Chapter 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past, Present and Future César Garcia, Cecília Sérgio, and James R. Shevock Abstract This chapter aims to present a review of the knowledge of the bryological flora for the São Tomé and Príncipe Islands (Gulf of Guinea). An updated catalogue is presented, as well as a brief overview of the first expeditions conducted by the University of Coimbra. The labels of the historical herbarium collections and correspondence were analyzed, which provides an important source of data contributing toward research in taxonomy and conservation of these oceanic islands. Since 2007, exploratory fieldwork was carried out in different habitats of this archipelago along an altitudinal gradient, aiming to improve the knowledge of the ecology and distribution patterns of its bryophyte flora. A total of 304 taxa of bryophytes (133 mosses, 164 liverworts and seven hornworts) are currently reported, of which 21 are endemic to São Tomé and Príncipe and 144 species are shared endemics with the African continent. Several vouchers, especially in the herbaria of the University of Lisbon and of the California Academy of Sciences, are still under study and will likely provide further insights and new discoveries. Keywords Africa · Biodiversity · Bryophytes · Conservation · Expeditions · Herbaria Introduction Bryophytes are a group of land plants that includes mosses, liverworts, and hornworts, and with over 20,000 described species, they are the second most speciose group of higher plants, after angiosperms (Patiño and Vanderpoorten 2018; Song C. Garcia (*) · C. Sérgio Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Lisbon, Portugal Centre for Ecology, Evolution and Environmental Changes (cE3c), CHANGE Associated Laboratory - Global Change and Sustainability Institute, Faculdade de Ciências, Universidade de Lisboa, Lisbon, Portugal e-mail: cgarcia@fc.ul.pt J. R. Shevock California Academy of Sciences, Department of Botany, San Francisco, USA © The Author(s) 2022 L. M. Pires Ceríaco et al. (eds.), Biodiversity of the Gulf of Guinea Oceanic Islands, https://doi.org/10.1007/978-3-031-06153-0_9 217 218 C. Garcia et al. et al. 2021). Bryophytes are a common component of tropical forests and provide important ecosystem functions. In tropical regions during rainstorms numerous bryophyte species can quickly absorb (and retain) many times their dry weight in water. The absorbed water is then slowly released over time back into the environment, thereby reducing the erosive effects of heavy rain and allowing other plants and animals to benefit from the rain and the humid environment for a longer period (Pócs 1982). Bryophytes are widely distributed in terrestrial ecosystems (St Martin and Mallik 2017), and islands provide an exceptional natural laboratory for ecological and evolutionary research in this group of terrestrial plants that is often poorly studied. Oceanic tropical islands usually host amazing bryophyte diversity, including endemic species, especially in the montane forests favoured by ideal climatic conditions, such as frequent rainfall and permanent fog (Ah-Peng et al. 2012). The first known bryophyte collections of São Tomé and Príncipe were made by Friedrich Welwitsch (1806–1872) in 1853 and 1860, as part of the expeditions supported by the Portuguese government to Angola (Dolezal 1974), followed by Charles Barter (1821–1859) and Gustav Mann (1836–1916), botanists with special interest in vascular flora (Sérgio and Garcia 2011). One of the greatest Portuguese mentors of Botany in Africa was Júlio Augusto Henriques (1838–1928) and he outlined a plan for studying the flora of São Tomé and Príncipe (Coutinho 1929–30). Throughout his life, Henriques remained deeply invested in understanding the botanical diversity of the archipelago, which motivated his tireless research as professor and director of the Botanical Garden of the University of Coimbra for more than 50 years. His initial interest in the botanical study of São Tomé and Príncipe was likely related to cultivation of the Cinchona tree (Chinchona spp.) and other medicinal plants at the Coimbra Botanical Garden (Perpétuo et al. 2012). At the age of 65, Henriques led an expedition to the island of São Tomé to study the island’s flora, departing from Lisbon on June 23, 1903. The research based on this visit culminated in an important publication (Henriques 1917). During his stay in the archipelago, Henriques was received by the owners of many farms (roças), taking notes and obtaining important data on the natural history of the island. Fernandes (1980, 1986) noted that when Henriques planned the study of the flora of São Tomé and Príncipe in the 1880s, he implemented a set of measures that proved to be extremely important for the enrichment of the University of Coimbra herbarium (currently, Herbário do Instituto Botânico de Coimbra, Coimbra, Portugal - COI). First, he promoted and intensified the development of the Coimbra herbarium and trained qualified botany specialists. It was to this end that he appointed Adolpho Frederico Möller (1842–1920), a renowned collector of flora in Portugal and later in São Tomé and Príncipe (four months in 1885), and in turn Francisco Joaquim Dias Quintas (1864–1909) in botanical field studies in São Tomé and Príncipe. Second, Henriques sent material collected on these expeditions to the greatest bryologists at the time. Their results were published in the “Boletim da Sociedade Broteriana”, a scientific journal dedicated to Botany and co-published by the University of Coimbra and the Broterian Society (Sociedade Broteriana). 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 219 Herbarium Specimens and Associated Documentation Numerous naturalists passing through or purposefully visiting São Tomé and Príncipe gathered biological collections, some of which were then published in different scientific outlets. The Austrian naturalist and explorer Friedrich Welwitsch visited São Tomé in 1853 and 1860, during stopovers at the beginning and end of his botanical survey of Angola (Sérgio and Garcia 2011). The major scientific expeditions during the nineteenth century were performed by Adolpho Möller in 1885 (Henriques 1917; Sérgio and Garcia 2011), Francisco Quintas from 1888 to 1889, and Francisco Newton (1864–1909) between 1885 and 1895 (Sérgio and Garcia 2011). Casual collections were made by different naturalists, for example, the French botanist Auguste Jean Baptiste Chevalier (1873–1956) in 1905 (Exell 1944). In all these field studies, botanical collections were organized, and duplicates were distributed to different herbaria. These included COI, and the herbaria of the Museu Nacional de História Natural e da Ciência da Universidade de Lisboa, Lisbon, Portugal (LISU), of the Natural History Museum, London, United Kingdom (BM), of the Muséum National d’Histoire Naturelle de Paris, France (PC), of the Instituto de Investigação Científica Tropical - ULisboa (LISC), and of the Conservatoire et Jardin botaniques de la Ville de Genève, Geneve Switzerland (G), and the Brotherus Herbarium (H-BR) of the Finnish Museum of Natural History University of Helsinki, Finland (H) (Herbaria acronyms according to Thiers 2016). The original herbarium collections, details presented in field notes, draft descriptions of species, and the extensive correspondence between the collectors and the specialists that studied the biological material are a valuable source of data and a base for modern studies regarding São Tomé and Príncipe flora. Presently, COI and LISU herbaria hold most of the bryophyte specimens cited in the bibliography for São Tomé and Príncipe, corresponding mostly to the collections of Friedrich Welwitsch, Möller, Quintas and Newton, in addition to the smaller collections of Júlio Henriques. However, duplicates of these collections are also found in other European herbaria (Sérgio and Garcia 2011). The bryological collections resulting from the expeditions organized by Júlio Henriques were studied by several experts. The liverworts were sent to Franz Stephani (1842–1927) between 1886 and 1913. The mosses were first sent to Carl Müller (1818–1889) in Halle (between 1885 and 1887) and later to Viktor Ferdinand Brotherus (1849–1929) in Helsinki (between 1889 and 1904). The letters that Henriques sent to Stephani and Brotherus (Biblioteca Digital de Botânica da Universidade de Coimbra 2021) list all the specimens exchanged. Thus, specimens originating from these collections were progressively divvied up and disseminated by several international herbaria at the discretion of the authors who studied them. In this study, by cross-checking the LISU database referring to the aforementioned studied herbaria, we were able to confirm where the reference material and most of the respective nomenclatural types are currently located (Sérgio and Garcia 2011). Additionally, we also gathered and compared all available information on 220 C. Garcia et al. Fig. 9.1 Two specimens (1, 2) of Bryum coronatum Schwägr., corresponding to the same Welwitsch specimen (n 126, Insula de S. Thomé loco called Monte Caffé) collected in December 1860, and (3, 4) Sendtnera mollis Steph. Typus, Slopes of Pico de São Tomé, 1500-2100 m, 1885, Adolpho Möller 23. (1) At BM herbarium with “Inter Angolense” labels, with original iconography; (2) At LISU herbarium without iconography but with a handwritten label by Welwitsch; (3) At COI herbarium with the Möller label; (4) At BM herbarium (BM000745048) of Stephani’s herbarium and handwritten data by the same author where the voucher specimens from São Tomé and Príncipe are currently housed (e.g., Figs. 9.1 and 9.2). Most of the material of each specimen in the COI herbarium is abundant and generally corresponds to isotypes found also in BM, G, H or PC. The labels in COI are generally not the original and must have been written by Möller or Quintas (Fig. 9.2), who organized the collections, with many duplicates sent to other herbaria. Arthur Wallis Exell (1901–1993) first landed on São Tomé Island in October 1932 to initiate a botanical expedition of the islands of the Gulf of Guinea (e.g., Fig. 9.3.1–2). He visited the four principal islands (São Tomé, Príncipe, Bioko 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 221 Fig. 9.2 Brotherus specimen identifications. Two specimens of Ectropothecium drepanophyllum corresponding to the same specimen from Quintas, n 23: (1) Holotypus in H herbarium (H-BR1415–022) with Brotherus handwriting; (2) Isotypus in COI herbarium with a handwritten label from Quintas and Annobón), and the results of this expedition were published in 1944, in the Catalogue of the Vascular Plants of São Tomé (Exell 1944). There are also numerous other specimens with labels based on the printed text of the work published by Carl Müller (1886a), such as the mosses collected by Adolpho Frederico Möller in 1885 and published in the Boletim da Sociedade Broteriana. We also verified the presence of specimens with original handwritten labels by Stephani and Brotherus in COI herbarium collections (Fig. 9.3.3–4), although some labels have two handwritings with the numbering of localities corresponding to Adolpho Möller manuscripts and the identifications handwritten by Stephani (Figs. 9.3.3–4 and 9.4) or Carl Müller. The historical specimens from São Tomé and Príncipe archived in the Stephani collection in the herbarium of Genève (G) (Geissler 1982) have duplicates at COI. However, some taxa collected by Newton and Quintas were not returned to Coimbra, at least those studied after 1900. These correspond to the specimen references indicated in the most recent volumes of the Index Hepaticarum (Stephani 1901–1906, 1905–1909, 1909–1912, 1912–1917, 1917–1925). Likewise, there are a considerable number of specimens collected during Júlio Henriques career in the herbarium of Paris (PC), either included in the collections of Jules Cardot (1860–1934), Ferdinand Renauld (1837–1910) or Robert Potier de La Varde (1878–1961), that are often cited in revisionary studies of bryophyte genera. 222 C. Garcia et al. Fig. 9.3 Arthur Exell (1, 2) and Francisco Quintas specimens (3, 4). (1) Octoblepharum albidum, Esperança, circa 350 ft., 1932, Exell 675 in the herbarium of Coimbra (COI); (2) Hygrolejeunea pulcherrima, Santa Maria, circa 4200 ft., 1932, Exell 197 in the London (BM) collections; (3) Isotypus of Metzgeria thomeensis (BM); (4) The same material in COI, isotypes. Both labels (3, 4) correspond to Stephani’s manuscripts At the Helsinki herbarium (H), a significant part of material from São Tomé was found in the Brotherus (H–BR) collections, particularly type specimens collected in this archipelago (Sérgio and Garcia 2011), corresponding almost exclusively to Quintas and Möller collections. However, some of the specimens originally studied by Brotherus are now at BM, PC or COI herbaria and were not found at H herbarium (e.g., Leucobryum homalophyllum Broth.). It should be noted that, contrary to Stephani, Brotherus returned all the material he studied to Coimbra, mostly with handwritten labels. 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 223 Fig. 9.4 (1) Original description of Lejeunea ramosissima Steph. included in the archives of the Botanical Institute of Coimbra Library as part of the letter dated 25 February 1886, and (2) Iconography of several species, including L. ramosissima Steph. (33 and 34) and Sendtneria mollis Steph. (22 and 23) by Stephani (1886) Historical Correspondence The correspondence between naturalists who studied bryophyte specimens collected by Möller, Quintas and Newton, is largely housed in the archives of the University of Coimbra at present. The documentation referring to Welwitsch’s correspondence, currently in Lisbon at the Museu Nacional de História Natural e da Ciência (MUHNAC) of the University of Lisbon, does not provide any mention of bryophytes, despite the existence of bryophyte herbarium specimens of Welwitsch’s expeditions in LISU herbarium (MUHNAC). Among this documentation are lists of the identified species and, in some cases, the original descriptions of species are also included. Although it was not possible to study all Júlio Henriques’ correspondence in some foreign institutions (except for the one in the library of the University of Helsinki and in the Botanical Garden of Geneva), we analyzed the correspondence exchanged between him and several specialists that is filed at the University of Coimbra (Biblioteca Digital de Botânica da Universidade de Coimbra). Some important parts of this correspondence are transcribed below. 224 C. Garcia et al. Correspondence from Júlio Henriques to Franz Stephani Henriques’ first letter referring sending material to Stephani, was on 8 January 1885. The first publication of Stephani concerning the liverworts of São Tomé (Stephani 1886) refers to the A. Moller 1885 collections, so the specimen identifications must have been rather hasty. It describes 19 new species, in addition to other taxa, such as those already described by Mitten from the Cameroon Mountains (Mitten 1863) (e.g., Radula bipinnata Mitt.). Among the first records for São Tomé and Príncipe there was, for example, Lejeunea ramosissima Steph. (Fig. 9.4), Plagiochila integerrima Steph., Sendtnera mollis Steph. and Anthoceros pinnatus Steph., all currently still considered as distinct species with valid names. In the letter of 25 February 1886, Stephani sent along a list referencing the figures with the iconographies that were included in the same publication of Stephani (1886). Correspondence from Júlio Henriques to Carl Müller The correspondence sent by C. Müller from Halle to Henriques is very sparse and consists only of four letters between 1885 and 1887. C. Müller’s second letter, dated 21 March 1886 (UC Digitalis 2021), is the most important as it includes a list of about 50 taxa, corresponding to the identification of the specimens cited in Müller (1886a, b). These specimens include more than 25 new species, some of them still recognized as species, as in the case of Funaria acicularis Müll.Hal and Leucobryum leucophanoides Müll.Hal. Correspondence from Júlio Henriques to Viktor Ferdinand Brotherus Sérgio and Garcia (2011) analyzed much of the correspondence between Henriques and Brotherus. Henriques’ first letter dated 31 January 1889, refers to sending (on 24 January 1889) a package including mosses from São Tomé to Copenhagen and then to Helsinki. The exchange of bryological material continued and Henriques must have sent a second package that also included material from Portugal. In Brotherus’s letter to Coimbra, sent on 19 August 1889, he states “J’ai reçu en bon état, il ya quelque jours, la quaisse avec des mousses du Portugal et j’irais à leurs déterminations et révision aussitôt qu’il me sera possible. Les mousses de l’île S. Thomé de votre second envoi j’ai déjà examinés et vous communique le nom des espèces. Sont-elles aussi recueillies par M. Quintas”? [“I received in good condition, a few days ago, the case with mosses from Portugal and I will make their identifications and revision as soon as possible. The mosses of the island S. Thomé of your second shipment I have already examined and communicated to 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 225 you the names of the species. Were they also collected by Mr. Quintas”?] (UC Digitalis 2021).The reprints referenced in these letters correspond to the 1890 article, published in the “Boletim da Sociedade Broteriana” (Brotherus 1890), where 29 new species are described, some still considered valid species, including several endemics, such as Leucobryum homalophyllum Broth. and Ectropothecium drepanophyllum Broth (Fig. 9.2). Correspondence from Júlio Henriques to Francisco Quintas Although there are no extant records of correspondence from Möller to Henriques, a set of letters from Quintas sent from São Tomé to Henriques in Coimbra are still extant (Biblioteca Digital de Botânica da Universidade de Coimbra 2021). From most of this correspondence, it is evident that Quintas kept Henriques apprised of his research and situation on the island, attaching lists of the material he sent to Coimbra. Although there is no specific numbering for the bryophyte specimens, there were indications of the boxes that contained cryptogams. For instance, the attachment to the letter issued on 21 July 1888 (UC Digitalis 2021). In that same letter, he also mentioned that mushroom specimens were listed separately. There are many bryophyte specimens collected by Quintas, which correspond to about 70 different bryophyte taxa, some of them corresponding to new species, such as Plagiochila flabellata Steph., P. amplifolia Steph. Among the liverworts and numerous species of mosses (ca. 31) described as new by Brotherus in 1890, we have the examples of Pilotrichella calomicra Broth., Porotrichum quintasii Broth., P. caudatum Broth., Trichosteleum dicranelloides Broth., among other new taxa. Correspondence from Francisco Newton to Júlio Henriques Some correspondence between Henriques and Newton is available in the historical archives of MUHNAC, but none of these letters has any reference to bryophytes. In Coimbra University, there is a letter sent by Newton to Henriques about his 1885 upcoming mission to Africa. Based on this letter, dated 23 August 1885, Newton makes Henriques aware of certain material from Angola (UC Digitalis 2021). He also confirms in that letter that he proposed to make a stop at Príncipe Island and then São Tomé. Ultimately, Newton arrived in São Tomé on 24 September 1885 (Guedes 2021). Strangely, most of the bryophyte material collected by Newton and found in the different herbaria (BM, FH, G, JE and M herbaria) corresponds to specimens collected in 1887 on Príncipe and only a few are indicated to be from São Tomé. In fact, in the different publications concerning Newton, specimens correspond to Príncipe Island, excluding two references corresponding to São Tomé, in Angolares (Stephani 1888a, b). It is also interesting to note that the bryophyte specimens 226 C. Garcia et al. collected on Príncipe Island were only shipped in September of 1885 (Newton 1885), after this letter, but Newton should have sent more material later. Apparently, Newton did not organize the numbering of his bryophyte specimens and the labels are very uninformative. The numbering of specimens was made when the plants arrived in Coimbra. However, it should be noted that based on Newton’s collections, some liverwort species were described by Stephani (1888a, b), as Microlejeunea africana Steph., Lejeunea newtonii Steph. (now included in Cheilolejeunea newtonii Steph. ex Schiffn.), Plagiochila thomeensis Steph. (currently a synonym for Plagiochila terebrans Nees et Mont. ex Lindenb); Cheilolejeunea principensis Steph. (synonymized to Cheilolejeunea serpentina (Mitt.) Mizut.), and Lophocolea newtonii Steph. (synonymized to Lophocolea martiana Nees). Historical Collecting Localities Based on the data associated with the aforementioned collections, the location (exact or approximate) of the historical collecting activities by the first naturalists dedicated to the study of bryophytes in São Tomé Island was georeferenced (Fig. 9.5), and used as a starting point for the most recent fieldwork performed by the authors. Exell’s 1944 plant catalogue (Exell 1944) includes all the species known at the time and new reports of some taxa for the islands (Figueiredo 1994, 2005; Figueiredo and Gascoigne 2001), including diverse bryophytes. He was based in Vanhulst (Macambrará), in the Roça Zampalma and collected most of the bryophyte material in this region. This bryophyte collection was the basis for two publications, in which about 40 taxa of liverworts and mosses are listed (Exell 1944). Most of Exell’s specimens are stored in BM, except for some specimens that are kept at COI. After the Exell expedition, other collections were obtained in 1956 by the French naturalist Théodore Monod (1902–2000) and C. A. Thorold (1906–1998) in São Tomé and Príncipe, mainly in Pico de Príncipe, during the “6th Conférence lnternationale des Africanistes de I’Ouest” (Monod 1960). Most of this material is hosted in PC and was the basis of the publication of Potier de la Varde (1959). Arnaldo Roseira also collected in the islands between 1954 and 1958, corresponding to 79 specimens of three taxa in the PO herbarium (Universidade do Porto) (Costa 2020). Recent Studies Since the middle of the twentieth century and after the works of Exell, the study of bryophytes of São Tomé and Príncipe came to a halt. Only more recently has a new effort emerged through the project Bryotome (Sérgio and Garcia 2011). During this project, the first author carried out fieldwork in São Tomé and Príncipe in 2007 and 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 227 Fig. 9.5 Historical collecting localities of naturalists who collected bryophytes specimens on São Tomé Island from the mid-nineteenth century to mid-twentieth century: (1) Auguste Jean Baptiste Chevalier (1873–1956), (2) Arthur Wallis Exell (1901–1993), (3) Gustav Mann (1836–1916), (4) Adolpho Frederico Möller (1842–1920), (5) Théodore Monod (1902–2000), (6) Francisco Newton (1864–1909), (7) Francisco Joaquim Dias Quintas (1864–1909), (8) Charles Aubrey Thorold (1906–1998), (9) Friedrich Welwitsch (1806–1872) 228 C. Garcia et al. Fig. 9.6 Study localities in 2007, 2008, 2010, 2013 and 2016 for the enrichment of the University of Lisbon (LISU) and the California Academy of Science (CAS) herbaria 2008, collecting about 6000 specimens at various altitudes, including Pico de São Tomé. This project enabled the study of herbarium specimens, georeferencing historical specimens, planning new fieldwork more effectively, and studying different substrates (epiphytic, epiphyllous, rupicolous, terricolous and humicolous) to better determine specific microhabitats of species across the islands. In 2010, 2013 and 2016, expeditions sponsored by the California Academy of Sciences were carried out (Fig. 9.6). During these new expeditions, several species were discovered, including new records for the archipelago and the African continent, as well as species new to science (Figs. 9.7 and 9.8). One of the new species discovered was particularly interesting: Dendroceros paivae is distinct from most other species of the genus in its ecology, gametophyte, and sporophyte characters, resembling only the Bornean D. foliicola J. Haseg. In comparison to the type material of D. foliicola, D. paivae has a narrower sporophyte diameter. The thallus of D. paivae does not form rosettes, the cuticle is weakly papillose, and apices are plane to undulate, while D. foliicola forms rosette-like patches with strong crispate margins, even at branches apices, and the cuticle is slightly papillose (Garcia et al. 2012). 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 229 Fig. 9.7 Liverworts and hornworts from São Tomé and Príncipe: (1, 2) Dendroceros paivae C.A. Garcia, Sérgio & J. C. Villarreal. (hornworts) at the type locality (LISU 237201) (Garcia et al. 2012); (3, 4) Megaceros flagellaris (Mitt,) Steph. (hornworts) growing on a tree trunk in the first known locality in São Tomé and Príncipe and the second one reported for the African Continent (LISU 237200); (5) Anthoceros pinnatus Steph. (hornworts); (6) Phaeoceros carolinianus (Michx.) Prosk. (hornworts); (7) Colura sp. (liverworts); (8) Cyathodium cavernarum Kunze (liverworts) 230 C. Garcia et al. Fig. 9.8 Liverworts and mosses from São Tomé and Príncipe. (1) Marchantia pappeana Lehm. subsp. pappeana (liverworts). (2), Plicanthus hirtellus (F. Weber) R.M. Schust. (liverworts). (3), Calymperes lonchophyllum Schwägr. (mosses). (4), Octoblepharum albidum Hedw. (mosses). (5), Orthostichella sp. (mosses). (6), Trematodon longicollis Michx. (mosses). (7), Macromitrium sulcatum var. sulcatum (Hook.) Brid. (mosses). (8), Calymperes palisotii Schwägr. (mosses) 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 231 Diversity, Composition and Endemism In recent years, several papers resulting from these last expeditions have been published. These included the study of secondary metabolism compounds (Figueiredo et al. 2010) and the description of new species and new genera of bryophytes (Enroth and Shevock 2011; Müller et al. 2011; Garcia and Sérgio 2012a, b, c, d, e; Shevock et al. 2013; Pócs et al. 2015; Sollman et al. 2016; Enroth and Shevock 2017a, b; Müller and Shevock 2018; Müller et al. 2019). These works allowed revising the number of species and endemics for each island individually and for the two islands combined (Table 9.1 and Appendix). One hundred and forty-four species of bryophytes occurring on the islands are currently considered endemic to Africa, 21 of which are endemic to the archipelago, including seven liverworts or hornworts and 14 mosses. One of these species is the hornwort Dendroceros paivae C.A. Garcia, Sérgio & J. C. Villarreal. (hornworts), endemic to São Tomé Island (Garcia et al. 2012) and found only in a single location, in a very restricted area. The known bryophyte species diversity of the islands has increased markedly as specimens of various families are critically examined. For example, prior to fieldwork by the authors, only three species of the moss genus Fissidens (Fissidentaceae) were reported (O’Shea 2006). Now, Fissidens Hedw. is the most species-rich bryophyte genus in the archipelago with 24 known species (Shevock et al. 2013). The liverwort genera, Lejeunea Lib. and Plagiochila (Dumort.) Dumort. are also quite diverse with 19 and 18 species respectively (Müller et al. 2011; Pócs et al. 2015). A similar story of species additions for the islands was provided in a recent study of the moss families Neckeraceae (Enroth and Shevock 2011, 2017a, b) and Table 9.1 Bryophyte species diversity and endemism for each island individually and for the two islands combined Príncipe São Tomé P&ST MARCHANTIOPHYTA and ANTHOCEROTOPHYTA Liverworts and hornworts Total species/taxa 108 138 171 Island endemism 4 4 7 African endemism 52 65 80 DIVISION BRYOPHYTA Mosses Total species/taxa 41 114 133 Island endemism 1 13 14 African endemism 16 59 64 TOTAL BRYOPHYTES Liverworts, hornworts and mosses Total species/taxa 149 252 304 Island endemism 5 17 21 African endemism 68 124 144 Liverworts and hornworts, according to Wigginton (2018), and mosses, according to O’Shea (2006) 232 C. Garcia et al. Pottiaceae (Sollman et al. 2016). Ongoing work on the moss family Calymperaceae also has discovered several new species for these islands. We anticipate species additions for these islands will continue for many years to come. Most studies were carried out on existing trails to reach higher areas, and several regions have not yet been surveyed (Figs. 9.5 and 9.6) due to the difficult terrain. Additional species will likely be documented and discovered as more remote cloud forest environments can be systematically surveyed. Our updated summary for the bryoflora of the islands reveals the documented diversity has increased significantly since the last reports of the mosses (O’Shea 2006) and of the liverworts and hornworts (Wigginton 2018). Thus, the 304 bryophytes documented for the archipelago at this time are likely a vast underestimate of the true diversity (Appendix). The species catalogue of the bryophyte flora of the islands of São Tomé and Príncipe presented in this work (Appendix) is based on all known published literature. All the literature on bryophytes of São Tomé and Príncipe Islands was surveyed, including liverworts, hornworts, and mosses. The delimitation of families follows the latest version of the Checklist of sub-Saharan Africa of Wigginton (2018) for liverworts and hornworts and O’Shea (2006) for mosses. Taxa are presented in alphabetical order of all the accepted names (including subspecies and varieties). Taxa with synonymies (homotypic and heterotypic synonyms) whose type locality corresponds to São Tomé and Príncipe are designated in a second column with respective authors and the year of publication for São Tomé and Príncipe. The most accurate information about the original description, as well as the relevant synonyms, were considered with general taxonomic criteria. The present table includes only records published up until June 2020 (unpublished data of the authors, including new species and localities, are not included). The catalogue is not a taxonomic document, and no new taxonomic nor nomenclatural acts are published here. Synopses of families and genera are placed alphabetically within each order. Final Remarks The known bryoflora of São Tomé and Príncipe includes at least 304 species. Based on our ongoing studies and the number of specimens still awaiting critical study, this number will increase in the coming years as the diversity of this group becomes more comprehensively documented. The apparently low number of known species may be explained by the logistical and practical difficulties of carrying out fieldwork in dense forests and in areas of rough terrain, a reality that affects most of the scientists working in the region. Bryophytes are also generally very small plants, and many species occur in small populations. Therefore, during fieldwork some species can easily be overlooked or may occupy exceedingly specialized microhabitats that are difficult to find (e.g., fine twigs in the tree canopy). Further difficulty originates from the fact that the taxonomy of different bryophyte families is not well developed in the tropics, and world experts in bryophyte taxonomy are also starting to become 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . 233 scarce, especially those dedicated to the study of tropical species. Many bryophyte species reported from Africa are known only from type specimens or based on a handful of collections. Almost 45 bryophyte taxa reported for São Tomé and Príncipe have not been resampled since the nineteenth century. For a large number of bryophyte species, the ecology and habitat specificity are not well known or not known at all, and also the distribution patterns, elevational range, and abundance for most species remain to be determined. The threats affecting forest habitats in São Tomé and Príncipe, such as habitat destruction or competition by invasive species, may affect the survival of bryophytes. One major threat to biodiversity conservation in the archipelago, particularly to the cryptogamic communities, forest structure, and habitat diversity along the altitudinal gradients is deforestation, especially that associated with the plantations of oil palm Elaeis guineesis Jacq. In the Emolve region (southern region of São Tomé), there is a monoculture of more than 600 ha of oil palm that is expected to continue growing, which would result in a significant loss in biodiversity (bryophytes and other taxonomic groups), especially to forests at lower elevations. By contrast, ancestral roças (old colonial farms), with Cofeea spp. and Theobroma cacao L. plantations, seem to preserve a high diversity of bryophyte species, mainly epiphytic taxa. New bryological studies are urgently needed, especially in areas that have never been surveyed. Together with the recently collected material that is currently being studied by the coauthors and other colleagues, these new surveys will continue to increase our knowledge of distributions, species diversity and the particularities of bryophyte endemism in the country. A more comprehensive and updated species list will be essential to inform a future IUCN Red List assessment of bryophytes from São Tomé and Príncipe and to designate priority areas for conservation. Acknowledgments The present study was partly supported by “Fundação para a Ciência e a Tecnologia” (POCTI/AFR/58699/2004 and SFRH/BPD/22304/2005) and by California Academy of Sciences Gulf of Guinea Funds (the expedition of 2016 performed by the first author and all those carried out by the last author). We are grateful to the curators of the following herbaria for kindly allowing us to study in situ the plant material and for the loan of specimens, including types: BM, G, H; PC, NICH and VIT. The authors are grateful to the são-tomenses, Eng. Salvador Pontes, Aurélio Espirito-Santo ({), Francisco Alamô, Estevão Soares, Mr. Lagoas ({), Sátiro Raúl José da Costa, Ostelino Conceição Rocha (Balú) e Júlio da Conceição Rocha for their help during the fieldwork. Thanks also to expedition photographer Andrew Stanbridge. 234 C. Garcia et al. Appendix Updated catalogue of bryophytes from São Tomé and Príncipe Islands. A: African endemics. E: Island endemism. Synonyms (basionyms) based in São Tomé and Príncipe collections Taxon species/subspecies/variety DIVISIONS Marchantiophyta and Anthocerotophyta Liverworts and Hornworts Acrolejeunea emergens (Mitt.) Steph. var. emergens Aneura pinguis (L.) Dumort. sens. lat. Aneura latissima Spruce Anthoceros pinnatus Steph. Bazzania decrescens subsp. molleri (Steph.) E.W.Jones Bazzania nitida (F.Weber) Grolle Brachiolejeunea laxifolia (Taylor) Schiffn. Calypogeia fissa (L.) Raddi Calypogeia peruviana Nees et Mont. Caudalejeunea africana (Steph.) Schiffn. Caudalejeunea dusenii Steph. Caudalejeunea hanningtonii (Mitt.) Schiffn. Caudalejeunea lehmanniana (Gottsche) A.Evans Ceratolejeunea cornuta (Lindenb.) Steph. Ceratolejeunea floribunda Steph. Cheilolejeunea intertexta (Lindenb.) Steph. Cheilolejeunea montagnei (Gottsche) R.M.Schust. Cheilolejeunea rigidula (Nees ex Mont.) R.M.Schust. Cheilolejeunea surrepens (Mitt.) E. W.Jones Anthoceros pinnatus Steph. 1886 Mastigobryum molleri Steph. 1886 P ST X X X X X X X A X X X 2015/ 2011 1956/ 1886 1888/ 2011 1886 2015/ 1886 2011 1912 X X 1970 1976 X X 2011 A A A 2015 2000/ 2000 2015 X E 1960/ 2015 1913 X 2015/ 1893 1863 Brachiolejeunea thomeensis Steph. 1912 X X Ceratolejeunea floribunda Steph. 2013 1913 Cheilolejeunea newtonii Steph. ex Schiffn. 1893 Euosmolejeunea thomeensis Steph. 1914 Cheilolejeunea principensis Steph. ex Paris 1888 Year of first ref. P/ST X X 1888 X 2015 (continued) 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . Taxon species/subspecies/variety Cheilolejeunea trifaria (Reinw. et al.) Mizut. Chiloscyphus difformis (Nees) J.J. Engel et R.M.Schust. Cololejeunea africana (Steph.) R. M.Schust. Cololejeunea cuneifolia Steph. Cololejeunea iradieri Infante et Heras Cololejeunea lanceolata E.W.Jones Cololejeunea leloutrei (E.W.Jones) R.M.Schust. Cololejeunea mocambiquensis S. W.Arnell Cololejeunea obliqua (Nees et Mont.) Schiffn. Cololejeunea obtusifolia (E.W. Jones) Tixier Cololejeunea papilliloba Steph. Cololejeunea platyneura (Spruce) A.Evans Cololejeunea pusilla Steph. Cololejeunea zenkeri (Steph.) E.W. Jones Colura calderae Pócs Colura digitalis (Mitt.) Steph. Synonyms (basionyms) based in São Tomé and Príncipe collections Lejeunea grandistipula Steph. 1886 Lophocolea molleri Steph. 1886 Physocolea africana Steph. 1915 X Year of first ref. P/ST 2015/ 1886 1886 A 1916 A 2015 2015 A 2015 1960 ST X A A A Cololejeunea crenatiflora Steph. 1891 X 2015 X A X X 2011/ 1891 2015 2015 2015 A A A A A A A X A A X E E Lophocolea devexa Mitt. 1863 X X Lophocolea newtonii Steph. 1907 X X 2015 2000/ 2015 2011 1958/ 1953 2015 2015 2015/ 1958 2015/ 2011 2015/ 1863 1953 X X A E 1952 1863 2010 2012 Colura hattoriana Pócs Colura obesa Jovet-Ast Colura tenuicornis (A.Evans) Steph. Colura thomeensis Pócs Conoscyphus trapezioides (Sande Lac.) Schiffn. Cryptolophocolea martiana (Nees) L.Söderstr., Crand.-Stotl. et Stotler subsp. martiana Cyathodium cavernarum Kunze Dendroceros crispatus Nees Dendroceros herasii M.Infante Dendroceros paivae C.Garcia, Sérgio & J.C. Villarreal P X 235 (continued) 236 Taxon species/subspecies/variety Dibrachiella africana (Steph.) X.Q. Shi, R.L.Zhu et Gradst. Dibrachiella autoica (Vanden Berghen) X.Q.Shi, R.L.Zhu et Gradst. Diplasiolejeunea cavifolia Steph. Drepanolejeunea capulata (Taylor) Steph. Drepanolejeunea cultrella (Mitt.) Steph. Drepanolejeunea physifolia (Gottsche) Pearson Dumortiera hirsuta (Sw.) Nees Folioceros incurvus (Steph.) D.C. Bharadwaj Fossombronia indica Steph. Fossombronia sp. Frullania angulata Mitt. var. angulata Frullania apicalis Mitt. Frullania apiculata (Reinw. et al.) Nees Frullania caffraria Steph. C. Garcia et al. Synonyms (basionyms) based in São Tomé and Príncipe collections Frullania purpurea Steph. Frullania rio-janeirensis (Raddi) Ångstr. Frullania serrata Gottsche var. serrata Frullania spongiosa Steph. Fuscocephaloziopsis connivens subsp. fissa (Steph.) Váňa et L. Soderstr. Herbertus dicranus (Taylor ex Gottsche, Lindenb. et Nees) Trevis. ST A Lejeunea cavifolia Steph. 1886 Drepanolejeunea molleri Steph. 1913 Prionolejeunea fissistipula Steph. 1913 2015 X A 1886 2011 A A A A X X A A 2015/ 1913 1960/ 1913 1960/ 1886 1888/ 1889 2019 2011 1863 X X Frullania angulata Mitt. 1863; F. subatrata Steph. 1911; F. cordifolia Steph. 1911 Frullania laceriloba Steph. 1911 A A X 2015/ 1911 2011 X 1894 X A X X X X X X X 1886 1886/ 1863 1910 2015/ 2004 1976 1976/ 1891 1886 Frullania molleri Steph. 1894 (Probably synonym) Frullania diptera (Lehm.) Drège Frullania ericoides (Nees) Mont. Frullania obscura (Sw.) Mont. Frullania obscurifolia Mitt. P A Frullania thomeensis Steph. 1910 Frullania africana Steph. 1891 A X Sendtnera mollis Steph. 1886 Year of first ref. P/ST 2000 X X X X X X 2011/ 2011 1988/ 1988 2011/ 1886 (continued) 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . Taxon species/subspecies/variety Heteroscyphus dubius (Gottsche) Schiffn. Heteroscyphus spectabilis (Steph.) Schiffn. Synonyms (basionyms) based in São Tomé and Príncipe collections 237 Year of first ref. P/ST 1888 P X ST A A 2015/ 1886 Lejeunea abyssinica (Gola) Cufod. A A Lejeunea acuta Mitt. A A Lejeunea anisophylla Mont. X X 2015/ 2015 2019/ 1960 2011/ 2011 2015 2011 2011 2015 1888/ 1960 1896 Isotachis perfoliata Steph. 1886 Chiloscyphus thomeensis Steph. 1893 nom. nud.? Lejeunea brenanii E.W.Jones Lejeunea cf. obtusata Gottsche Lejeunea conformis Nees et Mont. Lejeunea eckloniana Lindenb. Lejeunea flava (Sw.) Nees Lejeunea grossecristata (Steph.) E. W.Jones Lejeunea helenae Pearson Lejeunea ibadana A.J.Harr. et E.W. Jones Lejeunea jungneri (Steph.) Steph. Lejeunea lyratiflora Steph. Lejeunea papilionacea Prantl Lejeunea phyllobola Nees et Mont. Lejeunea pulchriflora (Pearson) G.E. Lee, Bechteler, Pócs, SchäfVerw. & Heinrichs Lejeunea ramosissima Steph. A X A X X X A Hygrolejeunea grossecristata Steph. 1896; Taxilejeunea longirostris Steph. 1914 X A A A A X X X Lejeunea ramosissima Steph. 1886 X X Lejeunea setacea (Steph.) Steph. A A Lejeunea tuberculosa Steph. Lepidozia succida Mitt. X A A Lepidozia ubangiensis Steph. Leptolejeunea astroidea (Mitt.) Steph. Leptolejeunea epiphylla (Mitt.) Steph. Leptolejeunea maculata (Mitt.) Schiffn. A A A X X X X Leptolejeunea quintasii Steph. 1891 Lejeunea thomeensis Steph. 1886; Drepanolejeunea gomphiae Steph. 1913 2015 2015/ 2015 1901 2015 2011 2011 2015 1996/ 1886 1969/ 1969 2011 2011/ 1891 2015 2015 2015/ 1891 2015/ 1886 (continued) 238 Taxon species/subspecies/variety Lopholejeunea nigricans (Lindenb.) Schiffn. Lopholejeunea subfusca (Nees) Schiffn. Marchantia debilis Goebel Marchantia pappeana Lehm. subsp. pappeana Marchesinia excavata (Mitt.) Schiffn. Marchesinia principensis Frank Müll. et Shevock Mastigophora diclados (Brid. ex F. Weber) Nees Megaceros flagellaris (Mitt.) Steph. Metalejeunea cucullata (Reinw. et al.) Grolle Metzgeria furcata (L.) Dumort. Metzgeria leptoneura Spruce Metzgeria lindbergii Schuffn. Microlejeunea africana Steph. Microlejeunea ankasica E.W. Jones Microlejeunea kamerunensis Steph. Neurolejeunea breutelii (Gottsche) A.Evans var. africana Pócs Notoscyphus lutescens (Lehm. et Lindenb.) Mitt. Odontolejeunea lunulata (F.Weber) Schiffn. Pallavicinia lyellii (Hook.) Carruth. Phaeoceros carolinianus (Michx.) Prosk. Plagiochila barteri Mitt. Plagiochila barteri var. valida (Steph.) Vanden Berghen Plagiochila brunneola Steph. Plagiochila divergens var. capensis (Steph.) E.W. Jones Plagiochila flabellata Steph. Plagiochila fusifera Taylor C. Garcia et al. Synonyms (basionyms) based in São Tomé and Príncipe collections A A Year of first ref. P/ST 2011/ 2011 2015/ 2011 2011 1886 A 1886 P X ST X X X Marchantia planiloba Steph. 1886 Homalolejeunea henriquesii Steph. 1888 A X 2018 X X X Metzgeria thomeensis Steph. 1891 X X Metzgeria recurva Steph. 1886 X X X A A A A A A Microlejeunea africana Steph. 1888 Microlejeunea cochlarifolia Steph 1888 (probably synonymy) E Odontolejeunea thomeensis Steph. 1912 Pallavicinia pilifera Steph. 1891 Plagiochila triangularis Steph. 1886; P. quintasii Steph. 1904 Plagiochila flabellata Steph. 1886; P. molleri Steph. 1886 Plagiochila amplifolia Steph. 1901 2015 X 2004/ 1912 1891 2011 A A A A A X A 2004/ 1891 2004/ 1886 2015 1888/ 1891 2015 1990/ 1888 2015 X X X Plagiochila brunneola Steph. 1904 2015/ 1886 2012 2015 A X 1962/ 1886 1981/ 1981 1904 1962 2011/ 1886 1901 (continued) 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . Taxon species/subspecies/variety Plagiochila gibbiflora Steph. Plagiochila heterostipa Steph. Plagiochila integerrima Steph. Synonyms (basionyms) based in São Tomé and Príncipe collections Plagiochila gibbiflora Steph. 1904 P 239 ST E A X X A A Plagiochila moenkemeyeri Steph. A A Plagiochila neckeroidea Mitt. A A Plagiochila pectinata Willd. ex Lindenb. Plagiochila pinniflora Steph. Plagiochila praemorsa Steph. A A X A X A A A A Plagiochila loloensis Steph. Plagiochila integerrima Steph. 1886 Plagiochila rotundifolia Steph. 1904 Plagiochila cacuminis Steph. 1918 Plagiochila sarmentosa (Lehm. et Lindenb.) Lindenb. Plagiochila strictifolia Steph. Requires confirmation Plagiochila terebrans Nees et Mont. ex Lindenb. Pleurozia gigantea (F.Weber) Lindb. Plicanthus hirtellus (F.Weber) R.M. Schust. Porella abyssinica var. hoehnelii (Steph.) Pócs. Porella subdentata (Mitt.) E.W. Jones var. subdentata Porella subdentata var. camerunensis E.W.Jones Prionolejeunea grata (Gottsche) Schiffn. Prionolejeunea principensis Vanden Berghen Radula ankefinensis Gottsche ex Steph. Radula appressa Mitt. Plagiochila thomeensis Steph. 1886 Radula boryana (F.Weber) Mont. Radula flaccida Lindenb. et Gottsche Madotheca thomeensis Steph. 1910 A X 1962/ 1962 2011/ 1886 1863 X 1886 A 2011 A 1963/ 1910 2011 A X X 1960/ 1960 1960 A A A A X X X X 2011/ 2015 1910/ 1886 1996/ 1863 1939/ 1939 E Radula angustata Steph. 1886; R. molleri Steph. 1910 Radula tamariscina Mitt. 1863; R. bipinnata Mitt. 1863 Year of first ref. P/ST 1904 2015 2011/ 1886 2011/ 1904 2011/ 1960 2011/ 1904 2011/ 1960 2011 1888/ 1918 (continued) 240 Taxon species/subspecies/variety Radula fulvifolia (Hook.f. et Taylor) Gottsche et al. Radula stenocalyx Mont. Riccardia amazonica (Spruce) Schiffn. ex Gradst. et Hekking Riccardia erosa (Steph.) E.W.Jones Riccardia limbata (Steph.) E.W. Jones Riccardia longispica (Steph.) Pearson Riccia congoana Steph. Riccia discolor Lehm. et Lindenb. Riccia lanceolata Steph. Riccia moenkemeyeri Steph. Riccia stricta (Lindenb.) Perold Schiffneriolejeunea occulta (Steph.) Gradst. Schiffneriolejeunea pappeana (Nees) Gradst. var. pappeana Schiffneriolejeunea polycarpa (Nees) Gradst. Solenostoma borgenii (Gottsche ex Pearson) Steph. Solenostoma dusenii (Steph.) Váňa, Hentschel et Heinrichs. Spruceanthus abbreviatus (Mitt.) X. Q.Shi, R.L.Zhu et Gradst. Spruceanthus floreus (Mitt.) Sukkharak et Gradst. Stictolejeunea balfourii (Mitt.) E. W.Jones Symphyogyna podophylla (Thunb.) Mont. et Nees Syzygiella manca (Mont.) Steph. Telaranea coactilis (Spruce) J.J. Engel et G.L.Merr. Telaranea nematodes (Gottsche ex Austin) M.Howe Thysananthus auriculatus (Wilson) Sukkharak et Gradst. var. auriculatus C. Garcia et al. Ptychocoleus quintasii Steph. 1912 A Year of first ref. P/ST 2001/ 2011 2015/ 1910 2015/ 2011 2011/ 1891 2011/ 1891 2015/ 2011 2012 2015 2015 2012 2012 2015/ 2011 1912 Phragmicoma amplectens Steph. 1886 ¼ P. molleri Steph. 1886 X 1886 A 1974 X X 2019/ 1974 2015 A 1891 X 2015 Synonyms (basionyms) based in São Tomé and Príncipe collections P X ST X X X X X Aneura erosa Steph. 1891 A A Aneura reticulata Steph. 1891 A A A A X X A A X Jungermannia geminiflolia Mitt. 1863 Lepidozia quintasii Steph. 1922 X A X A X 2011 X 1863 X 2011 X 1922 X 2011/ 1949 (continued) 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . Taxon species/subspecies/variety Thysananthus humilis (Gottsche) Sukkharak et Gradst. Thysananthus nigrus (Steph.) Sukkharak et Gradst. Thysananthus turgidus (Steph.) Sukkharak et Gradst. DIVISION Bryophyta Mosses Afrothamnium stipitatum (Mitt.) Enroth Anoectangium aestivum (Hedw.) Mitt. Anoectangium stracheyanum Mitt. Barbula cf. seramensis H.Akiyama Brachymenium leptophyllum (Bruch & Schimp. ex Müll.Hal.) Bruch & Schimp. ex A.Jaeger Brachymenium nepalense Hook. Brachymenium subuliferum (Mitt.) A.Jaeger Brachymitrion moritzianum (Müll. Hal.) A.K.Kop. Bryum apiculatum Schwägr. Bryum argenteum Hedw. var. argenteum Bryum coronatum Schwägr. Bryum huillense Welw. & Duby Bryum thomeanum P. de la Varde Caduciella mariei (Besch.) Enroth Callicostella brevipes (Broth.) Broth. Callicostella chionophylla (Müll. Hal.) Broth. Callicostella fissidentella (Besch.) Kindb. Callicostella perpapillata Broth. & P.de la Varde Callicostella salaziae (Besch.) Broth. Calymperes afzelii Sw. Calymperes lonchophyllum subsp. saxatile (Müll. Hal. ex Besch.) S.R. Edwards Synonyms (basionyms) based in São Tomé and Príncipe collections 241 P X ST X Mastigolejeunea nigra Steph. 1891 A X Mastigolejeunea turgida Steph. A X Bryum subuliferum Mitt. 1863; Bryum molleri Müll.Hal. 1886 Orthodon thomeanus Broth. 1890; Tayloria thomeana Broth. 1903 Bryum areoblastum Müll.Hal. 1886 Bryum squarripilum Müll.Hal. 1886 Bryum erythrostegum Müll.Hal. 1886 Bryum quintasii Broth.1890 Bryum thomeanum P.de la Varde 1959 X 1982 X 2016 X X X 2016 2016 1972 X A 1972 1863 X 1890 X 1886 X 1886 X 1886 X E 1890 1959 A 2017 1952 E 1886 A 1890/ 1890 1944 A 1890 X 1863 1944 X Hookeria chionophylla Müll.Hal. 1886 Hookeria thomeana Broth. 1890 A X Hookeria quintasi Broth. 1890 Calymperes quintasi Broth. 1890 A Year of first ref. P/ST 2014/ 1888 2015/ 1891 1983/ 1917 (continued) 242 Taxon species/subspecies/variety Calymperes palisotii Schwägr. Calymperes pintasii Müll.Hal. ex Besch. Calymperes tenerum Müll.Hal. Calyptothecium acutifolium var. breviusculum (Müll.Hal. ex Dusén) Argent Campylopus flexuosus (Hedw.) Brid var. flexuosus. Campylopus savannarum (Müll. Hal.) Mitt. Chionoloma bombayense (Müll. Hal.) P. Sollman Cyclodictyon filicuspis P.de la Varde Cyclodictyon laetevirens (Hook. & Taylor) Mitt. Deslooveria quintasii (Broth.) Enroth Deslooveria saotomensis (Enroth & Shevock) Enroth Dicranella falcularia Müll.Hal. ex Dusén Ectropothecium brevifalcatum (Müll.Hal.) Kindb. Ectropothecium diffusum (Mitt.) A. Jaeger Ectropothecium drepanophyllum Broth. Fissidens asplenioides Hedw. Fissidens borgenii Hampe C. Garcia et al. Synonyms (basionyms) based in São Tomé and Príncipe collections P X ST A Calymperes principis Broth. 1890 X Campylopus quintasii Broth. 1890 Dicranum divaricatum Mitt. 1863; Campylopus erythrocaulon Broth. 1890 X X A 1890/ 1959 2011 X 1890 X 1890/ 1863 X 2016 X 1944 X 1944 Porotrichum quintasii Broth. 1890 A 1890 Porotrichum saotomense Enroth and Shevock (2011) E 2011 A Hypnum brevifalcatum Müll.Hal. 1886 Stereodon diffusus Mitt. 1863 1944 A A E 1890 A X X X X 2013 2013/ 2013 2013/ 2013 2013/ 2013 2013 2013 2013/ 2013 2013/ 2013 1890/ 1890 Ectropothecium drepanophyllum Broth. 1890 X X A X A A Fissidens flaccidus Mitt. X X A A Fissidens subglaucissimus Broth. 1890 1888 1863 Fissidens crispulus Brid. var. crispulus Fissidens crispulus var. robinsonii (Broth.) Z. Iwats.& Z.-H. Li Fissidens crispus Mont. Fissidens darntyi Schimp. Fissidens enervis Sim Fissidens glaucissimus Welw. & Duby Year of first ref. P/ST 1987 1896 (continued) 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . Taxon species/subspecies/variety Fissidens intramarginatus (Hampe) A.Jaeger Fissidens metzgeria (Müll.Hal.) Broth. Fissidens microcarpus Mitt. Fissidens ovatus Brid. Fissidens pallidinervis Mitt. Fissidens palmatus Hedw Fissidens pellucidus Hornsch. Synonyms (basionyms) based in São Tomé and Príncipe collections P 243 ST X Year of first ref. P/ST 2013 A 2013 A A X X X X Fissidens porrectus Mitt. A A Fissidens punctulatus Sande Lac. Fissidens ramulosus Mitt. A X A A A Fissidens sciophyllus Mitt. Fissidens serratus Müll.Hal. var. serratus Fissidens submarginatus Bruch Fissidens usambaricus Broth. Fissidens zollingeri Mont. Floribundaria floribunda (Dozy & Molk.) M.Fleisch. Floribundaria vaginans (Welw. & Duby) Broth. Funaria acicularis Müll.Hal. Funaria hygrometrica Hedw. var. hygrometrica Gymnostomiella erosula (Müll.Hal. ex Dusén) Arts Gymnostomiella vernicosa (Hook.) M.Fleisch. Hydrogonium consanguineum (Thwaites & Mitt.) Hilp. Hydrogonium orientale (F. Weber) Kucera Hymenostylium recurvirostrum (Hedw.) Dixon var. recurvirostrum Hyophila involuta (Hook.) A.Jaeger Hypopterygium tamarisci (Sw. ex Sw.) Brid. ex Müll.Hal. Isopterygium nanoglobum (Müll. Hal.) Paris Fissidens purpureocaulis Müll. Hal. 1900 X X A X 2013 2013 2013 2011 A 1886 E X 1886 1901 A 2016 X 2016 X 2016/ 2016 2016/ 1987? 2016 X Papillaria patentissima Müll.Hal. 1886 Funaria acicularis Müll.Hal. 1886 X X X Hypopterygium brevifolium Broth. 1890 Hypnum nanoglobum Müll.Hal. 1886 2013 2013 2013 2013 2013/ 2013 2013/ 2013 1890 2013/ 2013 2013/ 2013 2013 X X X X E 2016/ 2016 1997/ 1863 1886 (continued) 244 Taxon species/subspecies/variety Lepidopilum lastii Mitt. Lepidopilum niveum (Müll.Hal.) Kindb. Leptodontium viticulosoides (P. Beauv.) Wijk & Margad. var. viticulosoides Leucobryum fouta-djalloni Paris & Cardot Leucobryum homalophyllum Broth. Leucobryum leucophanoides Müll. Hal. Leucoloma gracilescens Broth. Leucoloma secundifolium Mitt. Leucomium strumosum (Hornsch.) Mitt. Leucophanes molleri Müll.Hal. Leucophanes unguiculatum Mitt. Lopidium struthiopteris (Brid.) M. Fleisch. Macromitrium sulcatum (Hook.) Brid. var. sulcatum Mesonodon flavescens (Hook.) W.R. Buck Mittenothamnium leptoreptans (Broth.) Cardot Neckeromnion lepineanum (Mont.) S.Olsson, Enroth, Huttunen & D. Quandt Neckeropsis disticha (Hedw.) Kindb. Octoblepharum albidum Hedw. Orthostichella rigida (Müll. Hal.) B.H.Allen & Magill Orthostichella versicolor (Müll. Hal.) B.H. Allen & W.R. Buck Orthostichidium involutifolium subsp. thomeanum (Broth.) Argent Orthostichidium involutifolium (Mitt.) Broth. subsp. involutifolium. C. Garcia et al. Synonyms (basionyms) based in São Tomé and Príncipe collections P Hookeria niveum Müll.Hal. 1886 Leucobryum homolophyllum Broth. 1890 Leucobryum leucophanoides Müll. Hal. 1886 Leucoloma gracilescens Broth. 1890 Leucoloma secundifolium Mitt.1863 Leucophanes molleri Müll.Hal. 1886 Leucophanes unguiculatum Mitt.1863 Hypopterygium subtrichocladum Broth. 1890 Macromitrium undatifolium Müll. Hal. 1886 ST A A Year of first ref. P/ST 1944 1886 X 2016 A 1959 E 1890 E 1886 A 1890 A 1863 X 1944 X 1886 A 1863 X X X X 1890/ 1997 1917/ 1886 2011 E 1890 X X 2017/ 2017 X X X X X 2011/ 1993 1944/ 1959 1886 X 1886 A 1996/ 1890 A 1959 X Microthamnium leptoreptans Broth. 1890 Pilotrichella leptoclada Müll.Hal. 1886; P. calomicra Broth. 1890 Pilotrichella inflatifolia Müll.Hal. 1886 Hildebrandtiella thomeana Broth.1890; Orthostichidium thomeanum (Broth.) Broth. 1906 A (continued) 9 The Bryophyte Flora of São Tomé and Príncipe (Gulf of Guinea): Past,. . . Taxon species/subspecies/variety Philonotis nanothecia (Müll.Hal.) Kindb. Philonotis trichodonta (Müll.Hal.) Kindb. Pinnatella minuta (Mitt.) Broth. Pinnatidendron piniforme (Brid.) Enroth Plagiomnium rhynchophorum (Hook.) T.J.Kop. var. rhynchophorum. Pogonatum gracilifolium Besch. Pogonatum usambaricum (Broth.) Paris Pyrrhobryum spiniforme (Hedw.) Mitt. Racopilum orthocarpioides Broth. Racopilum thomeanum Broth. Radulina borbonica (Bél.) W.R. Buck Rhacopilopsis trinitensis (Müll. Hal.) E.Britton ex Dixon Rhizofabronia persoonii (Schwägr.) M.Fleisch var. persoonii. Rhynchostegium hopfferi (Welw. & Duby) A.Gepp Scabrellifolium elongatum (Welw. & Duby) Enroth Scabrellifolium substriatum (Hampe) Enroth Sematophyllum amblystegiocarpum (Müll.Hal.) Broth. Splachnobryum obtusum (Brid.) Müll.Hal. Symphyodon pygmaeus (Broth.) S. He & Snider Syrrhopodon gardneri (Hook.) Schwägr. Syrrhopodon lamprocarpus Mitt. Tayloria solitaria (Hedw.) T.J.Kop. & W.Weber Synonyms (basionyms) based in São Tomé and Príncipe collections Bartramia nanothecia Müll.Hal. 1886 Bartramia trichodonta Müll.Hal. 1886 Hypnum africanum Welw. & Duby 1872 245 ST A Year of first ref. P/ST 1886 E 1886 X X X X 1917/ 1872 2011/ 2011 1944 P X Polytrichum rubentiviride Müll. Hal. 1886; P. molleri Müll.Hal. 1886 Racopilum orthocarpioides Broth. 1890 Rhacopilum thomeanum Broth. 1890 Trichosteleum subpycnocylindricum Broth. 1890 Microthamnium subelegentulum Broth. 1890 Hypnum hopfferi Welw. & Duby 1872 Hypnum molleri Müll.Hal. 1886 Porotrichum caudatum Broth. 1890 Hypnum amblystegiocarpum Müll. Hal. 1886 Syrrhopodon quintasii Broth. 1890 A A 1944/ 1886 A 1989 X 1886 A 1890 A 1890 X 1890 X 1890 A 1863 E 1872 A 1886 X 1890 E 1886 X 2016 X 2011 X 1890 A A 1886 1972 (continued) 246 Taxon species/subspecies/variety Thamnobryum corticola (Kindb.) De Sloover Thuidium involvens subsp. thomeanum (Broth.) Touw Trachypodopsis serrulata (P. Beauv.) M.Fleisch. var. serrulata Trachypus bicolor var. viridulus (Mitt.) Zanten Trematodon divaricatus Bruch Trematodon longicollis Michx. Trichosteleum dicranelloides Broth. Vesicularia glaucula (Broth.) Broth. Vesicularia scaturigina (Brid.) Broth. Vesicularia strephomischos (Welw. & Duby) Broth. Wijkia monodii (P.de la Varde) H. Akiyama Wijkia trichocoleoides (Müll.Hal.) H.A.Crum C. Garcia et al. ST A Year of first ref. P/ST 1902 Thuidium thomeanum Broth. 1890 A 1890 Trachypodopsis quintasiana Broth. 1909 Papillaria molleri Müll.Hal. 1886 A 1909 X 1886 A 1886 X 1886 A 1890 A 1890 A 1863 A 1872 E 1959 A 1886 Synonyms (basionyms) based in São Tomé and Príncipe collections Trematodon flexifolius Müll.Hal. 1886?? Trematodon flexifolius Müll.Hal. 1886 Trichosteleum dicranelloides Broth. 1890 Ectropothecium glauculum Broth. 1890 Hypnum strephomischos Welw. & Duby 1872 Gollania monodii P.de la Varde 1959 Hypnum trichocoleoides Müll.Hal. 1886 P References Ah-Peng C, Wilding N, Kluge J et al. (2012) Bryophyte diversity and range size distribution along two altitudinal gradients: continent vs. Island. 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