synopsis and key to species of neotropical cololejeunea

Acta Botanica Hungarica 56(1–2), pp. 185–226, 2014 DOI: 10.1556/ABot.56.2014.1–2.14

SYNOPSIS AND KEY TO SPECIES OF NEOTROPICAL COLOLEJEUNEA (LEJEUNEACEAE) T. Pócs1, A. Bernecker2 and P. Tixier† 1

Institute of Biology, Department of Botany, Eszterházy Károly College H-3301 Eger, Pf. 43, Hungary; E-mail: [email protected] 2 Escuela de Biología, Universidad de Costa Rica, San Pedro, San José 2060, Costa Rica (Received 9 October, 2013; Accepted 30 November, 2013)

A revision of Neotropical members of the genus Cololejeunea resulted in 69 accepted species after amalgamation of the former genera or subgenera of Aphanolejeunea, Campylolejeunea, Leptocolea, Pedinolejeunea, Physocolea and Taeniolejeunea and the emendation of subgenus Protocolea, restricted to its former sectio Protocolea with Cololejeunea minutissima and related species. After the examination of types the new combinations Cololejeunea gracilis var. linearifolia (R.M. Schuster) Pócs, Cololejeunea subscariosa (Spruce) Pócs and the following new synonymy is established: Physocolea reflexiloba Steph. and Cololejeunea reflexifolia (Steph.) Tixier are new synonyms of Cololejeunea inflectens (Mitt.) Benedix; Lejeunea ensifolia Spruce, Physocolea ensifolia (Spruce) Steph., Physocolea elliptica Steph. and Cololejeunea elliptica (Steph.) S. W. Arnell of Cololejeunea camillii (Lehm.) A. Evans, Cololejeunea trinitensis Tixier of Cololejeunea spruceana Tixier; Cololejeunea katiae Tixier of Cololejeunea lanciloba Steph.; Physocolea chilensis Herzog, Cololejeunea chilensis (Herzog) S. Arnell., Cololejeunea coseguinana L. Clark, Lejeunea turbinifera Spruce, Cololejeunea turbinifera (Spruce) Léon-Yanez, Gradst. et Wegner, Cololejeunea uleana Steph., Physocolea uleana (Steph.) Steph. of Cololejeunea minutissima (Sm.) Steph.; Leptocolea manaosensis Herzog and Cololejeunea manaosensis (Herzog) O. Yano of Cololejeunea subscariosa (Spruce) R. M. Schust.; The latter species is newly lectotypified. Cololejeunea latilobula (Herzog) Tixier, Cololejeunea planissima (Mitt.) Abeyw. and Cololejeunea pseudofloccosa (Horik.) Benedix are new to the Neotropics.An illustrated identification key for all Neotropical Cololejeunea species is presented. Key words: Cololejeunea, Lejeuneaceae, Neotropics, new combinations, synonyms

INTRODUCTION The first species of Cololejeunea (Spruce) Steph. as currently defined were described under Lejeunea Lib. and of Jungermannia L. Spruce (1885) distinguished Colo-Lejeunea as a subgenus of Lejeunea and Stephani (1891) was the first to give it generic rank (Söderström et al. 2014). 0236–6495/$ 20.00 © 2014 Akadémiai Kiadó, Budapest

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The first species described from the Neotropics was Cololejeunea cardiocarpa (Mont.) A. Evans, published under the name of Lejeunea cardiocarpa Mont. in Ramón de la Sagra (1842) from Cuba. Further species were described under different generic names by Evans (1905, 1913, 1915, 1918); Gottsche in Gottsche et al. (1844–1847); Herzog (1942a, b, 1951, 1952); Jovet-Ast (1947, 1960); Lehmann (1857); Nees and Mont. in Montagne (1843); Robinson (1964); Schuster (1955, 1956); Spruce (1884) and by Stephani (1888, 1890, 1895, 1905, 1916). Bonner’s Index Hepaticarum II (1962), IV (1962–1963) and its Supplementum A–C (1977) counted 38 species names from the Neotropics belonging to the genus (several of which since have fallen into synonymy). Since then, new species were added by Bernecker-Lücking (1998), Bernecker-Lücking and Morales (1999), Dauphin et al. (2006), Gradstein et al. (2011), Morales and Dauphin (1998), Morales and Lücking (1995), Pócs (2002), Pócs and Bernecker (2013), Pócs and Schäfer-Verwimp (2012), ReinerDrehwald (1995), Schäfer-Verwimp (2012), Schuster (1978, 1980a, b), and Tixier (1980, 1991, 1995). Three species are unpublished new records for the Americas from the collections of G. Dauphin, A.-L. Ilkiu-Borges, A. SchäferVerwimp and R. M. Schuster, identified by T. Pócs. In addition new combinations and new names are made. As a result, 69 validly published Cololejeunea species are accepted from the Neotropics. Several genera segregated in the past from Cololejeunea (Aphanolejeunea A. Evans, Campylolejeunea S. Hatt., Chondriolejeunea (Benedix) G. Kis et Pócs, Leptocolea (Spruce) A. Evans, Pedinolejeunea (Benedix ex Mizut.) P. C. Chen et P. C. Wu, Physocolea (Spruce) Steph., Taeniolejeunea Zwickel and others), have been amalgamated in the genus Cololejeunea based on molecular evidence. Inclusion of all Aphanolejeunea species, supported by the molecular research of Gradstein et al. (2006) and Wilson et al. (2007), was done by Pócs and Bernecker (2009). It has recently been found by molecular investigations (Yu et al. 2013) that Cololejeunea minutissima (Sm.) Schiffn., the type of former subgenus Protocolea R. M. Schust., probably along with a few other related species and with the genus Myriocoleopsis Schiffn., form a clade separate from Cololejeunea, which can be characterised by its stoloniform main stem, very variable lobule joining the lobe without a real keel and not forming a water sack. Leaf cells are thin walled, mammillose, perianth exserted on a relatively long, stipulate neck and with 5 sharp keels with two rows of protruding cells. Until further molecular studies will clarify the position of this and related species, we keep the Neotropical members of the group within the subgenus Protocolea of genus Cololejeunea. Members of the genus Cololejeunea can be defined, as 1) relatively small, mostly epiphyllous leafy liverworts (occurring also on bark, twigs, decaying

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wood and on rocks); 2) stems generally with 5–7 rows of cortical and one row of medullary cells; 3) branching thecal (Lejeunea type) or athecal (Aphanolejeunea type), basiscopic, in certain species (subgen. Incalejeunea) both types occur; 4) leaves developing in two rows, from the lateral merophytes, uniform, dimorphic or trimorphic (lobulate, elobulate, reduced in subgen. Aphanolejeunea), usually appressed to the substrate, sometimes with hyaline margin for water retention under the lobe; lobulate leaves are attached to stem by a short U shaped line and consist of a larger lobe and a smaller lobule, which is often modified into a water retaining sac, usually with two marginal teeth; 5) at the leaf base usually a hyaline or green stylus is present; 6) on the ventral merophyte, instead of underleaves, only rhizoid tufts develop near the base of each leaf, hence the segmentation is pendular; 7) sexuality is autoicous, paroicous or dioicous; 8) sporophytes develop in an elaborated perianth subtended by two female bracts; 9) androecia usually develop on male branches with 1–2 antheridia in the axils of male bracts arranged in two rows; 10) spores usually released in protonematal, multicellular form; 11) vegetative reproduction is very common by disciform gemmae, which develop exogenously or endogenously (Pócs 2012) on one or both surfaces of the lobe, and can very soon develop into mature plants.

Contribution of the authors Pierre Tixier (1980, 1991, 1995) described 18 new species from different Neotropical collections, examined many types and compiled a draft manuscript of Cololejeunea for the Flora Neotropica, including 41 species. Tamás Pócs (1984a, b) started the worldwide revision of Aphanolejeunea. Andrea Bernecker (formerly A. Lücking, in publications also Andrea Bernecker-Lücking studied the epiphyllous liverworts of Costa Rica (Lücking, 1995, Bernecker-Lücking 1998), described 4 new species and started to revise the Neotropical Aphanolejeunea together with Tamás Pócs, who collected large material in Cuba, Dominican Republic and Venezuela and (partly with coauthors) described 4 new species. Andrea Bernecker also prepared very precise illustrations of former Aphanolejeunea species. In the meantime she has discontinued bryological research and has bequeathed all her collections, observations and drawings to Tamás Pócs to deal with. They published new combinations of all formerly described Aphanolejeunea species within the genus Cololejeunea (Pócs and Bernecker 2009). Following the decease of Pierre Tixier the editor of Flora Neotropica, Prof. Stephan Robbert Gradstein, asked Tamás Pócs to rewrite Tixier’s manuscript, with incorporation of new results. He also invited him to contribute the Aphanolejeunea and Cololejeunea treatments for the Brazilian and French Guinean


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hepatic floras (Pócs 2003a, b, 2009). Now Pócs is working on the completion of the revision of Neotropical Cololejeunea (including former Aphanolejeunea) for the Flora Neotropica. So far almost all types have been seen, thousands of specimens have been identified and the present synopsis and identification key for all species have been completed. The planned Flora Neotropica treatment will contain detailed descriptions of the taxa and their illustrations and distributional data with dot maps.

SYNOPSIS OF THE NEOTROPICAL SPECIES OF COLOLEJEUNEA (SPRUCE) STEPH. The synopsis contains all validly published Neotropical Cololejeunea taxa, with their records of description, type locality and all available synonyms. Present molecular investigations (Yu et al. 2013) claim that morphological characters hitherto used to distinguish the subgenera of Cololejeunea are mostly paraphyletic or polyphyletic as result of rampant homoplasy. Therefore we are using the previously accepted “traditional” subgeneric division (Asthana and Srivastava 2003, Benedix 1953, Mizutani 1961, Pócs and Piippo 2011, Schuster 1963, 1980b, Zhu and So 2001) with slight modifications.

Subgen. APHANOLEJEUNEA (A. Evans) Benedix Beih. Feddes Repert. 134: 15, 1953 Holotypus: Aphanolejeunea microscopica (Taylor) A. Evans; Bull. Torrey Bot. Club 38: 273, 1911. Basionym: Aphanolejeunea A. Evans, Bull. Torrey Bot. Club 38: 272, 1911. 1. Cololejeunea berneckerae Pócs in Pócs and Bernecker, Polish Bot. J. 54: 4, 2009. – Basionym: Aphanolejeunea pocsii Bernecker, Nova Hedwigia 66: 168–169. Typus: Bernecker 92-467, Costa Rica, Limón. Note: Andrea Bernecker used as author names also A. Lücking and Bern.-Lück. For the sake of conformity we use in this work always her maiden name Bernecker as auctor name, which is her actual surname as well, similarly as we use the auctor name Ast instead of the formerly used Jov.-Ast or S. J.-A.). 2. Cololejeunea cingens (Herzog) Bernecker et Pócs, Polish Bot. J. 54: 4, 2009. – Basionym: Aphanolejeunea cingens Herzog, Svensk Bot. Tidskr. 46: 104, 1952. Typus: Harling 228 pp., Ecuador, Santiago-Zamora, Mendez. 3. Cololejeunea cornutissima (R. M. Schust.) Stotler et Crand.-Stotl., Bryologist 80: 411, 1977. – Basionym: Aphanolejeunea cornutissima R. M. Schust. Bryologist 59: 217, 1956. Typus: Schuster 33701, USA: Florida. Synonym: Aphanolejeunea evansii R. M. Schust., J. Elisha Mitchell Sci. Soc. 71: 134, 1955, hom. illeg., non Aphanolejeunea evansii Herzog, Beih. Bot. Centralbl. 61: 583–584, 1942.



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4. Cololejeunea costaricensis (Bernecker) Bernecker et Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 5, 2009. – Basionym: Aphanolejeunea costaricensis Bernecker, Nova Hedwigia 66: 164, 1998. Typus: Bernecker 92-4471, Costa Rica, Limón. 5. Cololejeunea cubensis Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 5, 2009. – Basionym: Aphanolejeunea evansii Herzog, Beih. Bot. Zentralblatt, Abt. B, 61: 583, 1942. Typus: Böker s.n. 1937, Cuba, Sierra Rosario; non Cololejeunea evansii Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. 6. Cololejeunea gracilis (Ast) Pócs, in Dauphin et al., Cryptogamie, Bryol. 29: 233, 2008. – Basionym: Aphanolejeunea gracilis Ast, Rev. Bryol. Lichenol. 16: 21, 1947. Typus: Allorge s.n. 1936, Guadeloupe, Cascade Vauchelet. Synonyms: Aphanolejeunea heterophylla R. M. Schust., J. Elisha Mitchell Sci. Soc. 71: 128, 1955; Lejeunea heterophylla K. I. Goebel, Pflanzenbiol. Schild. 1: 178, 1889, hom. illeg., non Lejeunea heterophylla Sande Lac., Syn. Hep. Javan.: 69, 1856. Cololejeunea gracilis var. gracilis Cololejeunea gracilis var. linearifolia (R. M. Schust.) Pócs, comb. nova – Basionym: Aphanolejeunea gracilis Ast var. linearifolia R. M. Schust., Phytologia 45: 434, 1980. The type specimen, Schuster 67025 (F: C0171802) , Jamaica, Blue Mts., Hardwar Gap, has almost only (1–)2 cells wide, linear, reduced leaves with denticulate margin. 7. Cololejeunea microscopica (Taylor) Schiffn. – Basionym: Jungermannia microscopica Taylor in Mackay, Fl. Hibern. 2: 59, 1836. Typus: Ireland, Gortagarree near Killarney. Synonyms: Aphanolejeunea microscopica (Taylor) A. Evans, Bull. Torrey Bot. Club 38: 273, 1911; Aphanolejeunea diaphana Herzog, Nat. Hist. Juan Fernandez and Easter Islands 2: 748, 1942b; Cololejeunea fernandeziana Solari, J. Hattori Bot. Lab. 54: 546, 1983. Cololejeunea microscopica var. microscopica Cololejeunea microscopica var. exigua (A. Evans) Pócs, in Gradstein and Ilkiu-Borges, Mem. N. Y. Bot. Garden 76: 73, 2009. – Basionym: Aphanolejeunea exigua A. Evans, Bull. Torrey Bot. Club 38: 273, 1911. Typus: Evans 21 p.p., Puerto Rico: El Yunque. Synonym: Aphanolejeunea microscopica (Taylor) A. Evans var. exigua (A. Evans) Bernecker et Pócs, in Pócs and Bernecker, Haussknechtia 7: 26–27, 1997. Cololejeunea microscopica var. africana (Pócs) Pócs et Bernecker, in Dauphin et al., Cryptog. Bryol. 29: 234, 2008. – Basionym: Aphanolejeunea exigua A. Evans var. africana Pócs, Cryptogamie, Bryol. Lichénol. 5: 247, 1984. Typus: Pócs & Mwanjabe 6827/BC, Tanzania, Uluguru Mts, Lukwangule Plateau, 2350–2450 m. Synonyms: Aphanolejeunea microscopica (Taylor) A. Evans var. africana (Pócs) Pócs et Bernecker, Haussknechtia Beih. 7: 27, 1997; Aphanolejeunea asperrima Steph., Spec. Hep. 5: 859, 1916; Aphanolejeunea verrucosa Ast, Rev. Bryol. Lichenol. 16: 25, 1947.


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8. Cololejeunea minuscula Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 7, 2009. – Basionym: Aphanolejeunea minuta R. M. Schust., Hep. Anthoc. N. Amer. 4: 1310, 1980. Typus: Schuster 76-319, USA: Florida, Highlands Hammock; non Cololejeunea minuta Steph., Bull. Herb. Boissier 5: 79, 1915. 9. Cololejeunea papillosa (K. I. Goebel) Mizut., J. Hattori Bot. Lab. 29: 156, 1966. – Basionym: Physocolea (Aphanolejeunea) papillosa K. I. Goebel, Ann. Jard. Bot. Buitenzorg 39: 39, 1928. Typus: Goebel s.n. Sumatra: Fort Cock. Synonyms: Aphanolejeunea angustissima Steph., Spec. Hep. 5: 859, 1916; Aphanolejeunea papillosa (K. I. Goebel) Herzog, Trans. Brit. Bryol. Soc. 1: 325, 1949; Aphanolejeunea cyathiphylla Herzog, Svensk Bot. Tidskr. 46: 105, 1952. 10. Cololejeunea sicifolia (Gottsche ex Steph.) Pócs et Bernecker, Polish Bot. J. 54: 8, 2009 “sicaefolia”. – Basionym: Aphanolejeunea sicifolia (Gottsche ex Steph.) A. Evans, Bull. Torrey Bot. Club 38: 277, 1911 “sicaefolia”. Typus: Crüger s.n. 1847, Trinidad. The original specific epithet “sicaefolia” is orthographically corrected according to the ICBN Recommendation 60G. Synonym: Cololejeunea sicifolia Gottsche ex Steph., Hedwigia 27: 290, 1888, “sicaefolia” comb. inval. Cololejeunea sicifolia subsp. sicifolia Cololejeunea sicifolia subsp. jamaicensis (R. M. Schust.) Bernecker et Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 8, 2009. – Basionym: Aphanolejeunea jamaicensis R. M. Schust., Phytologia 45: 434, 1980. Typus: Schuster 67-019, Jamaica: Blue Mts, from Hardwar Gap to Caledonia Peak. Synonym: Aphanolejeunea sicifolia subsp. roraimensis Pócs et Bernecker, Haussknechtia 7: 26–27, 1997 “sicaefolia”. 11. Cololejeunea sintenisii (Steph.) Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 8, 2009. – Basionym: Aphanolejeunea sintenisii (Steph.) Steph., Spec. Hep. 5: 861, 1916. Typus: Sintenis 136 p.p. Puerto Rico, s.n. Synonyms: Cololejeunea sintenisii Steph., Hedwigia 27: 291, 1888, comb. inval.; Aphanolejeunea angustiloba Horik., J. Sci. Hiroshima Univ. ser. b, div. 2: 91, 1932. Cololejeunea angustiloba (Horik.) Mizut., Proc. Biol. Soc. Japan 6: 246, 1996. Aphanolejeunea ephemeroides R. M. Schust., J. Elisha Mitchell Sci. Soc. 71: 130, 1955; Aphanolejeunea teotonii V. Allorge et Ast, Rev. Bryol. Lichenol. 19: 21, 1950; Cololejeunea teotonii (V. Allorge et Ast) Grolle, Feddes Repert. 87: 190, 1976. 12. Cololejeunea subsphaeroidea (R. M. Schust.) Pócs, Cryptogamie, Bryol. 29: 235, 2008. – Basionym: Aphanolejeunea subsphaeroidea R. M. Schust., Phytologia 39: 431, 1978. Typus: Schuster & Ruiz-Teran 76-1500, Venezuela: Mérida: Sierra Nevada de Mérida, Rio Frias. 13. Cololejeunea winkleri (M. I. Morales et Bernecker) Bernecker et Pócs, in Gradstein and Ilkiu-Borges, Mem. N. Y. Bot. Garden 76(4): 78. 2009. – Basionym: Aphanolejeunea winkleri M. I. Morales et Bernecker, Nova Hedwigia 60: 120, 1995. Typus: Morales, Dauphin & Umana 2361, Costa Rica: Heredia, Finca la Selva. Acta Bot. Hung. 56, 2014


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Subgen. CHLOROCOLEA R. M. Schust. Beih. Nova Hedwigia 9: 178, 1963 Holotypus: Cololejeunea ciliatilobula (Horik.) R. M. Schust., Beih. Nova Hedwigia 9: 178, 1963 hom. illeg. (= C. desciscens Steph.) non C. ciliatilobula (Schiffn.) Schiffn. (= C inflectens (Mitt.) Benedix). 14. Cololejeunea linopteroides H. Rob., Bryologist 67: 457, 1964. – Typus: A. Smith 48/312, Costa Rica: Alajuela, San Antonia de Zarcero. 15. Cololejeunea standleyi Herzog, Rev. Bryol. Lichenol. 20: 205–206, 1951. – Typus: Standley 54.827, Honduras: Lancetilla Valley.

Subgen. CHLOROLEJEUNEA R. M. Schust. Beih. Nova Hedwigia 9: 176, 1963 Typus: Cololejeunea madothecoides (Steph.) Benedix, Beih. Feddes Repert. 134: 81, 1953. Synonym: subgen. Chlorolejeunea Benedix, Beih. Feddes Repert. 134: 81, 1953, no type designated. In fact, Schuster (1963) used the subgeneric delimitation differently from Benedix (1953), including sectio Lancilobae Schust. with a number of Asian and Neotropical species, but in a later work (Schuster and Inoue 1974) they restricted the subgenus to only three species in the present concept. 16. Cololejeunea stotleriana Gradst., Ilk.-Borg. et Vanderp., Bryologist 114: 13, 2011. – Typus: Gradstein 12073, Ecuador: Zamora-Chinchipe, Rio Nangaritza S of Guayzimi.

Subgen. CRYPTOLEJEUNEA Benedix Beih. Feddes Repert. 134: 77, 1953 Lectotypus: Cololejeunea inflectens (Mitt.) Benedix, designated by Zhu and So, Nova Hedwigia, Beih. 121: 282, 2001. (Doubtful in the Neotropics).

Subgen. COLOLEJEUNEA Lectotypus: Cololejeunea calcarea (Lib.) Steph., Bot. Gaz. 17: 171, 1892. Designated by A. Evans, Bull. Torrey Bot. Club 38: 252, 1911. 17. Cololejeunea elegans Steph., Hedwigia 30: 208, 1891. – Typus: Dusén 133, Cameroons: Bomana. Synonym: Physocolea elegans (Steph.) Steph., Spec. Hep. 5: 870, 1916. An African species. Note: The record from Brazil, Sao Paolo (Visnadi 1998) has to be controlled. 18. Cololejeunea kuciana Pócs et Schäf.-Verw., Polish Bot. J. 57: 51, 2012. – Typus: Schäfer-Verwimp & Nebel 32062, Ecuador: Zamora-Chinchipe, Res. Biol. San Francisco 35 km E of Loja.


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19. Cololejeunea platyneura (Spruce) A. Evans, Mem. Torrey Bot. Club 8: 172, 1902. – Basionym: Lejeunea platyneura Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 299, 1884. Typus: Spruce L 516, Brazil: Rio Negro, San Gabriel. Synonyms: Physocolea vittata Steph., Spec. Hep. 5: 873, 1916; Cololejeunea usambarica E. W. Jones, Trans. Brit. Bryol. Soc. 2: 434, 1954. 20. Cololejeunea tamasii Schäf.-Verw., Phytotaxa 60: 9–11, 2012. – Typus: Schäfer-Verwimp & Verwimp 30992, Panama, Chiquiri, Cordillera de Talamanca, Vulkan Baru. 21. Cololejeunea thiersiana Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. – Typus: Spruce s.n., Peru: Tarapoto, Mt. Guayrapurina.

Subgen. INCALEJEUNEA Tixier ex Pócs Acta Bryolichenol. Asiatica 4: 129 Holotypus: Cololejeunea erostrata (Herzog) Bernecker et Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 5, 2009. 22. Cololejeunea antillana Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 3, 2009. – Basionym: Aphanolejeunea longifolia Ast, Rev. Bryol. Lichenol. 16: 23, 1947, Typus: Allorge s.n., 1936, Guadeloupe, Étang des Grands Fonds, non Cololejeunea longifolia Schiffn., Hedwigia 39: 200, 1900. 23. Cololejeunea azorica V. Allorge et Ast, Mitt. Thüring. Bot. Ges. 1: 21, 1955. – Typus: P. & V. Allorge s.n., 1936, Azores, I. San Miguel, Parc Jose de Canto. Synonym: Aphanolejeunea azorica (V. Allorge et Ast) Bernecker et Pócs, Mem. N. Y. Bot. Garden 87: 109, 2003. 24. Cololejeunea camillii (Lehm.) A. Evans, Bryologist 15: 59, 1912. – Basionym: Lejeunea camillii Lehm., Pugillus 10: 15, 1857. Typus: Ex hb. Montagne s.n., s.d., Cuba. Synonyms: Lejeunea montagnei Lehm. ex Steph., Hedwigia 29: 96, non Lejeunea montagnei Gottsche ex Mont., Ann. Sci. Nat. Bot. ser. 2, 19: 261, 1843; Aphanolejeunea camillii (Lehm.) R. M. Schust., Hep. Anthoc. N. Amer. 4: 1297, 1980; Lejeunea liliputiana Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 297, 1884; Aphanolejeunea liliputiana (Spruce) R. M. Schust., J. Elisha Mitchell Soc. 71: 140, 1955, “liliputana”; Physocolea liliputiana (Spruce) Steph., Spec. Hep. 5: 878, 1916; Cololejeunea liliputiana (Spruce) S. W. Arnell, Österr. Akad. Wiss. Math.-Naturwiss. Kl. Denkschr. 111: 97, 1964. Syn. nova: Lejeunea ensifolia Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 297, 1884; Physocolea ensifolia (Spruce) Steph.; Physocolea elliptica Steph., Spec. Hep. 5: 876. 1916. The investigated lectotype (G 17856, Türckheim s.n., 1908, Guatemala, designated by Grolle in 1976) fully agrees with C. camillii except for a part of leaves has obtuse apex; Cololejeunea elliptica (Steph.) S. W. Arnell, Svensk Bot. Tidskr. 53: 503, 1959. nom. inval. 25. Cololejeunea clavatopapillata Steph., Hedwigia 34: 246, 1895. – Typus: Ule 371, Brazil, s.l. Synonyms: Physocolea clavatopapillata (Steph.) Steph. Acta Bot. Hung. 56, 2014


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Spec. Hep. 5: 875, 1916; Aphanolejeunea clavatopapillata (Steph.) M. E. Reiner, Trop. Bryol. 10: 29–30, 1995; Cololejeunea tuberculata A. Evans, Bryologist 18: 84, 1915; Aphanolejeunea tuberculata (A. Evans) R. M. Schust., Hep. Anthoc. N. Amer. 4: 1290, 1980; Cololejeunea cristallina Herzog, Feddes Repert. 55: 16, 1952; Cololejeunea fadenii Pócs, Acta Bot. Acad. Sci. Hung. 21: 353, 1975; Aphanolejeunea fadenii (Pócs) Pócs, J. Hattori Bot. Lab. 55: 309, 1984. 26. Cololejeunea contractiloba A. Evans, Amer. J. Bot. 5: 131, 1918. – Typus: Rapp, s.n., 1915, USA: Florida, Sanford. Synonym: Aphanolejeunea contractiloba (A. Evans) R. M. Schust., Hep. Anthoc. N. Amer. 4: 1314, 1980. 27. Cololejeunea crenata (A. Evans) Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 5, 2009. – Basionym: Aphanolejeunea crenata A. Evans, Bull. Torrey Bot. Club 38: 276, 1911. Typus: Evans 36, Puerto Rico, El Yunque. 28. Cololejeunea diaphana A. Evans, Bull. Torrey Bot. Club 32: 184, 1905. – Typus: Small & Carter 1365 p.p., USA: Florida betw. Cutler and Camp Longview. Synonyms: Aphanolejeunea diaphana (A. Evans) R. M. Schust. Hep. Anthoc. N. Amer. 4: 1294, 1980, hom. illeg. non Aphanolejeunea diaphana Herzog, Nat. Hist. Juan Fernandez 2: 748, 1942b; Cololejeunea diaphana A. Evans var. cristulata R. M. Schust., J. Elisha Mitchell Soc. 72: 112, 1956; Aphanolejeunea diaphana (A. Evans) R. M. Schust. var. cristulata (R. M. Schust.) R. M. Schust. Hep. Anthoc. N. Amer. 4: 1298, 1980; Aphanolejeunea subdiaphana (Ast) Pócs, J. Hattori Bot. Lab. 55: 311, 1984; Aphanolejeunea truncatifolia Horik., J. Sci. Hiroshima Univ. Ser. 3, Div. 2, 2: 284, 1934; Cololejeunea ninhbinhiana Tixier, Ann. Hist. Nat. Mus. Nat. Hung. 66: 89, 1974; Aphanolejeunea ninhbinhiana (Tixier) Tixier, in Pócs, J. Hattori Bot. Lab. 55: 310, 1984; Physocolea proboscoidea K. I. Goebel, Ann. Jard. Bot. Buitenzorg 39: 35, 1928; Aphanolejeunea proboscoidea (K. I. Goebel) R. M. Schust., Hep. Anthoc. N. Amer. 4: 1218, 1980; Aphanolejeunea proboscoidea (K. I. Goebel) Tixier, in Pócs, J. Hattori Bot. Lab. 55: 310, 1984; Cololejeunea ensifolia (Spruce) Steph. var. pygmaea (Spruce) Herzog, Rev. Bryol. Lichenol. 20: 174, 1951. 29. Cololejeunea erostrata (Herzog) Bernecker et Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 5, 2009. – Basionym: Physocolea erostrata Herzog, Beih. Bot. Centralblatt, Abt. B, 61: 582, 1942. Typus: Herzog 4100/a, Bolivia: Rio Tocorani. 30. Cololejeunea moralesiae (Bernecker) Bernecker et Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 8, 2009. – Basionym: Aphanolejeunea moralesiae Bernecker, Nova Hedwigia 66: 166, 1998. Typus: Bernecker 92-462, Costa Rica: Limón, “Braulio Carillo” N.P. 31. Cololejeunea papilliloba (Steph.) Steph., Hedwigia 34: 250, 1895. – Basionym: Lejeunea papilliloba Steph., Hedwigia 29: 73, 1890, non Lejeunea papilliloba Herzog, Feddes Repert. 55: 14, 1953 “1952”, nom. illeg. Typus: Ule s.n. et s.d., Brazil: Sao Francisco. Synonyms: Physocolea papilliloba Steph., Spec. Hep. 5: 881, 1916; Aphanolejeunea kunertiana Steph., Spec. Hep. 5: 860, 1916; CololejeActa Bot. Hung. 56, 2014

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unea pustulosa Ast, Rev. Bryol. Lichenol. 29: 35, 1960; Aphanolejeunea pustulosa (Ast) Bernecker et Pócs in Equihua and Pócs, Bryologist 102: 749, 1999. 32. Cololejeunea paucifolia (Spruce) Bernecker et Pócs, in Pócs and Bernecker, Polish Bot. J. 54: 8, 2009. – Basionym: Lejeunea paucifolia Spruce, Bull. Soc. Bot. France 36 (Suppl. Congrès Bot. 1889): 194, 1890 (Rev. Bryol. 15: 35, 1888 nom. inval.). Typus: Balansa 3722 p.p. Paraguay: Guarapi. Synonyms: Physocolea paucifolia (Spruce) Steph., Spec. Hep. 5: 881, 1916; Aphanolejeunea paucifolia (Spruce) M. E. Reiner, Trop. Bryol. 10: 33–35, 1995; Cololejeunea lignicola Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1996 “1995”. 33. Cololejeunea pterocolea Herzog, Svensk Bot. Tidskr. 46: 103, 1952. – Typus: Harling 2212a p.p. Ecuador: Santiago-Zamora, Mendez. Synonym: Aphanolejeunea pterocolea (Herzog) R. M. Schust., Beih. Nova Hedwigia 9: 183, 1963. 34. Cololejeunea spathulata Ast, Rev. Bryol. Lichenol. 29: 36, 1960. “spatulata” – Typus: Heim s.n., 1959, Mexico: Chiapas, Zacatepec. Syn. nov.: Cololejeunea parallelifolia R. M. Schust., Phytologia 45: 433–434, 1980. Typus: Schuster 67-025 (F: C0171843), Jamaica, Blue Mts: track from Hardwar Gap to waterfall, on Caledonia Peak track. The type material is an intricate mixture of Lejeuneaceae with scanty material of several species. Luckily Schuster (1984: 1031, fig. 85) gave an illustration, which helped to identify this taxon with C. spathulata. The only difference seems to be the length of female bracts on Schuster’s drawing, being much shorter than in the type of C. spathulata, but the examined type specimen of C. parallelifolia also had long female bracts. 35. Cololejeunea yelitzae Pócs et Bernecker, Acta Bot. Hung. 55: 386, figs 1–3, 2013. – Typus: S. & T. Pócs and R. Rico 9732/CA, Venezuela: Mérida, Cordillera del Norte, Las Correras de los González 4 km ENE of Jají.

Subgen. LEPTOCOLEA (Spruce) Schiffn. in Engler and Prantl: 122, 1893 Lectotypus: Evans (1911): Cololejeunea micrandroecia (Spruce) Schiffn. ex Mizut. Note: The lectotype now belongs to subgen. Pedinolejeunea Benedix ex Mizut. (1961). We treat members of Leptocolea together with the species removed from former subgen. Protocolea R. M. Schust. Subgen. Protocolea was a very poorly defined “rubbish bin” and can be used in a much narrower sense (see below). 36. Cololejeunea hildebrandii (Austin) Steph., Bull. Herb. Boissier, sér. 1, 5: 842, 1897 – Basionym: Lejeunea hildebrandi Austin, Bot. Gaz. 1: 35, 1876. Typus: Baldwin s.n., Hawaii: Oahu. Synonyms: Physocolea hillebrandii (Austin) Steph., Spec. Hep. 5: 895, 1916; Cololejeunea filicaulis Steph., Hedwigia 31: 162, 1892; Physocolea filicaulis (Steph.) Steph., Spec. Hep. 5: 871, 1916; Physocolea villaumei Steph., Spec. Hep. 5: 873, 1916. Note: Although Austin (1876) named it Lejeunea hildebrandi, the species is often called also C. hillebrandii, supposing



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(without proof) that the author intended to name it after the pioneer botanist of the Hawaiian Islands, Wilhelm Hillebrand, author of the first Hawaiian flora and collector of numerous Hawaiian bryophytes (source of biographic information: Frahm and Eggers 2001). 37. Cololejeunea obliqua (Nees et Mont.) Schiffn., Bot. Jahrb. Syst. 23: 586, 1897. – Basionym: Lejeunea obliqua Nees et Mont., Ann. Sci. Nat., Bot., ser. 2, 19: 264, 1843. Typus: Collector unknown, Brazil. Synonyms: Lejeunea scabriflora Gottsche, Abh. Naturwiss. Vereine Bremen 7: 362, 1882; Leptocolea scabriflora (Gottsche ex Steph.) A. Evans, Bull. Torrey Bot. Club 38: 262, 1911. A complete list of Asian synonyms is given by Zhu and So, Beih. Nova Hedwigia 121: 304–305, 2001. 38. Cololejeunea planiuscula Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. – Typus: Spruce s.n., Brazil: Amazonas, San Gabriel. 39. Cololejeunea skottsbergii Herzog, Nat. Hist. Juan Fernandez and Easter Islands 2: 748, 1942. – Typus: Skottsberg 165, Chile: Juan Fernandez Islands: Masafuera. 40. Cololejeunea spruceana Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. – Typus: Pearson s.n., Venezuela: Amazonas, S. Carlos, sub nomine Lejeunea obliqua var. elobulata Spruce (MANCH 18829). Syn. nov.: Cololejeunea trinitensis Tixier, Cryptogamie Bryol. Lichénol. 16: 229, 1995. Typus: Britton, Coker & Rowland 221, Trinidad. Note: Examining the type of C. spruceana Tixier (MANCH 18829) and of C. trinitensis (NY 221), Pócs could not find any real difference between the leaf shape of the two. 41. Cololejeunea setiloba A. Evans, Bryologist 16: 51, 1913. – Typus: Rapp 27, USA: Florida, Seminole, Sanford.

Subgen. PEDINOLEJEUNEA Benedix ex Mizut. J. Hattori Bot. Lab. 24: 240, 1961 Holoypus: Cololejeunea lanciloba Steph., Hedwigia 34: 250, 1895. 42. Cololejeunea bekkeri Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. – Typus: Lindeman et al. 874a, Surinam: Nickerie, Kabalebo. 43. Cololejeunea bischleriana Tixier, Bradea 3: 36, 1980. – Typus: Bischler 2297, Colombia: Cundinamarca, Salto de Tequendama. 44. Cololejeunea cardiocarpa (Mont.) A. Evans, Mem. Torrey Bot. Club 8: 172, 1902. – Basionym: Lejeunea cardiocarpa Mont. in Sagra, Hist. 9 (Bot., Pl. Cell.): 476, 1842. Typus: Sagra & Aubert, Cuba. Synonyms: Leptocolea cardiocarpa (Mont.) A. Evans, Bull. Torrey Bot. Club 38: 268, 1911; Lejeunea jooriana Austin, Bull. Torrey Bot. Club 6: 20, 1875; Cololejeunea jooriana (Austin) A. Evans, Mem. Torrey Bot. Club 8: 173, 1902; Leptocolea jooriana (Austin) A. Evans, Bull. Torrey Bot. Club 38: 170, 1911.


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45. Cololejeunea cremersii Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. – Typus: Cremers 5343, French Guyana, Île de Cayenne, ancienne piste du Camp des Annamites. Synonym: Cololejeunea fefeana Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. 46. Cololejeunea cristata (Steph.) R. M. Schust. Beih. Nova Hedwigia 9: 173, 1963. – Basionym: Physocolea cristata Steph., Spec. Hep. 5: 869, 1916. Typus: Leloutre, s.n., 1898. Madagascar (part of Corbière 122). Synonym: Leptocolea cristata (Steph.) E. W. Jones, Trans. Brit. Bryol. Soc. 2: 149, 1953. Note: Confirmed records of the species are only from East Africa. The Neotropical records (“Caribbean”) should be confirmed. 47. Cololejeunea ecuadoriensis Pócs, Acta Bot. Hung. 44: 372, 2002. – Typus: R. Lücking s.n., May 1996, Ecuador: Pichincha, Guajalito Biol. Station 45 km W of Quito. 48. Cololejeunea geissleriana Tixier, Bradea 3: 37, 1980. – Typus: Ex Hb. Nees in G 19602, Antilles. 49. Cololejeunea guadelupensis Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. – Typus: Ex Hb. Lehmann sub nom. Jungermannia marginata s.n., s.d., La Guadeloupe. 50. Cololejeunea lanciloba Steph., Hedwigia 34: 250, 1895. – Typus: Kurz 3917, India: Nicobar I., Katschall. Synonyms: Leptocolea lanciloba (Steph.) A. Evans, Bull. Torrey Bot. Club 38: 268, 1911; Cololejeunea tonkinensis Steph., Hedwigia 34: 252, 1895; Physocolea bolombensis Steph., Spec. Hep. 5: 868, 1916. Further Asian synonyms are enumerated in Zhu and So, Nova Hedwigia, Beih. 121: 276, 2001. Syn. nov.: Cololejeunea katiae Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. Typus: Katia Cavalcanti-Porto 1456, Brazil: Recife. The differences given to distinguish this species (biapiculate lobule apex, smooth cuticle) also often occur in the Asian populations of C. lanciloba. 51. Cololejeunea latilobula (Herzog) Tixier, Bryophyt. Bibl. 27: 156, 1985. – Basionym: Leptocolea latilobula Herzog in Handel-Mazzetti, Symb. Sin. 5: 54, 1930. Typus: Handel-Mazzetti 10171, China: Yunnan, “bei Djindjischan nächst Loping”. Synonyms: Leptocolea himalayensis Pandé et R. N. Misra, Proc. Nat. Acad. India 13: 26, 1957; Pedinolejeunea himalayensis (Pandé et R. N. Misra) P. C. Chen et P. C. Wu, Acta Phytotax. Sin. 9: 266, 1964. Further Asian synonyms in Zhu and So, Nova Hedwigia, Beih. 121: 275, 2001. Widespread in Asia and Africa, new to the Neotropics (Bolivia and Ecuador). 52. Cololejeunea micrandroecia (Spruce) Schiffn. ex Mizut., J. Hattori Bot. Lab. 24: 252, 1961. – Basionym: Lejeunea micrandroecia Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 298, 1884. Typus: Spruce L12, Peru, Tarapoto, Huallaga River. Synonym: Leptocolea micrandroecia (Spruce) A. Evans, Bull. Torrey Bot. Club 38: 262, 1911. 53. Cololejeunea minutilobula Herzog, Rev. Bryol. Lichenol. 20: 171–172, 1951. – Typus: Standley 26176, s.l., Costa Rica. Acta Bot. Hung. 56, 2014


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54. Cololejeunea nigerica (E. W. Jones) R. M. Schust., Nova Hedwigia, Beih. 9: 177, 1963. – Basionym: Leptocolea nigerica E. W. Jones, Trans. Brit. Bryol. Soc. 2: 150, 1953. Typus: Jones 70, Nigeria: Benin, Okomu Forest Res. Note: Yano (2004) reports it from Brazil. Due to its similarity to C. latilobula (Herzog) Tixier and to C. planifolia (A. Evans) R. M. Schust., the record of this African species in the Neotropics should be confirmed. 55. Cololejeunea panamensis G. Dauphin et Pócs in Dauphin et al., Trop. Bryol. 27: 76, 2006. – Typus: Dauphin s.n., Panamá: Canal Zone, I. Barro Colorado. 56. Cololejeunea planifolia (A. Evans) R. M. Schust., Beih. Nova Hedwigia 9: 177, 1963. – Basionym: Leptocolea planifolia A. Evans, Bull. Torrey Bot. Club 38: 265, 1911. Synonym: Cololejeunea evansii Tixier, Cryptogamie, Bryol. Lichénol. 16: 229, 1995. Typus: Britton & Marble 882 p.p., Puerto Rico, Utuado. 57. Cololejeunea planissima (Mitt.) Abeyw., Ceylon J. Sci., Bio. Sci. 2: 73, 1959. – Basionym: Lejeunea planissima Mitt., J. Proc. Linn. Soc., Bot. 5: 117, 1861 “1860”. Typus: Gardner 1393, Ceylon, Pt. Galle. Synonyms: Leptocolea aoshimensis Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 18, 193; Cololejeunea aoshimensis (Horik.) S. Hatt., J. Hattori Bot. Lab. 19: 138, 1958. Further Asian synonyms in Zhu and So, Nova Hedwigia Beih. 121: 279, 2001. New to the Neotropics (coll. Ilkiu-Borges in Brazil). 58. Cololejeunea schusteri Pócs, Acta Bot. Hung. 44: 376–377, 2002. – Typus: Schuster 79-20-1040, Brazil: Amazonia, along Rio Negro NW of Sao Gabriel de Cachoeira, near the mouth of Rio Uaupés. 59. Cololejeunea subcardiocarpa Tixier, Bradea 3: 39, 1980. – Typus: Le Gallo s.n., 1957, French Guyana: Cayenne, forêt de Matoury. Synonym: Cololejeunea yanoanae Tixier, Cryptogamie, Bryol. Lichénol. 16: 230, 1995. 60. Cololejeunea submarginata Tixier, Bradea 3: 41, 1980. – Typus: Le Gallo 732, La Guadeloupe: Moscou. 61. Cololejeunea subscariosa (Spruce) Pócs comb. nov. The combination of Schuster, Beih. Nova Hedwigia 9: 167, 1963, is invalid (ICBN, Art.33.4). – Basionym: Lejeunea subscariosa Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 300, 1884. Typus: Spruce s.n., Brazil: Rio Negro, S. Gabriel. Lectotypus nov. designated by T. Pócs: MANCH 18338 (EMu 73,055). There are three specimens in the Spruce herbarium, all from the same locality, as the typus of Lejeunea subscariosa. They belong to three different species of Lejeuneaceae. Morales-Zürcher (Cryptogamie, Bryol. 30: 272, 2009, figs 1–4) indicated as holotype and made a drawing of another specimen (MANCH 18839), which does not fit Spruce’s diagnosis. This specimen, or at least the part that she investigated, has finger-like hyaline cells at the lobe apex and belongs to Cololejeunea micrandroecia (Spruce) Schiffn., as it was compared with the holotype of Lejeunea micrandroecia Spruce (MANCH, EMu 73,020). Therefore, it was necessary to select a new lectotype specimen that matches the original diagnosis of Spruce. Syn. nov.: Cololejeunea manaosensis (Herzog) Herzog, Rev. Bryol. Acta Bot. Hung. 56, 2014

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Lichenol. 20: 174, 1951; Basionym: Leptocolea manaosensis Herzog, Hedwigia 71: 349, 1932 “1931”. Typus: Lützelburg s.n., Brazil, Amazonia, Manaus. The only difference is that its type has a little bit thinner cell walls than that of Lejeunea subscariosa. 62. Cololejeunea surinamensis Tixier, Bradea 3: 42, 1980. – Typus: R. Heim s.n., Surinam: “bord de Mana en venant d’Ougando”. Tixier indicates in the protologue, that the cuticle is papulose. Together with Dr Anna-Luisa Ilkiu-Borges we investigated the cuticle at higher magnification. It turned out to be an interesting reticulate pattern, unusual among Cololejeunea species. 63. Cololejeunea verwimpii Tixier, Cryptogamie, Bryol. Lichénol. 16: 230, 1995. – Typus: Schäfer-Verwimp 7606, Brazil, São Paolo, forêt à Embu Guaçu. 64. Cololejeunea vitaliana Tixier, Cryptogamie, Bryol. Lichénol. 16: 230, 1995. “vitalana” – The specific epithet should be “vitaliana” (ICBN art. 60C.1). Typus: Vital, 13166, Brazil, São Paolo, Ilha do Cordoso, Cauaucia.

Subgen. PROTOCOLEA R. M. Schust. Beih. Nova Hedwigia 9: 171, 1963 emend. Pócs 2013 Holotypus: Cololejeunea minutissima (Smith.) Steph., Bot. Gaz. 17: 171, 1892. Syn. nov.: Cololejeunea subgen. Protocolea Schust. sectio Protocolea Schust., Beih. Nova Hedwigia 9: 171. Subgenus Protocolea was an ill defined group containing several alien elements. Through molecular investigations (Wilson et al. 2007, Yu et al. 2013) it turned out that Cololejeunea vuquangensis Pócs and Cololejeunea minutissima (Smith.) Schiffn. itself form a clade together with Myriocoleopsis Schiffn., which is closer to Colura (Dumort.) Dumort. than to the rest of Cololejeunea (Spruce) Steph. Further molecular investigation of this group of species, including C. chuahiana Pócs, C. dauphinii R. L. Zhu and of C. minutissima (Smith) Steph. is in progress. Before its final results we keep them within the genus Cololejeunea, subgen. Protocolea, but without the alien elements, which were incorporated in this subgenus maybe just because of their small size. It means that the subgenus is reduced to its former sectio Protocolea Schust. 65. Cololejeunea dauphinii R. L. Zhu, J. Bryol. 28: 277, 2006. – Synonym: Cololejeunea tixieri (“tixierii”) M. I. Morales et G. Dauphin, Trop. Bryol. 14: 133, 1998, hom. illeg. non Cololejeunea tixieri Onraedt, Bull. Jard. Bot. Nat. Belg. 59: 436 (1989). Typus: sub nom. Cololejeunea tixierii: Dauphin, León et Matarrita, 1556, Panama, Prov. Bocas de Toro, Parc. Internat. Amistad, Cima del Cerro Fábrega, 3350 m. 66. Cololejeunea minutissima (Sm.) Steph., Bot. Gaz. 17: 171, 1892 (Stephani 1892e). – Basionym: Jungermannia minutissima Sm., Engl. Bot. 23: pl. 1633, 1806. Typus: Lyell, England, Hampshire, New Forest. Synonyms: Physocolea apiahyna Steph., Spec. Hep. 5: 874, 1916; Cololejeunea apiahyna (Steph.) Yano; Acta Bot. Hung. 56, 2014


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Lejeunea megalandra Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 294, 1884; Physocolea megalandra (Spruce) Steph., Spec. Hep. 5: 879, 1916; Cololejeunea megalandra (Spruce) Yano, J. Hattori Bot. Lab. 56: 573, 1984; Lejeunea myriandroecia Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 296, 1884; Physocolea myriandroecia (Spruce) Steph., Spec. Hep. 5: 879, 1916; Cololejeunea myriandroecia (Spruce) Yano, J. Hattori Bot. Lab. 56: 578, 1984; Lejeunea tambillensis Loitl., Diss. cl. Math. Phys. Acad. Litt. Cracow 234, 1894; Physocolea tambillensis (Loitl.) Steph., Spec. Hep. 5: 885, 1916; Further synonyms see in Schuster (1980b). Syn. nov.: Cololejeunea chilensis (Herzog) S. Arnell, Österr. Akad. Wiss. Math. Naturwiss. Kl. Denkschr. 111: 97, 1964; Physocolea chilensis Herzog, Arch. Escuela Farm. Cordoba 7: 31–32, 1938; Cololejeunea coseguinana Clark, Proc. California Acad. Sci. 27: 600; Lejeunea turbinifera Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 295, 1884; Cololejeunea turbinifera (Spruce) Léon-Yanez, Gradst. et Wegner, Hepaticas y Antoceros del Ecuador. Herbario QCA, Quito: 34, 2006; Cololejeunea uleana Steph., Hedwigia 44: 226, 1905; Physocolea uleana (Steph.) Steph., Spec. Hep. 5: 886, 1916. Cololejeunea minutissima subsp. minutissima Cololejeunea minutissima subsp. myriocarpa (Nees et Mont.) R. M. Schust., J. Elisha Mitchell Sci. Soc. 71: 232, 1955. – Basionym: Lejeunea myriocarpa Nees et Mont., Sagra, Hist. Phys. Cuba, Bot., Pl. Cell.: 473, 1842. Typus: Sagra, Cuba, La Habana. Synonyms: Cololejeunea myriocarpa Steph., Hedwigia 29: 89, 1890; Lejeunea epiphyta Gottsche in Gottsche et al. Syn. Hep. 391, 1845; Cololejeunea epiphyta (Gottsche) Steph., Hedwigia 29: 90, 1910; Lejeunea heteromorpha Spruce, J. Linn. Soc. Bot. 30: 348.

Subgen. TAENIOLEJEUNEA (Zwickel) Benedix Beih. Feddes Repert. 134: 21, 1953 Basionym: Taeniolejeunea Zwickel, Ann. Bryol. 6: 106, 1933. Lectotypus: Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. designated by Vanden Berghen, Lejeunia Mém. 6: 52. 67. Cololejeunea appressa (A. Evans) Benedix, Beih. Feddes Repert. 134: 31, 1953. – Basionym: Leptocolea appressa A. Evans, Bull. Torrey Bot. Club 39: 606, 1912. Typus: Evans 104, Jamaica: John Crow Peak. Synonym: Taeniolejeunea appressa (A. Evans) Zwickel, Ann. Bryol. 6: 107, 1933. 68. Cololejeunea falcata (Horik.) Benedix, Beih. Feddes Repert. 134: 29, 1953. – Basionym: Physocolea falcata Horik., J. Sci. Hiroshima Univ. Ser. B, Div. 2, Bot. 1: 22, 1931; Typus: Horikawa 1962a, Japan: Bonin I., Mt. Chuo. Synonyms: Taeniolejeunea falcata (Horik.) Zwickel, Ann. Bryol. 6: 107, 1933; Cololejeunea falcatoides Benedix, Beih. Feddes Repert. 134: 30, 1953. 69. Cololejeunea pseudofloccosa (Horik.) Benedix, Beih. Feddes Repert. 134: 36, 1953. – Basionym: Leptocolea pseudofloccosa Horik. J. Sci. Hiroshima Acta Bot. Hung. 56, 2014

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Univ. Ser. B, Div. 2, Bot. 1: 87, 1932; Typus: Noguchi s.n., China: Taiwan “Formosa”, Tainan, Numano-Daira, Arisan. Synonym: Taeniolejeunea pseudofloccosa (Horik.) S. Hatt., J. Jap. Bot. 17: 465, 1941. New to the Neotropics (coll. R. M. Schuster in Brazil).

Doubtful species 70. Cololejeunea inflectens (Mitt.) Benedix, Beih. Feddes Repert. 134: 79, 1953, as “C. inflectidens”. – Basionym: Lejeunea inflectens Mitt., J. Proc. Linn. Soc., Bot. 5: 117, 1861 “1860”. Typus: Gardner 36, Ceylon, Horton Plains. Synonyms: Physocolea “inflectidens” (Mitt.) Steph., Spec. Hep. 5: 896, 1916; Campylolejeunea inflectens (Mitt.) Mizut., J. Hattori Bot. Lab. 26: 184, 1963; Cololejeunea ciliatilobula (Schiffn.) Schiffn., Consp. Hep. Archip. Ind.: 242, 1898; Physocolea peculiaris Herzog, Mitt. Inst. Allgem. Bot. Hamburg 7: 216; (further synonyms are enumerated in Zhu and So, Beih. Nova Hedwigia, 121: 283, 2001). Syn. nov.: Physocolea reflexiloba Steph., Spec. Hep. 5: 883, 1916. Typus: unknown collector Hb. Jack G., Peru; Cololejeunea “reflexifolia” (Steph.) Tixier, C. R. Soc. Biogéogr. 65: 177, 1989. Note: Tixier (1977), although using mistakenly the species epithet, as “reflexifolia”, recognised its close affinity (“demeure voisin”) with C. inflectens. The specimen label of the type of Physocolea reflexifolia in the Stephani Herbarium indicates Peruvian origin. But on Stephani’s drawing (Stephani: Icones, No. 12269) states: “Peruvia, legit. Jack, in cortice China”. The words “legit Jack” have been written on the sheet later by Stephani. As Jack had never collected in Peru, although the specimen came from Jack’s herbarium, it could have a different origin. Either it really came from Peru from bark of china (Cinchona) trees, as Jack worked with Neotropical material collected by others (Frahm and Eggers 2001), or it simply means that it was collected on Cinchona bark somewhere else in tropical Asia, then Stephani’s indication in Species Hepaticarum of the origin of its type is a mistake. Although its identity with C. inflectens is correct, but the occurrence of this palaeotropic species (and subgenus) here is somehow doubtful and needs confirmation. Cololejeunea marginata (Lehm. et Lindenb.) Schiffn., Consp. Hepat. Archip. Indici 245, 1998. – Basionym: Jungermannia marginata Lehm. et Lindenb. in Lehmann, Nov. Min. Cogn. Stirp. Pugillus 5: 11, 1833. Typus: Hb. Nees, Mauritius. Synonyms: Lejeunea marginata (Lehm. et Lindenb.) Gottsche et al., Synop. Hep. 393, 1845; Leptocolea marginata (Lehm. et Lindenb.) A. Evans, Bull. Torrey Bot. Club 38: 262, 1911. Note: This species from subgen. Pedinolejeunea is widespread in Madagascar and the Mascarene Islands. But its holotype from the Lindenberg Herbarium (No. 6760), indicated as Mauritan in origin, contains Caribbean species of Diplasiolejeunea (Tixier 1985: 71). Grolle (1995: 48) discussed in detail the problem of this variously interpreted taxon. According to him all records from the Neotropics are wrong. Acta Bot. Hung. 56, 2014


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Species to be transferred to other genus Aphanolejeunea lancifera R. M. Schust., Phytologia 45: 434, 1980. – Typus: Schuster 67-342 (F: C0171855), Jamaica, Blue Mts: NW of Hardwar Gap below summit of Caledonia Peak. Note: The examined type specimen, even the small sample sorted out by Schuster himself, contains only Drepanolejeunea sp. with mostly detached underleaves, which could cause the confusion.

Species not seen Cololejeunea conchifolia (Gottsche) Gradstein, in Gradstein and Hekking, J. Hattoti Bot. Lab. 45: 109. “conchaefolia”. – Basionym: Lejeunea conchaefolia Gottsche, Ann. Sci. Nat. 5: 163, 1864. Typus: Lindig, Colombia, Cundinamarca, 2600 m. Synonym: Physocolea conchaefolia (Gottsche) Steph., Spec. Hep. 5: 876, 1916. Note: Judging from Stephani’s description and drawing (Stephani 1985: 12.242), the species belongs to subgen. Protocolea. Cololejeunea papulosa R. M. Schust., Phytologia 45: 433, 1980. – Invalid homonym: Cololejeunea papulosa Tixier, Ann. Fac. Sci. Phnom Penh 1: 59, 1968. Typus: Schuster 67-025a (F: C0171849), Jamaica: Blue Mts, NW of Hardwar Gap, track to waterfall on Caledonia Peak track. Note: In the type material investigated there was no specimen to fit the diagnosis given by Schuster. There is an indication of a slide, which could be traced now in F.

ILLUSTRATED KEY TO THE NEOTROPICAL SPECIES OF COLOLEJEUNEA Terms to describe the size of plants in all keys are defined with the shoot width of well developed specimens, as minuscule: < 0.6 mm, small: 0.6–0.9 mm, medium-sized: 0.9–1.2 mm, large: 1.2–2.3 mm and robust: > 2.3 mm width.

Key to subgenera occurring in the Neotropics 1a

Minuscule heterophyllous plants, i.e. there are fully developed leaves with urn-shaped or inflated lobule, elobulate leaves of full size, but without lobule or with lobules marked only by a narrow band and finally rudimentary leaves reduced into a small, 1–5 cells wide lamina. Branching athecal. Disciform gemmae only on the leaf margin subgen. Aphanolejeunea

1b

Leaves uniform in size and shape, at least part of the branches Lejeunea 2 type. Disciform gemmae also on the leaf surface


202


2a

Minuscule to small, whitish plants. Both athecal basiscopic and thecal Lejeunea type branches occur. Leaves distant, either elobulate (or with a lobule reduced to 1–4 cells), when only very few leaves, sometimes only the female bracts have normal lobule or in other cases most leaves uniformly lobulate, but lobule always narrower than 10 cells. Female bracts quite similar to average leaves. Stylus not visible subgen. Incalejeunea

2b

Small to large plants, only with thecal, Lejeunea type branching. Leaves distant to imbricate, almost always consisting of a larger lobe and of a much shorter, often inflated lobule, which is usually wider than 10 cells. Stylus normally visible, at least on the juvenile shoot apex 3

3a

Leaf lobe with a sharply delimited vitta consisting of several elongated, enlarged, translucent cells with one large, pellucide, botryoidal oil body in fresh state (vitta absent in C. pseudofloccosa). Small to medium-sized, yellowish or whitish green plants with an opalescent lustre. Epiphyllous, adherent to the substrate, do not shrivel when dry. Other median cells small with evenly thickened walls (diameter less than 10 μm), each with a small dorsal papilla subgen. Taeniolejeunea

3b

Leaves without vitta

4a

Leaf lobe bordered by a shiny hyaline margin or by translucent sigmoid cells. Medium to large-sized, mostly epiphyllous, pale green plants 5 strongly adherent to the substrate. Perianth strongly compressed

4b

Leaf lobe not bordered by hyaline cells

5a

Leaf lobe or at least its apex bordered by a sharply delimited, shiny hyaline margin consisting of 1–3 rows of quadrate or 1 row of finger-like, thin-walled, transparent cells or at least 1 or a few finger-like cells tip the leaf apex subgen. Pedinolejeunea

5b

Leaf lobe bordered by translucent, sigmoid cells which are not sharply delimited but gradually transient to the normal lobe cells subgen. Chlorocolea

6a

Stem in cross section composed by 9–20 cortical cells. Robust, fleshy, dark green plants with leaf cells fully packed by very many chloroplasts. Ventral merophyte 3–5 cells wide. Branches with clustered gynoecia. Plants living on irrigated riverbed rocks or epiphyllous on temporarily inundated riverbed shrubs subgen. Chlorolejeunea

6b

Stem in cross section with 5–7 (maximum 8) cortical cells. Medium to large-sized plants, leaves not fleshy, often pale green with much less


4

6


203

chlorophyll. On different substrates, but mostly epiphyllous, not occurring in temporarily inundated streambeds 7 7a

Lobules strongly inflated, large compared to the lobes with strongly involute free margin and ciliate teeth. Large, rigid plants, often brownish in colour even alive with a silky shine. Discoid gemmae without adherent cells. Corticolous or epiphyllous, never adherent to the substrate. Doubtful in the Neotropics subgen. Cryptolejeunea

7b

Lobules not strongly or only partly inflated, when fully inflated, the free margin not involute, teeth not ciliate. Small to robust, delicate, not rigid plants. When fresh, vivid or pale or yellowish green. Discoid gemmae with or without adherent cells. Mostly epiphyllous, loosely or strongly adherent to the substrate, but may occur also on bark or on rocks 8

8a

Each lobe cell with a distinct, mostly acute papilla. Lobe margin dentate, plants green. Adherent or loosely lying on the substrate subgen. Cololejeunea

8b

Cell smooth or if papillose, papillae are restricted to the marginal or sub9 marginal cells. Lobe margin entire, crenulate or denticulate

9a

Usually pale green, small to robust, mostly epiphyllous plants, appressed to the substrate, with imbricate to contiguous leaves. Lobule variable, often saccate, usually connected to the lobe with a well defined keel, which never exceeds the half length of the lobe. Lobe and lobule cells with various thickenings, walls usually not convex and protruding. Lobe margin entire or crenulate or denticulate or dentate. Gemmae often with adhesive cells. Lobule of female bracts often with keel. Perianths various, but without swollen neck and not stipulate subgen. Leptocolea

9b

Bright to yellowish green, small plants often occurring on bark or other substrates too, richly branching, often with a creeping, stoloniform main stem with erect branches and distant leaves. Lobule polymorphic, often reaching or exceeding 0.6 of lobe length, connected to the lobe by an ill defined, vestigial sinus and often erect, stiff or saccate, or represented only by a narrow fold. Real keel absent. Lobe and lobule cells thin walled, without thickenings, often mammillose with concave, protruding walls making the leaf margin crenulate. Gemmae without adhesive cells. Female bracts are, similarly to the leaves, without keels. Perianth obovoid or obpyriform, with swollen neck, often on a prolonged stalk, stipitate and long exserted with age, with 5 equal, sharp keels which are crest-like, formed by two rows of somewhat protruding cells subgen. Protocolea


204


Key to the Neotropical species of subgen. Aphanolejeunea (Evans) Benedix 1a

Fully developed leaves lanceolate, acute

2

1b

Fully developed leaves ovate with rounded or triangular apex

7

2a

1st lobule tooth bicellular, falcately incurved outwards. Lobe margin slightly inrolled. Gemmae angulate. Reduced leaves linear, 2–4 cells wide 2. C. cingens

2b

1st lobule tooth unicellular, lobe margin flat

3a

Perianth obconic, sharply 5-carinate, with keels sharply angulate above and conically papillate. Beak conspicuous 3. C. cornutissima

3b

Perianth obovate, its whole surface obtusely or acutely papillose. Beak inconspicuous 4

4a

1st lobule tooth inconspicuous, shorter than the 2nd tooth, rounded or triangular, projecting ventrally. Lobe dorsally papillose (subgen. Incalejeunea: C. contractiloba)

4b

1st tooth at least as long as the 2nd one, mostly falcate and cross each other 5

5a

Lobulate leaves with lobule exceeding 1/2 lobe length, with its width equalling the lobe width 4. C. costaricensis

5b

Lobulate leaves with lobule shorter than 1/2 lobe length and much narrower than the lobe 6

6a

Leaf cells at the keel and margin conically protuberant 10a. C. sicifolia subsp. sicifolia

6b

Leaf cells (except for the lobule) with acutely papillose 10b. C. sicifolia subsp. jamaicensis

7a

Lobe strongly contracted at the junction of the lobe and lobule. 1st lobule tooth slightly falcate covering the 2nd, obsolete tooth 13. C. winkleri

7b

Lobe not or only slightly contracted at the junction of lobe and lobule. Only the 1st lobule tooth is well developed, not covering the 2nd blunt and obsolete tooth 8

8a

Leaves and perianth smooth, cells not papillose but in cases mammillose. Lobe margin of fully developed leaves entire. Lobule 0.25–0.4 of lobe length 9


3


205

Plate 1. 1 = Cololejeunea berneckerae (habit and lobule), 2 = C. cingens (fully developed and reduced leaf), 3 = C. cornutissima (habit), 4 = C. costaricensis (fully developed and reduced leaf), 5 = C. cubensis (habit, fully developed and reduced leaf), 6 = C. gracilis (habit with fully developed and reduced leaf), 7a = C. microscopica var. microscopica, 7b = C. microscopica var. exigua, 7c = C. microscopica var. africana (fully developed leaves), 8 = C. minuscula (habit, stem sections and gemmaling). Figure numbers follow the numbering of taxa in the synopsis and in the keys. All pictures of habit and leaves are in ventral view. Acta Bot. Hung. 56, 2014

206


8b

Leaves and perianth more or less papillose or the cells conically protuberant. Lobe margin denticulate or crenate. Lobule 0.4–0.5 of lobe length 16

9a

Almost all leaves elobulate or reduced, only very few leaves are fully 10 developed with saccate lobule

9b

A greater part of leaves are fully developed, but also elobulate and reduced leaves occur. Stem always composed of 5 cortical and 1 medullar cell row 13

Plate 2. 9 = Cololejeunea papillosa (habit with reduced leaves), 9a = C. grossepapillosa (fully developed leaf), 10a = C. sicifolia ssp. sicifolia, 10b = C. sicifolia ssp. jamaicensis, 11 = C. sintenisii, 12 = C. subsphaeroidea, 13 = C. winkleri (fully developed leaves), 14 = C. linopteroides (habit, stem sections, extremely reduced lobule and lobe margin). Figure numbers follow the numbering of taxa in the synopsis and in the keys. Acta Bot. Hung. 56, 2014


207

10a Small plant living on decaying wood. Almost all leaves and even the female bracts are reduced into a cuneiform, lobulato-dentate, 5–7 cells long and 3–5 cells wide structure, having small lobule marked by a narrow band of 2–6 cells 5. C. cubensis 10b Minuscule plants forming a thin web on the substrate, similarly to filamentous algae. Almost all leaves reduced into two cells wide, linear structures 11 11a Stem in cross section reduced into 5 cortical cells, without medullary cell row. Leaves often grow in a long, ribbon-like gemmae. Discoid gemmae also present, producing similar ribbon-like, 2 cells wide gemmalings 8. C. minuscula 11b Stem is always composed of 5+l cell rows. Reduced leaves 4–6 cells long, 1–2 (usually 3) cells wide, linear, but not developing into long, ribbonlike gemmae 12 12a Reduced leaves with entire margin 12b Reduced leaves with dentate margin

9. C. papillosa 6. C. gracilis var. linearifolia

13a Lobule 0.6 to 0.95 of lobe length. 1st lobule tooth consists of two cells, slightly falcate. Cells mammillose 14 13b Lobule 0.25 to 0.4 of lobe length. 1st lobule tooth 1–2-celled, most cases falcate 15 14a Free part of lobe straight, obtuse or acutely triangular. Lobule 0.6 to 0.8 of lobe length, saccate with free opening 6a. C. microscopica var. microscopica 14b Free part of lobe apex rounded, deflexed-involute, partly covering the opening of lobule sac. The subglobose lobule reaches 0.9 of the lobe length 12. C. subsphaeroidea 15a Free part of lobe broad or narrow triangular. Reduced leaves common, lanceolate, 2–4 cells broad. Lobule about half of lobe length. Gemmae 20–22-celled. Perianth inflated, almost terete, smooth, with 5 week carinae papillose at edges 11. C. sintenisii 15b Free part of the lobe spathulate with rounded apex. Lobule only 0.25 of lobe length and much narrower than lobe width. 1st tooth 1–2-celled, strongly falcate, forming the lobule apex into lunular shape 1. C. berneckerae


208


16a Lobule of fully developed leaves only up to 0.5–0.6 of lobe length. Perianth and lobule with conical papillae 17 16b Lobule of fully developed leaves 0.6–0.8 of lobe length

18

17a Reduced leaves linear, 2–4 cells wide, with dentate margin. First lobule tooth straight 6. C. gracilis When practically all leaves are reduced

C. gracilis var. linearifolia

17b Reduced leaves broader, ovate in outline, 3–8 cells broad with entire margin. First lobule tooth falcate 11. C. sintenisii 18a First lobule tooth straight. Dorsal side of lobe and perianth densely covered by conical papillae. Many leaves reduced, linear, only 2 cells broad (species hitherto known only from the Palaeotropics) 9a. C. grossepapillosa 18b First lobule tooth falcate. A good number of leaves fully developed. Reduced leaves ovate or ligulate, 3–4 cells broad 19 19a Leaf (especially along the keel) and the perianth with conically protuberant cells 9b. C. microscopica var. exigua 19b Leaf and perianth with acute or spinose papillae 9c. C. microscopica var. africana

Key to the Neotropical species of subgen. Chlorocolea R. M. Schust. 1a

Small plant with 0.75 mm wide shoot and flattened, ribbon-like stem with 1–5 ventral merophytes and with 4 dorsal cortical cells. Lobule almost fully reduced, only one basal cell with a hyaline papilla (stylus) represents it. Female bract lobule setiform, narrow lanceolate. Gemmae without adhesive cells 14. C. linopteroides

1b

Large plant with 1.5 mm wide shoot and with almost terete stem. Lobule triangular-lanceolate, consisting of 6–9 cells of which 2–3 forms the uniseriate apex Female bract lobule broad ligulate. Gemmae with 3 adhesive cells 15. C. standleyi

Key to the Neotropical species of subgen. Chlorolejeunea Benedix This subgenus is represented only by one species in the Neotropics, although this recently described taxon is very distinct endemic of the Ecuadorian riv-



209

erbanks. A robust, fleshy plant with dark green with leaf cells fully packed by very many chloroplasts. Stem in cross section composed by 9–20 cortical cells. Ventral merophyte 3–5 cells wide. Branches with clustered gynoecia 16. C. stotleriana

Key to the Neotropical species of subgen. Cololejeunea 1a

Leaf triangular ovate with acute apex and with very long, digitally protuberant papillae at its margin and surface. Lobules of fully developed leaves ovate with truncate apex, saccate in their proximal half, bearing two obtuse, 1-celled teeth. Most leaves with reduced lobule consisting only of two enlarged marginal cells, the distal one tipped by a hyaline papilla 18. C. kuciana

1b

Leaves ovate to orbicular, with rounded or obtuse apex

2a

Lobe with vitta-like rows (pseudovitta) of larger cells not too sharply delimited of the other leaf cells. Dorsal leaf surface papillose, usually only in its outer third. Papillae small and low. Perianth smooth, 4 carinate 19. C. platyneura

2b

Lobe without pseudovitta, cells more or less equal

3a

Pale green, leaves almost orbicular (sometimes obtusely apiculate) with near entire, crenulate margin. Each lobe cells (except for the base) with 1 small papilla. Lobule reduced, consisting only of two inflated cells in a line, the distal one tipped by a small hyaline papilla. Stylus lacking. Small discoid gemmae abundant on both lobe surfaces, consisting of 16–18(–20) cells 20. C. tamasii

3b

Leaves ovate to suborbicular, margin strongly denticulate by protruding, acute papillae which cover also the leaf surface. Lobule well developed, with a 2-celled 1st and with a 1-celled 2nd tooth. Stylus present. Ventral merophyte as wide as the two neighbouring lateral merophyte cell rows 4

4a

Brownish green plants with almost symmetric ovate to suborbicular leaves. Keel smooth 21. C. thiersiana

4b

Pale green plants with very asymmetric ovate leaves. Keel with protruding papillae 17. C. elegans

2

3


210


The Neotropical species of subgen. Cryptolejeunea Benedix Robust, rigid corticolous or epiphyllous plant. Leaves brownish, parchment like with strongly involute, ciliate free lobule margin. Cell walls smooth, relatively thin with small trigones and intermediate thickenings. Female bracts almost as wide as long. Gemmae not observed. (A Palaeotropical species. Its occurrence in the Neotropics very doubtful) 70. C. inflectens

Plate 3. 15 = Cololejeunea standleyi (habit and lobe margin), 16 = C. stotleriana (habit and stem sections), 17 = C. elegans (habit and keel margin), 18 = C. kuciana (fully developed leaf), 19 = C. platyneura (leaf and lobule), 20 = C. tamasii (leaf and stem section), 21 = C. thierisana (leaf and lobe margin), 22 = C. antillana (habit with perianths), 23 = C. azorica (fully developed leaf). Figure numbers follow the numbering of taxa in the synopsis and in the keys.



211

Key to the Neotropical species of subgen. Incalejeunea Tixier ex Pócs 1a

Leaves spathulate or parallel sided, widest in their upper third with crenulate margin. Lobule well developed, with a straight, 1–2 cellular tooth. Perianth obovate with five tuberculate carinae 34. C. spathulata (the leaves of C. moralesiae in some cases can be also spathulate but this species is distinct by its five horned perianth)

1b

Leaves ovate to lanceolate, widest in their lower or median part

2a

All leaves acute, narrow lanceolate, length-width ratio more than 4 : 1, with lobule about 1/3 length of lobe. 1st lobule tooth 1-celled, short, blunt, equal with the 2nd tooth in size. Perianth terete without carinae 22. C. antillana

2b

At least the greater part of leaves ovate to ovato-lanceolate, length-width ratio 1.5–3 : 1 3

3a

Dorsal side of lobe and the whole perianth surface papillose

4

3b

Leaves and perianth smooth

7

4a

Most leaves with lobule reduced into a little fold of 1–6 cells, tipped by hyaline papilla. Well developed lobule almost as large as the lobe with a straight, bicellular tooth 25. C. clavatopapillata

4b

Fully developed lobules common, lobule reaching only 1/2 to 2/3 length of the lobe 5

5a

First lobule tooth 1-celled, obsolete, rounded or triangular, projecting ventrally. Leaves broad ovate with acute, triangular apex 26. C. contractiloba

5b

First lobule tooth well developed, straight, consisting of two cells

6a

Leaves ellipsoid-ovate, lobule at most 2/3 of lobe width. Lobe apex rounded, ending in more than one cell 31. C. papilliloba

6b

Leaves ovato-lanceolate to lanceolate, lobe apex acute ending in one triangular cell. Lobule almost as wide as the lobe 28a. C. diaphana var. cristulata

7a

A great part of leaves fully developed, lobulate

7b

Greatest part of leaves elobulate, lobulate leaves very rare

8a

Leaves broad ovate with lobule 2/3 of lobe length and width. Discoid gemmae round or reniform, devoid adhesive cells. Lobule tooth straight, bicellular 23. C. azorica

8b

Leaves ovate to lanceolate with lobule 1/3 to 1/2 of lobe length. Discoid gemmae (in C. diaphana and in C. yelitzae) reniform, with 3 adhesive cells 9

2

6

8 11


212 9a


Leaf width very variable from wide ovate to narrow lanceolate within the same plant. Lobule acute lanceolate with a wide gap between the 1st and 2nd teeth. Gemmae abundantly develop on leaf surface and even on the perianth 35. C. yelitzae

Plate 4. 24 = Cololejeunea camillii (leaf, discoid gemmae in front and in side view), 25 = C. clavatopapillata (habit with perianth), 26 = C. contractiloba (leaf and discoid gemma), 27 = C. crenata (leaf), 28 = C. diaphana var. diaphana (leaf and gemma), 28a = C. diaphana var. cristulata (leaf), 29 = C. erostrata (habit with perianths and gemmae), 30 = C. moralesiae (habit with perianth). Figure numbers follow the numbering of taxa in the synopsis and in the keys. Acta Bot. Hung. 56, 2014


9b

213

Leaf shape quite uniform. Lobule truncate with a small gap between the 1st and 2nd teeth 10

10a Leaves ovate to ovato-lanceolate. First lobule tooth long, straight, directed forward. Lobe apex more or less acute, ending in one cell. Lobule cells isodiametric 28. C. diaphana 10b Leaves ellipsoid to spathulato-ovate with crenulate margin. Lobe apex rounded. 1st tooth short, 1–2-celled, directed sideway. 2nd tooth blunt, obsolete. Lobule cells sigmoid 27. C. crenata 11a Leaves lanceolate with more or less acute apex

12

11b Leaves elliptic with more or less rounded apex

14

12a Leaves very asymmetric, falcate, with incurved, acuminate apex. Gemmae on the lobe surface 32. C. paucifolia 12b Leaves symmetric, ovato lanceolate, with acute apex

13

13a Perianth 5-carinate with a short beak, carinae tuberculate, leaf margin smooth to crenulate. Gemmae develop on the lobe margin, 16–20-celled 24. C. camillii 13b Perianth obconic with 5 auriculate wings

33. C. pteroccolea

13c Perianth inflate, ecarinate without a beak, with umbilicate mouth. Gemmae at the margin and on the surface of the lobe 29. C. erostrata 14a Perianth obovate, 5-carinate with tuberculate carinae. Leaves elliptic to narrow ovate 24. C. camillii 14b Perianth obconic with 5 long horns. Leaves elliptic to spathulate 30. C. moralesiae

The Neotropical species of subgen. Leptocolea (Spruce) Schiffn. and the species excluded from subgen. Protocolea sensu R. M. Schust. 1a

Lobule saccate, ovate and inflate, keel about 1/3 of lobe length

2

1b

Lobule small, triangular or lanceolate, directed diagonally, with an apex of several cells in line, keel much shorter 5

2a

Lobule at its distal end adherent to the lobe. Lobule of female bracts lanceolate, acute. Perianth terete, inflate, smooth. Lobule with two well developed teeth 38. C. planiuscula

2b

Lobule end not adherent to the lobe. Lobule of female bracts with obtuse or round apex. Perianth at least at its apex 5-carinate 3 Acta Bot. Hung. 56, 2014

214 3a


Small plants with lanceolate lobe, length-width ratio about 3 : 1. Perianth 4-carinate. Lobe and perianth cell walls without thickenings and papillae. Lobule unidentate 36.C. hildebrandii

Plate 5. 31 = Cololejeunea papilliloba (leaf), 32 = C. paucifolia (leaf), 33 = C. pterocolea (habit with perianth, leaf and spore), 34 = C. spathulata (habit and leaf), 35 = C. yelitzae (habit with gemmae, lobule), 36 = C. hildebrandii (leaf, lobule and lobe cells), 37 = C. obliqua (leaf, lobe margin, perianth with bracts), 38 = C. planiuscula (leaf and female bract lobules, perianth with bracts), 39 = C. skottsbergii (leaf, perianth with bracts), 40 = C. spruceana (leaf, lobules). Figure numbers follow the numbering of taxa in the synopsis and in the keys.



215

3b

Medium to large size plants with lobe asymmetric ovate, length-width ratio about 2 : 1. Lobule bidentate. Cell walls papillose or with intermediate thickenings 4

4a

Lobe at least near its border and the perianth papillose. 1st lobule tooth composed of two cells, longer than the 2nd, 1-celled tooth. Perianth obovate with 4 keels, truncate at apex 37. C. obliqua

4b

Lobe and perianth smooth, lobe cells with intermediate thickenings. Lobule teeth near equal, both consisting of two cells. Perianth elongated obconic 39. C. skottsbergii

5a

Medium to large plants, lobule reduced to a few cells consisting of a small triangular base of 6–8 cells with a 3–4 cells long uni-(or bi-) seriate tooth 40. C. spruceana

5b

Minuscule plants with lobules forming a narrow, 2–3 cells wide basal fold with a 2–4 cells long uniseriate tooth 41. C. setiloba

Key to the Neotropical species of subgen. Pedinolejeunea Benedix ex Mizut. 1a

Hyaline margin fimbriate or crenulate, composed of finger-like cells or at least one finger-like hyaline cell occur at the apex of certain leaves 2

1b

Hyaline margin entire, formed by 1–4 rows of rectangular cells

2a

Stylus filamentous, consisting of 3–8 seriate cells. Medium size plant. Leaves triangular ovate, a part of them tipped by one hyaline cell. Lobule about half of the lobe size with a straight, 2-celled tooth 47. C. ecuadoriensis

2b

Stylus unicellular or not visible. Small to medium size plants

3a

Hyaline cells only at the leaf apex, solitary or forming a group of 5–15, but a number of leaves may devoid them at all 4

3b

Hyaline cells form a continuous, crenulate margin extending to the dorsal lobe edge, where the finger-like cells gradually change into a row of rectangular ones 8

4a

Lobule setaceous formed by 4–6 uniseriate cells on a triangular base of 6–10 cells. Male branch with only 1–3 pairs of small bracts 52. C. micrandroecia

4b

Lobule not setaceous

9

3

5


216


5a

Lobule small, about 0.1 of the lobe length, consisting of only 8–18 cells, flat, rectangular or triangular, with unicelled tooth topped by one hyaline papilla. Leaves, male and female bracts with few or more hyaline cells at apex 53. C. minutilobula

5b

Lobule larger, at least one third of lobe length, inflated on almost all leaves 6

Plate 6. 41 = Cololejeunea setiloba (leaf and gemma), 42 = C. bekkeri (leaf and lobules), 43 = C. bischleriana (apex of lobules), 44 = C. cardiocarpa (leaf and lobe apex), 45 = C. cremersii (apex of female bract lobule, leaf lobule), 46 = C. cristata (habit, lobe margin), 47 = C. ecuadoriensis (habit, lobe apex), 48 = C. geissleriana (habit and apex of lobules). Figure numbers follow the numbering of taxa in the synopsis and in the keys.



217

6a

Leaf broad triangular ovate with only one finger-like or orbicular hyaline cell at the apex of a number of leaves, missing on others. Lobule teeth usually unicellular, triangular 55. C. panamensis

6b

Leaf elongato-ovate with usually more than 1 hyaline cell at lobe apex. Lobule 1/3 to 1/2 of lobe length, with elongate, 2-celled first tooth. Perianth wings smooth 7

7a

Leaves subsymmetric, ellipsoid ovate in outline. Lobule broad ovate, 20– 50 cells wide. Lobule cells isodiametric. 1st lobule tooth straight. Male bracts ovate with truncate apex 44. C. cardiocarpa

7b

Leaves asymmetric, subfalcate, ovato-ligulate. Lobule narrow ovate to lanceolate, 12–14 cells wide. Lobule cells elongate, 2–5 times longer than wide. 1st lobule tooth falcate, turned downwards. Male bracts lanceolate with acute apex 58. C. schusteri

8a

Lobules relatively large, at least 1/3 of lobe length, inflated. Cuticle smooth 59. C. subcardiocarpa

8b

Lobules small, less than 1/4 lobe length, inflated or flat and reduced to a small triangular or quadrate structure. Cuticle smooth to finely striate 63. C. verwimpii

9a

Lobules relatively large, ovate, inflate or ligulate, at least 1/3 of lobe length 10

9b

Lobules relatively small, shorter than 1/3 of lobe length

19

10a Lobules ovate, inflate, with well differentiated teeth (in C. guadelupensis ligulate lobules also occur) 11 10b Lobules flat or only partly inflated, ligulate or square, deltoid or triangular, sometimes alternating with ovate, inflated lobules. Lobule teeth weekly or not differentiated 16 11a Cuticle punctate. Lobule with two well differentiated teeth 49. C. guadelupensis 11b Cuticle smooth or finely striate

12

12a Dorsal lobe with auriculate base projecting across the stem

13

12b Dorsal lobe with rounded base, often projecting onto but not across the stem. Ventral merophyte one cell wide 14


218


Plate 7. 49 = Cololejeunea guadelupensis (leaves with different lobules, lobe apex), 50 = C. lanciloba (habit, leaf and lobules), 51 = C. latilobula (stem sections, leaves, lobule and lobe apex), 52 = C. microandroecia (leaf, lobules and lobe apex), 53 = C. minutiulobula ( habit with perianth, lobule and lobe apex), 54 = C. nigerica (leaves and lobules), 55 = C. panamensis (leaves), 56 = C. planifolia (habit, leaf). Figure numbers follow the numbering of taxa in the synopsis and in the keys.



219

13a Lobule with two prominent teeth. First tooth long, acute, bicellular, second tooth triangular. Cuticle striolate. Ventral merophyte of 2(–4) cell rows 64. C. vitaliana 13b Lobule with 3 teeth. The median one bicellular, short and blunt, while the 48. C. geissleriana two others obtuse and obsolete. Cuticle not striolate 14a Median lobule cells elongate, 3–4 times longer than wide. First lobule tooth often falcate and bent downwards. Female bract lobules with dentate apex 45. C. cremersii 14b Median lobule cells rectangular, only 1–2 times longer than wide. First lobule tooth always straight 15 15a Lobule teeth short, the 1st one 1–2-celled, blunt, while the 2nd short triangular, obsolete. Lobule of female bract entire 60. C. submarginata 15b Lobule teeth acute, the 1st one consists of 2–3 uniseriate cells, while the 2nd triangular or also uniseriate 43. C. bischleriana 16a Hyaline cells concentrated at the lobe apex, where they form an abruptly broadening, protruding, multiseriate group 54. C. nigerica 16b Hyaline cells all along the dorsal and distal lobe edge, forming an evenly broadening, relatively thin margin towards the apex 17 17a Lobule usually ligulate or triangular without any teeth on margin. An Ecuadorian specimen has finely striolate cuticle. Ventral merophyte in 2–5 rows 51. C. latilobula 17b Lobule very variable in shape, usually has an obsolete tooth on its lateral margin or two teeth when inflate, saccate lobule present 18 18a Medium size plant, leafy shoots 1 mm wide. Hyaline lobe margin 1–2 cells wide throughout. Cuticle smooth. Perianth smooth, inflated, without carinae or with a weak ventral fold 56. C. planifolia 18b Large plant, leafy shoots 1.2–2.4 mm wide. Hyaline margin 3–4 cells wide at the lobe apex, 2 cells wide at the dorsal margin 57. C. planissima 19a Lobules more or less ligulate, acute or blunt, directed forwards or diagonally, their axis much longer than the keel. Lobe base more or less auriculate, projecting across the stem. Hyaline margin continues to the base of ventral lobe margin, cuticle finely punctate. Lobule lanceolate with 1–2 blunt teeth at their proximal side and capped by a hyaline papilla 20 19b Lobules triangular, trapezoid or ovate, hyaline margin restricted to the distal part of lobe at least at its ventral edge 21


220


20a Lobule inserted to the keel

50. C. lanciloba

20b Lobule base partly adnate to the lobe

42. C. bekkeri

21a Cuticle conspicuously reticulate, hence lobe cells opaque 62. C. surinamensis 21b Cuticle smooth, lobe cells transparent

22

22a Lobule triangular-lanceolate, with 1–2 teeth at its proximal side and capped by a hyaline papilla 50. C. lanciloba 22b Lobule trapezoid or triangular, almost entire with 1–2 blunt or obsolete teeth 61. C. subscariosa

Key to the Neotropical species of subgen. Protocolea R. M. Schust. emend. Pócs (see characters in the subgeneric key) 1a

Lobe with acute apex. Lobules never inflated, with a fold of 0.5–0.6 lobe length, trapezoidal, perpendicular to it or coplanar with the lobe, with a triangular tooth tipped by the hyaline papilla. Female bracts similar to the leaves. Perianth on a 0.4–0.5 mm long stalk 65. C. dauphinii

1b

Lobe with rounded apex. Lobule either inflated, large (0.7–0.9 of lobe length), with bicellular 1st tooth and triangular, obtuse 2nd one (not well visible on not inflated lobules) or explanate. Hyaline papilla at the ental base of the 1st tooth. Female bracts obtuse. Perianth on a shorter stalk or not stipitate 2

2a

Lobule almost always inflated with two well developed teeth. Female bracts with well developed lobule 66a. C. minutissima subsp. minutissima

2b

Almost all lobules explanate, not inflate, often reduced into a narrow fold, teeth obsolete. Female bracts with rudimentary lobule 66b. C. minutissima subsp. myriocarpa

Key to the Neotropical species of subgen. Taeniolejeunea (Zwickel) Benedix 1a

Vitta sharply delimited from the rest of much smaller lobe cells. Hyaline papilla between the two unicelled teeth. 1st tooth falcate, 2nd one acute 2

1b

Vitta absent or not sharply delimited from the neighbouring cells


3


2a

221

Leaves ligulato-ovate, somewhat falcate. Vitta formed by two rows of elongated cells, which are 3–4 times longer than wide 68. C. falcata

Plate 8. 57 = Cololejeunea planissima (habit, lobule), 58 = C. schusteri (leaf), 59 = C. subcardiocarpa (leaf, lobe margin), 60 = C. submarginata (leaf, lobule), 61 = C. subscariosa (leaves, lobules), 62 = C. surinamensis (habit, lobe apex), 63 = C. verwimpii (leaf, lobule, lobe margin), 64 = C. vitaliana (leaf, lobule apex), 65 = C. dauphinii (folded and unfolded leaf). Figure numbers follow the numbering of taxa in the synopsis and in the keys.


222


2b

Leaves ovate. Vitta built up of one row of 3–4 oblong, quadrangular cells, which are only 2–3 times longer than wide 67. C. appressa

3a

Vitta absent. Stylus bicellular. Hyaline papilla hidden behind the two bicellular lobule teeth, which cross each other 69. C. pseudofloccosa

3b

Pseudovitta present, formed by 6–12 large basal cells arranged in 2–4 irregular rows and gradually transient into the smaller lobe cells. Stylus not seen. Lobule with two unicellular teeth with the hyaline papilla between them (subgen. Cololejeunea: C. platyneura)

A part of the small figures are original: 9, 10a, 10b, 12, 23, 24, 27, 31 and 35 were drawn by A. Bernecker; 50, 52, 56 and 61 by Tixier; 62 by A.-L. Ilkiu-Borges. The rest were extracted and modified from the publications of different authors, as Allorge and Jovet-Ast (1950): 11; Asthana and Srivastava (2003): 51, 57; Benedix (1953): 70; Bernecker-Lücking (1998): 1, 4, 30; Bernecker-Lücking

Plate 9. 66a = Cololejeunea minutissima ssp. minutissima (leaf), 66b = C. minutissima ssp. myriocarpa (habit), 67 = C. appressa (leaf and lobe apex), 68 = C. falcata (leaf and lobe apex), 69 = C. pseudofloccosa (leaf, lobule, lobe margin and spore), 70 = C. inflectens (habit, lobule, lobe cells). Figure numbers follow the numbering of taxa in the synopsis and in the keys.



223

and Morales (1999): 14; Dauphin and Pócs (Dauphin et al. 2006): 55; Gradstein et al. (2011): 16; Herzog (1942a): 5, 29; Herzog (1942b): 39; Herzog (1951): 15, 53; Herzog (1952): 32; Jones (1953): 54; Jones (1954): 67; Jones (1968): 17; Jovet-Ast (1947): 6, 22; Morales and Dauphin (1998): 65; Morales and Lücking (1998): 13; Pócs (1975): 19; Pócs (1980): 36; Pócs (1984a): 7a, 7b, 7c; Pócs (2002): 47, 58; Pócs and Piippo (1999): 9a; Pócs and Schäfer-Verwimp (2012): 18; Reiner-Drehwald (1995): 25; Schäfer-Verwimp (2012): 20; Schuster (1955); 3; Schuster (1980b): 8, 26, 28, 28a, 41, 44, 66a, 66b; Tixier (1980): 43, 48, 59, 60; Tixier (1985): 37; Tixier (1991): 21, 38, 40, 42, 45, 49, 63, 64; Zhu (1995): 68; Zhu and So (2001): 69. * Acknowledgements – T. Pócs is indebted to professors Attila Borhidi, Deysi Reyes Montoya, Yelitza León Vargas, Ricardo Rico for organising his collecting trips in Cuba and Venezuela, to the Hungarian Academy of Sciences, the Eszterházy College in Eger and to the Hungarian Research Fund (OTKA, Grant No. T38319) for sponsoring his research. He is grateful to Dénes Balázs, Andrea Bernecker, Alfons Schäfer-Verwimp, Gregorio Dauphin, Elena Reiner-Drehwald, Jens Eggers, Daniel H. Norris, Robert Lücking, Klaus Kalb and to Pierre Tixier for providing their collections for his investigations and to the herbaria of F, G, GOET, JE, MANCH, NY, PC, UTR for the loan of type materials. Finally thanks are due to Prof. Jochen Heinrichs, Anders Hagborg, Matt von Konrat for their multifold help and especially to Prof. S. Robbert Gradstein for his critical remarks and useful amendments and to Prof. Lars Söderström for his thorough nomenclatural and other corrections.

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synopsis and key to species of neotropical cololejeunea

synopsis and key to species of neotropical cololejeunea

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