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T. Pócs Department of Botany, Institute of Biology, Eszterházy Károly Catholic University H-3301 Eger, Pf. 43, Hungary

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This paper contains 142 Campylopoideae records from 10 collecting trips of the author with his colleagues in the East African islands. Among the 27 taxa 15 records were new to a certain island, of which 4 were known before only from continental Africa. With these the known number of species on the Indian Ocean islands raises from 30 to 34. Observations on the ecology, distribution and illustrations of most species are also given.

Abstract

This paper contains 142 Campylopoideae records from 10 collecting trips of the author with his colleagues in the East African islands. Among the 27 taxa 15 records were new to a certain island, of which 4 were known before only from continental Africa. With these the known number of species on the Indian Ocean islands raises from 30 to 34. Observations on the ecology, distribution and illustrations of most species are also given.

INTRODUCTION

The Campylopoideae collections made in tropical East Africa by the author and his colleagues were identified and published first by Bizot (in Bizot and Pócs 1974, 1979, Bizot et al. 1977, 1978), then by Ochyra (Ochyra and Pócs 1983, 1986 a, b) and later by Frahm in his monograph (1985 a) and in his posthumous paper (Frahm 2021). A large collection, mainly from the Indian Ocean islands of East Africa collected between 1987 and 1996, were sent also to Jan-Peter Frahm, which remained unidentified with their first specimens in EGR and duplicates were deposited after his death in the herbarium of the Botanic Garden and Botanical Museum Berlin-Dahlem (B). The author identified these and now publishes the detailed records of the less common species. From the 27 identified taxa 15 proved to be new to some of the islands and 4 of them were known before only in continental Africa. By the identification and establishing the up to date nomenclature of species the following references were used: Bizot and Kilbertus (1979), Frahm (1982 a, b, 1985 a, 1994, 2000), Padberg and Frahm (1985), Stech (1999) and Ziljstra (1998). To define the novelty of records and distribution of species apart from the above were used AhPeng and Bardat (2005), Crosby et al. (1983), Een (2000), Frahm (1985 b, 1993, 2021), Frahm et al. (2009), Frahm and Ho (2009), Frahm et al. (2012), O’Shea (1995, 2006), O’Shea et al. (1996), Tixier and Guého (1997).

We call East African or western Indian Ocean islands the group of isles eastwards from East Africa to the Rodrigues Island at the latitude of 63.5°E (see Fig. 1). These isles are geologically very different: the oldest are of Precambrian origin, as Madagascar and the Inner Seychelles, their base is built up of granite and gneiss, with different later deposits. Others are of volcanic origin, usually developing in a series subsequently above hot spots, like Comoro Islands and the Mascarenes, which built up mostly of basaltic lava. The third group are of more recent origin, either coral islands on different basement (like Aldabra or Agalega Islands) or sand keys on the continental shelf, as Mafia Island near the African coast (see more in details by Pócs 1997).

Fig. 1.
Fig. 1.

The East African or western Indian Ocean islands, where the Campylopus specimens were collected. Abbreviations see under the title of Enumeration

Citation: Acta Botanica Hungarica 64, 1-2; 10.1556/034.64.2022.1-2.9

ENUMERATION OF THE SPECIES

The following abbreviations are used: COM = Comoro Islands, MAD = Madagascar, MAF = Mafia Island, MAU = Mauritius, REU: = Réunion, SEY = Seychelles. The first two digits of collecting localities means the year of the collection. All specimens are deposited both in EGR and B, except for those, which are marked by EGR.

Atractylocarpus madagascariensis (Thér.) Padberg et J.-P. Frahm (Syn.: Metzlerella madagascariensis Thér., Metzleria madagascariensis (Thér.) J.-P. Frahm) – MAD: (AlaotraMangoro), Mantady Forest Reserve NE of Andasibe (Périnet), in submontane rainforest near the end of the graphite mining road at 1,030 m, Vojtkó 9485/CE. REU: E edge of Cirque de Mafate. Montane rainforest in Ravine Savon on the S side of Plateau Mahot, at 1,350– 1,550 m, Kis 9427/EU; Acacia heterophylla-Cyathea glauca woodland in Forêt de Bélouve, 1 km E from the Forest Station, at 1,450–1,500 m, Kis 9614/X. – Lemurian species in the sense of Tixier (1978) and Pócs (1997, 1999), indicating a distribution mainly on the Indian Ocean islands belonging biogeographically to Africa, eastwards to the Rodrigues Island at 63.5° E longitude, but with occurrence also on the Crystalline Arc and other East African mountains.

Campylopus arctocarpus (Hornsch.) Mitt. subsp. madecassus (Besch.) J.-P. Frahm (Syn.: Campylopus madecassus Besch., Campylopus madegassus Dix.) – REU: S slope of Piton Larde, at 1,450–1,650 m, G. Kis 9653/N. COM: Ngazidja (Grande Comore) Island. On 15 years old basaltic lava flow (from the 1977 eruption) above Singani village. 43° 19’ 16” E, Magill & Pócs MO 10911 (EGR 9361/D). – A Lemurian species, new to the Comoro Islands.

Campylopus aureonitens (Müll. Hal.) A. Jaeger – REU: Subalpine ericaceous bush on the N sloping plateau of “la Roche Ecrite”, S of St Denis, at 2,100 m, Vojtkó 9420/CO. – Southeast African species with an isolated occurrence in Angola.

Campylopus bartramiaceus (Müll. Hal.) Thér. – MAD: (Analamanga), Réserve Spéciale Ambohitantely E of Manankazo village (near Ankazazobe town), at 1,600 m alt. Relict sclerophyllous plateau forest with 10–12 m high canopy, Kis 9444/F. – Hitherto known from SE Africa and Réunion and New Zealand, new to Madagascar.

Campylopus bicolor (Müll. Hal.) Wils. subsp. atroluteus (Müll. Hal.) J.-P. Frahm – SEY: Mahé Island, Morne Seychellois Nat. Park. Copolia summit. Open, granite platform (glacis) at 480–510 m, Pócs 9335/R. – A South African species, new to the Seychelles.

Campylopus crateris Besch. – REU: Subalpine ericaceous bush on the N sloping plateau of “la Roche Ecrite”, S of St Denis, at 2,100 m, Kónya 9420/CG. – Lemurian subendemic with one occurrence in southern Kenya, not rare in Réunion.

Figs 2–5.
Figs 2–5.

Habit of different species. 2 = Campylopus flaccidus Ren. et Card. (from Pócs & Krog 89204/K). 3 = Campylopus. julaceus subsp. arbogastii (Ren. et Card.) J.-P. Frahm (from Vojtkó 9657/A). 4 = Campylopus flavicoma Müll. Hal. ex Broth. (from Orbán 9484/CB). All scale bars 20 µm. 5 = Campylopus trachyblepharon subsp. comatus (Ren. et Card.) J.-P. Frahm at the same magnification, as the previous (from Kis 9487/EG)

Citation: Acta Botanica Hungarica 64, 1-2; 10.1556/034.64.2022.1-2.9

Campylopus decaryi Thér. – MAD: Fianarantsroa prov., Andringitra Mts Nature Reserve. Mesic elfin forest dominated by bamboos, at 1,450–1,700 m alt., 6 km E from Pic Boby, Kis 9475/FX. REU: Ericaceous heath at and above the timberline on the S slope of Piton de la Fournaise volcano along the trail leading to Puys Ramond, 1,390–2,170 m, Vojtkó 9613/X. – Lemurian species, new to Réunion.

Campylopus flaccidus Ren. et Card. – MAF: SE of Kilindoni, near Kitemondo, Erica mafiensis heath (“Pori”), on open ground covers large areas of the white quartz sand, at 10–16 m. Pócs 87007/N, 89204/K. MAD: Isalo National Park. “Piscine Naturelle” 5 km W of Ranohira, sandstone canyon with Pandanus pulcher, at 800 m alt. Szabó & Orbán 9456/CE & CK. – A typical xerophyte of open habitats and temporarily dry substrate. Widespread in eastern and southern Africa from Sudan to Cape and in Madagascar, new to Mafia Island.

Campylopus flavicoma Müll. Hal. ex Broth. (Syn.: Bryohumbertia metzlerelloides P. de la Varde et Thér., Bryohumbertia flavicoma (Müll. Hal. ex Broth.) J.-P. Frahm, Campylopus flageyi Ren. et Card.) – REU: Plaine des Palmistes, lava flow at the E edge of Premier Villages Bas, at 900 m altitude. In heath-like vegetation on peaty ground, Kis & Orbán 9435/CE; Acacia heterophylla-Cyathea glauca woodland in Forêt de Belouve, 1 km E from the Forest Station, at 1,450–1,500 m, Kis 9614/V; Cirque de Cilaos, montane evergreen Forêt de la Mare à Joseph, E from le Bloc along the path at 1,370–1,380 m, Kis 9638/DX; MAD: (Haute Miasatra) Montane rainforest, bordered by Sphagnum marshes on the hill with radio tower, 10 km N of Fianarantsoa, at 1,230–1,360 m, Orbán 9461/CX; Andringitra Mts Nature Reserve. Montane rainforests on the W side of Korokoro River, at 750–1,000 m, Kis 9472/FC; Bog forest dominated by two Pandanus spp. in a small depression called “Akofa” on the ridge E from Korokoro River, at 920 m; Peat bogs with Eriocaulon, Drosera and Sphagnum spp. on the Ambavafatra Plateau at 1,270 m, Kis 9478/EH; (Alaotra-Mangoro) Mantady Forest Reserve NE of Andasibe (Périnet). Pandanus pulcher swamp forest in Saharanga Valley, at 995 m alt., Orbán 9484/CB; Central E Madagascar. Degraded, secondary montane evergreen forest 4 km E of Moramanga, near Antsahotsaka village, Kis 9491/CA. – Campylopus flavicoma is a shade tolerant, small species occurring on the ground of closed montane rainforests. A Lemurian species distributed in SE Africa and in the Indian Ocean islands. Closely related to the Neo-tropical Campylopus filifolius (Hornsch.) Mitt. (Frahm 1985, 2021).

Campylopus flexuosus (Hedw.) Brid. var. flexuosusCOM: Ngazidja (Grande Co-more) Island. Col du Dibwani, “Plateau des Fougères” between Bahani and Koimbani villages at 560 m, in remnants of dry forest, Pócs, Magill & Rupf 9263/AC. – Almost pantropical species distributed also in the Atlantic part of Europe. The commonest species of shady and half-shady habitats on all islands from the sea level to the upper forest limit, collected from 16 habitats in many specimens from the Indian Ocean islands, but proved to be new to the Comoros.

Campylopus flexuosus (Hedw.) Brid. var. incacorralis (Herz.) J.-P. Frahm (Syn.: Campylopus incacorralis Herz.) – COM: Ngazidja (Grande Comore) Island. Degraded rain-forest on the W slope of Kartala volcano, above Mvouni Village, at 840 m, Pócs & Manktelow 9150/R. MAD: (Sava) Reserve Integrale Nationale de Marojezy, Cyperaceae and mossy mires around the tarns on the main summit, at 2,050 m, Pócs, Magill & LaFarge-England 90018/F; (Fianarantsoa prov.) Andringitra Mts Nature Reserve, bog forest dominated by two Pandanus spp. in a small depression called “Akofa” on the ridge E from Korokoro River, at 920 m, Kis 9476/ER & ET; Ericaceous heath forest at the N foot of Mt Vohipia near Ambavafatra, at 1,270–1,350 m, Kis 9477/EO; (Vakinankaratra) Manjakatompo, Ericaceous heath-forest (3–4 m tall) on the SE slopes of Mt Andriandrahitokana, at 1,975 m. – An Afro-American disjunct, widespread in the Andes and in the East African mountains and known also from Réunion and Mauritius, but new to the flora of the Comoros and of Madagascar.

Campylopus hildebrandtii (Müll. Hal.) A. Jaeger – REU: E edge of Cirque de Mafate. Mossy elfin forest (2–4 m tall) on the SW ridge and on the summit of Piton Marmite at 1,820–1,878 m, Orbán 9424/CK; Ft de Belouve. Ravine Bringellier SSE from Gite Belouve, on shady volcanic cliff at 1,460 m, Rózsa 9647/D. – Scattered all over sub-Saharan Africa from Cameroon to Cape and to Réunion.

Campylopus introflexus (Hedw.) Brid. – MAD: (Sava) Reserve Integrale Nationale de Marojezy, Cyperaceae and mossy mires around the tarns on the main summit, at 2,050 m, Pócs, Magill & LaFarge-England 90018/B; (Ihorombe) Isalo National Park, dry, xeric vegetation with Pachypodium rosulatum on slopes of sandstone canyon above “Piscine Naturelle”, at 800–1,000 m, Orbán 9455/CX; (Fianarantsoa prov.) Andringitra Mts near Ambalavao village. Plateau forest with granite outcrops and cultivated patches at 1,012 m, Orbán 9459CI. SEY: Mahé Island, Morne Seychellois Nat. Park. On rock boulders on the ridge near Trois Frères Cross, at 640–695 m, Pócs 9320/AG & BL. – A species quite widespread in the southern temperate belt penetrating in tropics and introduced into Atlantic Europe. It is known in the southern part of Africa (Botswana, Lesotho, Namibia, South African Republic, Swaziland, and Zimbabwe) and from Cameroon, Chad, Ascension, St Helena, Réunion and Mauritius islands. It is new to Madagascar and the Seychelles.

Campylopus jamesonii (Hook.) A. Jaeger – REU: The lowermost occurrence, where our team collected, was Cirque de Salazie. E slope of “la Roche Ecrite”, N from Grand Ilet village. Open evergreen bushes on steep rocks at 1,500 m, Orbán 9416/CT. – An Andean-Afromontane disjunct, common also on Réunion Island from the timberline up to 2,700 m.

Campylopus johannis-meyeri (Müll. Hal.) Kindb. – SEY: Mahé Island, Morne Seychellois Nat. Park. Pandanus-palm thicket on the rocky summit (glacis) of Montagne Palm-iste at 240–280 m, Orbán 9338/S. – Afromontane-alpine species till now only known from the continent. New to the Indian Ocean islands at an unusual low altitude.

Campylopus julaceus Jaeger subsp. arbogastii (Ren. et Card.) J.-P. Frahm – MAD: (Haute Miasatra) Ranomafana National Park. Open, seeping gneiss rocks along the Fiana-rant soa–Ifanadiana road near Vohiparara, at 1,115 m, Orbán 9465/CF. REU: Montane rainforest with Pandanus montanus on the SW slopes of Piton de la Fournanaise, at 800–1,300, Szabó 9612/I, Vojtkó 9657/A; Cirque de Cilaos, at the junction of Bras Sec road and the tourist trail No. G.R.1, at 1,320–1,390 m, in Acacia heterophylla forest, Pócs 9681/J. SEY: Mahé Island. Montane evergreen forest on the NE escarpment and summit ridge of Mt Trois Frères, W of Victoria at 430–770 m, Pócs 9315/AF & AP, Orbán 9343/BH; NW ridge of New Savy summit N from USAF Satellite Tracking Station. Evergreen bush on open rocks at 520 m, Kis 9332/H & I. – Indian Ocean islands vicariant of Campylopus julaceus subsp. julaceus of eastern Brasil. It prefers the open places with shallow soil, in montane habitats from about 500 m to the forest limit.

Campylopus nanophyllus Müll. Hal. ex Broth. – MAD: (Analamanga) granite out-crops in secondary plateau grassland, 69 km NW of Antananarivo along the Mahajanga road, N of Soaniada-Nana village at 1,410 m, Orbán 9438/CB; Antanarivo prov., Mt Angavokely near Carion village, on flat topped granite rock with open Xerophyta (Velloziaceae) bush at 1,700 m, Orbán 9454/CJ & EJ; (Irhombe) Isalo National Park. “Piscine Naturelle” 5 km W of Ranohira. Sandstone canyon with Pandanus pulcher, at 800 m, Vojtkó 9456/CB. – A species with scattered distribution all over Subsaharan Africa, but seems to be rare. A tiny xerophytic species of sunhit rock surfaces.

Campylopus nivalis (Brid.) Brid. (Syn.: Campylopus chrismarii (Müll. Hal.) Mitt.) – REU: Cirque de Salazie. Pleine des Merles. Ericaceous woodland with thick (60 cm) moss layer of Sphagnum and Breutelia at 1,800 m, Orbán 9422/CF; E edge of Cirque de Mafate. Mossy elfin forest (2–4 m tall) on the SW ridge and on the summit of Piton Marmite at 1,820–1,878 m, Orbán 9424/CP; NE ridge of the Piton des Neiges summit. Subalpine ericaceous bush at 1,800–2,500 m, Vojtkó 9239/CF. – Campylopus nivalis is an Andean-African species restricted to the subalpine and alpine belts. Therefore it occurs only on the highest Indian Ocean islands, like Madagascar and Réunion.

Campylopus perichaetialis P. de la Varde et Thér. – REU: NE ridge of Piton des Neiges. Subalpine ericaceous bush at 1,800–2,500 m, Kónya 9639/C; Alpine semidesert on the ESE ridge of Piton des Neiges summit, at 2,700–3,000 m, Vojtkó 9440/CO. SEY: Mahé Island, Morne Seychellois Nat. Park. Mossy elfin forest on the summit ridge SW of Trois Frères rocks, at 740–770 m, Orbán 9343/BF, BG. – Afromontane species previously known only from the continent. New to the Indian Ocean islands.

Campylopus pilifer Brid. – It was collected at 19 localities on all islands. – Widely distributed in the tropical, subtropical and oceanic parts of America and Eurasia. Very common xerophyte occurring mostly on acidic rocky or sandy ground.

Figs 6–11.
Figs 6–11.

Details of different species. 6 = Leaves of Campylopus flaccidus Ren. et Card. (from Pócs & Krog 89204/K). 7 = Inner and outer basal lamina cells of Campylopus julaceus subsp. arbogastii (Ren. et Card.) J.-P. Frahm (from Vojtkó 9657/A). 8 = Obtuse leaf apex of the same specimen. 9 = Leaf of Campylopus nanophyllus Müll. Hal. ex Broth. (from Orbán 9438/CB). 10 = Leaf base of Campylopus trachyblepharon subsp. comatus (Ren. et Card.) J.-P. Frahm (from Kis 9487/EG). 11 = Acute leaf apex of Campylopus pilifer Brid. in contrast to the obtuse one of C. julaceus (from Vojtkó 9440/CO)

Citation: Acta Botanica Hungarica 64, 1-2; 10.1556/034.64.2022.1-2.9

Campylopus praetermissus J.-P. Frahm – MAD: Antananarivo prov., (Analamanga) Mt Angavokely near Carion village. Roadside vegetation among Eucalyptus and Pinus plantations at 1,400–1,650 m, Orbán 9453/CF. REU: Acacia heterophylla-Cyathea glauca wood-land in Forêt de Bélouve, 1 km E from the Forest Station, at 1,450–1,500 m, Gyarmati 9614/ DS. – A relatively rare (or overlooked) tropical African species new to Madagascar and the second locality in Réunion Island.

Campylopus pyriformis (Schultz) Brid. – MAD: (Fianarantsoa prov.) Andringitra Mts Nature Reserve near Antanifotsi. On the path to Pic Boby. Secondary grassland with ericaceous bush patches at 1,900–2,000 m, Orbán 9463/CF, CH; Same nature reserve, peat bogs with Eriocaulon, Drosera and Sphagnum spp. on the Ambavafatra Plateau at 1,270 m, Kis 9478/EI. REU: Cirque de Salazie. Pleine des Merles. Ericaceous woodland with thick (60 cm) moss layer of Sphagnum and Breutelia at 1,800 m, Orbán 9422/CG; MAU: Moka Region, Quartier Militaire distr., on stone heaps in sugar cane fields W of Melrose village, at 330 m, Pócs 9678/B. – Bipolar temperate species also in tropical mountains (in the Northern Hemisphere occurs only in Oceanic Europe). New to the flora of Madagascar and Mauritius.

Campylopus robillardei Besch. – MAU: Perrier Nature Reserve in the neighbour-hood of Mare aux Vacoas, in submontane Sideroxylon puberulum thicket on volcanic rocks, 540 m, Pócs 9529/X. REU: Cirque de Salazie. E slope of “la Roche Ecrite”, N of Grand Ilet village. Open, ericaceous heath on rocky slopes between 1,700 and 2,100 m, Kis 9418/CP. SEY: Mahé Island, Morne Seychellois Nat. Park, on the rocky summit (glacis) of Mt Palm-iste at 240–280 m, Kis 9338/S; Same island, mossy elfin forest on the summit ridge SW of Trois Frères rocks, at 740–770 m, Orbán 9443/BG. – Lemurian species.

Campylopus thwaitesii (Mitt.) A. Jaeger (Syn.: Campylopus cardotii Thér.) – It is widespread in the Indian Ocean islands. We collected in Madagascar and Réunion at 8 localities. – A Lemurian species.

Campylopus torrentis Thér. et P. de la Varde – REU: Forêt de Bélouve. Montane mossy forest along the trail from the Forest Station to Pic des Chèvres, at 1,550–1,610 m, Kis 9617/AC. – A rare species known only from Gabon and Nigeria in West Africa. New to the whole East Africa.

Figs 12–35.
Figs 12–35.

Details of different species. 12 = Campylopus aureonitens (Müll. Hal.) A. Jaeger. Upper lamina cells (from Vojtkó 9420/CO). 13 = Tip of hyaline awn from the same specimen. 14 = Campylopus bartramiaceus (Müll. Hal.) Thér. Proximal section of costa (from Kis 9444/EF). 15 = Campylopus flaccidus Ren. et Card. Basal lamina cells (from Vojtkó 9456/CE). 16 = Upper lamina cells (from Orbán 9463/H). 17 = Proximal section of costa (from Orbán 9463/HM). 18 = Campylopus flexuosus var. incacorralis (Herz.) J.-P. Frahm. Pitted basal lamina cells (from Pócs 9150/R). 19 = Proximal section of costa (from Kis 9476/ET). 20 = The same (from Kis 9477/ EO). 21 = Campylopus julaceus subsp. arbogastii (Ren. et Card.) J.-P. Frahm. Tip of hyaline awn (from Vojtkó 9657/A). 22 = Proximal section of costa (from Kis 9332/H). The same (from Vojtkó 9657/A). 24 = Campylopus jamesonii (Hook.) A. Jaeg. Proximal section of costa (from Orbán 9423/CC). 25 = Campylopus johannis-meyeri (Müll. Hal.) Par. Proximal section of leaf (from Orbán 9338/S). 26 = Campylopus nanophyllus Müll. Hal. ex Broth. Proximal section of leaf (from Orbán 9438/CB). 27 = Campylopus perichaetialis P. de la Varde et Thér. Awn-like tip of leaf apex (from Orbán 9443/BG). 28 = Thin walled basal lamina cells (from the same specimen). 29 = Proximal section of costa (from the same specimen). 30 = Campylopus pilifer Brid. Proximal section of costa (from Vojtkó 9659/A). 31 = Campylopus praetermissus J.-P. Frahm. Proximal section of leaf (from Szabó 9614/DS). 32 = Proximal section of costa (from the same specimen). 33 = Campylopus torrentis Thér. et P. de la Varde. Awn-like tip of leaf apex (from Kis 9617/AC). 34 = Campylopus trachyblepharon subsp. comatus (Ren. et Card.) J.-P. Frahm. Pitted basal lamina cells (from Kis & Pócs 9332/G). 35 = Proximal section of leaf (from Kis 9612/M).

Citation: Acta Botanica Hungarica 64, 1-2; 10.1556/034.64.2022.1-2.9

Campylopus trachyblepharon (Müll. Hal.) Mitt. subsp. comatus (Ren. et Card.) J.-P. Frahm – MAD: (Alaotra-Mangoro) montane rainforest on the rocky ridge of Mt Andrianavibe, 5 km ESE of Andasibe (Périnet), at 950–1,050 m, Kis 9487/EG. REU: Montane rainforest with Pandanus montanus on the S slopes of Piton de la Fournaise, along the path leading to Puys Ramond, at 800–960 m, Kis 9612/H. SEY: Mahé Island, Morne Seychellois Nat. Park. Mossy elfin forest with rock boulders on the ridge near Trois Frères Cross, at 640–770 m, Kis & Pócs 9320/AT, Orbán 9343/BG, BK; NW ridge of New Savy summit N from USAF Satellite Tracking Station. Evergreen bush on open rocks at 520 m, Kis & Pócs 9332/G; Morne Seychellois Nat. Park. Copolia summit. Open, granite platform (glacis) at 480–510 m, Kis 9335/V; Elfin forest on the summit ridge of Mt Le Niol at the W side of Bernard Peak, at 630–660 m, Orbán 9351/BF. – Vicariant of Campylopus trachyblepharon subsp. trachyblepharon of Brasil, endemic subspecies of the Indian Ocean islands and new to Seychelles.

DISCUSSION

Analysing the distribution of Campylopoideae species of the Indian Ocean islands, it turned out that only one strictly endemic taxon occurs there, what is a vicariant subspecies of a South American species. It is surprising, comparing to the vascular or cryptogamic floras of Madagascar and neigh-bouring islands. According to estimates, the Indian Ocean islands have 20% of liverwort endemics (Vanderpoorten and Hallingbäck 2008, based on Wigginton pers. comm.) and Réunion Island has 14% of moss endemics (Ah Peng et al. 2010). The low number of endemics among the Campylopus taxa of the area may be interpreted by the relatively small spore size: 7.2–15 µm (LuiziPronzo and Barth 1999) and the ability of vegetative reproduction of most Campylopideae, which both enhance long range air dispersal (Zanten and Pócs 1981). At the same time there are three bicontinental (Afro-American) disjuncts, all of them occur in altimontane habitats.

There is a good number (8) of subendemic, so called Lemurian taxa, which are widely distributed on the islands, but occur also on continental SE Africa, mainly in the Precambrian crystalline mountains. This seems to confirm a possibility of short range, step by step dipersal or the importance of land connection before the Cretaceous between mainland Africa and Madagascar (Pócs 1975, 1999). The distribution patterns of all known Indian Ocean island Campylopoideae taxa are the following:

Endemic, restricted only to the islands1Campylopus trachyblepharonsubsp. comatus
Subendemics occurring also in southeast tropical Africa8Atractylocarpus madagascariensis, C. arctocarpossubsp. madecassus, C. aureonitens, C. decaryi, C. pseudobicolor, C. robillardei, C. schmidii, C. thwaitesii
Disjuncts in S Africa and the Indian Ocean Islands4C. bartramiaceus, C. bicolorsubsp. atroluteus, C. crateris, C. julaceussubsp. arbogastii
Widespread Afromontane species4C. cambouei, C. flavicoma, C. johannis-meyeri, C. perichaetialis
Widespread tropical African species9C. arcuatus, C. aureonitens, C. flaccidus, C. nanophyllus, C. hildebrandtii, C. perpusillus, C. praetermissus, C. smaragdinus, C. torrentis
Afro-American bicontinental disjuncts3Campylopus flexuosusvar. incacorralis, C. jamesonii, C. nivalis
Bipolar temperate species2C. flexuosus, C. pyriformis
Southern temperate species2C. catarractilis, C. introflexus (introduced in Europe)
Cosmopolite2C. fragilis, C. pilifer

Acknowledgements

I wish to thank for the financial support of collecting trips the National Geographic Society (USA), the Missouri Botanical Garden, the Hungarian Scientific Research Fund OTKA and the Hungarian Academy of Sciences. I am grateful to my collecting companions: Robert E. Magill, Catherine LaFarge-England, Stephen Manktelow and Attila Rupf, the members of our Bryology Research Group and Botany Department: Gabriella Kis, Erika Kónya, Sándor Orbán, Andrea Sass-Gyarmati, Ibolya Sütő-Bakalár, András Szabó and András Vojtkó, to Sándor Rózsa (Aggtelek Nat. Park), Min Chuah Petiot (Nairobi University). I am much obliged also to our guide in the Andringitra Massif, Rzafindrabe Jean-Baptiste de la Salle from the Tsimbazaza Botanical and Zoological Garden (Antananarivo) and to Sonja Ribes, director of the Natural History Museum of Réunion (Saint Denis) for her multiple logistic help during our visits to this island, finally to Catherine Reeb (Sorbonne University and Natural History Museum, Paris) for the careful revision of my text.

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  • Ah-Peng, C., Bardat, J., Stamenoff, P., Hedderson, T. A. J. and Strasberg, D. (2010): Bryophytes de l’ile de La Réunion: diversité, endemicité et conservation.–Cryptog., Bryol. 31: 241270.

    • Search Google Scholar
    • Export Citation
  • Bizot, M. and Kilbertus, G. (1979): Les Campylopus africains subg. lucidus Biz. et Kilb. –Rev. Bryol. Lichénol. 45: 6195.

  • Bizot, M. and Pócs, T. (1974): East African bryophytes I. –Acta Acad.Paed. Agriensis n. ser. 12: 383449.

  • Bizot, M., Dury, M. N. and Pócs, T. (1977): East African Bryophytes, II. Collections made by L. Ryvarden in Malawi, SE Africa.–Acta Bot. Acad. Sci. Hung. 22: 18.

    • Search Google Scholar
    • Export Citation
  • Bizot, M., Friis, I., Lewinsky, J. and Pócs, T. (1978): East African Bryophytes IV. Danish collections.–Lindbergia 4: 259284.

  • Crosby, M. R., Schultze-Motel, U. and Schultze-Motel, W. (1983): Katalog der Laubermoose von Madagaskar und den umliegenden Inseln.–Willdenowia 13: 187255.

    • Search Google Scholar
    • Export Citation
  • Een, G. (2009): Moss flora of the Island of Mauritius. –Trop. Bryol. 30: 4571.

  • Een, G. (2000): Bryophytes collected in Mauritius by Gunnar Erdtman. –Trop. Bryol. 18: 8790.

  • Frahm, J.-P. (1980): Synopsis of the genus Campylopus in North America north of Mexico.–The Bryologist 83: 570588.

  • Frahm, J.-P. (1982a): Grossdisjunktionen von Arealen suedamerikanischer und afrikanischer Campylopus-Arten.–Lindbergia 8: 3645.

  • Frahm, J.-P. (1982b): A reinterpretation of Bryohumbertia P. d. Vard. et Thér.–Cryptog. Bryol. Lichénol. 3(4): 365369.

  • Frahm, J.-P. (1985a): Die afrikanischen Campylopus-Arten. –Bryoph. Bibl. 36: 1216.

  • Frahm, J.-P. (1985b): Campylopus aureus Bosch et Lac. in the Hawaiian islands. –The Bryologist 88: 359360.

  • Frahm, J.-P. (1993): Taxonomic results of the BRYOTROP expedition to Rwanda and Zaire. 17. Andreaeaceae, Bruchiaceae, Dicranaceae, Rhizogoniaceae, Bartramiaceae, Rhacocarpaceae, Hedwigiaceae, Cryphaeaceae, Leucodontaceae. – Trop. Bryol. 8: 153169.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Frahm, J.-P. (1994): Taxonomische Notizen zur Gattung Campylopus XVI.–Nova Hedwigia 59: 147155.

  • Frahm, J.-P. (2000): New combinations in the genera Atractylocarpus and Metzleria. –Trop. Bryol. 18: 115117.

  • Frahm, J.-P. (2021): East African Bryophytes XXXII. Campylopoideae records.–Frahmia 22: 15. (ed. T. Pócs).

  • Frahm, J.-P. and Ho, B.-C. (2009): A new contribution to the moss flora of the inner Seychelles (revised edition). – Arch. Bryol. 38: 116.

    • Search Google Scholar
    • Export Citation
  • Frahm, J.-P. and O’Shea, B. J. (1996): British Bryological Society expedition to Mulanje, Malawi. 4. Dicranaceae: Campylopodioideae (Atractylocarpus, Bryohumbertia, Campylopus, Microcampylopus). – J. Bryol. 19: 119134.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Frahm, J.-P., O’Shea, B. J. and Ho, B.-C. (2009): The moss flora of Mauritius. –Arch. Bryol. 51: 126.

  • Luizi-Ponzo, A. P. and Barth, O. M. (1999): Spore morphology of some Dicranaceae species (Bryophyta) from Brazil. –Grana 38(1): 4249.

  • Marline, L., Andriamiarisoa, R. L., Bardat, J., Chuah-Petiot, M., Hedderson, T. A. J., Reeb, C., Strasberg, D., Wilding, N. and Ah-Peng, C. (2012): Checklist of the bryophytes of Madagascar.–Cryptog., Bryol. 33: 199255.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Ochyra, R. and Pócs, T. (1983): East African Bryophytes, VI. Polish collections.–Acta Bot. Acad. Sci. Hung. 28: 361389.

  • Ochyra, R. and Pócs, T. (1986a): East African Bryophytes, XI. Dr. L. Péntek’s collection from Mozambique.–Abstracta Bot. 9, Suppl. 2: 6386.

    • Search Google Scholar
    • Export Citation
  • Ochyra, R. and Pócs, T. (1986b): East African Bryophytes, X. K. Norikoshi’s collectioon from Mahale Mountains National Park, Western Tanzania.–Hikobia 9: 387394.

    • Search Google Scholar
    • Export Citation
  • O’Shea, B. J. (1995): Checklist of the mosses of sub-Saharan Africa. –Tropic. Bryol. . 10: 91198.

  • O’Shea, B. J. (2006): Checklist of the mosses of sub-Saharan Africa (version 5, 12/06). –Tropic. Bryol. Research Reports 6: 1252.

    • Search Google Scholar
    • Export Citation
  • O’Shea, B. J., Frahm, J.-P. and Porembski, S. (1996): Die Laubmoosflora der Seychellen. –Trop. Bryol. 12: 169191.

  • Padberg, M. and Frahm, J.-P. (1985): Monographie der Gattung Atractylocarpus Mitt. –Cryptog. Bryol. Lichénol. 4: 315341.

  • Pócs, T. (1975): Affinities between the bryoflora of East Africa and Madagascar. In: Miège, J. and Stork, A. (eds): Origines des flores africaines et malgaches. Nature, spéciation. Comp.-rend. de la VIIIe réunion de l’AETFAT, 1. – Boissiera 24a: 125128.

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (1997): The distribution and origin of the foliicolous bryophyta in the Indian Ocean Islands. In Cryptogams in the Phyllosphaere: Systematics, distribution, ecology and use. Proceedings of the IAB and IAL Symposium on Foliicolous Cryptogams, 29 August–2 September 1995, Eger, Hungary.–Abstracta Bot. 21: 123134.

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (1999): Bryophyte speciation and diversity in the East African mountains.–Bryobrothera 5: 237245.

  • Stech, M. (1999): A reclassification of Dicranaceae (Bryopsida) based on non-coding cp-DNA sequence data. –J. Hattori Bot. Lab. 86: 137159.

    • Search Google Scholar
    • Export Citation
  • Tixier, P. (1978): La spéciation lemurienne et les Lejeuneacées. Le cas du genre Diplasiolejeunea. In Suire, C. (ed.): Congrés International de Bryologie, Bordeaux, 21–23 Novembre 1977, Comptes Rendus.–Bryophyt. Bibl. 13: 622645.

    • Search Google Scholar
    • Export Citation
  • Tixier, P. and Guého, J. (1997): Introduction to Mauritian bryology, a check list of mosses and liverworts. – Mauritius Sugar Industry Research Institute, Réduit, Mauritius, 233 pp.

    • Search Google Scholar
    • Export Citation
  • Vanderpoorten, A. and Hallingbäck, T. (2009): Conservation biology of bryophytes. – In Goffinet, B. and Shaw, A. J. (eds): Bryophyte Biology, 2nd ed. Cambridge Univ. Press, Cambridge, pp. 487533.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Zanten, B. O. and Pócs, T. (1981): Distribution and dispersal of bryophytes. – In Schultze-Motel, W. (ed.): Advances in Bryology 1. J. Cramer, Vaduz, pp. 479562.

    • Search Google Scholar
    • Export Citation
  • Zijlstra, G. (1998): Report of the Committee for Bryophyta: 4.–Taxon 47: 859862.

  • Ah-Peng, C. and Bardat, J. (2005): Checklist of the bryophytes of Reunion Island (France).–Bryoph. Div. Evol. (Trop. Bryol.) 26: 89118.

  • Ah-Peng, C., Bardat, J., Stamenoff, P., Hedderson, T. A. J. and Strasberg, D. (2010): Bryophytes de l’ile de La Réunion: diversité, endemicité et conservation.–Cryptog., Bryol. 31: 241270.

    • Search Google Scholar
    • Export Citation
  • Bizot, M. and Kilbertus, G. (1979): Les Campylopus africains subg. lucidus Biz. et Kilb. –Rev. Bryol. Lichénol. 45: 6195.

  • Bizot, M. and Pócs, T. (1974): East African bryophytes I. –Acta Acad.Paed. Agriensis n. ser. 12: 383449.

  • Bizot, M., Dury, M. N. and Pócs, T. (1977): East African Bryophytes, II. Collections made by L. Ryvarden in Malawi, SE Africa.–Acta Bot. Acad. Sci. Hung. 22: 18.

    • Search Google Scholar
    • Export Citation
  • Bizot, M., Friis, I., Lewinsky, J. and Pócs, T. (1978): East African Bryophytes IV. Danish collections.–Lindbergia 4: 259284.

  • Crosby, M. R., Schultze-Motel, U. and Schultze-Motel, W. (1983): Katalog der Laubermoose von Madagaskar und den umliegenden Inseln.–Willdenowia 13: 187255.

    • Search Google Scholar
    • Export Citation
  • Een, G. (2009): Moss flora of the Island of Mauritius. –Trop. Bryol. 30: 4571.

  • Een, G. (2000): Bryophytes collected in Mauritius by Gunnar Erdtman. –Trop. Bryol. 18: 8790.

  • Frahm, J.-P. (1980): Synopsis of the genus Campylopus in North America north of Mexico.–The Bryologist 83: 570588.

  • Frahm, J.-P. (1982a): Grossdisjunktionen von Arealen suedamerikanischer und afrikanischer Campylopus-Arten.–Lindbergia 8: 3645.

  • Frahm, J.-P. (1982b): A reinterpretation of Bryohumbertia P. d. Vard. et Thér.–Cryptog. Bryol. Lichénol. 3(4): 365369.

  • Frahm, J.-P. (1985a): Die afrikanischen Campylopus-Arten. –Bryoph. Bibl. 36: 1216.

  • Frahm, J.-P. (1985b): Campylopus aureus Bosch et Lac. in the Hawaiian islands. –The Bryologist 88: 359360.

  • Frahm, J.-P. (1993): Taxonomic results of the BRYOTROP expedition to Rwanda and Zaire. 17. Andreaeaceae, Bruchiaceae, Dicranaceae, Rhizogoniaceae, Bartramiaceae, Rhacocarpaceae, Hedwigiaceae, Cryphaeaceae, Leucodontaceae. – Trop. Bryol. 8: 153169.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Frahm, J.-P. (1994): Taxonomische Notizen zur Gattung Campylopus XVI.–Nova Hedwigia 59: 147155.

  • Frahm, J.-P. (2000): New combinations in the genera Atractylocarpus and Metzleria. –Trop. Bryol. 18: 115117.

  • Frahm, J.-P. (2021): East African Bryophytes XXXII. Campylopoideae records.–Frahmia 22: 15. (ed. T. Pócs).

  • Frahm, J.-P. and Ho, B.-C. (2009): A new contribution to the moss flora of the inner Seychelles (revised edition). – Arch. Bryol. 38: 116.

    • Search Google Scholar
    • Export Citation
  • Frahm, J.-P. and O’Shea, B. J. (1996): British Bryological Society expedition to Mulanje, Malawi. 4. Dicranaceae: Campylopodioideae (Atractylocarpus, Bryohumbertia, Campylopus, Microcampylopus). – J. Bryol. 19: 119134.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Frahm, J.-P., O’Shea, B. J. and Ho, B.-C. (2009): The moss flora of Mauritius. –Arch. Bryol. 51: 126.

  • Luizi-Ponzo, A. P. and Barth, O. M. (1999): Spore morphology of some Dicranaceae species (Bryophyta) from Brazil. –Grana 38(1): 4249.

  • Marline, L., Andriamiarisoa, R. L., Bardat, J., Chuah-Petiot, M., Hedderson, T. A. J., Reeb, C., Strasberg, D., Wilding, N. and Ah-Peng, C. (2012): Checklist of the bryophytes of Madagascar.–Cryptog., Bryol. 33: 199255.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Ochyra, R. and Pócs, T. (1983): East African Bryophytes, VI. Polish collections.–Acta Bot. Acad. Sci. Hung. 28: 361389.

  • Ochyra, R. and Pócs, T. (1986a): East African Bryophytes, XI. Dr. L. Péntek’s collection from Mozambique.–Abstracta Bot. 9, Suppl. 2: 6386.

    • Search Google Scholar
    • Export Citation
  • Ochyra, R. and Pócs, T. (1986b): East African Bryophytes, X. K. Norikoshi’s collectioon from Mahale Mountains National Park, Western Tanzania.–Hikobia 9: 387394.

    • Search Google Scholar
    • Export Citation
  • O’Shea, B. J. (1995): Checklist of the mosses of sub-Saharan Africa. –Tropic. Bryol. . 10: 91198.

  • O’Shea, B. J. (2006): Checklist of the mosses of sub-Saharan Africa (version 5, 12/06). –Tropic. Bryol. Research Reports 6: 1252.

    • Search Google Scholar
    • Export Citation
  • O’Shea, B. J., Frahm, J.-P. and Porembski, S. (1996): Die Laubmoosflora der Seychellen. –Trop. Bryol. 12: 169191.

  • Padberg, M. and Frahm, J.-P. (1985): Monographie der Gattung Atractylocarpus Mitt. –Cryptog. Bryol. Lichénol. 4: 315341.

  • Pócs, T. (1975): Affinities between the bryoflora of East Africa and Madagascar. In: Miège, J. and Stork, A. (eds): Origines des flores africaines et malgaches. Nature, spéciation. Comp.-rend. de la VIIIe réunion de l’AETFAT, 1. – Boissiera 24a: 125128.

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (1997): The distribution and origin of the foliicolous bryophyta in the Indian Ocean Islands. In Cryptogams in the Phyllosphaere: Systematics, distribution, ecology and use. Proceedings of the IAB and IAL Symposium on Foliicolous Cryptogams, 29 August–2 September 1995, Eger, Hungary.–Abstracta Bot. 21: 123134.

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (1999): Bryophyte speciation and diversity in the East African mountains.–Bryobrothera 5: 237245.

  • Stech, M. (1999): A reclassification of Dicranaceae (Bryopsida) based on non-coding cp-DNA sequence data. –J. Hattori Bot. Lab. 86: 137159.

    • Search Google Scholar
    • Export Citation
  • Tixier, P. (1978): La spéciation lemurienne et les Lejeuneacées. Le cas du genre Diplasiolejeunea. In Suire, C. (ed.): Congrés International de Bryologie, Bordeaux, 21–23 Novembre 1977, Comptes Rendus.–Bryophyt. Bibl. 13: 622645.

    • Search Google Scholar
    • Export Citation
  • Tixier, P. and Guého, J. (1997): Introduction to Mauritian bryology, a check list of mosses and liverworts. – Mauritius Sugar Industry Research Institute, Réduit, Mauritius, 233 pp.

    • Search Google Scholar
    • Export Citation
  • Vanderpoorten, A. and Hallingbäck, T. (2009): Conservation biology of bryophytes. – In Goffinet, B. and Shaw, A. J. (eds): Bryophyte Biology, 2nd ed. Cambridge Univ. Press, Cambridge, pp. 487533.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Zanten, B. O. and Pócs, T. (1981): Distribution and dispersal of bryophytes. – In Schultze-Motel, W. (ed.): Advances in Bryology 1. J. Cramer, Vaduz, pp. 479562.

    • Search Google Scholar
    • Export Citation
  • Zijlstra, G. (1998): Report of the Committee for Bryophyta: 4.–Taxon 47: 859862.

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Acta Botanica Hungarica
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