolle (1995) synonymized C. boschiana rank under C. cornuta. Mizutani (1981) differenti- Steph. with C. belangeriana, its taxonomic posi- ates C. maritima (= C. cornuta) from C. oceanica tion remains uncertain. (= belangeriana) by its highly infl ated, spherical lobules and its few-branched, usually long stems. SPECIES KNOWN FROM THE EAST AFRICAN C. belangeriana was previously known as wide- ISLANDS spread in the East African Islands only, but reports of its occurrence in the whole of Indomalaya and 1. Ceratolejeunea belangeriana (Gottsche) Oceania have been published more recently (Miller Steph. et al. 1983; Zhu et al. 2005). Spec. Hep. 5: 396. 1913. DISTRIBUTION. Comores, Madagascar, Sey- BASIONYM: Lejeunea belangeriana Gottsche in Gottsche chelles, Réunion, Mauritius, Indomalaya from et al., Syn. Hep.: 398. 1845. TYPE: Mauritius, Bélanger Thailand to the Philippines and New Guinea, s.n. (Fragment only in G. ex hb. Lehmann). Melanesia and Polynesia. IMPORTANT SYNONYMS SELECTED SPECIMENS WITH PERIANTHS INVESTI- Lejeunea mascarena Steph., Bot. Gazette 15: 284. 1890, GATED. MADAGASCAR: Prov. Antsiranana, Réserve Ceratolejeunea mascarena (Steph.) Steph., Spec. Hep. integrale Marojezy, montane rainforest on the sharp 5: 397. 1913 (fi de Grolle 1995). ridge N of Andampibe Falls at 780–1050 m a.s.l., on decaying wood, Pócs 90113/CN (EGR). Prov. Ant- Lejeunea mauritiana Steph., Bot. Gazette 15: 285 & 349. siranana, Réserve spéciale de Manongarivo Ambahatra, 1890, Ceratolejeunea mauritiana (Steph.) Steph., Hed- cours superior, camp 2, 1200 m a.s.l., on bark, Geissler wigia 31: 205. 1892 (fi de Grolle 1995). 19850 (CJB-G, dupl. EGR). RÉUNION: 12 km W of Lejeunea oceanica Mitt. in Seemann, Fl. Vitiensis: 414. Ste Anne, remnants of tropical rainforest alternating 1871, Ceratolejeunea oceanica (Mitt.) Steph., Bot. with secondary Psidium cattleyanum bushes around ‘le Jahrb. Syst. 23: 310. 1897 (fi de Zhu et al. 2005). Fur- Grand Étang’ reservoir, 500–550 m a.s.l., epiphyllous, ther synonymy of C. oceanica see in Mizutani (1981). Pócs, A. Szabó & Vojtkó 9433/AO, 29 Aug. 1994 (EGR). ILES DE LA SOCIETÉ: Mooréa, au-dessus du point ILLUSTRATIONS: Bonner (1953: 169, Fig. 8; 1953: 215, de vue de Belvédère, forêt du Inocarpus edulis, 300 m Fig. 58 under C. mascarena), Schiffner in Engler & Prantl a.s.l., sur tronc, De Sloover 20.979 (det. Grolle, NAM, (1893: 126 under C. mascarena), Stephani (1890: 284; dupl. EGR). Pl. xvii, fi g. 5), Bonner (1953: 169, Fig. 8 and 1953: 215, Fig. 58), Mizutani (1981: 306, Fig. 1, under C. oceanica), Grolle (1995) placed Ceratolejeunea boschiana Herzog in Herzog and Noguchi (1955: 47, Fig. 11h–m Steph. (1913) (lectotype: Bonner 1953: 168, Fig. 9) under C. exocellata Herz.) and Amakawa (1970: 180, in the synonymy of C. belangeriana. However, the Fig. 28 under C. ryukyuensis Amak.). lectotype specimen of C. boschiana has juvenile According to the protologue and the above- perianths only, which are much shorter than the cited illustrations, what distinguishes Ceratole- perichaetial leaves. At the time of Grolle’s pub- jeunea belangeriana from the more widespread lication, C. calabariensis (= synonym of C. cor- Ceratolejeunea cornuta is the length of its peri- nuta, with almost the same vegetative characters anth. The perianth of C. belangeriana reaches as C. belangeriana) was not known from the East up to 1.4 mm in length, as it has an attenuate, African Islands. On the basis of its juvenile peri- quasi-stalked, narrow conical base exserting it well anth, C. boschiana could belong to either of these above (more than half its length) the perichaetical two species, hence its taxonomic position remains leaves. Without the perianth it is diffi cult to dis- uncertain. T. PÓCS: EAST AFRICAN BRYOPHYTES XXIX. CERATOLEJEUNEA 133 2. Ceratolejeunea cornuta (Lindenb.) Steph. specimens and to the detailed descriptions and Figs 1–6 illustrations given by Dauphin (2003a) as well as those of the authors of the two species. I could in Engler, Pfl anzenwelt of Afrikas Theil C: 65. 1895. not fi nd any difference between the African pop- BASIONYM: Jungermannia cornuta Lindenb., Acta Nova ulations known under the name Ceratolejeunea Acad. Caes. Leop.-Carol. Suppl. 14: 23. 1829. TYPE: calabariensis Steph. and the highly variable Neo- Jamaica, on Grammitis serrulatus, Swartz s.n. (HOLO- tropical Ceratolejeunea cornuta (Lindenb.) Steph. TYPE W, fi de Dauphin 2003, not seen). populations. The original description of C. cala- AFRICAN SYNONYMS bariensis does not mention the utriculi (modifi ed infl ated lobules), while the detailed description of Ceratolejeunea calabariensis Steph., Hedwigia 34: 234. 1895, syn. nov. TYPE: West Africa, New Calabar (in the C. cornuta states that utriculi are rare, appear soli- Niger Delta of present-day southern Nigeria), on tree tary or in pairs at the base of lateral branches, and trunks, 8 Oct. 1884. ‘Mönkemeyer 5 sub ‘Lejeunea vari- are rounded to reniform in shape (Dauphin 2003a: abilis’ in Hb.G’, as indicated by Bonner 1963: 640. 44). I have, in fact, also found African plants with Ceratolejeunea jungneri Steph., Hedwigia 34: 234. the same type of utriculi. The perichaetial leaves of 1895. LECTOTYPE (selected by Bonner 1953: 205): Cam- African plants are described as dentate, while those eroons, Ekunda N’dene, 10 Sept. 1892, Dusén 936 (G), of the American plants as having ‘margins dentate seen by Vanden Berghen (1973: 381), who synonymized throughout or only distally’. The scattered, some- it with C. calabariensis Steph. times geminate basal leaf ocelli, the orbicular to Ceratolejeunea usambarensis Steph. in Brotherus, Denk. wide reniform bilobed underleaves with a narrow Kaiserl. Akad. Wissensch. 88: 731. 1913; Stephani, Sp. incision and often auriculate basis, the perianth Hep. 5: 446. 1913. LECTOTYPE (selected by Bonner with 4 short or longer conical horns and other fea- 1953: 247): East Africa, Usambara, Amani, 800 m a.s.l., tures of the African plants are all identical and fall July 1909, Brunnthaler 8 (G). Synonymized by Vanden Berghen (1973: 381) with C. calabariensis Steph. well within the high variability of the widespread Neotropical Ceratolejeunea cornuta as established Lejeunea variabilis Lindenb. in Gottsche et al., Synopsis by Dauphin (2003a). The presence of enlarged, Hepaticarum: 399. 1845, Ceratolejeunea variabilis (Lin- denb.) Schiffn. in Engler & Prantl, Nat. Pfl anzenfam. spherical lobules ‘throughout or here and there’ 1(3): 125. 1893. LECTOTYPE fi de Grolle in sched.: Saint is emphasized by Dauphin (2003b), which is also Kitts (W, ISOLECTOTYPES BM, M, G, S) according to true for certain African plants. Hence a typical Dauphin (2003a: 42). Although it was described from the case of Afro-American disjunction is at hand, as Neotropics, Grolle (1995) cited several references from encountered by Gradstein, Pócs and Váňa (1984). the East African Islands as well. For further American If further investigation proves C. belangeriana synonyms see Dauphin (2003a). to be only a subspecies of C. cornuta, the latter species should be considered a Pantropical fl ora ILLUSTRATIONS: Fulford [1945: 389, Figs 53–59 under C. maritima (Spruce) Steph., Figs 60–64 under C. valida element. A. Evans, Figs 69–73 under C. grandibracteata Fulford], In fact, Gottsche et al. (1845: 399) had already Bonner [1953: 205–206, Figs 47–48 under C. involvens reported C. cornuta from Madagascar, Stephani (Nees & Mont.) Steph., 207, Fig. 49 under C. jungneri (1895: 65) had reported it from Réunion, and Miller Steph. and under many other synonyms] and Vanden et al. (1983) had reported it from Polynesia (under Berghen (1951: 73, Fig. 26 and 1973: 382, Fig. 6) as the name C. maritima). Bonner (1953) and Grolle C. calabariensis. (1995) both examined the Madagascar specimen The plant known under the above synonyms from the Montagne Herbarium and identifi ed it as is widespread all over the wetter parts of tropical C. belangeriana. On that basis, Grolle (1995: 142) Africa, including the Indian Ocean Islands (Pócs concluded that the whole of the specimens reported 1995; Pócs & Geissler 2002). I have examined from Réunion ‘likewise has to be referred to C. be- a number of African plants and compared them langeriana’, but this has never been confi rmed. both to various American Cololejeunea cornuta Ceratolejeunea variabilis has on several occasions 134 POLISH BOTANICAL JOURNAL 56(2). 2011 Figs 1–6. Ceratolejeunea cornuta (Lindenb.) Schiffn. 1 – perichaetium with subgynoecial innovation (scale bar = 200 μm), 2 – part of shoot, ventral view (scale bar = 200 μm), 3 – leaf (scale bar = 250 μm), 4 – lobule (scale bar = 50 μm), 5 – under- leaf (scale bar = 200 μm), 6 – median lobe cells (scale bar 20 μm). Photographed from Pócs 6100/AK, Tanzania, Usambara Mts, Amani. T. PÓCS: EAST AFRICAN BRYOPHYTES XXIX. CERATOLEJEUNEA 135 Figs 7–14. Ceratolejeunea papulifl ora Steph. 7 – habit of fertile shoot (scale bar = 500 μm), 8 – shoot, dorsal view (scale bar = 250 μm), 9 – leaf base with lobule, seriate basal ocelli and scattered ocelli (scale bar = 100 μm), 10 – leaf (scale bar = 250 μm), 11–13 – lobe cells (11 – apical, 12 – median and 13 – basal; scale bars = 20 μm), 14 – serial basal ocelli (scale bar = 20 μm). 7 from Pócs et al. 90113/EW, Madagascar. Marojezy Reserve. 8 from Pócs & A. Szabó 9878/FE, Madagascar, Manananra Nord Biosphere Reserve, 9–14 from Pócs 9450/AB, Madagascar, Nosy Mangabe Island in Antongil Bay. 136 POLISH BOTANICAL JOURNAL 56(2). 2011 been reported from the East African Islands: from & Hall 1346, 11 Feb. 1971 (EGR). EQUATORIAL Madagascar (Gottsche 1882: 357), from Mauritius GUINEA: Muni, Cogo, on mangle trees in mangrove (Pearson 1892: 8) and from Réunion (Bescherelle at Utoche, along River Mitong, 5 m a.s.l., Heras VIT 367/96, 28 Aug. 1996 (VIT, dupl. EGR). D. R. CONGO 1895: 5) after Grolle (1995). All these records now (former Zaire): Prov. Kivu, Irangi Forest Station 110 km belong to C. cornuta according to the synonymy W of Bukavu, in submontane rainforest near waterfalls given by Dauphin (2003a). at 900 m a.s.l., on branches above water, Pócs 6813 (EGR). UGANDA: Kalangala District, Ssese Islands, DISTRIBUTION (see map in Fig. 81). Ubiquitous Jungo Forest near Mweno village, Bugala Islans, 1170 m all over Latin America from Cuba and Mexico to a.s.l., epiphyllous, Pócs, Lye & Samuela 97107/AH, southern Brazil (Dauphin 2003a), also widespread 7 Sept. 1997 (EGR). TANZANIA: East Usambara Mts, in the wetter parts of tropical Africa from Sierra Amani, on planted mango tree surrounded by submon- Leone and the Gulf of Guinea islands through tane rainforest, 900 m a.s.l., epiphyllous, Pócs 6100/K, Cameroon and Zaire to Tanzania and all East Af- Dec. 1969 (EGR). Uluguru Mts above Morogoro town, rican Islands. Mwere Valley, submontane rainforest, 1450–1550 m a.s.l., epiphyllous, Pócs 6100/AK, 29 Dec. 1969 (EGR). SELECTED SPECIMENS EXAMINED. PUERTO RICO: COMORES: Ndzuani (Anjouan) Island, Col de Moya, Sierra de Luquillo, Mt. Britton Loop Road, on bark. Ful- intercropped submontane rainforest, 700–800 m a.s.l., ford, Crandall & Stotler 334, 12–26 Feb. 1967 (EGR). on buttresses, Pócs 9166/X, 23 March 1991 (EGR). REP: DOMINICANA: Prov. Monseńor Nouel. Cordillera SEYCHELLES: Morne Seychellois Nat. Park, summit Central, degraded cloud forest E of Jima, 1200–1300 m ridge of Morne Blanc, elfi n forest, 590–669 m a.s.l., a.s.l., on decaying branch, S. & T. Pócs 03156/E, 22 epiphyllous, Pócs 9323/BA, 12 Aug. 1993 (EGR). Mahé Nov. 2003 (EGR). SURINAM: s.l., Lanjouw 831, 1933 Island, Victoria – Port Glaud, Sentier Morne Blanc, (NY, dupl. EGR). FRENCH GUIANA: Kourou, Mt. 450–667 m a.s.l., epiphyllous, Frahm SEY-032, 9 Oct. des Signes, ‘sentier botanique’ in mixed rainforest, alt. 2008 (Hb. Frahm, EGR). Praslin Island, Vallée de Mai ca 100 m a.s.l., epiphyllous on palm leaves, Gradstein Nat. Park, palm forest along Cascade River, 150–180 m 6271, 21 June 1986 (Bryophyta Neotropica Exsiccata a.s.l., on bark, Pócs 9358/AL (EGR). MADAGASCAR: No. 232, EGR). PANAMA: Prov. Panamá, nes summit Prov. Antsiranana, Réserve integrale Marojezy, montane of Cerro Jefe, alt. ca 1000 m a.s.l., on branches and rainforest on the sharp ridge N of Andampibe Falls at twigs in scrubby secondary vegetation, Salazar & Grad- 780–1050 m a.s.l., on decaying wood, Pócs 90113/CN stein 9405, 12 June 1991 (Bryophyta Neotropica Exsic- (EGR). Mananara Nord Biosphere Reserve. Mahavoho cata No. 280 under Ceratolejeunea maritima, EGR). Hill, lowland rainforest at 220–300 m a.s.l., streamside, ECUADOR: Galápagos, Santa Cruz Island, Miconia watered rocks, Pócs & A. Szabó 9878/AG (EGR, MO, scrub around Media Luna, on bark of a big avocado TAN). MAURITIUS: Le Pouce, forest halfway to the tree, 600–650 m. Gradstein & Weber H86, 1976 (U, top. On bark, Een, 7 Oct. 1962, Soc. D’Échange des dupl. EGR). SĂO TOMÉ: SE coast, W of Săo Jăo dos Muscinées No. 2988, under Ceratolejeunea renauldii Angolares, 20 m a.s.l., on palm trunk in grazed coconut Steph. (EGR). Black River National Park, Piton Sa- and breadfruit tree plantation, Pócs 00145/G, 27 Aug. vanne, rainforest remnants, 700 m a.s.l., on bark. Frahm 2000 (EGR). Around Cascata Monte Café in the valley MAU-010, 28 Sept. 2007 (EGR). of Rio Manuel Jorge, in submontane rainforest, at 820 m a.s.l., on bark. Pócs 00148/A, 28 Aug. 2000 (EGR). 3. Ceratolejeunea diversicornua (Steph.) Steph. PRINCIPE: E side of the island between Infante Hen- Figs 54–63 rique and Santo Antonio at the volcanic pinnacles called Twin Fingers, mixed hardwood forest, 115 m a.s.l., on Species Hepaticarum 5: 410. 1913. volcanic boulder, Shevock et al. 34633, 9 March 2010 (CAS, EGR). IVORY COAST: Soubré, Parc National BASIONYM: Lejeunea diversicornua Steph., Hedwigia de Taď, Aké-Assi 12641, 1977 (EGR, UCJ). Chemin des 30: 207. 1891. TYPE: Cameroons, s.l., s.d., Dusén inter 37 (G, seen and drawn by Vanden Berghen 1951). Avodirés, sur tronc couché dans marais, 10 m a.s.l., Assel RCI 183, 16 Feb. 1968 (EGR). En direction de M’Ponto Ceratolejeunea cornutissima Steph., Hedwigia 31: 166. en forèt claire, sur tronc demi-sec, Assel RCI 1046, 23 1892; Lejeunea cornutissima (Steph.) Steph., Hed- Sept. 1969 (EGR). GHANA: Central Region, Pra-Suhien wigia 31: XVI. 1892, fi de Jones in Wigginton et al. Forest Res., side of bough of fallen tree in gap, Jones (1996: 41). T. PÓCS: EAST AFRICAN BRYOPHYTES XXIX. CERATOLEJEUNEA 137 Figs 15–21. Ceratolejeunea papulifl ora Steph. 15 – male branch (scale bar = 250 μm), 16–17 – perianths (scale bars = 250 μm), 18 & 20 – serial basal ocelli (scale bars = 60 μm), 19 – scattered ocelli (scale bar = 50 μm), 21 – two-celled style (scale bar = 50 μm). 15–17 from Pócs et al. 90113/EW, Madagascar. Marojezy Reserve, 18–21 from Pócs 9450/AB, Madagascar, Nosy Mnagabe Island in Antongil Bay. 138 POLISH BOTANICAL JOURNAL 56(2). 2011 Figs 22–28. Ceratolejeunea andringitrae Pócs, sp. nov. 22 – habit with perichaetium (scale bar = 250 μm), 23 – shoot, ven- tral view (scale bar = 250 μm), 24 – androecium on branch apex (scale bar = 250 μm), 25 – autoicous branching system (scale bar = 1 mm), 26 – leaf with ocelli groups (scale bar = 250 μm), 27 – lobule (scale bar = 50 μm), 28 – underleaf (scale bar = 100 μm). All photographed from the type. T. PÓCS: EAST AFRICAN BRYOPHYTES XXIX. CERATOLEJEUNEA 139 Figs 29–41. Ceratolejeunea andringitrae Pócs, sp. nov. 29–37 – leaves with different arrangements of grouped ocelli (scale bars = 250 μm), 38 – pair of leaves (scale bar = 250 μm), 39 – ventral view of shoot with antheridia on short side branch (scale bar = 100 μm), 40 – dorsal view of shoot (scale bar = 100 μm), 41 – female bracts, dorsal view (scale bar = 200 μm). All photographs and SEM micrograph made from the type. 140 POLISH BOTANICAL JOURNAL 56(2). 2011 Figs 42–47. Ceratolejeunea moniliata Herz. for comparison. 42 – shoot, dorsal view (scale bar = 500 μm), 43 – perianth (scale bar = 200 μm), 44– 47 – leaves with moniliate ocelli (scale bars = 200 μm). Photographed from Streimann & Naoni 16480, Papua New Guinea: Central Province, Ehu Creek 13 km SW of Sogeri, det. R. Grolle (H ex CBG). ILLUSTRATIONS: Stephani (1892: 166, Tab. XII, fi gs 13, 180, Fig. 20 under C. cornutissima, 1953: 186, Fig. 27 14 under C. cornutissima), Vanden Berghen (1951: 67, under C. diversicornua). Fig. D under C. cornutissima, Fig. E and 1951: 69, Fig. 25A under C. diversicornua) and Bonner (1953: The distinctive characters of the species are