Bothalia - African Biodiversity & Conservaton ISSN: (Online) 2311-9284, (Print) 0006-8241

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Fify shades of red: Lost or threatened bryophytes in Africa

Authors: Background: A Red List of threatened bryophytes is lacking for Africa. The International 1,2 Jacques van Rooy Union for Conservation of Nature (IUCN) Species Survival Commission (SSC) Bryophyte Ariel Bergamini3 Irene Bisang4 Specialist Group has recently launched the ‘Top 10 Initiative’ to identify the 10 species on each continent that are at highest risk of extinction. Afliatons: 1Natonal Herbarium, South Objectives: The main aim of this paper was to highlight some of the lost or strongly threatened African Natonal Biodiversity bryophyte species in sub-Saharan Africa and the East African islands and to draw up a Top 10 Insttute, South Africa list for Africa.

2School of Animal, Method: Lost or threatened species have been identified with the help of experts on the Plant and Environmental bryoflora of Africa, global and regional Red Lists and taxonomic literature. Each species on Sciences, University of the this candidate list is discussed at the hand of its taxonomy, distribution, habitat, threat and Witwatersrand, South Africa current global or regional Red List status as far as previously assessed.

3 Department of Biodiversity Results: Fifty bryophyte species, representing 40 genera and 23 families, have been identified and Conservaton Biology, Swiss Federal Research as Top 10 candidates. Of these, 29 are endemic to Africa and 21 are restricted to the East African Insttute WSL, Switzerland islands. The majority of the candidate species occur in one of eight ‘biodiversity hotspots’ with most species (19) in the Madagascar and the Indian Ocean Islands hotspot. 4Research Division Directorate & Department Conclusion: This is the first list of lost or threatened bryophytes for Africa and the first Top 10 of Botany, Swedish Museum list of the IUCN Bryophyte Specialist Group. It represents an important step towards regional of Natural History, Sweden and global Red List assessment of bryophytes, thus meeting the targets of the Updated Global Corresponding author: Strategy for Plant Conservation 2011–2020 and priorities of The Shenzhen Declaration on Plant Jacques van Rooy, Sciences. [email protected]

Dates: Received: 18 Jan. 2018 Introducton Accepted: 23 Aug. 2018 The bryoflora of sub-Saharan Africa and the East African islands is poorly known and recent Published: 14 Jan. 2019 taxonomic and floristic activity is minimal in comparison to other regions of the world (Diop et al. How to cite this artcle: 2018; Hallingbäck & Hodgetts 2000; Magill 2010; O’Shea 2005; Wilding 2017). Recent, but still Van Rooy, J., Bergamini, A. & incomplete, floras and identification guides are available only for southern Africa (Magill 1981, Bisang, I., 2019, ‘Fify shades 1987; Magill & Van Rooy 1998; Perold 1999), tropical Africa (De Sloover 2003), Kenya (Chuah- of red: Lost or threatened Petiot 2003; Wilding et al. 2016), West Africa (Wigginton 2004) and Rwanda (Fischer 2013). bryophytes in Africa’, Bothalia 49(1), a2341. htps://doi.org/ However, a series of regional checklists with accepted names, synonyms, literature references and 10.4102/abc.v49i1.2341 geographical distribution by country, of which the moss checklist of O’Shea (2006) and the liverwort and hornwort checklist of Wigginton (2018) are the most recent, provide a sound basis Copyright: for research on the taxonomy, ecology and conservation of African bryophytes. © 2019. The Authors. Licensee: AOSIS. This work is licensed under the The documentation of plant diversity and its urgent conservation are priority objectives of the Creatve Commons Updated Global Strategy for Plant Conservation 2011–2020 (Convention on Biological Diversity Atributon License. [CBD] Secretariat 2017) and The Shenzhen Declaration on Plant Sciences (Crane et al. 2017). Several authors drew attention to the in general smaller and less spectacular, but to the same degree rare or threatened, bryophytes, especially in centres of bryophyte diversity, and emphasised the need for Red Lists (Hallingbäck & Tan 2010; Geffert et al. 2013; Longton & Hedderson 2000; Szabó & Pócs 2016).

The International Union for Conservation of Nature (IUCN) Red List Categories and Criteria provide an explicit framework for classifying species according to their extinction risk (IUCN Read online: 2012a). The IUCN Species Survival Commission (SSC) lists the expansion of geographic and Scan this QR taxonomic coverage of the IUCN Red Lists as its top priority (IUCN 2017b). The IUCN Red List of code with your smart phone or Threatened Species for bryophytes (IUCN 2017e; Tan et al. 2000) currently contains 102 species, mobile device including several from Africa. However, most of these assessments are more than 10 years old and to read online. thus in need of careful revision. There is currently no Red List for African bryophytes and the only

htp://www.abcjournal.org Open Access Page 2 of 7 Original Research regional Red List is the list of threatened liverwort and It is generally accepted that centres of diversity and endemism, hornwort species compiled for the East African island of or so-called ‘biodiversity hotspots’, are priority areas for Réunion (Ah-Peng et al. 2012). conserving biodiversity (Geffert et al. 2013; Marchese 2015). Conservation International, through the Critical Ecosystem To rigorously assess the estimated 18 000 bryophyte species Partnership Fund (CEPF) (2016a), recognises 36 global globally (Frey & Stech 2009), following IUCN methodology, biodiversity hotspots. A biodiversity hotspot is characterised which is now generally accepted and advocated by, for by high levels of vascular plant endemism (>1500 endemic example, Vanderpoorten and Hallingbäck (2009:488), is an species) and 30% or less of its original natural vegetation unsurmountable task given the current personnel and economic remaining (Conservation International 2017). Many of these resources devoted to bryophyte conservation. Such an are heavily threatened by habitat loss and other human endeavour of IUCN red-listing is currently underway for the activities (CEPF 2016a; Mittermeier et al. 2004). The hotspot ca. 1800 European bryophyte species (IUCN 2017c). To reinitiate system is used to assess global conservation priorities red-listing activities on a global scale, the IUCN SSC Bryophyte and to provide grants to non-governmental and private Specialist Group has launched the so-called ‘Top 10 Initiative’ sector organisations through the CEPF (2016b; Conservation (IUCN 2017d; IUCN SSC 2016) to identify a minimum of International 2017). 10 species at high risk of extinction from each continent and properly assign them to an IUCN Red List category (IUCN Eight of the global biodiversity hotspots are found in 2012a; IUCN Standards and Petition Subcommittee 2017). sub-Saharan Africa and the East African islands (Figure 1):

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0 800 1600 km 40 10 0 10 20 30 40 50 60

Source: Based on the Biodiversity hotspots map downloaded from htp://www.cepf.net/where_we_work/Pages/map.aspx, fgure licensed under the Creatve Commons Atributon-Share Alike 4.0 Internatonal license (Author: Conservaton Internatonal) km, kilometres. FIGURE 1: The number of lost or threatened bryophytes in Africa and the East African islands (Top 10 candidates) in each of the eight Biodiversity hotspots recognised by Conservaton Internatonal (CEPF 2016a). An additonal four candidate species are only known from inland areas of southern Africa outside the Biodiversity hotspots (not shown on map).

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Cape Floristic Region, Coastal Forests of Eastern Africa, proposal to search in the feld for species that have gone Eastern Afromontane, Guinean Forests of West Africa, unrecorded for years or decades (funding pendent). Horn of Africa, Madagascar and the Indian Ocean Islands, Maputaland–Pondoland–Albany and Succulent Inspired by these two initiatives, a list of lost and threatened Karoo biodiversity hotspots. species (Online Appendix 1) was compiled based on the experts’ and the frst author’s (J.v.R.) contributions. The Recent analyses of bryophyte distributions at a global, following criteria were applied in the selection of species: continental or regional scale (Geffert et al. 2013; Tan & • The species should be endemic to sub-Saharan Africa or Pòcs 2000; Vanderpoorten & Hallingbäck 2009; Van Rooy & the East African islands. Phephu 2016; Von Konrat et al. 2008) indicate that African • Known from a single or few localities with a narrow centres of bryophyte diversity and endemism partly coincide geographical distribution range. with the biodiversity hotspots defned on the basis of vascular • The habitat should be threatened and declining or the plants. The most species-rich areas are Madagascar, especially known records date from previous centuries despite more the lowland forests along the east coast, the mountains of recent collecting activity or searches in the area. eastern Africa, the Afromontane and Cape Floristic regions of South Africa, and West Africa. The list of 50 species includes all threatened African bryophytes listed in the global IUCN Red List of Threatened The aim of this paper is to highlight lost and threatened Species (IUCN 2017e; Tan et al. 2000), and the threatened bryophytes in sub-Saharan Africa and the East African African endemics identifed as ‘Critically Endangered’ in the islands and to draw up a Top 10 list of bryophytes in Africa Red List of liverworts and hornworts for Réunion (Ah-Peng that are at highest risk of extinction. The Top 10 species will et al. 2012). be subjected to a careful IUCN Red List assessment to be included in The IUCN Red List of Threatened Species, and In the Online Appendix 1, each species is presented at the they will be further evaluated (IUCN 2012b) to identify and hand of its taxonomy, geographical distribution, habitat design the most urgent conservation actions. We also test the (including substrate) and most likely threats. The current hypothesis that most lost and threatened bryophytes on our Red List status is indicated for those species that were list fall within the global biodiversity hotspots recognised by previously assessed, either at the global or at regional scales. Conservation International (CEPF 2016a). However, not all of these Red List assignments follow IUCN methodology consequently, and some of the global Red List Research method and design statuses are in need of revision. Family placement of the The area covered in this publication comprises sub-Saharan genera follows the classifcations of Goffnet, Buck and Shaw Africa, together with the Atlantic and Indian Ocean islands in (2009) and Frey and Stech (2009) for the mosses and the tropics (Figure 1). This corresponds to the area covered by Söderström et al. (2016) for the liverworts. Species author the checklists of O’Shea (2006) and Wigginton (2018), except citations follow Söderström et al. (2016) for the liverworts and for the islands of Cape Verde, Ascension and St. Helena, which Missouri Botanical Gardens’ TROPICOS database (Tropicos. are excluded here. org) for the mosses.

Experts on the taxonomy, ecology and conservation of The 10 species that are at highest risk of extinction, and most African bryophytes were invited to contribute to two likely to be categorised as ‘critically endangered’ when (re-) initiatives, namely the ‘Top 10 Initiative’ by the IUCN SSC assessed using the latest IUCN Red List criteria (IUCN Bryophyte Specialist Group (IUCN SSC 2016) and the ‘Search 2012a), were selected for the Top 10 list for Africa (Table 1). for Lost Species’ initiative (Global Wildlife Conservation Monotypic genera and subgenera received priority and 2014). Global Wildlife Conservation approached IUCN preference was given to species recognised and accepted Specialist Groups in 2014 to suggest candidate species for a in the latest world and African checklists of bryophytes

TABLE 1: The IUCN SSC Bryophyte Specialist Group Top 10 list of bryophytes in Africa that are at highest risk of extncton, compiled from the list of candidate species in the Online Appendix 1 where more details on the species’ ecology and threats are provided. Name Family Distributon Biodiversity hotspot† Bryopteris gaudichaudii Gotsche Lejeuneaceae Madagascar and Réunion Madagascar and the Indian Ocean Islands Cheilolejeunea ulugurica Malombe, Eb.Fisch. et Pócs Lejeuneaceae Endemic to Tanzania Eastern Afromontane Cololejeunea nosykombae A.Szabó & Pócs Lejeuneaceae Endemic to Madagascar Madagascar and the Indian Ocean Islands Ludorugbya springbokorum Hedd. & R.H.Zander Potaceae Endemic to South Africa Cape Floristc Region Neckeropsis pocsii Enroth & Magill Neckeraceae Endemic to the Comoros Madagascar and the Indian Ocean Islands Picobryum atomicum R.H.Zander & Hedd. Potaceae Endemic to South Africa Cape Floristc Region Pocsiella hydrogonioides Bizot Dicranaceae Endemic to Tanzania Eastern Afromontane Symbiezidium madagascariense Steph. Lejeuneaceae Madagascar and Seychelles Madagascar and the Indian Ocean Islands Xylolejeunea grolleana (Pócs) Xiao L.He et Grolle Lejeuneaceae Madagascar and Réunion Madagascar and the Indian Ocean Islands Xylolejeunea muricella Xiao L.He et Grolle Lejeuneaceae Endemic to the Seychelles Madagascar and the Indian Ocean Islands †, Biodiversity hotspots recognised by Conservaton Internatonal through the CEPF (2016a).

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(O’Shea 2006; Söderström et al. 2016; Tropicos.org) for which threatened bryophytes in Africa as well as on the Top 10 list precise information on localities is available. of bryophytes in Africa that are at highest risk of extinction. Not only is Lejeuneaceae the largest family of liverworts, Results but it contains more than 95% of all epiphyllous bryophytes (Gradstein 1994). The family displays high levels of diversity We have identifed a total of 50 species, representing 40 genera and endemism in moist tropical forests where they grow and 23 families of bryophytes, as lost and threatened in mainly on the bark of woody plants and on leaves at low the area (Online Appendix 1). Of these, 29 are endemic to and mid elevations (Gradstein 1992, 1994; Pócs 1996; Von mainland Africa, 19 to the East African islands and two are Konrat et al. 2008). Von Konrat et al. (2008:431) calculated restricted to West Africa as well as islands off the coast. that in Madagascar and some countries of central and West Twenty-nine species (in 19 genera and nine families) are Africa, between 50% and almost 90% of the liverwort fora liverworts and 21 species (in 21 genera and 14 families) are consist of Lejeuneaceae. mosses. The liverwort family Lejeuneaceae with 11 genera and 17 species is the largest family on the list whilst Cololejeunea (Spruce) Steph. and Riccia L. are the largest genera, represented Recent researches in tropical forests indicate that ancient by fve species each. uncut stands of forest, with original phorophyte diversity and intact forest canopies, are important drivers for high Altogether, the species’ known localities are found in 16 epiphytic and epiphyllous bryophyte diversity (Benítez, African countries (Online Appendix 1). Eighteen species Prieto & Aragón 2015; Malombe et al. 2016; Pócs & occur in the Flora of Southern Africa area (Magill 1981), with Tóthmérész 1997; Zartman 2003). The ongoing destruction, 15 species being endemic to South Africa. Of these, seven degradation and fragmentation of forests in the biodiversity species are restricted to the Fynbos Biome (Rebelo et al. 2006) hotspots of Africa (Aynekulu et al. 2016; CEPF 2016a; of the southwestern Cape. The Indian Ocean islands of Gradstein 1992; Green & Sussman 1990; IUCN 2017a; Réunion and Madagascar are well represented on the list with Malombe 2007) therefore pose serious threats to epiphytic 12 (six endemic) and 11 (four endemic) species, respectively. and epiphyllous bryophytes in general and species of Lejeuneaceae in particular. Six of the eight global biodiversity hotspots recognised in sub-Saharan Africa and the East African islands, based on The fact that 46 (92%) candidate species for the Top 10 list vascular plant species and vegetation, contain candidates fall within six of the eight biodiversity hotspots in the for the Top 10 list (Figure 1). The three hotspots with the region suggests that the African biodiversity hotspots most species are (1) Madagascar and the Indian Ocean designated on the basis of information from vascular plants Islands with 19 species, (2) Eastern Afromontane with nine may also represent hotspots for bryophytes. However, the species and (3) Cape Floristic Region with seven species. concentration of lost or threatened species in the biodiversity The remaining hotspots with threatened bryophytes are: hotspots may also be a consequence of collecting bias. Only Maputaland–Pondoland–Albany (four species), Succulent two of the global biodiversity hotspots in Africa, the Coastal Karoo (four species) and Guinean Forests of West Africa Forests of Eastern Africa and the Horn of Africa, are not (three species). represented on our list (Figure 1). Whilst these areas are under-explored (Hylander, Nemomissa & Hedenäs 2017; Only four species, all from inland areas of southern Africa, Wilding et al. 2016), lower bryophyte diversity in arid areas have not been reported from any of the global biodiversity such as in the Horn of Africa hotspot may further contribute hotspots. They are Anacamptodon marginata (Dixon) W.R.Buck to this outcome. Many African regions, for example in from Mashonaland West Province of Zimbabwe, Cryptomitrium Western Africa (Diop et al. 2018), are much less researched oreades Perold from the Highlands of Lesotho, Fissidens and thus still poorly understood in terms of plant diversity capriviensis Magill from the Caprivi Strip of Namibia and and threats to it. Thus, the detailed picture of the distribution Gymnostomum lingulatum Rehmann ex Sim from the of African bryophyte diversity is likely to change along with Woodbush–Haenertsburg area in the Limpopo Province of increasing knowledge and exploration, but we are confdent South Africa. that some overall patterns are recognisable based on this compilation. The majority of Top 10 species (six species) belong to the liverwort family Lejeuneaceae, and the Madagascar and the The biodiversity hotspot with the highest number of lost or Indian Ocean Islands hotspot is the biodiversity hotspot threatened species, as well as Top 10 species which are at with the highest number of species (six species) on the list highest risk of extinction in Africa, is Madagascar and the (Table 1). Xylolejeunea Xiao L.He et Grolle (Lejeuneaceae) is Indian Ocean Islands (Table 1, Figure 1). This hotspot is the only genus with more than one (two) species on the Top dominated by Madagascar, the fourth largest island 10 list for Africa. globally, and includes the Seychelles (including Aldabra), the Comoros, Mauritius (including Rodrigues) and the Discussion French overseas departments of Réunion, Mayotte (one of It is no coincidence that Lejeuneaceae is the liverwort family the Comoros) and the volcanic Iles Esparses around with the highest species number on the list of lost or Madagascar (Figure 1). The island of Madagascar, especially

htp://www.abcjournal.org Open Access Page 5 of 7 Original Research the eastern lowland rainforests, is known as a global centre threatened by invasive alien species, the disruption of fre of bryophyte diversity and endemism and the most species- regimes, plantation forestry and fragmentation. rich area of bryophytes in Africa (Geffert et al. 2013; Tan & Pócs 2000; Vanderpoorten & Hallingbäck 2009). It has also Conclusions been identifed as one of two major Lost Spots in Africa, This is the frst compilation of bryophytes for sub-Saharan countries where ‘lost species’, including a few mosses, Africa and the East African islands that are most probably are concentrated (Global Wildlife Conservation 2014). lost or threatened. Although it is not exhaustive, and many Agricultural activities, urbanisation, invasive alien species of the ‘threat categories’ reported herein are in need of and deforestation have devastated and fragmented habitats, revision and require to be scrutinised against the IUCN Red especially in the lowlands, throughout this hotspot (CEPF List criteria (IUCN 2012a), it represents a frst approximation 2016a). At least one of the endemic bryophyte species from of the threats to bryophytes in Africa. the Seychelles island of Mahé, Xylolejeunea muricella Xiao L.He et Grolle, may already be extinct (Online Appendix 1). The species on the list of lost or threatened bryophytes There is also concern about the recent marked increase in served as candidates for the selection of the Top 10 species in illegal logging in the remaining forests of Madagascar Africa at high risk of extinction. Therefore, it is a crucial (Green & Sussman 1990; IUCN 2017a). contribution towards the Top 10 Initiative of the IUCN Bryophyte Specialist Group (IUCN SSC 2015) and thus to the The second-highest number of lost or threatened species per (global) IUCN Red List of Threatened Species (IUCN 2017e), biodiversity hotspot are found in the Eastern Afromontane as well as towards an assessment of the extinction risk of hotspot (Figure 1). This hotspot is characterised by a series of bryophytes on the African continent. Hence, it is a signifcant montane islands and extensive plateaus along the eastern step towards meeting the targets of the Updated Global edge of Africa, from Ethiopia in the north to Zimbabwe in Strategy for Plant Conservation 2011–2020 (CBD Secretariat the south. It consists mainly of three ancient massifs: the 2017) and priorities of The Shenzhen Declaration on Plant Eastern Arc Mountains and Southern Rift, the Albertine Rift Sciences (Crane et al. 2017). and the Ethiopian Highlands. Eastern Africa has also been suggested as one of the main centres of bryophyte diversity The vast majority of threatened species on our list, and all and endemism in Africa (Geffert et al. 2013; Von Konrat et al. Top 10 species, are restricted to global biodiversity hotspots 2008) and Tanzania is another major Lost Spot in the region in Africa, areas of high endemism of vascular plants under for a variety of organisms (Global Wildlife Conservation severe threat of habitat loss (CEPF 2016a; Conservation 2014). Degradation and fragmentation of habitats and International 2017). Investment in biodiversity conservation unsustainable exploitation of natural resources are the main through the hotspots concept will therefore beneft threatened threats in this hotspot (CEPF 2016a). This is caused by African bryophytes as well. expanding agriculture, plantation forestry, logging, fres, invasive alien plants, mining, infrastructure development The Madagascar and Indian Ocean Islands biodiversity and gathering of frewood. hotspot, which is also considered as a global centre of bryophyte diversity and endemism (Vanderpoorten & A substantial number amongst the lost or threatened Hallingbäck 2009; Von Konrat et al. 2008), contains a bryophytes occur in the southwestern Cape, a region that particularly large number of threatened bryophytes. The contains the highest concentration of threatened vascular recent increase in logging poses a serious threat to the plants and plants of conservation concern in South Africa remaining forest habitats on Madagascar and other Indian (Raimondo & Van Staden 2009). The Fynbos Biome of the Ocean islands. The Eastern Afromontane and Cape Floristic Cape Floristic Region biodiversity hotspot is one of the most Region biodiversity hotspots also stand out as hotspots of threatened biomes in southern Africa, classifed as critically lost or threatened bryophytes in Africa. endangered (Driver et al. 2012). The southwestern Cape has been suggested as a global and regional centre of relatively The degradation and fragmentation of forests throughout high bryophyte species richness and endemism (Geffert et al. sub-Saharan Africa and the East African islands have a visible 2013; Tan & Pócs 2000). In an analysis of moss distributions in impact on epiphytic bryophytes, epiphyllous liverworts of southern Africa, Van Rooy and Phephu (2016:29) recognised the family Lejeuneaceae in particular. Bryological exploration a Southwestern Cape Centre of Moss Diversity and found of Africa should be stepped up to increase our knowledge of that the Cape Town-Table Mountain area is the most species- species occurrences and threats and to facilitate the red-listing rich in the region. The Cape Floristic Region is under and the conservation of rare and threatened bryophytes. increasing population pressure and much of the Fynbos and Renosterveld vegetation types of the lowlands have been destroyed or transformed by agriculture and urbanisation Acknowledgements (CEPF 2016a; Rebelo et al. 2006; Von Hase et al. 2003). The We would like to thank Tamás Pócs, Frank Müller, Itambo lowland Renosterveld areas have been identifed as top Malombe and Claudine Ah-Peng for their contributions. We conservation priorities by the Cape Action Plan for the are grateful to Hester Steyn and Elizma Fouche for editing Environment (Von Hase et al. 2003). The remaining Fynbos is the map.

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Hallingbäck, T. & Tan, B.C., 2010, ‘Past and present actvites and future strategy Competng interests of bryophyte conservaton’, Phytotaxa 9, 266–274. htps://doi.org/10.11646/ phytotaxa.​9.1.15 The authors declare that they have no fnancial or personal Hylander, K., Nemomissa, S. & Hedenäs, L., 2017, ‘Mosses of southwest Ethiopian relationships that may have inappropriately infuenced them montane forests – Notes on their occurrence patern and many new country records’, Journal of Bryology 39(4), 342–352. htps://doi.org/10.1080/03736687. in writing this article. 2017.1329793 Internatonal Union for Conservaton of Nature (IUCN), 2012a, IUCN red list categories and criteria: Version 3.1, 2nd edn., IUCN, Gland, Switzerland and Cambridge, UK, Authors’ contributons viewed 05 December 2017, from htp://www.iucnredlist.org/technical-documents/​ red-list-documents J.v.R. compiled the species accounts, analysed and interpreted Internatonal Union for Conservaton of Nature (IUCN), 2012b, Guidelines for the data and drafted the article. I.B. and A.B. conceptualised applicaton of IUCN red list criteria at regional and natonal levels: Version 4.0, IUCN, Gland, Switzerland and Cambridge, UK, viewed 05 December 2017, from and initiated the project ‘Top 10 Initiative’ and commented htp://www.iucnredlist.org/technical-documents/red-list-documents on manuscript versions. All the authors contributed to the Internatonal Union for Conservaton of Nature (IUCN), 2017a, Illegal actvites threaten natural World Heritage, pushing the vaquita to the brink and depletng fnal version of the article. forests, viewed 10 July 2017, from htps://www.iucn.org/news/iucn-41whc/​ 201706/illegal-activities-threaten-natural-world-heritage-pushing-vaquita- brink-and-depleting-forests-%E2%80%93-iucn?dm_i=2GI3,​13OT1,​40ELQB,​ 3C6Q3,1 References Internatonal Union for Conservaton of Nature (IUCN), 2017b, IUCN species strategic plan 2013–2016 Ah-Peng, C., Bardat, J., Pócs, T., Söderström, L., Staménof, P. & Strasbergah, D., 2012, , viewed 11 October 2017, from htps://cmsdata.iucn.org/ ‘Red list of liverworts and hornworts for Réunion (Mascarene archipelago)’, downloads/2013_2016_species_strategic_plan_fnal.pdf Phytotaxa 68, 1–23. htps://doi.org/10.11646/phytotaxa.68.1.1 Internatonal Union for Conservaton of Nature (IUCN), 2017c, Europe: LIFE European red lists Aynekulu, E., Aerts, R., Denich, M., Negussie, A., Friis, I., Demissew, S. et al., 2016, , viewed 19 October 2017, from htps://www.iucn.org/regions/europe/ ‘Plant diversity and regeneraton in a disturbed isolated dry Afromontane forest in projects/life-european-red-lists northern Ethiopia’, Folia Geobotanica 51(2), 115–127. htps://doi.org/10.1007/ Internatonal Union for Conservaton of Nature (IUCN), 2017d, SSC Groups: Plants & s12224-016-9247-y fungi, viewed 19 October 2017, from htps://www.iucn.org/ssc-groups/plants- Benítez, L., Prieto, M. & Aragón, G., 2015, ‘Large trees and dense canopies: Key factors fungi for maintaining high epiphytc diversity on trunk bases (bryophytes and lichens) in Internatonal Union for Conservaton of Nature (IUCN), 2017e, The IUCN red list of Tropical Montane forests’, Forestry 88(5), 521–527. htps://doi.org/10.1093/ threatened species. Version 2017–2, viewed 23 October 2017, from htp://www. forestry/cpv022 iucnredlist.org/ Conventon on Biological Diversity Secretariat, 2017, Updated global strategy for plant IUCN Species Survival Commission (SSC) (ed.), 2016, ‘Annual report of the species conservaton 2011–2020, viewed 16 August 2017, from htps://www.cbd.int/ survival commission and the global species programme’, Bryophyte Specialist gspc/strategy.shtml Group, Species 57, 70–71, viewed 08 November 2017, from htps://portals.iucn. org/library/sites/library/fles/documents/2017-008.pdf Chuah-Petot, M.S., 2003, Mosses, liverworts and hornworts of Kenya: An illustrated guide with descriptons and fgures of over 300 species and keys for identfcaton, IUCN Standards and Petton Subcommitee, 2017, Guidelines for using the IUCN red Chuah- Petot, Nairobi. list categories and criteria, Version 13, Prepared by the Standards and Pettons Hotspots Subcommitee, viewed 19 October 2017, from htp://cmsdocs.s3.amazonaws. Conservaton Internatonal, 2017, , viewed 03 January 2017, from htps:// com/RedListGuidelines.pdf www.conservaton.org/How/Pages/Hotspots.aspx. Longton, R.E. & Hedderson, T.A., 2000, ‘What are rare species and why conserve Crane, P.R., Ge, S., Yuan, D.-Y., Huang, H.-W., Knapp, S., Mooney, H. et al., 2017, ‘The them?’, Lindbergia 25, 53–61. Shenzhen declaraton on plant sciences: Unitng plant sciences and society to build a green, sustainable Earth’, Journal of Systematcs and Evoluton 55(5), 415–416. Magill, R.E., 1981, Flora of Southern Africa: Bryophyta. Part 1 Mosses. Fascicle 1 htps://doi.org/10.1111/jse.12283 Sphagnaceae–Grimmiaceae, Botanical Research Insttute, Pretoria, South Africa. Critcal Ecosystem Partnership Fund (CEPF), 2016a, The biodiversity hotspots, viewed 20 Magill, R.E., 1987, Flora of Southern Africa: Bryophyta. Part 1 Mosses. Fascicle 2 October 2017, from htp://www.cepf.net/resources/hotspots/Pages/default.aspx Gigaspermaceae–Bartramiaceae, Botanical Research Insttute, Pretoria, South Africa. Critcal Ecosystem Partnership Fund (CEPF), 2016b, About CEPF, viewed 07 September 2017, from htp://www.cepf.net/about_cepf/Pages/default.aspx Magill, R.E., 2010, ‘Moss diversity: New look at old numbers’ Phytotaxa 9, 167–174. htps://doi.org/10.11646/phytotaxa.9.1.9 De Sloover, J.L. (ed.), 2003, Illustratons de mousses Africaines, Scripta Botanica Belgica, Natonal Botanic Garden of Belgium, Meise. Magill, R.E. & Van Rooy, J., 1998, Flora of Southern Africa: Bryophyta. Part 1 Musci. Fascicle 3 Erpodiaceae–Hookeriaceae, Natonal Botanical Insttute, Pretoria. Diop, D., Diop, D., Bruggeman-Nannenga, M.A., Mbaye, M.S., Noba, K., Hedenäs, L. et al., 2018, ‘Bryophytes of Kédougou (Eastern Senegal), with a key to the Fissidens Malombe, I., 2007, ‘Systematcs of Cheilolejeunea (Spruce) Schifn. (Lejeuneaceae) in of Senegal’, Journal of Bryology 40(1), 62–67. htps://doi.org/10.1080/03736687. contnental Africa and its ecological signifcance in conservaton of Kakamega and 2017.1415662 Budongo rainforests’, PhD thesis, University of Koblenz-Landau, Germany. Driver, A., Sink, K.J., Nel, J.L., Holness, S.H., Van Niekerk, L., Daniels, F. et al., 2012, Malombe, I., Matheka, K.W., Pócs, T. & Patño, J., 2016, ‘Edge efect on epiphyllous Natonal biodiversity assessment 2011: An assessment of South Africa’s bryophytes in Taita Hills fragmented afromontane forests’, Journal of Bryology biodiversity and ecosystems, Synthesis Report, South African Natonal Biodiversity 38(1), 33–46. htps://doi.org/10.1179/1743282015Y.0000000015 Insttute & Dept. of Environmental Afairs, Pretoria. Marchese, C., 2015, ‘Biodiversity hotspots: A shortcut for a more complicated Fischer, E., 2013, ‘Liverworts and Hornworts of Rwanda’, Abc Taxa 14, 552 pp. concept’, Global Ecology and Conservaton 3, 297–309. htps://doi.org/10.1016/j. gecco.2014.12.008 Frey, W. & Stech, M., 2009, ‘Marchantophyta, Bryophyta, Anthocerotophyta’, in W. Frey, M. Stech & E. Fischer (eds.), Syllabus of plant families 3, 13th edn., pp. Mitermeier, R.A., Gil, P.R., Hofmann, M., Pilgrim, J., Brooks, T., Mitermeier, C.G. et 9–259, Schweizerbart, Stutgart. al., 2004, Hotspots revisited: Earth’s biologically richest and most endangered ecoregions, CEMEX, Mexico City. Gefert, J.L., Frahm, J.‐P., Barthlot, W. & Mutke, J., 2013, ‘Global moss diversity: Spatal and taxonomic paterns of species richness’, Journal of Bryology 35(1), 1–11. O’Shea, B.J., 2005, ‘Building a bryological framework – Getng over the threshold’, htps://doi.org/10.1179/1743282012Y.0000000038 Journal of the Hatori Botanical Laboratory 97, 281–285. Global Wildlife Conservaton, 2014, The search for lost species, viewed 20 October O’Shea, B.J., 2006, ‘Checklist of the mosses of sub-Saharan Africa’, Tropical Bryology 2017, from htps://www.globalwildlife.org/our-work/regions/global/the-search- Research Reports 6, 1–255. for-lost-species/ Perold, S.M., 1999, Flora of Southern Africa: Hepatophyta. Part 1: Marchantopsida. Gofnet, B., Buck, W.R. & Shaw, A.J., 2009, ‘Morphology and classifcaton of the Fascicle 1: Marchantidae, Natonal Botanical Insttute, Pretoria. Bryophyta’, in B. Gofnet & A.J. Shaw (eds.), Bryophyte biology, 2nd edn., pp. 55–138, Cambridge University Press, Cambridge. Pócs, T., 1996, ‘Epiphyllous liverwort diversity at worldwide level and its threat and conservaton’, Anales del Insttuto de Biología, Universidad Nacional Autónoma de Gradstein, S.R., 1992, ‘The vanishing tropical rain forest as an environment for México, Serie Botánica 67(1), 109–127. bryophytes and lichens’, in J.W. Bates & A.M. Farmer (eds.), Bryophytes and lichens in a changing environment, pp. 234–258, Clarendon Press, Oxford. Pócs, T. & Tóthmérész, B., 1997, ‘Foliicolous bryophyte diversity in tropical rainforest’, Abstracta Botanica 21(1), 135–44. Gradstein, S.R., 1994, Lejeuneaceae: Ptychantheae, Brachiolejeuneae, Flora Neotropica Monograph 62, New York Botanical Garden, New York. Raimondo, D. & Von Staden, L., 2009, ‘Paterns and trends in the Red List of South African plants’, in D. Raimondo, L. von Staden, W. Foden, J.E. Victor, N.A. Helme, Green, G.M. & Sussman, R.W., 1990, ‘Deforestaton history of the eastern rain forests R.C. Turner, et al. (eds.), Red List of South African plants, pp. 19–25, Strelitzia 25, of Madagascar’, Science 248(4952), 212–215. htps://doi.org/10.1126/science.​ South African Natonal Biodiversity Insttute, Pretoria. 248.4952.212 Rebelo, A.G., Boucher, C., Helme, N., Mucina, L. & Rutherford, M.C., 2006, ‘Fynbos Hallingbäck, T. & Hodgets, N., (compilers), 2000, Mosses, liverworts, and hornworts. biome’, in L. Mucina & M.C. Rutherford (eds.), The vegetaton of South Africa, Status survey and conservaton acton plan for bryophytes, IUCN/SSC Bryophyte Lesotho and Swaziland, pp. 53–219, Strelitzia 19, South African Natonal Specialist Group, IUCN, Gland, Switzerland and Cambridge, UK. Biodiversity Insttute, Pretoria, South Africa.

htp://www.abcjournal.org Open Access Page 7 of 7 Original Research

Söderström, L., Hagborg, A., Von Konrat, M., Bartholomew-Began, S., Bell, D., Briscoe, Von Hase, A., Rouget, M., Maze, K. & Helme, N., 2003, A fne-scale conservaton plan L. et al., 2016, ‘World checklist of hornworts and liverworts’, PhytoKeys 59, 1–828. for Cape Lowlands Renosterveld: Technical report, Report CCU2/03, Botanical htps://doi.org/10.3897/phytokeys.59.6261 Society of South Africa, Kirstenbosch. Szabó, A. & Pócs, T., 2016, ‘New or litle known epiphyllous liverworts, XX Cololejeunea Von Konrat, M., Hagborg, A., Söderström, L., Mutke, J., Renner, M., Gradstein, S.R. et al., nosykombae A.Szabó & Pócs sp.nov. from Madagascar’, Journal of Bryology 38(4), 2008, ‘Early land plants today: Global paterns of liverwort diversity, distributon and 302–307. htps://doi.org/10.1080/03736687.2016.1156357 foristc knowledge’, in H.B.B. Mohamed, A. Baki, A. Nasrulhaq-Boyce & P.K.Y. Lee (eds.), Bryology in the new Millennium, pp. 425–38, University of Malaya, Kuala Tan, B.C. & Pócs, T., 2000, ‘Bryogeography and conservaton of bryophytes’, in A.J. Lumpur. Shaw & B. Gofnet (eds.), Bryophyte Biology, pp. 401–448, Cambridge University Press, Cambridge. Wigginton, M.J. (ed.), 2004, E. W. Jones’s Liverwort and Hornwort Flora of West Africa, Scripta Botanica Belgica 30, Natonal Botanic Garden (Belgium), Meise. Tan, B., Geissler, P., Hallingbäck, T. & Söderström, L., 2000, ‘The 2000 IUCN World Red List of Bryophytes’, in T. Hallingbäck & N. Hodgets (compilers), Mosses, liverworts, Wigginton, M., 2018, ‘Checklist and distributon of the liverworts and hornworts of sub-Saharan Africa, including the East African Islands (editon 4, 25 June 2018)’, and hornworts. Status survey and conservaton acton plan for bryophytes, Tropical Bryology Research Reports 9, 1–138. pp. 77–90, IUCN/SSC Bryophyte Specialist Group, IUCN, Gland, Switzerland and Cambridge, UK. Wilding, N., 2017, ‘Three new species of Entosthodon Schwägr. (Bryopsida, Funariaceae) from sub-Saharan Africa’ Phytotaxa 312(1), 103–110. htps://doi.org/10.11646/ Tropicos.org, Missouri Botanical Garden, viewed 15 November 2017, from htp:// phytotaxa.312.1.8 www.tropicos.org/ Wilding, N., Hedderson, T., Ah-Peng, C. & Malombe, T., 2016, Bryophytes of Kenya’s Van Rooy, J. & Phephu, N., 2016, ‘Centres of moss diversity in southern Africa’, Bryophyte coastal forests: A guide to the common species, Indian Ocean Commission, Diversity and Evoluton 38(1), 27–39. htps://doi.org/10.11646/bde.38.1.3 Biodiversity Project. Vanderpoorten, A. & Hallingbäck, T., 2009, ‘Conservaton biology of bryophytes’, in B. Zartman, C.E., 2003, ‘Habitat fragmentaton impacts on epiphyllous bryophyte Gofnet & A.J. Shaw (eds.), Bryophyte Biology, 2nd edn., pp. 487–533, Cambridge communites in central Amazonia’, Ecology 84(4), 948–954. htps://doi.org/​ University Press, Cambridge. 10.1890/0012-9658(2003)084[0948:HFIOEB]2.0.CO;2

htp://www.abcjournal.org Open Access ONLINE APPENDIX 1

TABLE 1: Fifty lost or threatened bryophytes in sub-Saharan Africa and the East African islands, which served as candidates for the IUCN SSC Bryophyte Specialist Group Top 10 list of bryophytes in Africa that are at highest risk of extinction, arranged alphabetically by genus.

Name and taxonomy Distribution and habitat Threat and global or regional Sources threat category as far as assigned† 1. Actinodontium Cameroon and Bioko. For more than a century (since Müller and Pócs (2007); streptopogoneum Broth. in 1890), the species was known only Critical Ecosystem Botanische Jahrbücher für Epiphytic on tree trunks and from two sites around Mt Partnership Fund (2016). Systematik, Pflanzengeschichte twigs in forest, up to 1300 m Cameroon, which are now und Pflanzengeographie 24: 260 a.s.l. intensively cultivated. It was (1897). rediscovered in Africa on Bioko in 2002, where it grew on solitary Pilotrichaceae. The only species of trees in a pasture land. The trees at Actinodontium present in Africa. Its this site are heavily used (e.g. for relationship to American and Asian firewood). The species is thus species needs investigation. likely to be highly threatened. The Guinean Forests of West Africa biodiversity hotspot is one of the most critically fragmented habitats in the world.

2. Anacamptodon marginatus Endemic to Zimbabwe. Only known from a single Dixon and Wager (1929); (Dixon) W.R.Buck in Journal of Collected by the explorer collection made in 1870. The Buck (1980); Van Rooy the Hattori Botanical Laboratory Thomas Bains on Hartley Hill, habitat is under threat from (2011); Anonymous (2017). 47: 52 (1980). Zimbo (Chimbo) River, S. artisanal mining, cultivation and African Goldfields. overgrazing by livestock. Amblystegiaceae. Originally described Zimbabwe is bryologically poorly as Hypnofabronia marginata, the Epiphytic on trees in the known and this area has probably species was later transferred to Anacamptodon. The wiry stems and Savanna Biome, 230 m a.s.l. not been searched for bryophytes distant leaves with a strong and variable since the original collection. border, single costa and small, quadrate alar cells are distinctive.

3. Andreaea camerunensis Endemic to Cameroon: Only known from a single Richards (1952); CEPF P.W.Richards in Transactions of ‘Cameroon Mountain, Vefondi gathering made in 1948. Mount (2016). the British Bryological Society 2: Peak’. Cameroon is an active volcano and 66 (1952). the most recent eruptions were in On tall, pillar-like lava blocks February 2012. The habitat is Andreaeaceae. The species differs from in grassland at 3170 m a.s.l. threatened by lava flows. Tropical the closely related A. firma Müll.Hal. in rain forest loss and fragmentation, the broader leaves, broader, cucullate ecosystem degradation and limited leaf apices, shorter and wider basal local capacity for conservation are cells, less incrassate and slightly longer the major threats in the Guinean upper laminal cells. Forests of West Africa biodiversity hotspot.

4. Archidium andersonianum Endemic to the Western Cape This species is represented by a Snider (1975); Magill (1981); Snider in The Bryologist 78: 158 Province of South Africa single collection from the Cape Rebelo et al. (2006); CEPF (1975), fig. 26–36, 39. where it was collected on the Floristic Region, a global (2016). Stellenbosch flats by Almborn biodiversity hotspot. The vegetation Archidiaceae. This small but in 1966. of the Cape lowlands has been conspicuous species is similar in habit highly impacted by agriculture and to A. capense and A. muellerianum. It On sandy or gravelly soil in the urban development and alien plant differs in the deltoid innovation leaves Fynbos Biome. species have invaded most of the with costa ending in a strong hair-point remaining natural habitats. It is and the larger, broadly oblong-ovate highly threatened. perichaetial leaves with narrower costae.

5. Bazzania konratiana Gyarmati Endemic to Madagascar: Only known from two localities in CEPF (2016); IUCN (2017a); in Cryptogamie, Bryologie 38(2): Toamasina Prov., Mananara the Mananara Nord Biosphere Wigginton (2018); Sass- 120 (2017), fig. 1–8. Nord Biosphere Reserve and Reserve where it was collected in Gyarmati (2017). National Park. 1998. The remaining habitat in this Lepidoziaceae. The species can be biosphere reserve is threatened by separated from the Seychelles Corticolous in lowland illegal exploitation of forest endemic B. approximata Onr. by the rainforest with tree ferns, resources. Of the 15 species of larger plants, thick-walled leaf cells palms and Pandanus ssp., on E Bazzania recognised in Africa, 11 with large, bulging trigones and the and NW slopes, 250–300 m are restricted to the Madagascar proportion of hyaline and a.s.l. and the Indian Ocean Islands chlorophyllose cells in the biodiversity hotspot and six are underleaves. endemic to the island of Madagascar.

6. Bruchia eckloniana Müll.Hal. in Collected by Ecklon in the Only known from the original Magill (1981); Rebelo et al. Synopsis Muscorum Cape (probably around Cape locality in the Fynbos and (2006); CEPF (2016). Frondosorum omnium hucusque Town), without a precise Renosterveld vegetation of the Cognitorum 1: 19 (1848). locality (‘Prom. Bon. Spei’), Cape Floristic Region biodiversity between 1823 and 1848. hotspot, which is threatened by Bruchiaceae. The small calyptra, plane, urban development, agriculture and entire leaf margins and papillose to On soil in the Fynbos Biome. invasive alien plant species. The vermiculate spores will identify the Fynbos Biome is critically species. endangered.

7. Bryopteris gaudichaudii Madagascar and Réunion. Found only once since 1900 (in Hartmann et al. (2006); Gottsche in Annales des Sciences 1996) on Réunion and known from Wigginton (2018); Ah-Peng Naturelles; Botanique, sér. 4 8: Coastal ericaceous heath and only two localities on Madagascar, et al. (2012); CEPF (2016); 340 (1857). montane rainforest. about 75 km apart. Cultivation, IUCN (2017a). livestock grazing and deforestation Lejeuneaceae. Molecular evidence have devastated habitats, especially shows B. gaudichaudii as one of the in the lowlands, throughout the major clades in the genus. Madagascar and the Indian Ocean Islands biodiversity hotspot. Despite recent exhaustive searches on Réunion, this species could not be found and it is probably extinct on the island, as a result of the disappearance of its habitat. There is concern about the recent marked increase in illegal logging in the remaining forests of Madagascar. The species is facing an extremely high risk of worldwide extinction in the immediate future.

Critically Endangered globally: B1+2c (ver 2.3) (IUCN 2017b).

Regionally extinct on Réunion: (Ah-Peng et al. 2012)

8. Caudalejeunea grolleana Endemic to Madagascar where Only known from two localities in Green and Sussman (1990); Gradst. in Acta Botanica it is known from Nossi-Bé the Madagascar and the Indian Gradstein (2001a), CEPF Neerlandica 23: 334 (1974), pl. island and Antsohy Forest. Ocean Islands biodiversity hotspot (2016). 1. and not collected since 1973. The On bark of stems and dead remaining lowland rain forests of Lejeuneaceae. Within Caudalejeunea it wood in undisturbed lowland Madagascar are threatened with constitutes a separate, monotypic rainforest. destruction. subgenus. Tiny epiphytic plants, growing in small, dull green mats. Endangered globally: B1+2cd (ver 2.3) (IUCN 2017b).

9. Cheilolejeunea ulugurica Endemic to Tanzania, Uluguru Only known from a single Malombe, Fischer and Pócs Malombe, Eb.Fisch. et Pócs in Mts, N side of Bondwa peak. collection despite intensive (2010); CEPF (2016); Acta Biologica Plantarum bryological exploration in the Wigginton (2018). Agriensis 1: 24 (2010). Ramicolous on stems of Eastern Arc Mountains, which ericaceous shrubs at the edge forms part of the Eastern Lejeuneaceae. Distinguished by the of montane forest, 1650–1800 obovate-spathulate lobe with a lobule m a.s.l. Afromontane biodiversity hotspot. occupying ⅔ to ¾ its length with a Like vascular plants, any rare or strongly incurved keel apex and endemic bryophyte found in the forming a very narrow sinus with the Ulugurus at altitudes lower than postical lobe margin. Possibly conspecific with C. chenii R.L.Zhu et 1800 m is under serious threat from M.L.So from China. encroachment of habitat and deforestation. This species may be Critically Endangered.

10. Cololejeunea bosseriana Tixier Endemic to Réunion, Forêt de Known only from the type locality Ah-Peng et al. (2012); CEPF in Bulletin Trimestriel de Bébour. in the Madagascar and the Indian (2016); Söderström et al. l'Académie Malgache 55: 235 Ocean Islands biodiversity hotspot (2016). (1977)[1979], fig. 39. Epiphyll on ferns in humid and last collected by Bosser in forest at 1300 m a.s.l. 1972. Lejeuneaceae. According to the World checklist of hornworts and liverworts Critically Endangered on (Söderström et al. 2016), the species is Réunion: [B2ab(iii)] (Ah-Peng et not well known. al. 2012).

11. Cololejeunea decemplicata Madagascar and Réunion Urbanisation, agriculture and Ah-Peng et al. (2012); CEPF (Steph.) Tixier in Bulletin (three localities). invasive alien species have resulted (2016); Söderström et al. Trimestriel de l'Académie in the degradation and (2016). Malgache 55: 213 (1977)[1979]. Corticolous on trees like fragmentation of natural habitats Acacia sp., in forest. throughout the Madagascar and the Lejeuneaceae. The species is not well Indian Ocean Islands biodiversity known (Söderström et al. 2016). hotspot. On the Plaine des Cafres of Réunion, the known natural habitats of the species have been replaced by agriculture.

Critically Endangered on Réunion [B1b(iii)] (Ah-Peng et al. 2012) and most likely on Madagascar as well.

12. Cololejeunea jonesii Pócs in Kimboza Forest Reserve in Only known from two localities in Pócs (1975); Malombe et al. Acta Botanica Academiae Tanzania and Ngangao Forest the Eastern Afromontane (2016); CEPF (2016). Scientiarum Hungaricae 21: 361 in the Taita Hills of Kenya. biodiversity hotspot. The type (1975), fig. 35–43. locality in Tanzania is in a lowland Epiphyllous in lowland rain rainforest of the Kimboza Forest Lejeuneaceae. The number of cells in forest (300 m a.s.l.) and Reserve at the eastern foothills of the discoid gemmae is much higher montane tropical forest (2149 the Uluguru Mountains, seriously than in related species. The species will m a.s.l.). encroached by illegal ruby mining also be distinguished by the presence of and log felling. Recently recorded claviform styles and the biserial ventral in Ngangao Forest, a forest merophytes. fragment in the Taita Hills of Kenya, where only one specimen was found despite comprehensive collection of bryophytes. The species should be ranked as ‘Endangered’ because of few collections, rarity and fragmented or threatened forests.

13. Cololejeunea nosykombae Endemic to the volcanic Nosy The locality falls within the Szabó and Pócs (2016); CEPF A.Szabó & Pócs in Journal of Komba island of Madagascar. Madagascar and the Indian Ocean (2016). Bryology 38(4): 302 (2016). Islands biodiversity hotspot. The Epiphyllous in a submontane species is known only from a 150 Lejeuneaceae. The species differs from rainforest fragment, mostly on m × 50 m area in a forest remnant other species in the genus mainly by its Marattia fern leaflets, but also surrounded by cultivation. It is swallow-tail shaped perianth. Two on the fern Tectaria sp. and a considered to be ‘critically varieties are recognised: C. nosykombae broadleaved shrub, at an endangered’ because of its tiny var. laevis A.Szabó & Pócs as well as altitude of 570–580 m a.s.l. population size and very small area the typical variety. Associated with other of occurrence in a habitat under epiphyllous Lejeuneaceae. severe threat.

14. Cololejeunea takamakae Tixier Endemic to Réunion. Less than five localities, threatened Ah-Peng et al. (2012); in Bryophytorum Bibliotheca 27: with habitat degradation. A very Söderström et al. (2016); 319 (1985), fig. 73. Epiphyll at ±1000 m a.s.l. restricted endemic in the CEPF (2016). Madagascar and the Indian Ocean Lejeuneaceae. There are doubts about Islands biodiversity hotspot. the taxonomy of the species and it is possibly conspecific with C. Critically Endangered on angustiflora (Steph.) Mizut., which is Réunion: [B1b(iii)] (Ah-Peng et al. known also from Malaysia, Borneo, 2012). China, Taiwan and the Philippines.

15. Colura heimii Jovet-Ast in Comoros, Madagascar, Urbanisation, agriculture and Wigginton (2018); Ah-Peng Revue Bryologique et Mauritius and Réunion. invasive alien species have resulted et al. (2012); Söderström et al. Lichénologique 22: 275 (1953). in the degradation and (2016); CEPF (2016). On Réunion it grows on twigs fragmentation of natural habitats Lejeuneaceae. This is a recognised in lowland and upland forests. throughout the Madagascar and the species and currently accepted in the Indian Ocean Islands biodiversity World checklist of hornworts and hotspot. On Réunion the habitat is liverworts (Söderström et al. 2016). also threatened by lava flows.

Critically Endangered on Réunion [B2b(iii)] (Ah-Peng et al. 2012) and most likely throughout the region.

16. Cryptomitrium oreades Perold Endemic to the Oxbow area of Only known from two localities, Perold (1994, 1999); Olson in Bothalia 24: 149 (1994), fig. Lesotho. ±5 km apart. Heavy grazing and Dinerstein (2002). 1–2. pressure by livestock (carrying On soil over rock under basalt capacity is exceeded by up to Aytoniaceae. The genus is only boulders, in seepage areas in 300%) is a serious threat to the represented by one species in Africa, the high altitude Drakensberg Drakensberg alti-montane grassland alti-montane grasslands, at and woodland. It alters the which differs from the other two ±2550 m a.s.l. ecosystem and makes the existing species in the genus mainly by the lateral position of the stalk. habitats vulnerable to encroachment by Karoo vegetation. 17. Cygnicollum immersum Fife & Endemic to the Northern Cape This monotypic genus has only Magill (1987); Rebelo et al. Magill in The Bryologist 85: 99 Province of South Africa and been collected once (in 1977) and (2006); Goffinet (2013); (1982), fig. 1–19. only known from the type recent efforts to re-collect it were CEPF (2016). locality on the upper slopes of unsuccessful. However, the plants Funariaceae. This distinct, monotypic Vanrhyns Pass. are very small (stems up to 1 mm genus is recognised by its size (very long) and can easily be overlooked. small), bulbiform habit and completely It grew on sandy soil in small The Vanrhynsdorp Shale enclosed, pendent, cleistocarpic open areas amongst shrub at ±800 m a.s.l. Renosterveld vegetation is capsule. classified as least threatened, but overgrazing by livestock, cultivation and alien invasive species are major threats in the surrounding Succulent Karoo biodiversity hotspot. This particular locality is threatened by roadworks in the Vanrhyns Pass.

18. Drepanolejeunea helenae Pócs Endemic to Réunion. Known from only two localities, Pócs (1997); Ah-Peng et al. in Cryptogamie: Bryologie, where the type specimens were (2012); CEPF (2016). Lichénologie 18: 198 (1997), pl. On the bark of Psidium collected in 1994 and 1995. Habitat cattleyanum Sabine and 1–4. epiphyllous in remnants of destruction is the main threat to this tropical rainforest, ±500– very restricted endemic of the Lejeuneaceae. The species differs from 750 m a.s.l. Madagascar and the Indian Ocean D. ankasica E.W.Jones in the many Islands biodiversity hotspot. ocelli in the leaf lobe and the presence of ocelli in the perianth, amongst other Critically Endangered on characters. Réunion: [B2ab(iii)] (Ah-Peng et al. 2012).

19. Entosthodon heddersonii Endemic to Tanzania. Recently The species is known only from the Frontier.ac.uk (2014); CEPF N.Wilding in Phytotaxa 312(1): described from the type specimen collected in 1988. 103–110 (2017). Lukwangule plateau in the The main threats to this part of the (2016); Wilding (2017). Uluguru Mountains. Eastern Afromontane biodiversity Funariaceae. The narrowly oblong- hotspot are annual fires on the obovate, aristate leaves, bordered and Collected at an elevation of Lukwangule plateau that may ±2360 m a.s.l. along a rocky, toothed in the upper ⅔ and the change the ecology of the zygomorphic, peristomate capsules will peaty stream bank on the eastern edge of the afromontane grassland and spread differentiate E. heddersonii from other Lukwangule Plateau. into the forest, firewood collection, species in the genus. encroachment into the reserves and spread of the invasive Rubus sp.

20. Ephemerum diversifolium Mitt., Endemic to the Eastern Cape Only known from the type locality Magill (1987); Mucina et al. Harvey in Thesaurus Capensis 1: Province of South Africa: where it was collected during the (2006a), CEPF (2016). 63 (1859), pl. 100, fig. A. Uitenhage, Zwartkop’s River. first half of the 19th century by Zeyher. The plants are minute, Ephemeraceae. A very distinct species On soil in shrublands. making it difficult to find. with highly differentiated perichaetial Cultivation, invasive alien plants, leaves. urbanisation, pollution and mining activities are responsible for ongoing loss and degradation of habitat along the Swartkops River between Port Elizabeth and Uitenhage. The Albany Alluvial Vegetation is endangered and falls within the Maputaland–Pondoland– Albany biodiversity hotspot.

21. Fissidens capriviensis Magill in Endemic to Namibia. Collected Only known from the type locality Magill (1981); Van Rooy and Flora of Southern Africa, by Hans Vahrmeijer in the where it was found in 1980. Phephu (2016). Bryophyta 1: 45 (1981), fig. 9: Caprivi strip, Kongola area, Although the Kwando swamps are 1–7. Kwando swamps. relatively undisturbed and the area under-explored, agriculture is Fissidentaceae. The corticolous In gallery forest, on lower encroaching on the habitat in the substrate, well-defined limbidia, large trunk of Boscia sp. Kongola area. leaf cells and distinct costa distinguish the species.

22. Fossombronia nyikaensis Perold Endemic to the Nyika Plateau Only known from specimens Perold (2001a), The Nyika in Bothalia 31(1): 48 (2001). in northern Malawi. collected at the type locality in Vwaza (UK) Trust (2016); 2000. Although the locality is CEPF (2016). Fossombroniaceae. The ornamentation The species was found in protected in the Nyika National on the distal face of the spore is grassland, on soil in a rock Park, the native habitats on the distinctive. overhang, at 2227 m a.s.l. plateau are threatened by wildfires that fragment the forests, and invasive alien plants such as pine trees and the Himalayan raspberry. The Nyika Plateau is part of the Eastern Afromontane biodiversity hotspot.

23. Geocalyx orientalis Besch. et Endemic to Réunion: Cirque de Only known from one locality on Ah-Peng et al. (2012); Spruce in Bulletin de la Société Cilaos area. the island. The area of occupancy is Söderström et al. 2016, CEPF Botanique de France 36(suppl.): <4 km². Major threats are habitat (2016). 189 (1890). On soil and rotten wood in destruction and urbanisation in the montane forest, 1300–1500 m populated Cirque de Cilaos area. Geocalycaceae. According to the World a.s.l. The locality falls within the checklist of hornworts and liverworts Madagascar and the Indian Ocean (Söderström et al. 2016), the species is Islands biodiversity hotspot. not well known. Critically Endangered on Réunion: [B2ab(iii)] (Ah-Peng et al. (2012).

24. Gymnostomum lingulatum Endemic to the Limpopo Only known from the type locality Dixon and Gepp (1923); Rehmann ex Sim in Transactions Province of South Africa and not collected since 1879/80. Magill (1981); Mucina et al. of the Royal Society of South where it was collected ‘in The remaining indigenous forests Africa 15: 260 (1926). mont. Lechlaba ad Houtbosch’ and grassland in the Woodbush- (2006). by Antoni Rehmann and Haenertsburg area are threatened by Pottiaceae. The lanceolate to triangular, distributed as no. 437 in his afforestation, cultivation, invasive unistratose leaves with entire margins Musci Austro-Africani cont. alien plants, bush encroachment, and low, simple, scattered leaf cell exsiccatae. A specimen that overgrazing by livestock and urban papillae distinguish the species from resembles this species was development. The Woodbush others in southern Africa. collected near Dwesa in the Granite Grassland is critically Eastern Cape Province in 1985. endangered.

It probably grew on rock or soil in grassland, at ±1400 m a.s.l.

25. Haplolejeunea sticta Grolle in Réunion and Madagascar. The species is restricted to the Ah-Peng et al. (2012); Journal of the Hattori Botanical Madagascar and the Indian Ocean Söderström et al. (2016); Laboratory 39: 205 (1975), fig. On Réunion it grows on rotten Islands biodiversity hotspot. On CEPF (2016). 1–2. wood in lowlands, between Réunion, the species is known only 175 and 700 m a.s.l. from two localities in threatened Lejeuneaceae. According to the World lowland habitats. Deforestation and checklist of hornworts and liverworts forest degradation are major threats (Söderström et al. 2016), the species is to lowland habitats in Madagascar. not well known. Critically Endangered on Réunion [B2ab(iii)] (Ah-Peng et al. 2012) and probably on Madagascar as well.

26. Heteroscyphus spectabilis São Tomé and Príncipe, For more than 100 years this rare Wigginton (2004); Sérgio and (Steph.) Schiffn. in Cameroon. species was only known from São Garcia (2011); Pócs, Müller Oesterreichische Botanische Tomé and Cameroon. On São and Shevock (2015); CEPF Zeitschrift 60: 172 (1910). On tree bark in forest. On São Tomé it was collected by Moller in (2016). Tomé and Príncipe it was 1885 and in Cameroon the records Lophocoleaceae. The large trigones of found at elevations of 500 and date back to two specimens the leaf cells distinguish this species 1150 m a.s.l. collected by Dusén in the 1890s. It from others in West Africa. has recently (in 2012 and 2013) been found on the island of Príncipe by Shevock. Forest cover on São Tomé and Príncipe remains substantial, but growing population pressure in the rest of the Guinean Forests of West Africa biodiversity hotspot and the resulting demand for forested land is a serious threat to the remaining habitats.

27. Lejeunea gradsteiniana Pócs in Endemic to the Usambara Known from only two localities, 6 Pócs (2011); CEPF (2016). Acta Biologica Plantarum Mountains of Tanzania. km apart, in the Kwamkoro Forest Agriensis 1: 55 (2010)[2011]. Reserve of East Usambara, where it Nom. nov. pro Ceratolejeunea Epiphyllous on large was collected in 1911 and 1982. aberrans Steph. in Species Trichomanes sp. in dense, wet, The habitat is declining because of Hepaticarum 6: 399 (1923). tropical rain forest, 900– deforestation, intercropping and the 1000 m a.s.l. Associated with introduction of alien trees in the Lejeuneaceae. Previously other rare and endemic forest canopy. The reserve falls Cladolejeunea aberrans (Steph.) liverworts. within the Eastern Afromontane Zwickel. The only African species in biodiversity hotspot. the ‘Lejeunea eckloniana’ group with horned perianth keels. Endangered globally: B1+2cd (ver 2.3) (as Cladolejeunea aberrans) (IUCN 2017b).

28. Leucoloma crosbyi La Farge- Endemic to northern Only known from a couple of La Farge-England (1992, Engl. in Novon 2: 119 (1992). Madagascar: Diego Suarez, specimens collected about 5 km 1998); Jenkins et al. (2011); Parc National de Montagne apart by Marshall and Carol Crosby CEPF (2016). Dicranaceae. The species is d'Ambre. in 1972. Madagascar is the centre characterised by its robust habit, of diversity and endemism for narrow, opaque juxtacostal bands Corticolous and ramicolous in tapering to basal region, costal-laminal rainforest, 900–1200 m a.s.l. Leucoloma and the genus transition zone with 2–9 multi- to represents about 5% of the moss bistratose rows, interior cells forming flora of the Madagascar and the broad scarious region, narrow hyaline Indian Ocean Islands biodiversity margin and longitudinally thick-walled hotspot. The habitat in Parc alar cells. National de Montagne d'Ambre is under threat from encroachment as a result of charcoal extraction, cattle grazing, slash-and-burn agriculture, and rosewood collection.

29. Leucoperichaetium Endemic to Witputz in This monotypic genus is only Magill (1981); Van Rooy eremophilum Magill in Flora of southern Namibia, where it known from a single locality in the (2004); Legal Assistance Southern Africa, Bryophyta 1: was collected in 1978. Succulent Karoo biodiversity Centre of Namibia (2009); 273 (1981), fig. 81: 1–18. hotspot. The locality is threatened CEPF (2016). The plants grew on fine sand by mining activities which are Grimmiaceae. The species is unique over a quartzite outcrop in expanding and take precedence within the family because of the stark dwarf succulent shrublands, over all other land uses in Namibia, contrast between the small, bistratose ±1000 m a.s.l. The locality is in protected areas as well as private vegetative leaves and the large, hair- situated on the Namib land. pointed and colourless perichaetial escarpment, a transition zone leaves. between the Namib Desert and Vulnerable globally: D2 (ver 2.3) the central plateau. It is an arid, (IUCN 2017b). winter rainfall area.

30. Lopholejeunea minima Vanden Endemic to Réunion. The species was known from two Vanden Berghen (1984); Ah- Berghen in Bulletin du Jardin localities that were destroyed Peng et al. (2012); Botanique National de Belgique Epiphytic on tree trunks and during the 2007 volcanic eruption Söderström et al. (2016); 54: 437 (1984), fig. 21. fern fronds in lowland forest, and the species has not been CEPF (2016). 100–220 m a.s.l. observed since. It is restricted to the Lejeuneaceae. Vanden Berghen (1984) Madagascar and the Indian Ocean provided a key to African species of Lopholejeunea, but according to the Islands biodiversity hotspot. World checklist of hornworts and liverworts (Söderström et al. 2016), the Critically Endangered on species is not well known. Réunion: [B2ab(iii)] (Ah-Peng et al. (2012).

31. Lopholejeunea multilacera Madagascar, Mauritius and On Réunion the species is only Vanden Berghen (1984); Ah- Steph. in Botanical Gazette 15: Réunion. known from two localities. The Peng et al. (2012); 285 (1890). habitat is threatened with Söderström et al. (2016); On Réunion it is corticolous in destruction and degradation CEPF (2016). Lejeuneaceae. Vanden Berghen (1984) forest, 800–1350 m a.s.l. throughout the Madagascar and the provided a key to African species of Indian Ocean Islands biodiversity Lopholejeunea, but the broader hotspot. taxonomic relationships of L. multilacera are uncertain. Critically Endangered on Réunion [B2ab(iii)] (Ah-Peng et al. 2012), but most likely on Madagascar and Mauritius as well.

32. Ludorugbya springbokorum Endemic to the Western Cape This monotypic genus is only Hedderson and Zander Hedd. & R.H.Zander in Journal Province of South Africa. known from a few remnant (2007); CEPF (2016). of Bryology 29: 222 (2007). Renosterveld fragments in the On clay banks in Renosterveld Swartland region, in an area about Pottiaceae. The strongly differentiated fragments of the Fynbos the size of a half degree grid perichaetial leaves, immersed, urceolate Biome, at 150–250 m a.s.l. square. This vegetation type is one capsules with a very long, conic-mitrate of the most threatened in South operculum and a persistent, evertable Africa and falls within the Cape annulus are diagnostic features of the Floristic Region biodiversity species. hotspot. Ludorugbya springbokorum is suggested to be amongst the most threatened mosses in the world (Hedderson & Zander 2007).

33. Meiothecium fuscescens (A. Endemic to South Africa and Not collected since the early 1800s. Sim (1926); Driver et al. Jaeger ex Paris) Broth. in Die only known from the original The Fynbos, where this species was (2012); CEPF (2016). Natürlichen Pflanzenfamilien material, collected in the Cape found, is one of the most threatened I(3): 1103 (1908). (locality not precise) by Carl biomes in South Africa, classified Heinrich Bergius between as critically endangered. It is part of Sematophyllaceae. The smooth leaf 1815 and 1817. the Cape Floristic Region cells and single peristome place plants biodiversity hotspot. in Meiothecium, but the species has not Probably epiphytic in wooded been restudied recently. areas.

34. Neckeropsis pocsii Enroth & Endemic to the Comoros island Known only from the type locality Enroth and Magill (1994); Magill in The Bryologist 97: 171 of Mayotte. where it was collected in 1992. The Enroth (2002); CEPF (2016). (1994), fig. 1–9. Comoros belong to the Madagascar Growing on boulders in mesic and the Indian Ocean Islands Neckeraceae. The species resembles the evergreen forest, ±400 m a.s.l. biodiversity hotspot. The area of widely distributed N. disticha (Hedw.) occupancy is less than 10 km² and Kindb. in habit, but differs in the wider the habitat is declining as a result of leaf bases, stronger serrulate to crenate excessive logging. leaf apices, exserted capsules and filiform post-fertilisation paraphyses. Critically Endangered globally: B1+2c (ver 2.3) (IUCN 2017b).

35. Picobryum atomicum Endemic to South Africa and Restricted to the Olifants River Rebelo et al. (2006); Zander R.H.Zander & Hedd. in Journal only known from Beaverlac in section of Leipoldtville Sand and Hedderson (2011); CEPF of Bryology 33(2): 130 (2011). the Oliphants River Mountains Fynbos, another lowland vegetation (2016). of the Western Cape Province. type of the Fynbos Biome (Cape Pottiaceae. Plants of this monotypic Floristic Region biodiversity genus are very small with a Found in disturbed areas and hotspot) under severe threat. The cleistocarpic, globose, short apiculate patches of clayey mineral soil habitat is endangered by agriculture capsule and mitrate calyptra. It differs in Fynbos at ±300 m a.s.l. and invasive alien species. from Syntrichia Brid. in several respects.

36. Pocsiella hydrogonioides Bizot Endemic to Tanzania: This monotypic genus is known Frey and Stech (2009); CEPF in Cryptogamie: Bryologie, Kilimanjaro, ‘Bivouac Rocks’ from one locality only, where (2016) Lichénologie 1: 424 (1980), pl. along the Umbwe trail. mountaineers bivouac under the 1: fig. 2. overhanging cliffs, making fire. In large turfs on dripping The species was last observed in Dicranaceae. The leaves are flexuose, porphyry lava cliffs at 2900 m 1989 and is regarded as Critically ligulate and long-decurrent. The a.s.l. Endangered. Kilimanjaro falls laminal cells are large and smooth and within the Eastern Afromontane the alar cells undifferentiated. The biodiversity hotspot. peristome teeth are smooth.

37. Porella prolixa (Gottsche ex Endemic to Réunion, without a The species is known only from the Jones (1963); Ah-Peng et al. Steph.) E.W.Jones in precise locality. type collection. Urbanisation and (2012); CEPF (2016). Transactions of the British invasive alien species are the main Bryological Society 4: 460 Habitat unknown. The genus is threats to biodiversity on Réunion, (1963). rare in Africa, but species on which forms part of the the continent are rupicolous or Madagascar and the Indian Ocean Porellaceae. Porella prolixa appears to corticolous and occur at high Islands biodiversity hotspot. be distinct from other African species, altitude, > 2000 m a.s.l. none of which has apiculate leaves. Critically Endangered on Réunion: B2ab(i) (Ah-Peng et al. 2012).

38. Pterobryopsis rehmannii Magill Endemic to the KwaZulu-Natal The plants of P. rehmannii are Magill and Van Rooy (1998); in Memoirs of the Botanical Province of South Africa, large and like other members of the CEPF (2016). Survey of South Africa 43: 5 without precise locality, where genus probably cover large areas on (1979), fig. 33–41. it was collected by Buchanan trees and rocks in closed forests, between 1861 and 1874. which are protected in several Pterobryaceae. The broadly ovate to reserves. It is therefore curious that oval leaves with costa extending to From substrate fragments in the species has not been recollected above midleaf and upper margins the specimen, it appears that during many field trips to the area. inflexed and serrulate, distinguish this species from others in the region. the species grew on tree bark. Habitat in the Maputaland– Pondoland–Albany biodiversity hotspot is threatened by cultivation, plantation forestry and urbanisation.

39. Renauldia lycopodioides Bizot Endemic to the Eastern Arc The Eastern Arc mountains form Pócs (2000); Bytebier and ex Pócs in Folia Historico- mountains and known from the part of the Eastern Afromontane Chuah-Petiot (2002); CEPF naturalia Musei Matraensis 4: Western Usambara, Ukagura biodiversity hotspot, characterised (2016). 33 (1976)[1977]. and Udzungwa mountains in by a series of montane ‘islands’ and Tanzania and the Taita Hills in plateaus. The habitat is seriously Pterobryaceae. This moss is described south-eastern Kenya. threatened by deforestation for as spectacular, with tail-like stems agricultural land and firewood as about 10 cm long and 4 mm thick. The Epiphytic on tree branches in well as large-scale logging, leaves are ovate to panduriform, moist mossy elfin forest, 750– resulting in forest degradation and concave, with a short, often channelled 2200 m a.s.l. fragmentation. apiculus. The capsules are yellow, almost sessile and hang from the Endangered globally: B1+2cd (ver branches. 2.3) (IUCN 2017b).

40. Riccia alatospora O.H.Volk et Endemic to the Western Cape Since its initial collection in 1929, Perold (1999, 2008); Rebelo Perold in Bothalia 15: 534 Province, South Africa. the species is only known from et al. (2006); CEPF (2016). (1985), fig. 4–6. Platklip near Stellenbosch. Fynbos Biome. Annual on Although the Platklip locality is Ricciaceae. The species is distinguished shallow, sandy soil in hollows locally protected, much of the from the closely related R. duthieae on a dome-shaped granite surrounding Renosterveld O.H.Volk & Perold by its smaller thalli outcrop, ±80 m a.s.l. vegetation has been destroyed by but larger spores with very broad and urban development, viniculture and often perforated and eroded wings. alien invasive plants. This area is part of the Cape Floristic Region biodiversity hotspot.

Vulnerable globally: D2 (ver 3.1) (IUCN 2017b).

41. Riccia hantamensis Perold in Endemic to the Northern Cape Only known from two localities, Perold (1999); Van der Bothalia 19: 157 (1989), fig. 2– Province of South Africa. ±260 km apart, in the Succulent Merwe, Van Rooyen and Van 4: A–F: a. Originally described from the Karoo, a global biodiversity Rooyen (2008); CEPF (2016). Hantamsberge at Calvinia and hotspot. Overgrazing as a result of tentatively reported from the Ricciaceae. The robust size of the inadequate farming practices and plants will distinguish it from the Carolusberg near Springbok. In the Hantam, the species grows invasive alien plants are the closely related R. alatospora. on neutral to somewhat greatest threats to the vegetation in alkaline soil, together with the Hantam-Roggeveld region, small Crassula spp. and identified as a priority area for mosses, for example, conservation. Didymodon tophaceus (Brid.) Lisa, in an area with predominantly Fynbos vegetation. Altitude of ±1500 m a.s.l. with winter rainfall of less than 200 mm p.a.

42. Riccia hirsuta O.H.Volk et Endemic to the Kamiesberge of Only known from collections made Volk and Perold (1990); Perold in Bothalia 16: 187 Namakwaland in the Northern between 1977 and 1988, at two Perold (1999); CEPF (2016). (1986), fig. 1–3. Cape Province of South Africa. localities ±20 km apart. The Namaqualand Uplands is one of the Ricciaceae. Riccia hirsuta is It grows on sandy soil amongst priority areas for conservation in distinguished from other species in bushes in seasonally damp the Succulent Karoo biodiversity section Pilifer by the very tall, shiny places or seepage areas, ±1160 hotspot. Overgrazing by livestock dorsal cell pillars and by the triangular m a.s.l. in a winter rainfall area and cultivation along water courses scales, apically split into filamentous, with precipitation less than 200 are significant threats to the habitat. short and loose cellular strands. mm p.a.

43. Riccia rubricollis Garside et Endemic to Knysna in the This rare endemic is only known Perold (1991, 1999); Rebelo A.V.Duthie ex Perold in Western Cape Province of from a couple of sites on opposite et al. (2006); CEPF (2016). sides of the Knysna lagoon, ±3 km Bothalia 21(1): 51 (1991). South Africa. apart, collected by A.V. Duthie in the late 1920s. The Knysna Sand Ricciaceae. The species can be The species grows at low Fynbos as well as Garden Route distinguished by the conspicuous, altitude on turf in damp, shady Shale Fynbos vegetation types are mostly dark purple antheridial necks of places, frequently mixed with endangered and the habitat is the male plant and the spores with other bryophytes. threatened by urban sprawl, large, deep-set areolae on the distal encroachment from invasive alien face. vegetation, plantation forestry and agricultural activities. It falls within the Cape Floristic Region biodiversity hotspot.

44. Riccia sibayenii Perold in Endemic to Mpumalanga Known only from the type locality, Perold (2001b); Rutherford et Bothalia 31(1): 151 (2001). Province of South Africa. where it was found in 2000. al. (2006); CEPF (2016). Although the Tshokwane-Hlane Ricciaceae. The species is more The species grew on black Basalt Lowveld vegetation is delicate than R. rubricollis and lacks loamy soil of the Savanna classified as least threatened, the purple coloration. The spore Biome at an altitude of 270 m habitat at the type locality is ornamentation also differs. a.s.l. threatened by cultivation and border patrols in South Africa, and overgrazing by livestock across the border in Mozambique. The locality falls within the Maputaland–Pondoland–Albany biodiversity hotspot.

45. Sauteria nyikaensis Perold in Endemic to the Nyika Plateau The species is known only from the Perold (2003); Söderström et Bothalia 33(2): 167 (2003). of northern Malawi. type locality, where it was collected al. (2016); The Nyika Vwaza in 2000. Although the locality is (UK) Trust (2016); CEPF Cleveaceae. The species is It grew on calcareous soil protected in the Nyika National (2016). distinguished from others in the genus containing slivers of mica, in a Park, the ecology of the plateau is by a combination of thallus, ventral small, cave-like cavity at the threatened by wildfires that scale, stolon, and male and female base of a large rock, at altitude fragment the forests as well as inflorescence characters. However, it is 2343 m a.s.l. Associated with invasive alien plants such as pine treated as a doubtful species in the Lunularia cruciata (L.) trees and the Himalayan raspberry. World checklist of hornworts and Dumort. ex Lindb. and The Nyika Plateau forms part of the liverworts (Söderström et al. 2016). Plagiochasma eximium Eastern Afromontane biodiversity (Schiffn.) Steph. hotspot.

46. Sematophyllum rheophyticum Endemic to Rwanda: Gisakura, Known from a single locality where Crawford (2012); Buck and W.R.Buck & Hedd. in Bothalia Nyungwe National Park. it was collected in 2012. The Hedderson (2016); CEPF 46(1) (2016). species is one of few in the genus (2016). Grows on highly mineralised (two in Africa) that occurs in Sematophyllaceae. The species is schistose rock in a swift rheophytic habitats. The type distinguishable from others in Africa by flowing mountain stream material was associated with the a unique combination of characters below a waterfall, at 1845 m two rare species Bryocrumia including the concave, abruptly a.s.l. vivicolor (Broth. & Dixon) acuminate leaves, the alar areolation W.R.Buck and Scopelophila and the rheophytic habitat. ligulata (Spruce) Spruce. Threats to the Nyungwe Forest include forest fires, deforestation for firewood and construction materials as well as infrastructure development. The locality is situated in the Eastern Afromontane biodiversity hotspot.

47. Streptocalypta pulchriretis Endemic to the Drakensberg Known only from the type locality Dixon and Wager (1929); (Dixon) R.H.Zander in Bulletin Mountains of KwaZulu-Natal, where it was collected by H.A. Magill (1981); CEPF (2016). of the Buffalo Society of Natural South Africa. Wager in 1918. This locality is Sciences 32: 96 (1993). protected in the UKhahlamba Collected on dry banks in the Drakensberg Park (Royal Natal) Pottiaceae. Originally described as Gudu forest. and it is curious that this species Weisiopsis pulchriretis and later has not been recollected during transferred to Streptocalypta, the many trips to the area. Habitat in species is distinguished by the narrowly the Maputaland–Pondoland– elliptical leaves, extension of basal leaf Albany biodiversity hotspot is cells up the margins and paroicous threatened by cultivation, plantation sexual condition. forestry and urbanisation.

48. Symbiezidium madagascariense Endemic to the East African Not sighted on Madagascar since Gradstein (2001b); CEPF Steph. in Species Hepaticarum 5: islands of Madagascar (near St. 1904, notwithstanding intensive (2016); IUCN (2017a). 99 (1912). Marie) and the Seychelles collecting in recent years. In the (Mahé). Seychelles it is known from a Lejeuneaceae. The only species of the single collection made on Mahé in genus in Africa and it constitutes a Plants grow in small, brown- the 1960's. The remaining lowland separate subgenus, Eosymbiezidium. green mats on bark in lowland rainforest on Madagascar and the This species is morphologically quite rainforest. Seychelles, part of the Madagascar distinct from the neotropical species. and the Indian Ocean Islands biodiversity hotspot, is under severe threat of deforestation.

Endangered globally: B1+2cd (Gradstein 2001b).

49. Xylolejeunea grolleana (Pócs) Endemic to the East African Known from only two lowland Pócs (1999); He and Grolle Xiao L.He et Grolle in Annales islands of Madagascar and rainforest localities in the (2001); Ah-Peng et al. (2012); Botanici Fennici 38: 32 (2001). Réunion. Madagascar and the Indian Ocean CEPF (2016). Islands biodiversity hotspot. In Lejeuneaceae. Originally described in In Madagascar it grows in Madagascar the habitat is severely Trachylejeunea (Spruce) Steph., the humid lowland rainforest at threatened with deforestation and in species is easily recognised by its small low elevation, 220–260 m a.s.l. Réunion the species is very rare and leaf lobe with two transverse rows of In Réunion it was found on the endangered as a result of forest considerably enlarged and elongated southern slope of the active fragmentation. cells and ocelli at the base, and high, volcano Piton de La Fournaise. finger-like papillae on the dorsal Critically Endangered on Réunion surface. [B2ab(iii)] (Ah-Peng et al. 2012) and most likely on Madagascar as well.

50. Xylolejeunea muricella Xiao Endemic to the Seychelles Only known from the type locality He and Grolle (2001); CEPF L.He et Grolle in Annales island of Mahé. where it could not be re-found (2016). Botanici Fennici 38: 34 (2001). during a thorough search of the It grew on shaded rocks, mixed area. There is almost no rainforest Lejeuneaceae. The species differs from with Bazzania approximata left on the Seychelles, and the the closely related X. grolleana in the Onr., at an altitude of 40 m species may already be extinct. The small, spherical papillae on the dorsal a.s.l. surface of the leaf lobe, conspicuously island forms part of the Madagascar denticulate leaf margins, 3 or 4 lateral and the Indian Ocean Islands marginal cells of the lobule and biodiversity hotspot. orbicular underleaves. †, Note that not all of the assignments follow IUCN methodology consequently; and that many of the global assignments are in need of revision.

References

Ah-Peng, C., Bardat, J., Pócs, T., Söderström, L., Staménoff, P. & Strasbergh, D., 2012, ‘Red List of liverworts and hornworts for Réunion (Mascarene archipelago)’, Phytotaxa 68, 1–23. Anonymous, 2017, Fort Hill and the Cemetery at Hartley Hills goldfield, ZimFieldGuide.com, viewed 18 August 2017, from http://zimfieldguide.com/mashonaland-west/fort-hill-and-cemetery-hartley-hills-goldfield. Buck, W.R., 1980, ‘A re-interpretation of the Fabroniaceae: Additions and corrections’, Journal of the Hattori Botanical Laboratory 47, 45–55. Buck, W.R. & Hedderson, T.A.J., 2016, ‘Sematophyllum rheophyticum (Bryopsida, Sematophyllaceae), a new rheophytic species from Rwanda’, Bothalia 46(1), a2044. https://doi.org/10.4102/abc.v46i1.2044 Bytebier, B. & Chuah-Petiot, M., 2002, ‘A preliminary checklist of the bryoflora of the Taita Hills, Kenya’, Tropical Bryology 22, 55–66. Crawford, A., 2012, Conflict-Sensitive Conservation in Nyungwe National Park: Conflict analysis, The International Institute for Sustainable Development (IISD) Repost, viewed 18 August 2017, from http://www.iisd.org/pdf/2012/csc_nyungwe_conflict_analysis.pdf. Critical Ecosystem Partnership Fund (CEPF), 2016, The Biodiversity Hotspots, viewed 20 October 2017, from http://www.cepf.net/resources/hotspots/Pages/default.aspx. Dixon, H.N. & Gepp, A., 1923, ‘Rehmann’s South African mosses’, Kew Bulletin 6, 193–238. Dixon, H.N. & Wager, H.A., 1929, ‘New and Noteworthy Mosses from South Africa’, Transactions of the Royal Society of South Africa 18, 247– 261. Driver, A., Sink, K.J., Nel, J.L., Holness, S.H., Van Niekerk, L., Daniels, F. et al., 2012, ‘National Biodiversity Assessment 2011: An assessment of South Africa’s biodiversity and ecosystems’ Synthesis report, South African National Biodiversity Institute & Dept. of Environmental Affairs, Pretoria. Enroth, J., 2002, Neckeropsis pocsii Enroth & Magill, IUCN SSC Bryophyte Specialist Group fact sheet, viewed 30 October 2017, from http://www.slu.se/globalassets/ew/subw/artd/bryophyte-conservation/bryoconservation/pdf-artfaktablad/faktablad/neckeropsis_pocsii.pdf. Enroth, J. & Magill, R.E., 1994, ‘Neckeropsis pocsii (Neckeraceae, Musci) a new species from Comoro Islands’ The Bryologist 97(2), 171–173. Frey, W. & Stech, M., 2009, ‘Marchantiophyta, Bryophyta, Anthocerotophyta’ in W. Frey, M. Stech & E. Fischer (eds.), Syllabus of Plant Families, vol. 3, 13th edn., pp. 9–259, Schweizerbart, Stuttgart. Frontier.ac.uk, 2014, Uluguru Biodiversity Assessment Project 2004–2005, viewed 29 May 2017, from https://www.frontier.ac.uk/About- Us/Research/Past-Research-Projects.aspx. Gradstein, S.R., 2001a, Caudalejeunea grolleana Gradst, IUCN SSC Bryophyte Specialist Group fact sheet, viewed 08 August 2017, from http://www.slu.se/globalassets/ew/subw/artd/bryophyte-conservation/bryoconservation/pdf- artfaktablad/faktablad/caudalejeunea_grolleana.pdf. Gradstein, S.R., 2001b, Symbiezidium madagascariense Steph., IUCN SSC Bryophyte Specialist Group fact sheet, viewed 13 July 2017, from https://www.slu.se/globalassets/ew/org/centrb/artd/bryophyte-conservation/bryoconservation/pdf- artfaktablad/faktablad/symbiezidium_madagascariense.pdf. Green, G.M. & Sussman, R.W., 1990, ‘Deforestation history of the eastern rain forests of Madagascar’ Science 248(4952), 212–215. Hartmann, F.A., Wilson, R., Gradstein, S.R., Schneider, H. & Heinrichs, J., 2006, ‘Testing hypotheses on species delimitations and disjunctions in the liverwort Bryopteris (Jungermanniopsida: Lejeuneaceae), International Journal of Plant Sciences 167(6), 1205–1214. He, X. & Grolle, R., 2001, ‘Xylolejeunea, a new genus of the Lejeuneaceae (Hepaticae) from the neotropics, Madagascar and the Seychelles’, Annales Botanici Fennici 38, 25–44. Hedderson, T.A. & Zander, R.H., 2007, ‘Ludorugbya springbokorum (Pottiaceae) a new moss genus and species from the Western Cape Province of South Africa’, Journal of Bryology 29(4), 222–227. International Union for Conservation of Nature (IUCN), 2017a, Illegal activities threaten natural World Heritage, pushing the vaquita to the brink and depleting forests, viewed 10 July 2017, from https://www.iucn.org/news/iucn-41whc/201706/illegal-activities-threaten-natural-world- heritage-pushing-vaquita-brink-and-depleting-forests-%E2%80%93-iucn?dm_i=2GI3,13OT1,40ELQB,3C6Q3,1. International Union for Conservation of Nature (IUCN), 2017b, The IUCN Red List of Threatened Species. Version 2017–2, viewed 23 October 2017, from http://www.iucnredlist.org/. Jenkins, R.K.B., Andreone, F., Andriamazava, A., Anjeriniaina, M., Glaw, F., Rabibisoa, N. et al., 2011, Brookesia ambreensis, The IUCN Red List of Threatened Species 2011: e.T172930A6943410, viewed 13 November 2017, from https://doi.org/10.2305/IUCN.UK.2011- 2.RLTS.T172930A6943410.en. Jones, E.W., 1963, ‘African hepatics. XVI. Porella in tropical Africa’, Transactions of the British Bryological Society 4, 446–461. La Farge-England, C., 1992, ‘Leucoloma crosbyi (Dicranaceae), a new species endemic to northern Madagascar’, Novon 2, 119–122. La Farge-England, C., 1998, ‘The infrageneric phylogeny, classification, and phytogeography of Leucoloma (Dicranaceae, Bryopsida)’, The Bryologist 101(2), 181–220. Legal Assistance Centre of Namibia, 2009, Striking a better balance: An investigation of mining practices in Namibia’s protected areas, Standford Law School, John Meinert Printing, Windhoek. Magill, R.E., 1981, Flora of Southern Africa: Bryophyta. Part 1 Mosses. Fascicle 1 Sphagnaceae–Grimmiaceae, Botanical Research Institute, South Africa. Magill, R.E, 1987, Flora of Southern Africa: Bryophyta. Part 1 Mosses. Fascicle 2 Gigaspermaceae–Bartramiaceae, Botanical Research Institute, South Africa. Magill, R.E. & Van Rooy, J., 1998, Flora of Southern Africa: Bryophyta. Part 1 Musci. Fascicle 3 Erpodiaceae–Hookeriaceae, National Botanical Institute, Pretoria. Malombe, I., Fischer, E. & Pócs, T., 2010, ‘Cheilolejeunea ulugurica (Lejeuneaceae, Marchantiophyta), a new species from Tanzania’, Acta Biologica Plantarum Agriensis 1, 23–28. Mucina, L., Hoare, D.B., Lötter, M.C., du Preez, J., Rutherford, M.C., Scott-Shaw, C.R. et al., 2006, ‘Grassland Biome’, in L. Mucina & M.C. Rutherford (eds.), The vegetation of South Africa, Lesotho and Swaziland, pp. 348–437, Strelitzia 19, South African National Biodiversity Institute, Pretoria, South Africa. Mucina, L., Rutherford, M.C., W. Powrie, L.W., Gerber, J., Bezuidenhout, H., Sieben, E.J.J. et al., 2006a, ‘Inland azonal vegetation’, in L. Mucina & M.C. Rutherford (eds.), The vegetation of South Africa, Lesotho and Swaziland, pp. 616–657, Strelitzia 19, South African National Biodiversity Institute, Pretoria, South Africa. Müller, F. & Pócs, T., 2007, ‘A contribution to the knowledge of epiphyllous bryophytes of Bioko Island (Equatorial Guinea), including additional remarks on non-epiphyllous species’, Journal of Bryology 29(2), 81–94. Olson, D.M. & Dinerstein, E., 2002, ‘The Global 200: Priority ecoregions for global conservation’, Annals of the Missouri Botanical Garden 89(2), 199–224. Perold, S.M., 1991, ‘Studies in the genus Riccia (Marchantiales) from southern Africa. 22. R. rubricollis, now validated, typified and described’, Bothalia 21(1), 51–54. Perold, S.M., 1994, ‘Studies in the Marchantiales (Hepaticae) from southern Africa. 7. The genus Cryptomitrium (Aytoniaceae) and C. oreades sp. nov.’, Bothalia 24(2), 149–152. Perold, S.M., 1999, Flora of Southern Africa: Hepatophyta. Part 1: Marchantiopsida. Fascicle 1: Marchantiidae, National Botanical Institute, Pretoria. Perold, S.M., 2001a, ‘Notes on African Plants. Fossombronia nyikaensis, a new species from Malawi’, Bothalia 31(1), 48–52. Perold, S.M., 2001b, ‘Studies in the genus Riccia (Marchantiales) from southern Africa. 25. A new species in subgenus Ricciella, section Ricciella’, Bothalia 31(2), 151–154. Perold, S.M., 2003, ‘Notes on African Plants: Cleveaceae –Marchantiales. Sauteria nyikaensis, a new liverwort species from Malawi’, Bothalia 33(2), 165–171. Perold, S.M., 2008, Riccia alatospora, The IUCN Red List of Threatened Species 2008: e.T133734A3876785, viewed 14 November 2017, from https://doi.org/10.2305/IUCN.UK.2008.RLTS.T133734A3876785.en. Pócs, T., 1975, ‘New or little known epiphyllous liverworts. I. Cololejeunea from tropical Africa’, Acta Botanica Academiae Scientiarum Hungaricae 21, 353–375. Pócs T., 1997, ‘New or little known epiphyllous liverworts, VII. Two new Lejeuneaceae species from the Mascarene Islands’, Cryptogamie, Bryologie-Lichénologie 18(3), 195–205. Pócs T., 1999, ‘Trachylejeunea grolleana, a new representative of the Neotropical subgenus Hygrolejeuneopsis in Madagascar’, Haussknechtia Beiheft 9, 283–290. Pócs, T., 2000, Renauldia lycopodioides Bizot ex Pócs, IUCN SSC Bryophyte Specialist Group fact sheet, viewed 14 November 2017, from https://www.slu.se/globalassets/ew/subw/artd/bryophyte-conservation/bryoconservation/pdf- artfaktablad/faktablad/renauldia_lycopodioides.pdf. Pócs, T., 2011, ‘What is Cladolejeunea Zwickel? New or little known epiphyllous liverworts, XV’, Acta Biologica Plantarum Agriensis 1, 53–62. Pócs, T., Müller, F. & Shevock, J.R., 2015, ‘Additions to the liverwort and hornwort flora of São Tomé and Príncipe II, with Neurolejeunea, a genus new to Africa’, Herzogia 28(1), 50–69. Rebelo, A.G., Boucher, C., Helme, N., Mucina, L. & Rutherford, M.C., 2006, ‘Fynbos Biome’, in L. Mucina & M.C. Rutherford (eds.), The Vegetation of South Africa, Lesotho and Swaziland, pp. 53–219, Strelitzia 19, South African National Biodiversity Institute, Pretoria. Richards, P.W., 1952, ‘Notes on African mosses. I. Andreaea camerunensis sp. nov. and other mosses mostly from Nigeria and the British Cameroons’, Transactions of the British Bryological Society 2, 66–70. Rutherford, M.C., Mucina, L., Lotter, M.C., Bredenkamp, G.J., Smit, J.H.L., Scott-Shaw, A. et al., 2006, ‘Savanna Biome’, in L. Mucina & M.C. Rutherford (eds.), The vegetation of South Africa, Lesotho and Swaziland, pp. 438–539, Strelitzia 19, South African National Biodiversity Institute, Pretoria. Sass-Gyarmati, A., 2017, ‘Bazzania konratiana sp. nov. (Marchantiophyta: Lepidoziaceae) from Madagascar’, Cryptogamie, Bryologie 38(2), 119– 124. Sérgio, C. & Garcia, C., 2011, ‘Bryophyte flora of São Tomé e Príncipe Archipelago (West Africa): Annotated catalogue’, Cryptogamie, Bryologie 32(2), 145–196. Sim, T.R., 1926, ‘The Bryophyta of South Africa’, Transactions of the Royal Society of South Africa 15, The Royal Society of South Africa, Cape Town. Snider, J.A., 1975, ‘Three New Species of Archidium from Africa’, The Bryologist 78(2), 152–159. Söderström, L., Hagborg, A., von Konrat, M., Bartholomew-Began, S., Bell, D., Briscoe, L. et al., 2016, ‘World checklist of hornworts and liverworts’, PhytoKeys 59, 1–828. Szabó, A. & Pócs, T., 2016, ‘New or little known epiphyllous liverworts, XX Cololejeunea nosykombae A.Szabó & Pócs sp.nov. from Madagascar’, Journal of Bryology 38(4), 302–307. The Nyika Vwaza (UK) Trust, 2016, Nyika National Park, viewed 20 July 2017, from http://www.nyika-vwaza-trust.org/nyika/nyika.html. Vanden Berghen, C., 1984, ‘Le genre Lopholejeunea (Spruce) Schiffn. en Afrique’, Bulletin du Jardin Botanique National de Belgique 54, 393–464. Van der Merwe, H., Van Rooyen, M.W. & Van Rooyen, N., 2008, ‘Vegetation of the Hantam-Tanqua-Roggeveld subregion, South Africa Part 2: Succulent Karoo Biome related vegetation’, Koedoe 50(1), 160–183. Van Rooy, J., 2004, Leucoperichaetium eremophilum Magill, IUCN SSC Bryophyte Specialist Group fact sheet, viewed 05 May 2017, from https://www.slu.se/globalassets/ew/org/centrb/artd/bryophyte-conservation/bryoconservation/pdf- artfaktablad/faktablad/leucoperichaetium_eremophilum.pdf. Van Rooy, J., 2011, ‘The type locality of Hypnofabronia marginata Dixon’, Journal of Bryology 33(4), 327–328. Van Rooy, J. & Phephu, N., 2016, ‘Centres of moss diversity in southern Africa’, Bryophyte Diversity and Evolution 38(1), 27–39. Volk, O.H. & Perold, S.M., 1990, ‘Studies in the genus Riccia (Marchantiales) from southern Africa. 15. R. hirsuta and R. tomentosa, sp. nov., two distinct species previously treated as one’, Bothalia 20(1), 23–29. Wigginton, M.J. (ed.), 2004, ‘E. W. Jones's Liverwort and Hornwort Flora of West Africa’, Scripta Botanica Belgica 30, National Botanic Garden (Belgium), Meise. Wigginton, M., 2018, ‘Checklist and distribution of the liverworts and hornworts of sub-Saharan Africa, including the East African Islands (edition 4, 25 June 2018)’, Tropical Bryology Research Reports 9, 1–138. Wilding, N., 2017, ‘Three new species of Entosthodon Schwägr. (Bryopsida, Funariaceae) from sub-Saharan Africa’, Phytotaxa 312(1), 103–110. Zander, R.H. & Hedderson, T.A., 2011, ‘Picobryum, a new genus of Pottiaceae (Bryophyta) from South Africa, and an erratum for Acaulonopsis’, Journal of Bryology 33(2), 130–133.