View More View Less
  • 1 Botany Department, Institute of Biology, Eszterházy Károly University H-3301 Eger, Pf. 43, Hungary;
Open access

Drepanolejeunea clavicornis and D. friesii were previously synonymised with D. physaefolia or all of them with D. vesiculosa. In the meantime, Drepanolejeunea vandenberghenii was described from the same species group, as new. In this paper many African specimens are compared with the original descriptions of the above species. Morphological investigations of these and their distributional patterns suggested that the former synonymisation was not justified. In addition, a new, rheophytic species from the same group: Drepanolejeunea vanderpoortenii, is described, as new to science. As a result, from the taxa related to Drepanolejeunea vesiculosa, now six species are recognised from Africa, including its Indian Ocean islands. For these 6 morphotaxa an identification key is provided. The results need confirmation by a future molecular analysis.

Abstract

Drepanolejeunea clavicornis and D. friesii were previously synonymised with D. physaefolia or all of them with D. vesiculosa. In the meantime, Drepanolejeunea vandenberghenii was described from the same species group, as new. In this paper many African specimens are compared with the original descriptions of the above species. Morphological investigations of these and their distributional patterns suggested that the former synonymisation was not justified. In addition, a new, rheophytic species from the same group: Drepanolejeunea vanderpoortenii, is described, as new to science. As a result, from the taxa related to Drepanolejeunea vesiculosa, now six species are recognised from Africa, including its Indian Ocean islands. For these 6 morphotaxa an identification key is provided. The results need confirmation by a future molecular analysis.

INTRODUCTION

I already observed during our first collecting trip to Madagascar (March 1990) that most specimens identified as Drepanolejeunea physaefolia (Gottsche) Steph. or more properly D. physifolia (Gottsche) Pearson differ from the specimens collected in continental East Africa under the name of D. clavicornis Steph. and D. friesii Vanden Berghen, which were synonymised later with D. vesiculosa (Mitt.) Steph. (Tixier 1995, Wigginton and Grolle 1996) or with D. physifolia (Jones ex Wigginton 2004, formally not yet done). The main difference of Madagascan specimens is their dentate, sometimes serrulate leaf border and their ciliato-dentate perichaetial leaves.

I came to the conclusion, that the former synonymisations of the above Drepanolejeunea species were not well-founded. To clarify the matter, I started to study the more than 60 related African Drepanolejeunea specimens from our herbarium (EGR) and compare them with the original descriptions, illustrations and type specimens. I could successfully classify them into the species originally distinguished as Drepanolejeunea clavicornis, D. friesii, D. physifolia, D.vesiculosa and to the not long ago described D. vandenberghenii. At the same time, I tried to establish their different distribution pattern. Further molecular investigation has to confirm their separate existence at the species level.

Recently I received very interesting rheophytic specimens collected in Madagascar by prof. Alain Vanderpoorten (Liège University), which we started to study together with prof. Stephen Robbert Gradstein (Göttingen University), comparing it with Drepanolejeunea vandenberghenii Buchbinder et D.Fischer, not long ago described from dripping rocks in Rwanda and we describe it as Drepanolejeunea vanderpoortenii Gradstein et Pócs, a species new to science.

AFRICAN TAXA RELATED TO DREPANOLEJEUNEA VESICULOSA

Drepanolejeunea clavicornis Steph., Hedwigia 35: 81 (Stephani 1896). – (Figs 1–8) – Type: Insula São Tomé, Moller (holotype G!).

Figs 1–14. 1–8:
Figs 1–14. 1–8:

Drepanolejeunea clavicornis Steph. – 1–2 = habit, (from Shevock 42066, photo: Pócs); 3 = leaf; 4 = female perichaetium, 5 = bract; 6 = perianths with 4 or 5 globose horns; 7 = leaf margin; 8 = underleaves (slightly modified, from Vanden Berghen 1960, under the name of D. claviformis). 9–13: Drepanolejeunea friesii Vanden Berghen – 9 = habit, ventral view, 10 = lobule; 11 = underleaves; 12 = leaf; 13 = female perichaetium; 14 = gynoecium (9–11 and 14 modified from Buchbender and Fischer 2004, under the name of D. physaefolia, 12 from Vanden Beghen 1961, 13 from Vanden Berghen 1951)

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

The species was distinguished on the base of the inflated, globose perianth horns, which according to Stephani (1896: 82) is a unique and very interesting shape in the genus. The description and illustration (except for the misspelling of species name as D. claviformis) supplied by Vanden Berghen (1960: 61–62, fig. 7, shown in our Figs 3–8) agrees with, and more detailed than that of Stephani. Investigating many African and Asian specimens of this species group, I have never found transitions between this globose horned and the usual 4–5 keeled perianths and consider this character adequate to distinguish it at the species level.

The distribution of this taxon is also special, as it is restricted to West Africa (Fig. 15) except for one record from Madagascar (Cremers 3422 in Wigginton). It is recorded from São Tomé (Moller 385, holotype G) and Príncipe Islands (Monod 12114, BR in Vanden Berghen 1960), Camerun (Dusén, G). Our specimen (Figs 1–2) originates from Príncipe Island, along trail of Pico Papagaio above roça Santa Trinidade along summit ridge. NAD 83, 01° 36’ 38.6” N, 07° 23’ 32” E, 680 m, in mixed hardwood forest. On small diameter hardwood trunk. Shevock 42066 (CAS!, EGR!, previously identified by me, as D. physaefolia).

Fig. 15.
Fig. 15.

Distribution of Drepanolejeunea clavicornis (stars); of D. friesii (thick line) and of D. vesiculosa in Africa (dots)

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Drepanolejeunea friesii Vanden Berghen, Svensk Bot. Tidskr. 45: 366 (1951). – (Figs 9–13). – Type: Kenya, Mount Kenya occid: regio bambusina inferior in truncis Arundinariae alpinae, ca 2,500 m, 28 Jan. 1922, R. et T. Fries (holotype BR!).

When Vanden Berghen (1951) described it, underlined its similarity to Drepanolejeunea claviformis Steph., distinguishing by the missing ocelli and by the peculiar perianth shape. As ocelli of Drepanolejeunea species are often not observable on old herbarium specimens (Jones and Harrington 1983, Jones in Wigginton 2004), this is not a good character. In addition, when in fresh material they are still visible, in all species of this group 2–3 (to –5) suprabasal ocelli can form a short vitta. But the perianth shape remains as a useful distinguishing feature, as it is so strikingly different. Jones & Harrington (1983) mention another difference, apart from the perianth, in the shape of female bracts, which has only a few coarse teeth in D. friesii and laciniately dentate in D. clavicornis. They only tentatively identified their specimens from Sierra Leone and from Ghana, as D. friesii. The specimens examined by me from a great part of East Africa confirm their opinion. I have found on several specimens the same perianth type and perichaetial leaf shape, as described and illustrated by them. Buchbender and Fischer (2004: 279, fig. 5A, B, reproduced on our Figs 9–11 and 14) also published good illustrations of Drepeanolejeunea friesii (under the name of D. physaefolia). The differences from the real D. physifolia will be discussed under the latter species.

Drepanolejeunea friesii is the most widespread species of the group related to D. vesicaria all over the mountainous areas of continental Africa, and sporadically also on the Indian Ocean Islands (Fig. 15). It occurs on very different substrates, but prefers living leaves, ericaceous bark and twigs. It is published from Sierra Leone: Loma Mts., on sheltered boles, 1,680 m, Jones 1474 p.p., 1477 p.p., 1650 m, Jones 1491 (Jones and Harrington 1983). Ghana: Atewa Hills D. R., boughs in crown of Cassipourea, Jones1299d; boughs in crown of Lophira, Jones 1319a; upper part of bole of Antiaris, Jones 1320b (Jones and Harrington 1983). Rwanda: 27 km E of Shangugu, on roadcut surface, 1930 m, Symoens 5366 (BR); 30 km E of Shangugu, on the Kamiranzovu bog, 1,950 m, Erica bark, Symoens 5380 (Vanden Berghen. 1961). Kenya: Mt Kenya, bamboo belt, 2,500 m, R. & T. Fries 1922 (holotype, BR!); Tanzania: Mt Kilimanjaro (Vanden Berghen 1961); former Zaïre: Kivu (Vanden Berghen 1961).

Specimens seen and revised in our herbarium (EGR), which previously were identified mostly as D. physaefolia or as D. vesiculosa: former Zaïre: Kivu Prov., Kahuzi-Biega Nat. Park, dry Hagenia forest near Kaziruziru park gate, 2,300–2,370 m, on Agauria bark, Pócs 7268. – Kenya: Mt Kasigau, on the lower slopes of the peak, epiphyllous in mist forest, 1,400–1,600 m, Faden, Evans, Kariuki & Smeenk sn. 1971. Aberdare National Park (Nyandarua Range), around Chania Falls, 3,040–3,080 m, on Erica bark, Pócs 02030/AB. – Tanzania: Mt Meru, SW ridge along Engare Narok, 2,700–3,100 m, on Erica bark, Pócs, Mnyonga & Nsolomo 89194/W. Mt Kilimanjaro along Marangu Route, 2,130– 2,650 m, on Erica bark, Jones & Pócs 6365/K, Pócs & Orbán 89142/W, S.& T. Pócs 6240/O; Umbwe Route 2,850 m, on Erica twigs, Pócs 6929/NA; above Nkweseko, 1,800 m, Jones & Pócs 6359/E. W-Usambara Mts, NW slope of Magamba N of Lushoto, Ocotea-Podocarpus forest at 2,000–2,150 m, on bark, Pócs 6953/O; SE slope of Mt Kwagoroto, on shady rocks creeping on other liverworts, 1,750 m, Pócs 6957/V; Mazumbai, University Forest Reserve, mossy forest on Sagarav ridge, 1,750–1,980 m, on Erica bark and epiphyllous, Pócs & Jones 6372/DU, Pócs 6960/Z; Balangai West For. Res., Secondary heath, 1,550 m, on Ericaceae bark, Pócs 8509/D. N-Uluguru Mts, Lupanga ridge above Mbete village, elfin forest, 1,850–2,000 m, on twigs and epiphyllous, Pócs, Nchimbi 6285/BQ, 6287/N; Bondwa Peak above Morogoro town, elfin forest, 2,100 m, epiphyllous, Pócs 6233/N; on the top of Magari peak above Mzinga village, elfin forest, 2,340 m, epiphytic on Frullania, Pócs, Kondela, Nchimbi 06300/AS. S-Uluguru Mts, in bamboo (Sinarundinaria alpina) forest on the W slopes of Lukwangule Plateau, 2,250 m, on bamboo stem. Nguru Mts, Mt Kanga, S slope, 136 m at waterfalls, epiphyllous. Pócs 60140/AH, AI. – Réunion: Mare Longue Réserve Naturelle, Sentier Botanique, lowland rainforest on 4–500 years old lava flow, 280–320 m, on bark, Pócs 08056/B. W edge of Basse Vallée, along trail to Piton Larde, secondary Psidium cattleyanum bush, 750–985 m, on bark, Kis 9651/CG. S slope of Piton de la Fournaise volcano, alternating Pandanus montanus bush and mossy forest, 1,000–1,390 m, on palm trunk, Pócs 9612/CV; E edge of Cirque de Mafate, elfin forest on the SW ridge and summit of Piton Marmite, 1,820–1,878 m, on Erica bark, Szabó 9424/T. – Madagascar: Prov. Antsiranana, Réserve Integrale Nationale de Marojezy. Elfin forest on the SE ridge near Camp III, 1,830 m, epiphyllous. Pócs, Magill & LaFarge-England 90114/KG.

Drepanolejeunea physifolia (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892 (8): 8. – (Figs 1630) – Basionym: Lejeunea physaefolia Gottsche, Abh. Naturwiss. Vereins Bremen 7: 357 (1882). Type: Madagascar, in silva Ambatondrazaka sterilem inter alias Lejeunias repentem. Rutenberg 6. Dec, 1877. = Lejeunea securifolia Gottsche, Abh. Naturwiss. Vereins Bremen 7: 360 (1882). Type: Madagascar, in silva Ambatondrazaka inter alias Lejeunias. Rutenberg, probably at the same date. = Lejeunea intorta Besch. et Spruce, Bull. Soc. Bot. France 36, Suppl. CLXXXVIII. “1889” (1890). ≡ Drepanolejeunea intorta (Besch. et Spruce) Steph., Spec. Hep. 5: 323 (1913). Type: Réunion, Richard 641 ex Hb. Spruce (isotype G-17348).

Figs 16–24.
Figs 16–24.

Drepanolejeunea physifolia (Gottsche) Pearson – 16 = habit and 18 = gynoecium (after Stephani Icones 1985: 2447 and 2448, as D. intorta); 17 = habit (after Stephani Icones 2450. type of D. physaefolia); 19 = leaves; 20 = male branches; 21 = habit with gynoecium (after Arnell 1963); 22 = plants creeping on ericaceous twig (from Pócs, Orbán & Szabó 9486/H); 23 = habit, ventral view (from Pócs 08071/A); 24 =leaf (from Pócs, Orbán & Szabó 9486/H)

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Fig. 25
Fig. 25

Distribution of Drepanolejeunea physifolia (stars) and of D. vandenberghenii (squares)

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Figs 26–32.
Figs 26–32.

Drepanolejeunea physifolia (Gottsche) Pearson – 26 = underleaves (from S. & T. Pócs, Szabó 9889/AC); 27 = median lobe cells (from Pócs 08061/L); 28 = suprabasal ocelli (Pócs & Kis 9449/AT); 29 = female perichaetium (2 bracts + bracteole on the right, from S. Pócs 9890/ CJ); 30 = perianth (Pócs & Kis 9449/AT). – Figs. 31–32. Drepanolejeunea vesiculosa (Mitt.) Steph. s. str., habit with caduceus leaves – 31 (Pócs, Magill & Rupf 9274/S); 32 (Pócs 9188/A)

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Lejeunea physifolia (originally described, as “physaefolia”) sensu stricto is a species different from the previous taxa described from the African mainland and Sao Tomé Island. As the type material would have been destroyed during the 2nd World War in B (Grolle 1995) we can only relay on Stephani’s Icones (1985, figs 002450 and 002451), which depict both Drepanolejeunea physaefolia and D. securifolia based most probably on Gottsche’s types, as in Stephani’s time no other specimens were known. The illustration of D. intorta (Stephani 1985, fig. 002448, copied to our Figs 16 and 18) confirms Grolle’s concept synonymising it with D. physifolia. Both the Stephani Icones (1985) and the pictures by Arnell (1963: 181, fig. 132) illustrate the same species: a plant with obviously dentate lobe margin and with pinnately ciliato-laciniate perichaetial leaves. In fact, the acute papillae are turning outwards at the lobe margin forming acuminate teeth. Even the perianth keels are finely dentate as can be seen both on Arnell’s picture and on our microphoto (Figs 1921 and 2324). On the contrary, the lobe margins of Drepanolejeunea friesii (and of D. vesiculosa) are smooth or only minutely crenulate and the perichaetial leaves and perianth keels are almost entire or have a few coarse teeth (Figs 9–14). Concerning its distribution, D. physiformis s. str. in the original sense was published only from Southeast Africa and the Indian Ocean islands: Mozambique, Ngorongosa Mts. (Arnell 1957). South Africa: Transvaal, Pietersburg (now Polokwane); Basutoland, Woodbush; Natal, Ngome Forest (Arnell 1963). Comoros, Ndzouani island (Johanna or Anjouan, Herzog 1947); Ngazidja and Dzouani (Pócs 1993, 1995). Madagascar: type of Lejeunea physaefolia Gottsche (1882), Pearson (1892), and of L. securifolia (Grolle 1995), Pócs (1997). Réunion: Bescherelle and Spruce (1890) as type of Lejeunea intorta, W of Plaine des Palmistes, forest with tree ferns, above St. Paul, ca 2,000 m (Arnell 1965); Mauritius: Perrier Nature Reserve near Mare aux Vacoas; Le Pouce, rocks on old way between St. Pierre and Port Louis, ca 1,800 ft; Mt Cocotte, the top; the plateau N of Mt Cocotte (Arnell 1965). Seychelles: Mahé Island 100–800 m, not rare (Arnell 1957, Norkett, Onraedt in Grolle 1978); Praslin Island (Norkett in Grolle 1978).

Specimens seen and revised in our herbarium (EGR, duplicates of Madagascar specimens are deposited in TAN): I have found sporadic occurrence also in the mountains of East Africa, similarly to many other Lemurian flora elements: Kenya: Mt. Kenya E slope along Chogoria Trail, 2070 m, montane rainforest, on mosses, S. & T. Pócs, Chuah-Petiot 04002/X. – Tanzania: N-Uluguru Mts, NW slope and SW ridge of Lupanga above Morogoro town, mossy cloud forest at 1,700–2,000 m, on bark, Nchimbi & Pócs 6130/P, 6285/BY; NNE slope of Mnyera , below its western peak, mossy cloud forest, 1,900 m, epiphyllous, Pócs 6870/BF. S-Uluguru Mts, eastern ridge above Kibungo Mission, montane rainforest, 1,500–1,700 m, epiphyllous, Pócs & Lundqvist 6475/AD. WUsambara Mts, Kambi falls, 1,680 m, on bark, Pócs et al. 8528/Q. – The further records are all from the Indian Ocean islands: Comoro, Ndzouani Island, SE summit ridge of Mt Ntringi (N’Tingui), mossy elfin forest, 1,200–1,593 m, epiphyllous. – Madagascar: Antsiranana Province, Réserve spéciale de Manongarivo Ambahatra, cours supérieur, crête entre les deux bras de l’Ambahatra, forêt montagnarde, 1,250–1,300 m, sur tronc, Geissler 19698/2; Réserve Integrale Nationale de Marojezy, elfin forests on the SE ridge, 1,830 m, Pócs, Magill, LaFarge-England 90114/GQ and HN; Sava Region, Masoala National Park, summit ridge SE of Ambanizana village, 660–720 m, montane rainforest, on Pandanus leaf, Pócs & Kis 9449/AN, AT; Antanarivo Prov., forêt primaire dégradée, au lac de Mantasoa, 1,400 m, sur l’écorce, Onraedt, 74.M.2281; granite summits at the N end of Lake Mantasoa, near Anjozoro village, S of Manjakandriana town, ericaceous heath, 1,500 m, on bark, Pócs, Szabó, Ranaivojanoa 9851/B; Toamasina Prov., Andasibe-Mantadia Nat. Park, rainforest E of Andasibe village, 900–970 m, on branches, S. & T. Pócs, A. Szabó 9889/AC; rocky ridge 5 km ESE of Andasibe, montane rainforest, 950–1,050 m, epiphyllous, Szabó 9487/JL; NE of Andasibe, rainforest in a rocky streambed along tributary of Saharanga stream, 1,000 m, on twigs, Pócs, Orbán, Szabó 9486/H; Toamasina Prov., Maromizaha Forest Res. S of Andasibe Nat. Park, 2 km SW of Anevoka village, summit ridge of Mt Maromizaha, montane rainforest wish bamboo (Nastus sp.), 1,080–1,214 m, on Pandanus leaves, S. & T. Pócs 9890/CJ; Prov. Fianarantsoa, Ranomafana Nat. Park, near Ambodiamontana settlement, in the Valley of Namorona River, degraded submontane rainforest with relatively low (8– 18 m tall) canopy, at 865–1,015 m, epiphyllous, S. & T. Pócs et al. 04118/CL; W slope of Mt Vatolampy 2.5 km W of Ambatolahy village, montane rainforest, 1,080–1,200 m, on branches, S. & T. Pócs 04128/AB, 04129/BA; Haute Matsiatra Reg., Andringitra Mts and Nat. Park, W side of Korokoro River around Camp II, 750–1,000 m, on twigs, Kis 9472/BQ; N–S ridge running 6 km E from Pic Bory, near camp IV, mesic forest dominated by bamboos, 1,450–1,700 m, on decaying bamboo stem, Pócs 9475/AH. – Réunion: Plaine des fougères 3 km from Bois Rouge village, on the N slope of Piton Bé Massoune, ericaceous heath alternating with mossy elfin forest, 1,200–1,320 m, on decaying wood, Pócs 08071/EA; Cirque de Cilaos, forêt de la Mare a Josepf, E from Le Bloc, 1,370–1,380 m, on bark, Szabó, 9638/CA; Piton de la Fournaise, ericaceous elfin forest on the S slope of Piton Larde, 1,450–1,650 m, on twig; Forêt de Belouve, 1 km E from the Forest Station, Acacia heterophylla–Cyathea glauca woodland, 1,450–1,500 m, Szabó 9614/CJ; Forêt de Belouve, cliff at the SE edge of Cirque de Salazie, Acacia heterophylla–Cyathea glauca woodland, 1,500–1,555 m, Pócs 08061/L; Forêt de Belouve, montane mossy forest along the trail from the Forest Station to Pic de Chèvres, 1,560–1,610 m, on decaying wood, Gyarmati & Kis 9617/AF.

Drepanolejeunea vandenberghenii Buchbender et Eb. Fisch., J. Bryol. 26: 273 (2004). – (Fig. 25) – Type: Rwanda: Forêt de Rugege (= Nyungwe), rochers suinants à Sphagnum et Breutelia, 2,020 m, 20 Décembre 1971, J. L. De Sloover 12535 (holotype BR, paratypes KOBL, EGR!).

This species can be separated from Drepanolejeunea friesii and of D. vesicaria by its widely obtusely rounded leaf and underleaf lobe apices and by its fully edentate bracts. It was described from dripping rocks, growing among Sphagnum and Breutelia, accompanied by detailed illustrations. The detailed and well-illustrated protologue describes 5 other paratypes near to the type locality collected from different substrates in Nyungwe Forest by Buchbender, Fischer, Solga and Pócs. Later on it was reported also from Malawi (Pócs 2008).

Additional specimen in EGR: Réunion, SSW slope of Piton de la Fournaise volcano. N of Piton Larde, 1,650–1,850 m, open montane evergreen forest with Acacia heterophylla in the canopy, on bark. Szabó 9655/CO.

Drepanolejeunea vesiculosa (Mitt.) Steph. s. str., Spec. Hep. 5: 356 (1913). – (Figs 15, 3031) – Basionym: Lejeunea vesiculosa Mitten, J. Proc. Linn. Soc. Bot. 5: 116 (1861). Type: Insula Ceylon, ad Macromitrias, Gardner (NY).

I consider the African synonyms of D. vesiculosa enumerated in Wigginton and Grolle (1996: 73), insufficiently justified, based on Tixier’s synonymisation (Tixier 1995). Although only Drepanolejeunea friesii is similar in all other aspects, Drepanolejeunea vesiculosa has a character, the caduceus leaves, which do not occur in the majority of species described from Africa. But it is still a problem, whether this character is stable or not by the type and by all Asian specimens of D. vesiculosa. If any, then only Drepanolejeunea friesii of the African species can be synonymous with it. This finally can be decided after molecular investigation will have been done on many populations (not on one or two specimens).

For the moment I kept D. vesiculosa and D. friesii separated and was checking, whether any African specimens have caducous leaves. If this only specific distinction is right, I have found a few of them corresponding to this character, with caducous leaves, as follows:

Supposed African D. vesiculosa specimens in the herbarium EGR: Tanzania: N-Uluguru Mountains, elfin forest on Bondwa peak above Morogoro town, 2,120 m, on bark, Pócs 6889/M. – Malawi: W part of Mulanje Mts, fire effected, rocky Agauria–Protea woodland on the steep NW slopes of Likabula Valley along Lichenya path, on the N slope of Mt Chilemba, 1,800 m, on Agauria bark. – Comoro: Ngazidja Island, SE ridge of Mt Karthala above Nioumamilima village, submontane rainforest, 750 m, epiphyllous, with perianth, Pócs, Magill, Rupf 9266/BL; Ndzouani island, montane rainforests on the SE ridge of Mt Ntringi (Tingui) on the N side of Lake Dzialandze, 920–1,200 m, epiphyllous and corticolous, Pócs, Magill, Rupf 9273/I and 9273/BL. – Madagascar: Prov. Antsiranana, Réserve Integrale Nationale de Marojezy on the ridge N of Andampibe Falls, montane rainforest, 780–1,050 m, epiphyllous. Pócs, Magill, LaFarge-England 90113/FY.

Drepanolejeunea vanderpoortenii Gradstein et Pócs, spec. nova (Figs 3347)

Close to D. vandenberghenii , but well different by its blackish pigmentation, julaceous habit, thick-walled stems and, especially, the large preapical tooth almost touching the apical (= first) lobule tooth. The blackish pigmentation of the plants and the thick-walled stems may be interpreted as adaptations to the rheophytic habitat (Gradstein and Vidal 1975, Pócs 2010).

Type: Madagascar, Toamasine Province, Andasibe (Périnet) Mantadia National Park. Chutes Sacrées 4, 1,029 m, 18° 49’ 52” S, 48° 26’ 12” E, on flooded stone in stream in submontane rainforest, October 2018, A. Vanderpoorten MD 67 (holotype EGR!, isotype GOET!).

Forms dense, shiny blackish brown, sepia or rusty mats on streambed stones often under water. Shoots irregularly branching, 3–5 mm long, arcuately ascendent, the leaves even in wet stage upwards incurved, laying over the stem, forming an almost julaceous shoot of 2–300 µm width (with open leaves up to 450 µm wide). Stem diameter 50–80 µm, with 3–4 medullary and 7–8 cortical cells, with evenly incrassate (2–2.5 µm thick) and brownish pigmented walls.

Leaves strongly convex, asymmetrically triangular-ovate and falcate, 400–500 µm long and 300 µm wide when flattened, spreading at an angle of about 40° to the stem. Lobe with entire or finely crenulate margins and obtuse apex. Median lobe cells 25–29 µm, apical lobe cells 22–24 µm, isodiametric, with thin to moderately incrassate walls, at the base with trigones and on their dorsal side slightly mamillose-papillose with a low, acute papilla. Ocelli 2–3 suprabasal, forming a short vitta, cells of ocelli 50–55 × 22–25 µm. Lobules 100–150 µm long and 70–90 µm wide, apical (first) lobule tooth 1-celled, falcate, acute, lobule mouth semi-circular, distal end of the lobule with a large, 20–25 µm long, subglobose pre-apical tooth that is almost touching the apical tooth, the pre-apical tooth with chloroplasts and with walls of similar thickness to the other lobule cells (Figs 37, 39); keel curved, crenate in the upper half, angle between keel and ventral lobe margin sharp. Underleaves hardly wider than the stem, 50–75 × 70–80 µm, with an obcuneate to subquadrate lamina, a U-shaped sinus and two short, 1-cell wide, diverging lobes, the lobes two cells long and two cells wide at the base, ending in one cell. Female bract about 300 µm long, bilobate to its 2/3 length, acute with a few obtuse teeth. Bracteole 240 µm long, bilobate to its half-length. Perianth and androecium not seen. Vegetative reproduction not known.

Figs 33–39.
Figs 33–39.

Drepanolejeunea vandenpoortenii Gradstein et Pócs, spec. nova – 33 = habit, dorsal view (photo: Pócs); 34 = habit, side view; 35 = habit, ventral view, 36 = leaf, ventral view; 37 = lobule, ventral view; 38 = cross section of stem; 39 = perichaetium, bracteole on upper right side. (33–34, 36–39 from the holotype, Vanderpoorten MD 67, 35 from paratype, Pócs & Szabó 9878/AB)

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Figs 40–45.
Figs 40–45.

Drepanolejeunea vandenpoortenii Gradstein et Pócs, spec. nova – 40 = chloroplasts are shown in the pretooth; 41 = lobule with pre-tooth, behind its antical margin the ocelli are visible; 42= lobule apex with pre-tooth and hyaline papilla; 43 = underleaves; 44 = median lobe cells; 45 = lobe apex (40, 44 and 45 drawn from paratype, Vanderpoorten MD 62; 41–43 from paratype, Pócs & Szabó 9878/AB).

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Fig. 46.
Fig. 46.

The type locality of Drepanolejeunea vanderpoortenii, Chutes Sacrées in Andasibe-Mandtadia National Park (photo: Andrea Sass-Gyarmati)

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Fig. 47
Fig. 47

Distribution of Drepanolejeunea vanderpoortenii

Citation: Acta Botanica Hungarica ABot 63, 1-2; 10.1556/034.63.2021.1-2.11

Etymology: The species is dedicated to its collector, Prof. Alain Vanderpoorten of Liège University, renown expert on tropical mosses and bryophyte conservation.

Additional specimens examined: Madagascar, Toamasina Prov., Andasibe-Mantadia National Park, Chutes Sacrées 4, 1,029 m, 18° 49’ 52” S, 48° 26’ 12” E, on wet boulder in stream in submontane rainforest, October 2018, A. Vanderpoorten MD 62 (paratype EGR!, isoparatype GOET!). Toamasina Prov., Mananara Nord Biosphere Reserve and National Park. Lowland rainforest on the E slopes of Mahavoho Hill, On streambed stones of Mahavoho River, at 220 m alt. 16° 27’ S, 49° 47.00’ E, T. Pócs & A. Szabó 9878/AB, 14 Aug. 1978 (paratype EGR, isoparatype GOET! TAN!).

DISCUSSION

The group of taxa I summed up under the name of ‘Drepanolejeunea vesiculosa group’ exemplifies very well the problem arising from the difficulty to distinguish infraspecific variability (often influenced by environmental conditions) from the species level of genetical isolation. We have to take in account the morphological malleability, the distribution pattern and the cryptic species formation. Doubtless, molecular investigation is an answer, but only, if before the DNA investigation and its statistical analysis we try to define well the units to be investigated. I tried to do this, by the help of careful morpho-taxonomic and area-geographical investigation of the primarily described species before their premature and unjustified synonymisation. A future molecular analysis can prove the rightness of the formerly defined species concepts.

In case of Drepanolejeunea clavicornis I am quite sure about the good definition of this species, being allopatric and having a very different perianth shape. The latter property is not so much exposed to the environmental conditions than the vegetative body, therefore I attach great significance to it.

Drepanolejeunea physifolia in strict sense is more or less an allopatric vicariant of D. vesiculosa or D. friesii. It probably evolved when Madagascar was still united to mainland Africa or isolated later in the Indian Ocean islands and reached the continent by air dispersal. It is an interesting fact, that most plants are female with perichaetium, very rarely fertilised, as perianths and sporophytes are extremely rare.

Drepanolejeunea vanderpoortenii is an allopatric and at the same time a peripatric species. The niche isolation seems to act very well, because rheophytic life form is not known among other Drepanolejeunea species. At the same time this taxon occurs only in Madagascar and has two morphological properties (lobule pre-tooth and blackish pigmentation) unknown by its relatives.

The cases of Drepanolejeunea friesii and Drepanolejeunea vandenberghenii are not so clearcut. They are sympatric and their morphological differences against the very widespread Drepanolejeunea vesiculosa are not so sharp and some transitions seem to occur between them. In their case indeed only molecular investigation can clarify if they can be distinguished at the species level or not. The distribution of these three species, if they are really distinct, in Africa is still inadequately known.

Identification key

Based on the above morphological characters, I composed an identification key for the concerned taxa:

1a Perianth inflated, terete with 4–5 globose horns. Leaf margin crenulated. West African species. In all other aspects similar to Drepanolejeunea friesiiD. clavicornis

1b Perianth in its upper half with five equal keels each ending up in an apiculate or rounded rostrum2

2a Lobe margin with leaf margin dentate by the outward turning, high, acute papillae on (almost) each marginal cells. Female perichaetium leaves pinnately ciliato-laciniate, with teeth in length reaching or exceeding the half width of the bract or bracteole and ending in 2–3 uniseriate cells. Perianth keels regulately dentate at least on their rostrum.D. physifolia

2b Lobe margin entire or finely crenulated, acute papillae restricted to the lobe surface. Female perichaetium leaves almost entire or coarsely dentate with teeth shorter than half width of the bracts or bracteoles and not ending in uniseriate ciliae. Perianth keels entire or coarsely dentate 3

3a Leaves and underleaf lobes acuminate or acute. Female bracts coarsely toothed4

3b Leaves and underleaf lobes with broad, rounded apex. Female bracts with almost entire margins5

4a Leaves usually caducous, imbricate to distant. Denudate stems with remaining underleaves and fallen leaves for vegetative propagation often seen. Palaeotropical species, widespread in tropical Asia and the Pacific, scattered in AfricaD. vesiculosa

4b Leaves never caducous, leaves densely imbricate, vegetative reproduction unknown. The commonest species in the mountainous areas of AfricaD. friesii

5a Plants yellowish to light brownish-green, without blackish pigmentation. Stem with moderately thicked, colourless cell walls. Leaves imbricate, at apex often incurved, but the shoot never julaceous. Underleaves trapezoid in shape. Only the first lobular tooth is well developed, acute, falcate, one celled. There is no pretooth present. Although prefers wet habitat, never occurs in running waterD. vandenberghenii

5b Plants with blackish pigmentation, sepia or dark rust brown. Stem with incrassate (2 µm), brownish pigmented walls. Leaves densely imbricate, fully incurved towards the stem, forming julaceous habit. The apical (first) lobule tooth 1 celled, falcate. On each lobule a 20–25 µm large, almost globose pretooth is attached to the last cell at the distal end of lobule margin, almost touching the apical tooth. Rheophytic species, occurring on stones in running water or on irrigated streambankD. vanderpoortenii

Acknowledgements

I wish to thank for the financial support of collecting trips to the Sokoine University of Agriculture, Morogoro and to the Vice President’s Office in Tanzania, to the National Geographic Society (USA), to the Missouri Botanical Garden and to the ALCOA Foundation (USA), to the Swedish-Hungarian-Tanzanian Usambara Rainforest Program sponsored by the SAREC agency, to the Hungarian Scientific Research Fund OTKA, the Hungarian Academy of Sciences and the Hungarian Ministry of Education and Culture. I am grateful to my collecting companions Robert E. Magill, Catherine LaFarge-England and my late wife Sarolta, the staff of the Sokoine University of Agriculture, Morogoro and to members of our Bryology Research Group and Botany Department: Gabriella Kis, Sándor Orbán, Andrea Sass-Gyarmati, András Szabó and András Vojtkó, to the Head of Matuka David Kabala Research Centre of the Mananara-Nord Biosphere Reserve in Antanambe and to our experienced guide there, Mr Jean Baptiste Radriaka. Thanks are due to the directors and curators of the Meise Botanic Garden Herbarium (BR) and to the Geneva Herbarium (G) for the loan of valuable type specimens. Finally I am grateful to the review made by, Prof. S. R. Gradstein (GOET) and for his useful advise and corrections.

References

  • Arnell, S. W. (1957): Notes on some South African Hepatics V. – Bot. Not. 110: 399405.

  • Arnell, S. W. (1963): Hepaticae of South Africa. – Stockholm,411 pp.

  • Arnell, S. W. (1965): Hepaticae collected by Mr. Gillis Een in Mauritius and Réunion in 1962. – Svensk Bot. Tidsk.59: 6584.

  • Bescherelle, E. and Spruce, R. (1889): Hépatiques nouvelles des colonies Françaises. – Bull. Soc.Bot.France 36: clxxvii–clxxxix.

  • Buchbender, V. and Fischer, E. (2004): Drepanolejeunea vandenberghenii Jungermanni-opsida: Lejeuneaceae), a previously overlooked new species from Rwanda includ-ing comments on D. physaefolia and Harpalejeunea fischeri. – J. Bryol. 26: 273283. https://bibli.cbnbl.org/index.php?lvl=notice_display&id=606591

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Gottsche, C. M. (1882): Reliquiae Rutenbergianae Lebermoose. – Abh. Naturwiss.Vereine Bremen 7: 338365.

  • Gradstein, S. R. and Vidal, D. M. (1975): On Myriocoleopsis Schiffn. (Lejeuneaaceae). – Lindbergia 3: 3945.

  • Grolle, R. (1978): Die Lebermoose der Seychellen. – Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. R.27: 717.

  • Herzog, T. (1947): Hepaticae von der Comoreninsel Johanna. – Bot. Not. 101: 317334.

  • Jones, E. W. and Harrington, A. J. (1983): The Hepaticae of Sierra Leone and Ghana. – Bull. British Mus. (Nat. Hist.), Bot. ser.11: 215289.

    • Search Google Scholar
    • Export Citation
  • Mitten, W. (1860): Hepaticae Indiae Orientalis. – J. Proc. Linn. Soc. Bot.5(18): 89128.

  • Pearson, W. H. (1892): Lejeuneae madagascarienses. – Forh. Videns.-Selsk. Kristiania 8: 39.

  • Pócs, T. (1993): New or little known epiphyllous liverworts, IV. Two new Cololejeuneoideae from the Comoro Archipelago. – J. Hattori Bot. Lab.74: 4557.

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (1995): East African Bryophytes, XIV. Hepaticae from the Indian Ocean Islands. – Fragm. Flor. Geobot.40: 251277.

  • Pócs, T. (1997): The distribution and origin of the foliicolous bryophyta in the Indian Ocean Is-lands. –Abstracta Bot. 21: 123134. https://www.jstor.org/stable/43519212

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (2008): British bryological society expedition to Mulanje Mountain, Malawi. 19. Metzgeriaceae and other unpublished records. – Trop. Bryol.29: 13.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Pócs, T. (2010): Myriocoleopsis in Southeast Asia. – Trop. Bryol. 31: 123125.

  • Stephani, F. (1896): Hepticarum species novae IX. – Hedwigia 35(3).

  • Stephani, F. (1912): Species Hepaticarum 5. – George & Cie, Gèneve & Bâle, 1176.

  • Stephani, F. (1916): Species Hepaticarum 5. – George & Cie, Genève & Bâle, 8331008.

  • Stephani, F. (1985): Icones Hepaticarum. – Jardin Botanique de Genève, Zug, 1231 5 pp. (165 microfiches).

  • Tixier, P. (1995): Résultats taxonomiques de l’éxpédition Bryotrop au Zaire et Rwanda. 30. – Bryophytes épiphylles (récoltes de E. Fischer). – Trop. Bryol.11: 1176.

    • Search Google Scholar
    • Export Citation
  • Vanden Berghen, C. (1951): Note sur quelques hépatiques récoltées par R. E. Fries, en 1922. su Mont Kénia. – Svensk Bot. Tidsk.45(H.2): 362367.

    • Search Google Scholar
    • Export Citation
  • Vanden Berghen, C. (1960): Hépatiques récoltées en Afrique par M. Monod. – Rev. Bryol. Lichénol.29(1–2): 5067.

  • Vanden Berghen, C. (1961): Hépatiques récoltées par Dr J.-J. Symoens dans la région péri-tanganyikaise. –Bull. Soc. Roy. Bot. Belg.93: 5574.

    • Search Google Scholar
    • Export Citation
  • Wigginton, M. J. (2004): Checklist and distribution of the liverworts and hornworts of subSaharan Africa, including the East African Islands (edition 2, September 2004). – Trop. Bryol. Res. Reports 5: 1104.

    • Search Google Scholar
    • Export Citation
  • Wigginton, M. J. (ed.) (2004): E. W. Jones’s liverwort and hornwort flora of West Africa. – Scripta Bot. Belg.30: 1443.

  • Wigginton, M. J. and Grolle, R. (1996): Catalogue of the Hepaticae and Anthocerotae of Sub-saharan Africa. – Bryophyt. Bibl. 50: 1267.

    • Search Google Scholar
    • Export Citation
  • Arnell, S. W. (1957): Notes on some South African Hepatics V. – Bot. Not. 110: 399405.

  • Arnell, S. W. (1963): Hepaticae of South Africa. – Stockholm,411 pp.

  • Arnell, S. W. (1965): Hepaticae collected by Mr. Gillis Een in Mauritius and Réunion in 1962. – Svensk Bot. Tidsk.59: 6584.

  • Bescherelle, E. and Spruce, R. (1889): Hépatiques nouvelles des colonies Françaises. – Bull. Soc.Bot.France 36: clxxvii–clxxxix.

  • Buchbender, V. and Fischer, E. (2004): Drepanolejeunea vandenberghenii Jungermanni-opsida: Lejeuneaceae), a previously overlooked new species from Rwanda includ-ing comments on D. physaefolia and Harpalejeunea fischeri. – J. Bryol. 26: 273283. https://bibli.cbnbl.org/index.php?lvl=notice_display&id=606591

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Gottsche, C. M. (1882): Reliquiae Rutenbergianae Lebermoose. – Abh. Naturwiss.Vereine Bremen 7: 338365.

  • Gradstein, S. R. and Vidal, D. M. (1975): On Myriocoleopsis Schiffn. (Lejeuneaaceae). – Lindbergia 3: 3945.

  • Grolle, R. (1978): Die Lebermoose der Seychellen. – Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. R.27: 717.

  • Herzog, T. (1947): Hepaticae von der Comoreninsel Johanna. – Bot. Not. 101: 317334.

  • Jones, E. W. and Harrington, A. J. (1983): The Hepaticae of Sierra Leone and Ghana. – Bull. British Mus. (Nat. Hist.), Bot. ser.11: 215289.

    • Search Google Scholar
    • Export Citation
  • Mitten, W. (1860): Hepaticae Indiae Orientalis. – J. Proc. Linn. Soc. Bot.5(18): 89128.

  • Pearson, W. H. (1892): Lejeuneae madagascarienses. – Forh. Videns.-Selsk. Kristiania 8: 39.

  • Pócs, T. (1993): New or little known epiphyllous liverworts, IV. Two new Cololejeuneoideae from the Comoro Archipelago. – J. Hattori Bot. Lab.74: 4557.

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (1995): East African Bryophytes, XIV. Hepaticae from the Indian Ocean Islands. – Fragm. Flor. Geobot.40: 251277.

  • Pócs, T. (1997): The distribution and origin of the foliicolous bryophyta in the Indian Ocean Is-lands. –Abstracta Bot. 21: 123134. https://www.jstor.org/stable/43519212

    • Search Google Scholar
    • Export Citation
  • Pócs, T. (2008): British bryological society expedition to Mulanje Mountain, Malawi. 19. Metzgeriaceae and other unpublished records. – Trop. Bryol.29: 13.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Pócs, T. (2010): Myriocoleopsis in Southeast Asia. – Trop. Bryol. 31: 123125.

  • Stephani, F. (1896): Hepticarum species novae IX. – Hedwigia 35(3).

  • Stephani, F. (1912): Species Hepaticarum 5. – George & Cie, Gèneve & Bâle, 1176.

  • Stephani, F. (1916): Species Hepaticarum 5. – George & Cie, Genève & Bâle, 8331008.

  • Stephani, F. (1985): Icones Hepaticarum. – Jardin Botanique de Genève, Zug, 1231 5 pp. (165 microfiches).

  • Tixier, P. (1995): Résultats taxonomiques de l’éxpédition Bryotrop au Zaire et Rwanda. 30. – Bryophytes épiphylles (récoltes de E. Fischer). – Trop. Bryol.11: 1176.

    • Search Google Scholar
    • Export Citation
  • Vanden Berghen, C. (1951): Note sur quelques hépatiques récoltées par R. E. Fries, en 1922. su Mont Kénia. – Svensk Bot. Tidsk.45(H.2): 362367.

    • Search Google Scholar
    • Export Citation
  • Vanden Berghen, C. (1960): Hépatiques récoltées en Afrique par M. Monod. – Rev. Bryol. Lichénol.29(1–2): 5067.

  • Vanden Berghen, C. (1961): Hépatiques récoltées par Dr J.-J. Symoens dans la région péri-tanganyikaise. –Bull. Soc. Roy. Bot. Belg.93: 5574.

    • Search Google Scholar
    • Export Citation
  • Wigginton, M. J. (2004): Checklist and distribution of the liverworts and hornworts of subSaharan Africa, including the East African Islands (edition 2, September 2004). – Trop. Bryol. Res. Reports 5: 1104.

    • Search Google Scholar
    • Export Citation
  • Wigginton, M. J. (ed.) (2004): E. W. Jones’s liverwort and hornwort flora of West Africa. – Scripta Bot. Belg.30: 1443.

  • Wigginton, M. J. and Grolle, R. (1996): Catalogue of the Hepaticae and Anthocerotae of Sub-saharan Africa. – Bryophyt. Bibl. 50: 1267.

    • Search Google Scholar
    • Export Citation
All Time Past Year Past 30 Days
Abstract Views 0 0 0
Full Text Views 44 45 45
PDF Downloads 32 32 32