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  • 1 Department of Botany, , the Zamorin’s Guruvayurappan College (affiliated to the University of Calicut), GA College P. O., Kozhikode, Kerala-673014, , India;
  • | 2 Department of Botany, , Institute of Biology, Eszterházy University H-3301 Eger, Pf. 43, , Hungary;
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The genus Diplasiolejeunea is added to the liverwort flora of India from the Kerala part of the Western Ghats reporting two species: Diplasiolejeunea cavifolia Steph. and D. cobrensis Gottsche ex Steph. The genus is known from tropical Asia with eight species. Among them Diplasiolejeunea cavifolia is widespread, but still the genus was not yet recorded from India.

Abstract

The genus Diplasiolejeunea is added to the liverwort flora of India from the Kerala part of the Western Ghats reporting two species: Diplasiolejeunea cavifolia Steph. and D. cobrensis Gottsche ex Steph. The genus is known from tropical Asia with eight species. Among them Diplasiolejeunea cavifolia is widespread, but still the genus was not yet recorded from India.

INTRODUCTION

Diplasiolejeunea (Spruce) Schiffn. is a pantropical genus of the liverwort family Lejeuneaceae represented by approximately 75 valid species (Prudêncio et al. 2018, Söderström et al. 2016) around the world. Most of the species are epiphytes and occur in a good number in the American and African continents, but from Asia this genus is represented only by eight species including the pantropical Diplasiolejeunea cavifolia Steph., D. cobrensis Gottsche ex Steph. and D. rudolphiana Steph. while D. ingekarolae Schäf.-Verw., D. jovet-astiae Grolle, D. longilobula Herzog, D. onraedtii Grolle, and D. patelligera Herzog are restricted to Asia (Grolle 1995, He 1997, Lai et al. 2008, Piippo 1990, 1994, Schäfer-Verwimp 2004, 2006, Zhu and So 2001).

The genus Diplasiolejeunea is characterised by the development of under-leaves along every lateral leaf (similar condition is found only in the genus of Colura (Dumort.) Dumort.). Leaf lobes are large, usually loosely imbricate, apices are broad and rounded, leaf cells of lobes plane or with distinct trigones and intermediate thickenings and often with ocelli, hyaline papilla is situated at the base of the tooth on the inner surface of the lobule, underleaves deeply bifid with sharp or blunt apices and the perianth is sharply five keeled. The species of the genus are epiphytes (epiphyllous or ramicolous, growing on living leaves or on twigs) in the wet tropics.

During our recent investigation of the family Lejeuneaceae of Kerala, some interesting specimens were observed, among these two species belonging to the genus Diplasiolejeunea, viz. D. cavifolia and D. cobrensis. The majority of Diplasiolejeunea species are recorded from a narrow range of distribution, except for D. cavifolia, D. cobrensis and D. rudolphiana, which are pantropical (Dong et al. 2012). However, the present study recorded for the first time the genus along with these two species from India. A detailed description of both species with photo plates, illustration and the distribution map of D. cobrensis are provided.

DESCRIPTION OF SPECIES

Diplasiolejeunea cavifolia Steph. (Figs 1–2)

Fig. 1.
Fig. 1.

Diplasiolejeunea cavifolia Steph. a = portion of plant with male branch; b = portion of plant with androecial and gynoecial branches; c = single leaf with lobule; d = plant ventral view with lobule and underleaf; e, f = lobules; g = t-shaped lobule with basal hyaline papilla; h = underleaf; i = underleaf lobe cells; j = leaf lobe cells (after Chandini 10515b)

Citation: Acta Botanica Hungarica 63, 3-4; 10.1556/034.63.2021.3-4.2

Fig. 2.
Fig. 2.

Diplasiolejeunea cavifolia Steph. a = plant with male branch; b = portion of plant with male and female branch; c = male branch; d = perianth; e and g = plant ventral view with lobule and underleaf; f = portion of plant; h, i, j, k, l = different leaf lobules; m, n = leaf lobe; o, p = leaf lobe marginal cells; q = leaf lobe median cells; r = leaf lobe basal cells; s = portion of stem with underleaves; t = single underleaf lobule (after Chandini 10515b)

Citation: Acta Botanica Hungarica 63, 3-4; 10.1556/034.63.2021.3-4.2

Diplasiolejeunea cavifolia Steph., Bot. Jahrb. Syst. 20: 318 (1895); Piippo, Trop. Bryol. 9: 55 (1994); Schäfer-Verwimp, Cryptogamie, Bryol. 25(1): 13 (2004). – Bas.: Lejeunea cavifolia Steph., Bot. Jahrb. Syst. 8: 89 (1886). = Diplasiolejeunea brachyclada A. Evans, Bull. Torr. Bot. Club, 39: 216 (1912). = D. javanica Steph. Sp. Hepat. (Stephani) 5: 928 (1916). = D. ocellata Steph., Sp. Hepat. (Stephani) 5: 920 (1916). = D. vandenberghenii Grolle, Rev. Bryol. Lichénol. 29: 208 (1960), nom. illeg.

Diplasiolejeunea cavifolia was described by Stepani under different names as D. javanica Steph., D. ocellata Steph. and Lejeunea cavifolia Steph. (Blocked synonym Lejeunea cavifolia (Ehrh.) Lindb.; later homonym), many other workers also reported this species under different names. Unknowingly, Evans (1912) also reported this species as D. brachyclada Evans from Puerto Rico with detailed description and illustrations. Later, all these names were synonymised under Diplasiolejeunea cavifolia (Grolle 1978, Jones and Harrington 1983).

Plants pale green to yellowish, scattered, 4–8 mm long, 1.25–1.56 mm wide including leaves, stem widely branching; rhizoids hyaline, clustered at the base of underleaves. Leaves loosely imbricated, lobes widely spreading, not firmly appressed to the substratum, broadly ovate, plane, 0.75–1.0 × 0.5–0.6 mm, dorsal margin rounded and extending across the stem, slightly curved or with wavy margin, margin plane, apex rounded or slightly obtuse in some leaves. Cells plane, without trigones and intermediate thickenings, thick walled, marginal cells rectangular to polygonal, 9.2–15.3 × 10.2–16.6 µm, median cells polygonal, 8.4–23.6 µm, basal cells elongated, polygonal, 15.8– 35.5 × 11.2–21.1 µm, ocelli scattered in the lobe; lobule ovate, 0.4–0.5 × 0.2–0.28 mm, inflated, 1/2 as long as the leaf lobe, keel arched, free margin involute up to near the base, first tooth acute (median), truncate or T-shaped, three to four cells long, one or two cells wide at base, proximal tooth inflexed, shorter than first tooth, lies along the ventral margin of leaf lobe, acute, hyaline papilla seen on the base of the first tooth, indistinct. Underleaves distant, rounded to subcordate at base, deeply bifid, 0.22–0.30 × 0.18–0.20 mm, three times wider than the stem, sinus wide, lobes 6–10 cells wide at base, acute, margin entire. Monoecious, observed as autoicous, female inflorescence present on very short branch, perianth oblong or oblong ovate, rounded to truncate at apex, very short beak present, 0.76–0.12 × 0.46–0.85 mm, bracts obliquely spreading, rounded at apex, as long as leaf lobe; male gametoecia on very short branches, bracts imbricate, in 3–4 pairs, mature sporophyte is not seen by us (Figs 1 and 3). Spori, as observed by Weis (2001), are 45–55 × 20–30 µm in size, with rosettes of 5–7 µm diameter decorated by obtuse spines.

Fig. 3.
Fig. 3.

Diplasiolejeunea cobrensis Gottsche ex Steph. a = portion of plant ventral view; b = single leaf (ventral); c = single leaf (dorsal); d, e = leaf lobules; f, g, h = underleaves; i = underleaf lobe cells; j = leaf lobe median cells with trigones and intermediate thickenings; k = marginal cells; l = leaf lobe basal cells with ocelli; m = leaf cells with oil bodies; n = stem c. s. (after Vinjusha 13182b)

Citation: Acta Botanica Hungarica 63, 3-4; 10.1556/034.63.2021.3-4.2

Specimen examined: India, Kerala, Palakkad district, Silent Valley National Park, Parathode, way to Poochippara (1,000 m +), coll.: Chandini (10515b), 11.01.2018 (ZGC).

Habitat: Epiphyllous, associated with Lejeunea cocoes Mitt., Leptolejeunea elliptica (Lehm. et Lindenb.) Schiffn. in evergreen forest above 1,000 m.

Distribution: Cambodia, China (Hainan), Java, Sumatra, Luzon, Malaysia, Sri Lanka, Taiwan, New Caledonia (Schäfer-Verwimp 2006), Vietnam, Laos and Thailand, Philippines (Lai et al. 2008, Miller et al. 1983, Söderström et al. 2020), Australia (McCarthy 2003), Melanesia (Piippo 1990), West Indies (Evans 1912, Reyes 1982, incl. distribution map), Central America, Mexico, South America (Schäfer-Verwimp 2004), all over tropical Africa (Wigginton 2018), Madagascar and the Mascarene islands (Marline et al. 2012, Wigginton 2018).

Diplasiolejeunea cobrensis Gottsche ex Steph. (Figs 3–5)

Fig. 4.
Fig. 4.

Diplasiolejeunea cobrensis Gottsche ex Steph. a = portion of plant with branches; b = portion of plant, ventral view; c = leaf arrangement on stem; d, g = leaf lobule; e = leaf lobe; f = leaf with lobule; h, i = underleaves; j = leaf lobe maginal cells; k = leaf lobe median cells; l = leaf lobe basal cells with ocelli; m = stem c. s. (after Vinjusha 13182b)

Citation: Acta Botanica Hungarica 63, 3-4; 10.1556/034.63.2021.3-4.2

Fig. 5.
Fig. 5.

Worldwide distribution of Diplasiolejeunea cobrensis Gottsche ex Steph. Dots = known distribution; asterisk = the new locality in India

Citation: Acta Botanica Hungarica 63, 3-4; 10.1556/034.63.2021.3-4.2

Diplasiolejeunea cobrensis Gottsche ex Steph., Spec. Hepat. 5: 923 (1916) subsp. cobrensis = D. harpaphylla Steph., Sp. Hepat. 5: 919 (1916). = D. incurvata Jovet-Ast et Tixier, Rev. Bryol. Lichénol. 31: 29 (1962).

Plant small, light green, loosely attached to the substratum, 2–4 mm long, 0.85–1.1 mm wide including leaves, irregularly branched, branches up to 1 mm long, stem c.s. with 3 medullary and 7 cortical cells, 82.9 µm in diameter; rhizoids hyaline, fasciculate. Leaves imbricate, obliquely spreading, lobe convex in dorsal margin, asymmetric, semi-circular, broadly ovate, 0.6–0.67 × 0.48–0.60 mm, ventral margin curved to nearly straight, entire, cells with prominent trigones and intermediate nodular thickenings, usually one per cell wall, marginal cells rectangular to pentagonal, 6.9–11.6 × 8.2–15.4 µm, median cells polygonal, 13.2–25.6 ×10–25.9 µm, basal cells more irregular, 20.8– 35.6 µm in size; ocelli scattered, 8–15, basal ocellus larger than companion cells, usually one, rarely two, 53.2–60.4 × 30.2–37.5 µm; oil bodies 2–5 per cells, rounded, granular, scattered in lobes, lobules and underleaves; lobules inflated, tubulate,1/3–1/2 as long as the leaf lobe, 0.17–0.19 × 0.07–0.09 mm, first tooth normally well developed, consisting of 1–2 cells, single cell observed as superimposed in some lobules, proximal tooth one celled, indistinct, lies along the ventral margin, hyaline papilla indistinct at base of lobule teeth. Underleaves small, not much wider than the stem, deeply bilobed nearly to the base, sinus U- or V-shaped, 0.12–0.22 × 0.05–0.09 mm, lobes 2–3 cells wide at base; biseriate, 5–6 cells long, ending with 1–3 uniserial cells at tip, sexual and asexual reproductive structures not seen (Figs 2 and 4).

Habitat: Epiphyllous, associated with Leptolejeunea balansae Steph. in semi-evergreen forests

Specimen examined: India, Kerala, Wayanad district, Kuruva dweep (750 m), approx. 11° 49’ N, 76° 06’ E, coll.: Vinjusha (13182b), 07.12.2018 (ZGC).

Distribution: Cuba (type from El Cobre, coll.: Wright 1173) (Reyes 1982); Guyana (Gradstein and Hekking 1989); Brasil (Germano and Pôrto 2004, Prudêncio et al. 2018, Schäfer-Verwimp 1992); West Africa (Ghana – Jones 1973; Sierra Leone – Jones and Harrington 1983, East African islands (Mafia – Pócs and Váňa 2015; Mayotte – Pócs 2010; Madagascar – Grolle 1966, Tixier 1986 (several localities under D. harpaphylla); Pócs 2001 (also the ssp. nov. andringitrae); Dong et al. 2012, Marline et al. 2012), China (Hainan – Zhu and So 2001), Vietnam (Jovet-Ast and Tixier 1962 (as D. incurvata sp. nov.)); Thailand (Tixier 1973, 1986 (as D. incurvata), Schäfer-Verwimp 2006, Lai et al. 2008); Malaysia (Sabah, Mt Kinabalu – Mizutani 1966 (mistaken as D. brachyclada, corrected in 1970 and a Sabah east coast record added). There is an unpublished record new to the Andes: Peru, Dept. Pasco, prov. Oxapampa, Parque Nacional Yanachaga-Chemillén, epiphyllous in cloud forest at 2,415 m elevation. Coll. J. Larraín & C. Rothfels (40272/AB), 22 May 2016, det. Pócs (EGR!).

DISCUSSION

Even though, Diplasiolejeunea cavifolia is described as widely distributed one, not yet reported any species of Diplasiolejeunea from India. This may be due to the inadequate survey of the family Lejeuneaceae in our area. Compared to other genera of Lejeuneaceae of Kerala Diplasiolejeunea are represented only with these collections. Both taxa are epiphyllous along with the other members of the same family and prefer montane evergreen forest patches. They are easily identifiable by its double number of underleaves and not firmly appressed nature of the thallus to the substratum. D. cavifolia is distinguished by its leaf lobules with mostly three to four cells long linear or T-shaped first tooth. Lobe cells plane and mostly hexagonal in shape, distant underleaves with triangular, acute lobes. D. cobrensis can be distinguished by half circle shaped lobes, its cells with trigones and intermediate nodular thickenings, with 2–5 oil bodies, scattered ocelli with large basal ones. The lobule is tubular with usually two celled first tooth and with an obsolete second tooth. Underleaves small and deeply bilobed nearly to the base with 2–3 basal cells, biseriate lobes ending at 2–3 celled uniseriate apex. D. cobrensis has similarities by its tubular lobule and small underleaves with 1–2 cells wide segment to Diplasiolejeunea cornuta Steph., widespread in Africa. But differs from it by its entire lobe margin and by its scattered ocelli (D. cornuta has acuminate lobes with dentate margin and only basal ocelli). It is remarkable, how different the two Indian species are in their way of dispersal. While D. cavifolia is very widespread and common in all wet tropics, D. cobrensis, although also pantropical, in general is very rare and has a scattered distribution (see map on Fig. 5). The explanation of this difference is not easy, as both are monoecious, have no organs of vegetative reproduction and rarely produce sporophytes. The spori of D. cavifolia are too large for long range air dispersal and the spori of D. cobrensis are not yet known. Either historical factors cause the difference or they are able to reproduce themselves at different level of fragmentation and so aerial dispersal can be counted to different extent. Dong et al. (2012) claimed on the base of their phylogram of maximum likelihood analysis, that D. cavifolia evolved in the Neotropis and from there dispersed to the Paleotropis and D. cobrensis, vice versa, evolved in Africa and migrated from there to the other continents.

The present discovery is a new generic record for the Indian flora. Over the years several partial treatment and floristical approaches appeared, but a comprehensive world treatment of this genus was not yet published.

Acknowledgements

The authors are thankful to the authorities of the Zamorin’s Guruvayurappan College for providing facilities and encouragement. The financial assistance from Kerala State Council for Science Technology and Environment (KSCSTE), Thiruvananthapuram is also acknowledged. We sincerely acknowledge the help rendered by the officials of Kerala Forest department for providing permission to collect bryophyte specimens from the study area.

References

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  • Dong, S. , Schäfer-Verwimp, A. , Meinecke, P. , Feldberg, K. , Bombosch, A. , Pócs, T. , Schmidt, A. R. , Reitner, J. , Schneider, H. and Heinrichs, J. (2012): Tramps, narrow endemics and morphologically cryptic species in the epiphyllous liverwort Diplasiolejeunea. –Mol. Phyl. Evol. 65(2): 582594.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Evans, A. W. (1912): Hepaticae of Puerto Rico. XI. Diplasiolejeunea. –Bull. Torrey Bot. Club 39(5): 209225.

  • Germano, S. R. and Pôrto, K. C. (2004): Novos registros de briófitas para Pernambuco, Brasil.–Acta Bot. Bras. 18(2): 343350.

  • Gradstein, S. R. and Hekking, W. H. A. (1989): A catalogue of the bryophytes of the Guyanas. I. Hepaticae and Anthocerotae. –J. Hattori Bot. Lab. 66: 197230.

    • Search Google Scholar
    • Export Citation
  • Grolle, R. (1966): Über Diplasiolejeunea in Asien.–Feddes Repert. 73: 7889.

  • Grolle, R. (1978): Eine neue Diplasiolejeunea-Art aus Sri Lanka.–Feddes Repert. 89: 301305.

  • Grolle, R. (1995): The Hepaticae and Anthocerotae of the East African Islands. An annotated catalogue. –Bryophyt. Bibl. 48: 1178.

  • He, X.-L. (1997): A review and checklist of the Lejeuneaceae in China.–Abstracta Bot. 21: 6977.

  • Jones, E. W. (1973): African Hepatics XXIV. Lejeuneaceae: some new or little-known species, and extensions of range. –J. Bryol. 7: 545561.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Jones, E. W. and Harrington, A. J. (1983): The hepatics of Sierra Leone and Ghana. –Bull. Brit. Mus. (Nat. Hist.), Bot. 11(3): 215289.

    • Search Google Scholar
    • Export Citation
  • Jovet-Ast, S. and Tixier, P. (1962): Hepatiques du Viet-Nam, II. –Rev. Bryol. Lichénol. 31(1–2): 2333.

  • Lai, M.-J. , Zhu, R.-L. and Chantanaorrapint, S. (2008): Liverworts and hornworts of Thailand: an updated checklist and bryofloristic accounts. –Ann.Bot. Fenn. 45: 321341.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • Marline, L. , Andriamiarisoa, R. L. , Bardat, J. , Chuah-Petiot, M. , Hedderson, T. A. , Reeb, C. , Strasberg, D. , Wilding, N. and Ah-Peng, C. (2012): Checklist of the bryophytes of Madagascar.–Cryptogamie, Bryol. 33(3): 199255.

    • Crossref
    • Search Google Scholar
    • Export Citation
  • McCarthy, P. M. (2003): Catalogue of Australian liverworts and hornworts. – Flora of Australia, Suppl. Series 21. Australian Biological Resources Study, Canberra, 137 pp.

    • Search Google Scholar
    • Export Citation
  • Miller, H. A. , Whittier, H. O. and Whittier, B. A. (1983): Prodromus Florae Hepaticarum Polynesiae. –Bryophyt. Bibl. 25: 1423.

  • Mizutani, M. (1966): Epiphyllous species of Lejeuneaceae from Sabah (North Borneo). –J. Hattori Bot. Lab. 29: 153170.

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  • Gy. BORBÉLY (Debrecen)
  • A. ČARNY (Ljubljana)
  • A. CSERGŐ (Dublin)
  • B. CZÚCZ (Paris)
  • M. HÖHN (Budapest)
  • K. T. KISS (Budapest)
  • A. KUZEMKO (Uman)
  • Z. LOSOSOVÁ (Brno)
  • I. MÁTHÉ (Szeged)
  • E. MIHALIK (Szeged)
  • S. ORBÁN (Eger)
  • R. PÁL (Butte)
  • Gy. PINKE (Mosonmagyaróvár)
  • T. PÓCS (Eger)
  • K. PRACH (České Budejovice)
  • E. S. RAUSCHERT (Cleveland)
  • E. RUPRECHT (Cluj Napoca)
  • G. SRAMKÓ (Debrecen)
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  • É. SZŐKE (Budapest)
  • B. TOKARSKA-GUZIK (Katowice)
  • B. TÓTHMÉRÉSZ (Debrecen)
  • P. TÖRÖK (Debrecen)

Botta-Dukát, Zoltán
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or

Lőkös, László
E-mail: acta@bot.nhmus.hu
Institute: Botanical Department, Hungarian Natural History Museum
Address: Könyves K. krt. 40. H-1097 Budapest, Hungary

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2020  
Scimago
H-index
19
Scimago
Journal Rank
0,417
Scimago
Quartile Score
Plant Science Q2
Ecology, Evolution, Behavior and Systematics Q3
Scopus
Cite Score
155/89=1,7
Scopus
Cite Score Rank
Plant Science 221/445 (Q2)
Ecology, Evolution, Behavior and Systematics 374/647 (Q3)
Scopus
SNIP
0,838
Scopus
Cites
260
Scopus
Documents
22
Days from submission to acceptance 127
Days from acceptance to publication 132
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Rate
36%

 

2019  
Scimago
H-index
17
Scimago
Journal Rank
0,404
Scimago
Quartile Score
Plant Science Q2
Ecology, Evolution, Behavior and Systematics Q3
Scopus
Cite Score
164/91=1,8
Scopus
Cite Score Rank
Plant Science 209/431 (Q2)
Ecology, Evolution, Behavior and Systematics 358/629 (Q3)
Scopus
SNIP
0,699
Scopus
Cites
215
Scopus
Documents
23
Acceptance
Rate
30%

 

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Acta Botanica Hungarica
Language English
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German
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Size B5
Year of
Foundation
1954
Publication
Programme
2021 Volume 63
Volumes
per Year
1
Issues
per Year
4
Founder Magyar Tudományos Akadémia
Founder's
Address
H-1051 Budapest, Hungary, Széchenyi István tér 9.
Publisher Akadémiai Kiadó
Publisher's
Address
H-1117 Budapest, Hungary 1516 Budapest, PO Box 245.
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Publisher
Chief Executive Officer, Akadémiai Kiadó
ISSN 0236-6495 (Print)
ISSN 1588-2578 (Online)

 

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