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Bothalia - African Biodiversity & Conservation
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Authors:
Jacques van Rooy1,2 Ariel Bergamini3 Irene Bisang4 Affiliations:
1National Herbarium, South African National Biodiversity Institute, South Africa
2School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, South Africa
3Department of Biodiversity and Conservation Biology, Swiss Federal Research Institute WSL, Switzerland
4Research Division Directorate & Department of Botany, Swedish Museum of Natural History, Sweden Corresponding author:
Jacques van Rooy, j.vanrooy@sanbi.org.za Dates:
Received: 18 Jan. 2018 Accepted: 23 Aug. 2018 Published: 14 Jan. 2019 How to cite this article:
Van Rooy, J., Bergamini, A. &
Bisang, I., 2019, ‘Fifty shades of red: Lost or threatened bryophytes in Africa’, Bothalia 49(1), a2341. https://doi.org/
10.4102/abc.v49i1.2341 Copyright:
© 2019. The Authors.
Licensee: AOSIS. This work is licensed under the Creative Commons Attribution License.
Introduction
The bryoflora of sub-Saharan Africa and the East African islands is poorly known and recent taxonomic and floristic activity is minimal in comparison to other regions of the world (Diop et al.
2018; Hallingbäck & Hodgetts 2000; Magill 2010; O’Shea 2005; Wilding 2017). Recent, but still incomplete, floras and identification guides are available only for southern Africa (Magill 1981, 1987; Magill & Van Rooy 1998; Perold 1999), tropical Africa (De Sloover 2003), Kenya (Chuah- Petiot 2003; Wilding et al. 2016), West Africa (Wigginton 2004) and Rwanda (Fischer 2013).
However, a series of regional checklists with accepted names, synonyms, literature references and 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 for research on the taxonomy, ecology and conservation of African bryophytes.
The documentation of plant diversity and its urgent conservation are priority objectives of the Updated Global Strategy for Plant Conservation 2011–2020 (Convention on Biological Diversity [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 2012a). The IUCN Species Survival Commission (SSC) lists the expansion of geographic and taxonomic coverage of the IUCN Red Lists as its top priority (IUCN 2017b). The IUCN Red List of Threatened Species for bryophytes (IUCN 2017e; Tan et al. 2000) currently contains 102 species, including several from Africa. However, most of these assessments are more than 10 years old and thus in need of careful revision. There is currently no Red List for African bryophytes and the only Background: A Red List of threatened bryophytes is lacking for Africa. The International Union for Conservation of Nature (IUCN) Species Survival Commission (SSC) Bryophyte Specialist Group has recently launched the ‘Top 10 Initiative’ to identify the 10 species on each continent that are at highest risk of extinction.
Objectives: The main aim of this paper was to highlight some of the lost or strongly threatened bryophyte species in sub-Saharan Africa and the East African islands and to draw up a Top 10 list for Africa.
Method: Lost or threatened species have been identified with the help of experts on the bryoflora of Africa, global and regional Red Lists and taxonomic literature. Each species on this candidate list is discussed at the hand of its taxonomy, distribution, habitat, threat and current global or regional Red List status as far as previously assessed.
Results: Fifty bryophyte species, representing 40 genera and 23 families, have been identified as Top 10 candidates. Of these, 29 are endemic to Africa and 21 are restricted to the East African 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.
Conclusion: This is the first list of lost or threatened bryophytes for Africa and the first Top 10 list of the IUCN Bryophyte Specialist Group. It represents an important step towards regional and global Red List assessment of bryophytes, thus meeting the targets of the Updated Global Strategy for Plant Conservation 2011–2020 and priorities of The Shenzhen Declaration on Plant Sciences.
Fifty shades of red: Lost or threatened bryophytes in Africa
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http://www.abcjournal.org Open Access regional Red List is the list of threatened liverwort and
hornwort species compiled for the East African island of Réunion (Ah-Peng et al. 2012).
To rigorously assess the estimated 18 000 bryophyte species globally (Frey & Stech 2009), following IUCN methodology, which is now generally accepted and advocated by, for example, Vanderpoorten and Hallingbäck (2009:488), is an unsurmountable task given the current personnel and economic resources devoted to bryophyte conservation. Such an endeavour of IUCN red-listing is currently underway for the ca. 1800 European bryophyte species (IUCN 2017c). To reinitiate red-listing activities on a global scale, the IUCN SSC Bryophyte Specialist Group has launched the so-called ‘Top 10 Initiative’
(IUCN 2017d; IUCN SSC 2016) to identify a minimum of 10 species at high risk of extinction from each continent and properly assign them to an IUCN Red List category (IUCN 2012a; IUCN Standards and Petition Subcommittee 2017).
It is generally accepted that centres of diversity and endemism, or so-called ‘biodiversity hotspots’, are priority areas for conserving biodiversity (Geffert et al. 2013; Marchese 2015).
Conservation International, through the Critical Ecosystem Partnership Fund (CEPF) (2016a), recognises 36 global biodiversity hotspots. A biodiversity hotspot is characterised by high levels of vascular plant endemism (>1500 endemic species) and 30% or less of its original natural vegetation remaining (Conservation International 2017). Many of these are heavily threatened by habitat loss and other human activities (CEPF 2016a; Mittermeier et al. 2004). The hotspot system is used to assess global conservation priorities and to provide grants to non-governmental and private sector organisations through the CEPF (2016b; Conservation International 2017).
Eight of the global biodiversity hotspots are found in sub-Saharan Africa and the East African islands (Figure 1):
Source: Based on the Biodiversity hotspots map downloaded from http://www.cepf.net/where_we_work/Pages/map.aspx, figure licensed under the Creative Commons Attribution-Share Alike 4.0 International license (Author: Conservation International)
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 Conservation International (CEPF 2016a). An additional four candidate species are only known from inland areas of southern Africa outside the Biodiversity hotspots (not shown on map).
40
10 10 20 30 40 50 60
0 800 1600 km
0 30
20 10 0 10 20 30
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http://www.abcjournal.org Open Access Cape Floristic Region, Coastal Forests of Eastern Africa,
Eastern Afromontane, Guinean Forests of West Africa, Horn of Africa, Madagascar and the Indian Ocean Islands, Maputaland–Pondoland–Albany and Succulent Karoo biodiversity hotspots.
Recent analyses of bryophyte distributions at a global, continental or regional scale (Geffert et al. 2013; Tan &
Pòcs 2000; Vanderpoorten & Hallingbäck 2009; Van Rooy &
Phephu 2016; Von Konrat et al. 2008) indicate that African centres of bryophyte diversity and endemism partly coincide with the biodiversity hotspots defined on the basis of vascular plants. The most species-rich areas are Madagascar, especially the lowland forests along the east coast, the mountains of eastern Africa, the Afromontane and Cape Floristic regions of South Africa, and West Africa.
The aim of this paper is to highlight lost and threatened bryophytes in sub-Saharan Africa and the East African islands and to draw up a Top 10 list of bryophytes in Africa that are at highest risk of extinction. The Top 10 species will be subjected to a careful IUCN Red List assessment to be included in The IUCN Red List of Threatened Species, and they will be further evaluated (IUCN 2012b) to identify and design the most urgent conservation actions. We also test the hypothesis that most lost and threatened bryophytes on our list fall within the global biodiversity hotspots recognised by Conservation International (CEPF 2016a).
Research method and design
The area covered in this publication comprises sub-Saharan Africa, together with the Atlantic and Indian Ocean islands in the tropics (Figure 1). This corresponds to the area covered by the checklists of O’Shea (2006) and Wigginton (2018), except for the islands of Cape Verde, Ascension and St. Helena, which are excluded here.
Experts on the taxonomy, ecology and conservation of African bryophytes were invited to contribute to two initiatives, namely the ‘Top 10 Initiative’ by the IUCN SSC Bryophyte Specialist Group (IUCN SSC 2016) and the ‘Search for Lost Species’ initiative (Global Wildlife Conservation 2014). Global Wildlife Conservation approached IUCN Specialist Groups in 2014 to suggest candidate species for a
proposal to search in the field for species that have gone unrecorded for years or decades (funding pendent).
Inspired by these two initiatives, a list of lost and threatened species (Online Appendix 1) was compiled based on the experts’ and the first author’s (J.v.R.) contributions. The following criteria were applied in the selection of species:
• The species should be endemic to sub-Saharan Africa or the East African islands.
• Known from a single or few localities with a narrow geographical distribution range.
• The habitat should be threatened and declining or the known records date from previous centuries despite more recent collecting activity or searches in the area.
The list of 50 species includes all threatened African bryophytes listed in the global IUCN Red List of Threatened Species (IUCN 2017e; Tan et al. 2000), and the threatened African endemics identified as ‘Critically Endangered’ in the Red List of liverworts and hornworts for Réunion (Ah-Peng et al. 2012).
In the Online Appendix 1, each species is presented at the hand of its taxonomy, geographical distribution, habitat (including substrate) and most likely threats. The current Red List status is indicated for those species that were previously assessed, either at the global or at regional scales.
However, not all of these Red List assignments follow IUCN methodology consequently, and some of the global Red List statuses are in need of revision. Family placement of the genera follows the classifications of Goffinet, Buck and Shaw (2009) and Frey and Stech (2009) for the mosses and Söderström et al. (2016) for the liverworts. Species author citations follow Söderström et al. (2016) for the liverworts and Missouri Botanical Gardens’ TROPICOS database (Tropicos.
org) for the mosses.
The 10 species that are at highest risk of extinction, and most likely to be categorised as ‘critically endangered’ when (re-) assessed using the latest IUCN Red List criteria (IUCN 2012a), were selected for the Top 10 list for Africa (Table 1).
Monotypic genera and subgenera received priority and preference was given to species recognised and accepted 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 extinction, 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 Distribution Biodiversity hotspot†
Bryopteris gaudichaudii Gottsche 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 Pottiaceae Endemic to South Africa Cape Floristic Region
Neckeropsis pocsii Enroth & Magill Neckeraceae Endemic to the Comoros Madagascar and the Indian Ocean Islands Picobryum atomicum R.H.Zander & Hedd. Pottiaceae Endemic to South Africa Cape Floristic 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 Conservation International through the CEPF (2016a).
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http://www.abcjournal.org Open Access (O’Shea 2006; Söderström et al. 2016; Tropicos.org) for which
precise information on localities is available.
Results
We have identified a total of 50 species, representing 40 genera and 23 families of bryophytes, as lost and threatened in the area (Online Appendix 1). Of these, 29 are endemic to mainland Africa, 19 to the East African islands and two are restricted to West Africa as well as islands off the coast.
Twenty-nine species (in 19 genera and nine families) are liverworts and 21 species (in 21 genera and 14 families) are 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 by five species each.
Altogether, the species’ known localities are found in 16 African countries (Online Appendix 1). Eighteen species occur in the Flora of Southern Africa area (Magill 1981), with 15 species being endemic to South Africa. Of these, seven species are restricted to the Fynbos Biome (Rebelo et al. 2006) of the southwestern Cape. The Indian Ocean islands of Réunion and Madagascar are well represented on the list with 12 (six endemic) and 11 (four endemic) species, respectively.
Six of the eight global biodiversity hotspots recognised in sub-Saharan Africa and the East African islands, based on vascular plant species and vegetation, contain candidates for the Top 10 list (Figure 1). The three hotspots with the most species are (1) Madagascar and the Indian Ocean Islands with 19 species, (2) Eastern Afromontane with nine species and (3) Cape Floristic Region with seven species.
The remaining hotspots with threatened bryophytes are:
Maputaland–Pondoland–Albany (four species), Succulent Karoo (four species) and Guinean Forests of West Africa (three species).
Only four species, all from inland areas of southern Africa, have not been reported from any of the global biodiversity hotspots. They are Anacamptodon marginata (Dixon) W.R.Buck from Mashonaland West Province of Zimbabwe, Cryptomitrium oreades Perold from the Highlands of Lesotho, Fissidens capriviensis Magill from the Caprivi Strip of Namibia and Gymnostomum lingulatum Rehmann ex Sim from the Woodbush–Haenertsburg area in the Limpopo Province of South Africa.
The majority of Top 10 species (six species) belong to the liverwort family Lejeuneaceae, and the Madagascar and the Indian Ocean Islands hotspot is the biodiversity hotspot with the highest number of species (six species) on the list (Table 1). Xylolejeunea Xiao L.He et Grolle (Lejeuneaceae) is the only genus with more than one (two) species on the Top 10 list for Africa.
Discussion
It is no coincidence that Lejeuneaceae is the liverwort family with the highest species number on the list of lost or
threatened bryophytes in Africa as well as on the Top 10 list of bryophytes in Africa that are at highest risk of extinction.
Not only is Lejeuneaceae the largest family of liverworts, but it contains more than 95% of all epiphyllous bryophytes (Gradstein 1994). The family displays high levels of diversity and endemism in moist tropical forests where they grow mainly on the bark of woody plants and on leaves at low and mid elevations (Gradstein 1992, 1994; Pócs 1996; Von Konrat et al. 2008). Von Konrat et al. (2008:431) calculated that in Madagascar and some countries of central and West Africa, between 50% and almost 90% of the liverwort flora consist of Lejeuneaceae.
Recent researches in tropical forests indicate that ancient uncut stands of forest, with original phorophyte diversity and intact forest canopies, are important drivers for high epiphytic and epiphyllous bryophyte diversity (Benítez, Prieto & Aragón 2015; Malombe et al. 2016; Pócs &
Tóthmérész 1997; Zartman 2003). The ongoing destruction, degradation and fragmentation of forests in the biodiversity hotspots of Africa (Aynekulu et al. 2016; CEPF 2016a;
Gradstein 1992; Green & Sussman 1990; IUCN 2017a;
Malombe 2007) therefore pose serious threats to epiphytic and epiphyllous bryophytes in general and species of Lejeuneaceae in particular.
The fact that 46 (92%) candidate species for the Top 10 list fall within six of the eight biodiversity hotspots in the region suggests that the African biodiversity hotspots designated on the basis of information from vascular plants may also represent hotspots for bryophytes. However, the concentration of lost or threatened species in the biodiversity hotspots may also be a consequence of collecting bias. Only two of the global biodiversity hotspots in Africa, the Coastal Forests of Eastern Africa and the Horn of Africa, are not represented on our list (Figure 1). Whilst these areas are under-explored (Hylander, Nemomissa & Hedenäs 2017;
Wilding et al. 2016), lower bryophyte diversity in arid areas such as in the Horn of Africa hotspot may further contribute to this outcome. Many African regions, for example in Western Africa (Diop et al. 2018), are much less researched and thus still poorly understood in terms of plant diversity and threats to it. Thus, the detailed picture of the distribution of African bryophyte diversity is likely to change along with increasing knowledge and exploration, but we are confident that some overall patterns are recognisable based on this compilation.
The biodiversity hotspot with the highest number of lost or threatened species, as well as Top 10 species which are at highest risk of extinction in Africa, is Madagascar and the Indian Ocean Islands (Table 1, Figure 1). This hotspot is dominated by Madagascar, the fourth largest island globally, and includes the Seychelles (including Aldabra), the Comoros, Mauritius (including Rodrigues) and the French overseas departments of Réunion, Mayotte (one of the Comoros) and the volcanic Iles Esparses around Madagascar (Figure 1). The island of Madagascar, especially
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of bryophyte diversity and endemism and the most species- rich area of bryophytes in Africa (Geffert et al. 2013; Tan &
Pócs 2000; Vanderpoorten & Hallingbäck 2009). It has also been identified as one of two major Lost Spots in Africa, countries where ‘lost species’, including a few mosses, are concentrated (Global Wildlife Conservation 2014).
Agricultural activities, urbanisation, invasive alien species and deforestation have devastated and fragmented habitats, especially in the lowlands, throughout this hotspot (CEPF 2016a). At least one of the endemic bryophyte species from the Seychelles island of Mahé, Xylolejeunea muricella Xiao L.He et Grolle, may already be extinct (Online Appendix 1).
There is also concern about the recent marked increase in illegal logging in the remaining forests of Madagascar (Green & Sussman 1990; IUCN 2017a).
The second-highest number of lost or threatened species per biodiversity hotspot are found in the Eastern Afromontane hotspot (Figure 1). This hotspot is characterised by a series of montane islands and extensive plateaus along the eastern edge of Africa, from Ethiopia in the north to Zimbabwe in the south. It consists mainly of three ancient massifs: the Eastern Arc Mountains and Southern Rift, the Albertine Rift and the Ethiopian Highlands. Eastern Africa has also been suggested as one of the main centres of bryophyte diversity and endemism in Africa (Geffert et al. 2013; Von Konrat et al.
2008) and Tanzania is another major Lost Spot in the region for a variety of organisms (Global Wildlife Conservation 2014). Degradation and fragmentation of habitats and unsustainable exploitation of natural resources are the main threats in this hotspot (CEPF 2016a). This is caused by expanding agriculture, plantation forestry, logging, fires, invasive alien plants, mining, infrastructure development and gathering of firewood.
A substantial number amongst the lost or threatened bryophytes occur in the southwestern Cape, a region that contains the highest concentration of threatened vascular plants and plants of conservation concern in South Africa (Raimondo & Van Staden 2009). The Fynbos Biome of the Cape Floristic Region biodiversity hotspot is one of the most threatened biomes in southern Africa, classified as critically endangered (Driver et al. 2012). The southwestern Cape has been suggested as a global and regional centre of relatively high bryophyte species richness and endemism (Geffert et al.
2013; Tan & Pócs 2000). In an analysis of moss distributions in southern Africa, Van Rooy and Phephu (2016:29) recognised a Southwestern Cape Centre of Moss Diversity and found that the Cape Town-Table Mountain area is the most species- rich in the region. The Cape Floristic Region is under increasing population pressure and much of the Fynbos and Renosterveld vegetation types of the lowlands have been destroyed or transformed by agriculture and urbanisation (CEPF 2016a; Rebelo et al. 2006; Von Hase et al. 2003). The lowland Renosterveld areas have been identified as top conservation priorities by the Cape Action Plan for the Environment (Von Hase et al. 2003). The remaining Fynbos is
threatened by invasive alien species, the disruption of fire regimes, plantation forestry and fragmentation.
Conclusions
This is the first compilation of bryophytes for sub-Saharan Africa and the East African islands that are most probably lost or threatened. Although it is not exhaustive, and many of the ‘threat categories’ reported herein are in need of revision and require to be scrutinised against the IUCN Red List criteria (IUCN 2012a), it represents a first approximation of the threats to bryophytes in Africa.
The species on the list of lost or threatened bryophytes served as candidates for the selection of the Top 10 species in Africa at high risk of extinction. Therefore, it is a crucial contribution towards the Top 10 Initiative of the IUCN Bryophyte Specialist Group (IUCN SSC 2015) and thus to the (global) IUCN Red List of Threatened Species (IUCN 2017e), as well as towards an assessment of the extinction risk of bryophytes on the African continent. Hence, it is a significant step towards meeting the targets of the Updated Global Strategy for Plant Conservation 2011–2020 (CBD Secretariat 2017) and priorities of The Shenzhen Declaration on Plant Sciences (Crane et al. 2017).
The vast majority of threatened species on our list, and all Top 10 species, are restricted to global biodiversity hotspots in Africa, areas of high endemism of vascular plants under severe threat of habitat loss (CEPF 2016a; Conservation International 2017). Investment in biodiversity conservation through the hotspots concept will therefore benefit threatened African bryophytes as well.
The Madagascar and Indian Ocean Islands biodiversity hotspot, which is also considered as a global centre of bryophyte diversity and endemism (Vanderpoorten &
Hallingbäck 2009; Von Konrat et al. 2008), contains a particularly large number of threatened bryophytes. The recent increase in logging poses a serious threat to the remaining forest habitats on Madagascar and other Indian Ocean islands. The Eastern Afromontane and Cape Floristic Region biodiversity hotspots also stand out as hotspots of lost or threatened bryophytes in Africa.
The degradation and fragmentation of forests throughout sub-Saharan Africa and the East African islands have a visible impact on epiphytic bryophytes, epiphyllous liverworts of the family Lejeuneaceae in particular. Bryological exploration of Africa should be stepped up to increase our knowledge of species occurrences and threats and to facilitate the red-listing and the conservation of rare and threatened bryophytes.
Acknowledgements
We would like to thank Tamás Pócs, Frank Müller, Itambo Malombe and Claudine Ah-Peng for their contributions. We are grateful to Hester Steyn and Elizma Fouche for editing the map.
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Competing interests
The authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article.
Authors’ contributions
J.v.R. compiled the species accounts, analysed and interpreted the data and drafted the article. I.B. and A.B. conceptualised and initiated the project ‘Top 10 Initiative’ and commented on manuscript versions. All the authors contributed to the final version of the article.
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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 threat category as far as assigned†
Sources
1. Actinodontium
streptopogoneum Broth. in Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 24: 260 (1897).
Pilotrichaceae. The only species of Actinodontium present in Africa. Its relationship to American and Asian species needs investigation.
Cameroon and Bioko.
Epiphytic on tree trunks and twigs in forest, up to 1300 m a.s.l.
For more than a century (since 1890), the species was known only from two sites around Mt
Cameroon, which are now intensively cultivated. It was rediscovered in Africa on Bioko in 2002, where it grew on solitary trees in a pasture land. The trees at this site are heavily used (e.g. for firewood). The species is thus likely to be highly threatened. The Guinean Forests of West Africa biodiversity hotspot is one of the most critically fragmented habitats in the world.
Müller and Pócs (2007);
Critical Ecosystem Partnership Fund (2016).
2. Anacamptodon marginatus (Dixon) W.R.Buck in Journal of the Hattori Botanical Laboratory 47: 52 (1980).
Amblystegiaceae. Originally described as Hypnofabronia marginata, the species was later transferred to
Endemic to Zimbabwe.
Collected by the explorer Thomas Bains on Hartley Hill, Zimbo (Chimbo) River, S.
African Goldfields.
Epiphytic on trees in the
Only known from a single collection made in 1870. The habitat is under threat from artisanal mining, cultivation and overgrazing by livestock.
Zimbabwe is bryologically poorly known and this area has probably
Dixon and Wager (1929);
Buck (1980); Van Rooy (2011); Anonymous (2017).
Anacamptodon. The wiry stems and distant leaves with a strong and variable border, single costa and small, quadrate alar cells are distinctive.
Savanna Biome, 230 m a.s.l. not been searched for bryophytes since the original collection.
3. Andreaea camerunensis
P.W.Richards in Transactions of the British Bryological Society 2:
66 (1952).
Andreaeaceae. The species differs from the closely related A. firma Müll.Hal. in the broader leaves, broader, cucullate leaf apices, shorter and wider basal cells, less incrassate and slightly longer upper laminal cells.
Endemic to Cameroon:
‘Cameroon Mountain, Vefondi Peak’.
On tall, pillar-like lava blocks in grassland at 3170 m a.s.l.
Only known from a single gathering made in 1948. Mount Cameroon is an active volcano and the most recent eruptions were in February 2012. The habitat is threatened by lava flows. Tropical rain forest loss and fragmentation, ecosystem degradation and limited local capacity for conservation are the major threats in the Guinean Forests of West Africa biodiversity hotspot.
Richards (1952); CEPF (2016).
4. Archidium andersonianum Snider in The Bryologist 78: 158 (1975), fig. 26–36, 39.
Archidiaceae. This small but
conspicuous species is similar in habit to A. capense and A. muellerianum. It differs in the deltoid innovation leaves with costa ending in a strong hair-point and the larger, broadly oblong-ovate perichaetial leaves with narrower costae.
Endemic to the Western Cape Province of South Africa where it was collected on the Stellenbosch flats by Almborn in 1966.
On sandy or gravelly soil in the Fynbos Biome.
This species is represented by a single collection from the Cape Floristic Region, a global
biodiversity hotspot. The vegetation of the Cape lowlands has been highly impacted by agriculture and urban development and alien plant species have invaded most of the remaining natural habitats. It is highly threatened.
Snider (1975); Magill (1981);
Rebelo et al. (2006); CEPF (2016).
5. Bazzania konratiana Gyarmati in Cryptogamie, Bryologie 38(2):
Endemic to Madagascar:
Toamasina Prov., Mananara
Only known from two localities in the Mananara Nord Biosphere
CEPF (2016); IUCN (2017a);
Wigginton (2018); Sass-
120 (2017), fig. 1–8.
Lepidoziaceae. The species can be separated from the Seychelles endemic B. approximata Onr. by the larger plants, thick-walled leaf cells with large, bulging trigones and the proportion of hyaline and
chlorophyllose cells in the underleaves.
Nord Biosphere Reserve and National Park.
Corticolous in lowland rainforest with tree ferns, palms and Pandanus ssp., on E and NW slopes, 250–300 m a.s.l.
Reserve where it was collected in 1998. The remaining habitat in this biosphere reserve is threatened by illegal exploitation of forest resources. Of the 15 species of Bazzania recognised in Africa, 11 are restricted to the Madagascar and the Indian Ocean Islands biodiversity hotspot and six are endemic to the island of
Madagascar.
Gyarmati (2017).
6. Bruchia eckloniana Müll.Hal. in Synopsis Muscorum
Frondosorum omnium hucusque Cognitorum 1: 19 (1848).
Bruchiaceae. The small calyptra, plane, entire leaf margins and papillose to vermiculate spores will identify the species.
Collected by Ecklon in the Cape (probably around Cape Town), without a precise locality (‘Prom. Bon. Spei’), between 1823 and 1848.
On soil in the Fynbos Biome.
Only known from the original locality in the Fynbos and Renosterveld vegetation of the Cape Floristic Region biodiversity hotspot, which is threatened by urban development, agriculture and invasive alien plant species. The Fynbos Biome is critically endangered.
Magill (1981); Rebelo et al.
(2006); CEPF (2016).
7. Bryopteris gaudichaudii
Gottsche in Annales des Sciences Naturelles; Botanique, sér. 4 8:
340 (1857).
Lejeuneaceae. Molecular evidence shows B. gaudichaudii as one of the major clades in the genus.
Madagascar and Réunion.
Coastal ericaceous heath and montane rainforest.
Found only once since 1900 (in 1996) on Réunion and known from only two localities on Madagascar, about 75 km apart. Cultivation, livestock grazing and deforestation have devastated habitats, especially in the lowlands, throughout the Madagascar and the Indian Ocean Islands biodiversity hotspot.
Despite recent exhaustive searches
Hartmann et al. (2006);
Wigginton (2018); Ah-Peng et al. (2012); CEPF (2016);
IUCN (2017a).
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
Gradst. in Acta Botanica Neerlandica 23: 334 (1974), pl.
1.
Lejeuneaceae. Within Caudalejeunea it constitutes a separate, monotypic subgenus. Tiny epiphytic plants, growing in small, dull green mats.
Endemic to Madagascar where it is known from Nossi-Bé island and Antsohy Forest.
On bark of stems and dead wood in undisturbed lowland rainforest.
Only known from two localities in the Madagascar and the Indian Ocean Islands biodiversity hotspot and not collected since 1973. The remaining lowland rain forests of Madagascar are threatened with destruction.
Endangered globally: B1+2cd (ver 2.3) (IUCN 2017b).
Green and Sussman (1990);
Gradstein (2001a), CEPF (2016).
9. Cheilolejeunea ulugurica Malombe, Eb.Fisch. et Pócs in Acta Biologica Plantarum Agriensis 1: 24 (2010).
Lejeuneaceae. Distinguished by the
Endemic to Tanzania, Uluguru Mts, N side of Bondwa peak.
Ramicolous on stems of ericaceous shrubs at the edge of montane forest, 1650–1800
Only known from a single collection despite intensive bryological exploration in the Eastern Arc Mountains, which forms part of the Eastern
Malombe, Fischer and Pócs (2010); CEPF (2016);
Wigginton (2018).
obovate-spathulate lobe with a lobule occupying ⅔ to ¾ its length with a strongly incurved keel apex and forming a very narrow sinus with the postical lobe margin. Possibly
conspecific with C. chenii R.L.Zhu et M.L.So from China.
m a.s.l. Afromontane biodiversity hotspot.
Like vascular plants, any rare or endemic bryophyte found in the Ulugurus at altitudes lower than 1800 m is under serious threat from encroachment of habitat and
deforestation. This species may be Critically Endangered.
10. Cololejeunea bosseriana Tixier in Bulletin Trimestriel de l'Académie Malgache 55: 235 (1977)[1979], fig. 39.
Lejeuneaceae. According to the World checklist of hornworts and liverworts (Söderström et al. 2016), the species is not well known.
Endemic to Réunion, Forêt de Bébour.
Epiphyll on ferns in humid forest at 1300 m a.s.l.
Known only from the type locality in the Madagascar and the Indian Ocean Islands biodiversity hotspot and last collected by Bosser in 1972.
Critically Endangered on Réunion: [B2ab(iii)] (Ah-Peng et al. 2012).
Ah-Peng et al. (2012); CEPF (2016); Söderström et al.
(2016).
11. Cololejeunea decemplicata (Steph.) Tixier in Bulletin Trimestriel de l'Académie
Malgache 55: 213 (1977)[1979].
Lejeuneaceae. The species is not well known (Söderström et al. 2016).
Madagascar and Réunion (three localities).
Corticolous on trees like Acacia sp., in forest.
Urbanisation, agriculture and invasive alien species have resulted in the degradation and
fragmentation of natural habitats throughout the Madagascar and the Indian Ocean Islands biodiversity 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
Ah-Peng et al. (2012); CEPF (2016); Söderström et al.
(2016).
most likely on Madagascar as well.
12. Cololejeunea jonesii Pócs in Acta Botanica Academiae
Scientiarum Hungaricae 21: 361 (1975), fig. 35–43.
Lejeuneaceae. The number of cells in the discoid gemmae is much higher than in related species. The species will also be distinguished by the presence of claviform styles and the biserial ventral merophytes.
Kimboza Forest Reserve in Tanzania and Ngangao Forest in the Taita Hills of Kenya.
Epiphyllous in lowland rain forest (300 m a.s.l.) and montane tropical forest (2149 m a.s.l.).
Only known from two localities in the Eastern Afromontane
biodiversity hotspot. The type locality in Tanzania is in a lowland rainforest of the Kimboza Forest Reserve at the eastern foothills of the Uluguru Mountains, seriously encroached by illegal ruby mining and log felling. Recently recorded in Ngangao Forest, a forest 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.
Pócs (1975); Malombe et al.
(2016); CEPF (2016).
13. Cololejeunea nosykombae A.Szabó & Pócs in Journal of Bryology 38(4): 302 (2016).
Lejeuneaceae. The species differs from other species in the genus mainly by its swallow-tail shaped perianth. Two varieties are recognised: C. nosykombae var. laevis A.Szabó & Pócs as well as the typical variety.
Endemic to the volcanic Nosy Komba island of Madagascar.
Epiphyllous in a submontane rainforest fragment, mostly on Marattia fern leaflets, but also on the fern Tectaria sp. and a broadleaved shrub, at an altitude of 570–580 m a.s.l.
Associated with other epiphyllous Lejeuneaceae.
The locality falls within the Madagascar and the Indian Ocean Islands biodiversity hotspot. The species is known only from a 150 m × 50 m area in a forest remnant surrounded by cultivation. It is considered to be ‘critically endangered’ because of its tiny population size and very small area of occurrence in a habitat under
Szabó and Pócs (2016); CEPF (2016).
severe threat.
14. Cololejeunea takamakae Tixier in Bryophytorum Bibliotheca 27:
319 (1985), fig. 73.
Lejeuneaceae. There are doubts about the taxonomy of the species and it is possibly conspecific with C.
angustiflora (Steph.) Mizut., which is known also from Malaysia, Borneo, China, Taiwan and the Philippines.
Endemic to Réunion.
Epiphyll at ±1000 m a.s.l.
Less than five localities, threatened with habitat degradation. A very restricted endemic in the
Madagascar and the Indian Ocean Islands biodiversity hotspot.
Critically Endangered on
Réunion: [B1b(iii)] (Ah-Peng et al.
2012).
Ah-Peng et al. (2012);
Söderström et al. (2016);
CEPF (2016).
15. Colura heimii Jovet-Ast in Revue Bryologique et
Lichénologique 22: 275 (1953).
Lejeuneaceae. This is a recognised species and currently accepted in the World checklist of hornworts and liverworts (Söderström et al. 2016).
Comoros, Madagascar, Mauritius and Réunion.
On Réunion it grows on twigs in lowland and upland forests.
Urbanisation, agriculture and invasive alien species have resulted in the degradation and
fragmentation of natural habitats throughout the Madagascar and the Indian Ocean Islands biodiversity hotspot. On Réunion the habitat is also threatened by lava flows.
Critically Endangered on Réunion [B2b(iii)] (Ah-Peng et al. 2012) and most likely throughout the region.
Wigginton (2018); Ah-Peng et al. (2012); Söderström et al.
(2016); CEPF (2016).
16. Cryptomitrium oreades Perold in Bothalia 24: 149 (1994), fig.
1–2.
Aytoniaceae. The genus is only represented by one species in Africa, which differs from the other two species in the genus mainly by the
Endemic to the Oxbow area of Lesotho.
On soil over rock under basalt boulders, in seepage areas in the high altitude Drakensberg alti-montane grasslands, at
±2550 m a.s.l.
Only known from two localities,
±5 km apart. Heavy grazing pressure by livestock (carrying capacity is exceeded by up to 300%) is a serious threat to the Drakensberg alti-montane grassland and woodland. It alters the
ecosystem and makes the existing
Perold (1994, 1999); Olson and Dinerstein (2002).
lateral position of the stalk. habitats vulnerable to
encroachment by Karoo vegetation.
17. Cygnicollum immersum Fife &
Magill in The Bryologist 85: 99 (1982), fig. 1–19.
Funariaceae. This distinct, monotypic genus is recognised by its size (very small), bulbiform habit and completely enclosed, pendent, cleistocarpic
capsule.
Endemic to the Northern Cape Province of South Africa and only known from the type locality on the upper slopes of Vanrhyns Pass.
It grew on sandy soil in small open areas amongst shrub at
±800 m a.s.l.
This monotypic genus has only been collected once (in 1977) and recent efforts to re-collect it were unsuccessful. However, the plants are very small (stems up to 1 mm long) and can easily be overlooked.
The Vanrhynsdorp Shale Renosterveld vegetation is 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.
Magill (1987); Rebelo et al.
(2006); Goffinet (2013);
CEPF (2016).
18. Drepanolejeunea helenae Pócs in Cryptogamie: Bryologie, Lichénologie 18: 198 (1997), pl.
1–4.
Lejeuneaceae. The species differs from D. ankasica E.W.Jones in the many ocelli in the leaf lobe and the presence of ocelli in the perianth, amongst other characters.
Endemic to Réunion.
On the bark of Psidium cattleyanum Sabine and epiphyllous in remnants of tropical rainforest, ±500–
750 m a.s.l.
Known from only two localities, where the type specimens were collected in 1994 and 1995. Habitat destruction is the main threat to this very restricted endemic of the Madagascar and the Indian Ocean Islands biodiversity hotspot.
Critically Endangered on Réunion: [B2ab(iii)] (Ah-Peng et al. 2012).
Pócs (1997); Ah-Peng et al.
(2012); CEPF (2016).
19. Entosthodon heddersonii N.Wilding in Phytotaxa 312(1):
Endemic to Tanzania. Recently described from the
The species is known only from the type specimen collected in 1988.
Frontier.ac.uk (2014); CEPF
103–110 (2017).
Funariaceae. The narrowly oblong- obovate, aristate leaves, bordered and toothed in the upper ⅔ and the
zygomorphic, peristomate capsules will differentiate E. heddersonii from other species in the genus.
Lukwangule plateau in the Uluguru Mountains.
Collected at an elevation of
±2360 m a.s.l. along a rocky, peaty stream bank on the eastern edge of the Lukwangule Plateau.
The main threats to this part of the Eastern Afromontane biodiversity hotspot are annual fires on the Lukwangule plateau that may change the ecology of the
afromontane grassland and spread into the forest, firewood collection, encroachment into the reserves and spread of the invasive Rubus sp.
(2016); Wilding (2017).
20. Ephemerum diversifolium Mitt., Harvey in Thesaurus Capensis 1:
63 (1859), pl. 100, fig. A.
Ephemeraceae. A very distinct species with highly differentiated perichaetial leaves.
Endemic to the Eastern Cape Province of South Africa:
Uitenhage, Zwartkop’s River.
On soil in shrublands.
Only known from the type locality where it was collected during the first half of the 19th century by Zeyher. The plants are minute, making it difficult to find.
Cultivation, invasive alien plants, 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.
Magill (1987); Mucina et al.
(2006a), CEPF (2016).
21. Fissidens capriviensis Magill in Flora of Southern Africa,
Bryophyta 1: 45 (1981), fig. 9:
1–7.
Fissidentaceae. The corticolous
Endemic to Namibia. Collected by Hans Vahrmeijer in the Caprivi strip, Kongola area, Kwando swamps.
In gallery forest, on lower
Only known from the type locality where it was found in 1980.
Although the Kwando swamps are relatively undisturbed and the area under-explored, agriculture is encroaching on the habitat in the
Magill (1981); Van Rooy and Phephu (2016).
