Tropical Atlantic Oceanic Islands and Archipelagos: Physical structures, plant diversity, and affinities of the bryofloras of northern and southern islands

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Cluster analysis was used to determine similarities among the bryofloras of ten islands and archipelagos. We examined the bryophyte species to address the following questions: (1) How many bryophyte species are there in the total area and on each island and archipelago? (2) Do mosses and liverworts share patterns of diversity? (3) What are their species distribution patterns? (4) How many endemic species are there on the islands? (5) Is there a high similarity among those bryophyte floras? We encountered 1,647 species, 325 genera, and 204 families, with the richest families (Lejeuneaceae, Lophocoleaceae, Plagiochilaceae, Brachytheciaceae, Bryaceae, Fissidentaceae, and Pottiaceae), representing 35% of all species. Over 12% of the bryophytes are African elements, 8% have global distributions, approximately 3% are Macaronesia elements, 5% are endemic, and 34% are included in other distributions. We present a checklist of 1,647 bryophyte species, of which 91 are endemic. Bryophyte richness and diversity differed significantly among the ten islands and archipelagos studied, and their low similarities illustrate their high degrees of heterogeneity. This paper is the first evaluation of bryophyte species on ten islands and archipelagos of the Atlantic Ocean, giving insight into their species richness, endemism and similarities, and providing information for analyzing species turnover, migration, invasion, etc., representing an increase in our general knowledge of the bryofloras of oceanic islands and archipelagos, contributing to the targets of the Global Strategy for Plant Conservation. Atlantic Oceanic Islands Bryophytes Distribution Endemism Similarity Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Scientists have long been fascinated by the unique nature of remote oceanic islands, especially after Charles Darwin’s study of the Galapagos Islands. Young oceanic islands of volcanic origin represent the summits of large volcanoes that rose from the bottom of the ocean. They are isolated from other landmasses by deep seas and have never been connected to continental landmasses so their environments are unique products of trans-oceanic dispersal and eventual speciation (Wace & Holdgate 1976; Cowie and Holland 2006). Islands therefore represent natural laboratories that simplify the complexities of the natural world (Whittaker and Fernandez-Palacios 2007). The introduction of non-native species, however, is a constant threat to an island’s biota, and it is generally known that warmer temperatures can improve the chances of alien invasive species settling, reproducing, and successfully outcompeting the native biota, and can lead to significant biodiversity losses. There are numerous islands and archipelagos in the Atlantic Ocean: including the archipelagos of the Azores and Madeira within the territorial waters of Portugal; the archipelago of the Canary Islands (Spain); the archipelagos of Cape Verde and São Tomé and Príncipe (Africa); Fernando de Noronha, Trindade and Martin Vaz (Brazil); and the islands of St Helena, Ascension and the archipelago of Tristan da Cunha (England) (Fig. 1). The bryofloras of the ten islands and archipelagos were inventoried: Azores, Madeira, São Tomé e Príncipe, Cape Verde, and Canary islands (Northern Atlantic) and Trindade, Fernando de Noronha, Ascension, St Helena and Tristan da Cunha (Southern Atlantic) to: (1) provide an overview of their bryofloras; (2) describe the species richness of mosses and liverworts and incorporate new reports of bryophyte taxa; (3) analyze the distribution patterns and endemism of those species; (4) analyze the similarities among northern and southern islands; and (5) establish a diversity baseline for comparisons. We describe here the diversity, endemism, and similarities of mosses and liverworts on ten islands and archipelagos located in the Atlantic Ocean. Studied Islands and Archipelagos (physical structures and bryofloras) The Portuguese archipelagos are of volcanic origin and represent important geological sites. The Azores archipelago comprises nine islands. It is located ca. 1600 km from the European coast and represents a chain of submarine volcanic islands with sources of fresh water. The archipelago harbors a total of 436 species of bryophytes, with the diversity among the different islands varying greatly, from 104-324 species; only 5% of the archipelago is covered by forest vegetation. The Madeira archipelago is located 978 km from Portugal and ca. 700 km from Africa. It is also of volcanic origin, with two main islands (Madeira and Porto Santo), as well as two groups of uninhabited islands (Desertas and Selvagens). The highest points there are the Pico Ruivo (1862 m), Pico das Torres (1851 m), and Pico do Arieiro (1818 m). The mountainous landscape and their exposure to constant winds mean that the small islands have different microclimates and higher precipitation rates on the northern coast; few places have original intact vegetation, and they have no sources of fresh water. The Madeira archipelago has 577 species of bryophytes (Table 1). Table 1 General data on the ten islands and archipelagos studied. AZO = Azores, MAD = Madeira, STP = São Tomé and Príncipe, CV = Cape Verde, CAN = Canary Islands, FN = Fernando de Noronha, TRI = Trindade, ASC = Ascension, STH = St Helena, TRC = Tristan da Cunha, END = Endemic AREA ORIGIN CLIMATE HABITAT MOSSES LIVERWORT HORNWORT TOTAL AZO Volcanic (9 islands), 0.3-0.8 mya, 2,351 Km 2 , population: 236,440 permanent residents, highest point is Monte do Pico (2,351 m) Subtropical oceanic, 17°C, 1,026 mm precipitation Laurissilva forest 270 spp. 160 spp. 6 spp. 436 spp. (40 END – 10%) MAD Volcanic (4 islands), 5 mya, 724 Km 2 , population: 262,456 permanent residents (Madeira Island), highest peak is Pico Ruivo (1,862 m) Mediterranean and Temperate in the highest parts, 20°C, 2,000 mm precipitation Laurissilva forest (best preserved evergreen forests in Macaronesia), with a great diversity of bryophytes covering the trunks and branches of trees and shrubs 389 spp. 182 spp. 6 spp. 577 spp. (68 END – 12%) STP Volcanic (3 islands), 31 mya, 991 Km 2 , population: 213.000 permanent residents, highest peak is Pico de São Tomé (2024 m) Equatorial oceanic (hot and wet), 22-30°C, 214 mm precipitation Tropical moist broadleaf forest 133 spp. 170 spp. 7 spp. 310 spp. (16 END – 7%) CV Volcanic (10 islands, and one active vulcan in the Fogo Island), 8-20 mya, 4,033 Km 2 , population: 561.901 (nine islands have permanent residents), highest peak is Pico do Fogo (2829 m) Arid to semi-arid, 22-27°C, 214 mm precipitation Savanna or steppe, and tropical climate depending on elevation 168 spp. 54 spp. 2 spp. 224 spp. (5 END – 2%) CAN Volcanic (7 islands), 8-20 mya, 7,493 Km 2 , population: 2.2 million permanent residents, highest peak is Monte Teide (3715 m) Warm subtropical and semi-desertic Subtropical, with xerophytic, humid forest, the laurel forest, fayal-brezal, the pine forest and high mountain vegetation 352 spp. 143 spp. 6 spp. 501 spp. (54 END – 11%) FN Volcanic (21 islands), 30 mya, 26 Km 2 , population: 3,100 permanent residents, highest peak is Desejado (323 m) Tropical wet and dry, 26,5°C, 1,350 mm precipitation Seasonal deciduous forest, subxerophytic (species typical of the northeastern agreste ). Dry Forest (Ponta da Sapata), 25% of shrubs and trees on the main island. Mangrove (only occurrence of an insular mangrove in the South Atlantic Ocean) is located in the Sueste Bay. Creepers covering native bushes and trees during the rainy season. 23 spp. 7 spp. 2 spp. 32 spp. TRI Volcanic (5 islands), 3 mya, 10.4 Km 2 , population: 32 soldiers + 8 researches (no permanent residents), highest peak is Pico do Desejado (620 m) Tropical oceanic, 25°C, 923 mm precipitation Exotic plants (100-150 m), lowland (up to 400 m) grass fields, Giant Fern Forest or Giant Ferns Nebular Forest (> 400 m). Less than 5% of the island is covered by forest and ca. 60% by herbaceous vegetation 14 spp. 20 spp. 1 sp. 35 spp. (NO END) ASC Volcanic, 1 mya, 91 Km 2 , population: 800 inhabitants (no permanent residents), highest peak is Green Mountain (859 m) Dry tropical oceanic, 20-31°C, 130-680 mm precipitation Dryland (0-300 m), mid-altitude area on Green Mountain (330-600 m), high-altitude with cloud forest area top of Green Mountain (> 660 m) 51 spp. 26 spp. 4 spp. 81 spp. (16 END – 20%) STH Volcanic (one island), 7 mya, 122 Km 2 , population: 4,500 permanent residents, highest peak is Diana’s Peak (818 m) Tropical, 17-28°C, 750-1,000 mm precipitation Tree fern forest, pastures and coastal zones, dry and eroded, and middle elevations now destroyed 62 spp. 35 spp. 2 spp. 99 spp. (26 END – 28 %) TRC Volcanic (6 small islands -Nightingale – >18 mya, Inaccessible – 3-4 mya, Tristan da Cunha – 200,000 mya; Gough – 3-5 mya), 207 Km 2 , population: 264 permanent residents (only in Tristan da Cunha), highest peak is Queen Mary's Peak (2,062 m) Cool temperate, 16-25°C, 1670 mm precipitation Tussock grassland, pastures, fern bush, wet heath, feldmark and alpine, bogs and other wetland, and lava field 158 spp. 142 spp. 4 spp. 305 spp. (86 END – 22%) The African archipelagos are likewise of volcanic origin and include two islands (São Tomé and Príncipe) and some islets. São Tomé is the largest island in the archipelago and consists of two islands formed during the tertiary era. The archipelago is located off the coast of central Africa (ca. 250 km distant), close to the equator. The island of São Tomé is the capital of the archipelago, with an area of ca. 1001 km². The climate there is equatorial, hot, and humid, with temperatures varying according to altitude and exposure (22-30°C). The island is steep, with a plain zone, high mountains to the south and west, a flat landscape to the north, and relatively fertile soil. It holds a total of 310 species of bryophytes. Cape Verde is an island country that is part of an archipelago formed by 10 islands covering ca. 4033 km², located off the coast of West Africa. A total of 224 species of bryophytes have been recorded there, with the islands of Santo Antão, Santiago, and Fogo having the highest numbers of plant species. Santo Antão has the highest species richness of bryophytes, with 111 species (73% of the archipelago's total bryoflora), (Table 1). The Spanish archipelago of the Canary Islands is of volcanic origin that emerged 8-12 million years ago and covers an area of 7747 km 2 . It composes part of Macaronesia along with Madeira, Azores, and Cape Verde. A total of 501 species of bryophytes are recognized there (Table 1). The Brazilian archipelagos are of volcanic origin and represent important national and international geological sites. The Trindade archipelago is located ca. 1100 km from the Brazilian coast. Its mountainous islands belong to a chain of submarine volcanoes, with a source of fresh water. Most of the vegetation on the archipelago is herbaceous, with only 5% of the vegetation being forest; it harbors 35 species of bryophytes. The archipelago of Fernando de Noronha is formed by 21 small islands of volcanic origin, located ca. 350 km from the Brazilian coast. There are few places with original vegetation, and no source of fresh water; most of the vegetation is dry forest similar to Caatinga, with small trees, shrubs, and grasses. There are 32 species of bryophytes growing there, but neither Brazilian archipelago has any endemic species (Table 1). The two islands and the English archipelago are also of volcanic origin. Ascension is a small island covering only 91 km², 1660 km from Africa and 2250 km from South America; it is approximately 1 million years old, and has a dry tropical oceanic climate with little seasonal variation; its highest peak is Green Mountain (859 m). The island has three vegetation zones related to altitude: a bare and rocky dry zone at 0-330 m; the base surroundings of the mountain at altitudes between 330-660 m, with few bryophytes species; and the top of Green Mountain, altitude > 660 m, with a cloud forest and vegetation dominated by ferns and bryophytes (81 species of bryophytes, 4 hornworts, and 22 liverworts and 55 mosses, with 12 endemic species). The Island of Santa Helena is a volcanic island covering 122 km 2 , 4,000 km east of Rio de Janeiro (Brazil) and 1,950 km west of Africa; it is approximately 7 million years old, has a rugged terrain, and is one of the most remote regions in the world. The climate there is tropical, marine, and mild, with continuous winds, with an average annual temperature of 17-28° C. The highest point is Pico Diana (818 m), whose high altitude areas concentrate endemic species. It has three vegetation zones (tree fern forests; dry and eroded pastures and coastal zones; medium elevations with highly altered vegetation), and a total of 99 bryophyte species. Tristan da Cunha is an archipelago of volcanic origin formed by three islands: Gough, Inaccessible, and Nightingale, with a total of 305 species of bryophytes (61 being endemic), (Table 1). Methods Data collection A dataset was compiled (Excel spreadsheet) containing the information on species growing on the archipelagos of Azores, Madeira, São Tomé and Príncipe, Cape Verde, and the Canary Islands. The plant names were updated, with information concerning their distributions and endemism, and conservation data, with a total of 1520 recorded species. The data set also included information concerning the bryophyte species found on the islands and archipelagos of Trindade, Fernando de Noronha, Ascension, St Helena, and Tristan da Cunha, with a final total of 1647 species (675 hornworts plus liverworts, and 972 mosses). Herbarium collections Subsequently, ca. 600 specimens from the archipelago of São Tomé and Príncipe, ca. 80 from the archipelago of Cape Verde, and ca. 330 samples from the Azores archipelago were identified at the LISU herbarium. The nomenclature and synonyms for each species were checked using databases such as TROPICOS and some new literature. Data analysis ( diversity and floristic affinities) To determine bryophyte diversity on the ten islands and archipelagos, a floristic matrix was prepared using incidence data (presence/absence) to quantify diversity (Table 1 Supplementary Material). Rare species (occurring on only a single island or archipelago) were included. We used this matrix to calculate the Jaccard index, verify floristic similarities, and build a dendrogram of plant diversity among the ten islands and archipelagos. These statistical analyses were performed using PAST program version 4.12 (Hammer et al. 2001). The phytogeographic patterns of the taxa were characterized based on their global distributions, according to information available in the scientific literature and databases (Wigginton 2004, 2018; O’Shea 2006; Patiño-Llorente et al. 2005; TROPICOS database; and the general literature). These datasets included the more important literature and were used to analyze differences in the bryofloras of the ten islands and archipelagos in the Atlantic Ocean in relation to their total numbers of taxa, endemic species, exotic species, and possibly extinct species (Table 2), as well as the main families and any exclusive ones (Table 3). Results And Discussion Diversity of the ten islands and archipelagos of the Atlantic Ocean (alpha, beta, and gama diversity) A total of 1647 species ( gama diversity ) were listed for the ten islands and archipelagos in the Northern and Southern Atlantic Ocean , including 1152 species of mosses and 495 species of liverworts and hornworts. Of these, 197 species are endemic (63 species of liverworts and hornworts and 134 species of mosses), representing 12% of all species surveyed (Table 2); only 10 species were shared by all five archipelagos. The number of endemic species in the five North Atlantic Ocean archipelagos (197 spp.) was higher than the five islands and archipelagos of the South Atlantic Ocean , demonstrating their importance to the conservation of bryophyte biodiversity on Atlantic Ocean islands. This is especially critical because they will soon suffer the consequences of climate change – added to continuous pressure from tourism, pollution, population growth, the introduction of alien species, and habitat destruction. Table 2 Species diversity data for each studied island and archipelago. Spp = Total number of species, Ext = Extinct species, End = Endemic species, Exo = Exotic species Islands and archipelagos Spp. Ext End Exo Main references Azores 436 ? 40 1 Corley et al. (1981), Grolle (1983), and results of this study Madeira 577 ? 68 1 Persson (1939), Sjögren (1975, 2001), Sérgio et al. (2006), plus results of this study São Tomé and Príncipe 310 0 16 0 Sérgio and Garcia (2011), Garcia et al. (2012), and results of this study Cape Verde 224 ? 5 ? Leyens and Lobin (1996); Arechavaleta et al. (2005); Medina & Gomes (2015), and results of this study Canary Islands 501 ? 54 1 Persson (1939), Eggers (1982), Dirkse et al. (1993), Gonzáles et al. (2008) Trindade 35 ? 0 ? Serafin et al. (2010), Faria et al. (2012), Costa and Rezende (2022) Fernando de Noronha 32 ? 0 ? Vital et al. (1991), Serafin et al. (2010), Pereira and Câmara (2015), Costa et al. (2021) St Helena Island 99 0 26 24 Wigginton (2012) Ascension Island 81 4 16 3 Pressel et al. (2014) Tristan da Cunha 305 0 86 ? Dixon (1960), Wace (1961), Wace and Dickson (1965), Váña and Engell (2013) Finally, with the inclusion of species from the South Atlantic Ocean islands and archipelagos (Fernando de Noronha, Trindade, St Helena, Ascension, and Tristan da Cunha) to analyze and compare them with those of the North Atlantic Ocean ( gama diversity ), the total number of species increased to 1647 species (11 hornworts, 664 liverworts, and 972 mosses). Species richness varied strongly among the ten southern Atlantic islands and archipelagos, ranging from 32 species on Fernando de Noronha, to 577 on Madeira (Table 2). The bryophyte diversity on the ten islands and archipelagos studied here ( gama diversity) was high when compared to Africa, Tropical America, and the world (Gradstein et al. 2001; Shaw and Goffinet 2000; O’Shea 2006; Wigginton 2018), (Fig. 2). Phytogeographical patterns and endemism We analyzed the islands and archipelagos of the North and South Atlantic Oceans separately, and then together; the results are presented in Tables 3, 4. The phytogeographical patterns of the North Atlantic Ocean islands and archipelagos were combined into ten categories for this analysis: 1) Global (130 taxa; 23 liverworts and 107 mosses); 2) Africa (194 taxa; 117 liverworts and 23 mosses); 3) Endemic (91 taxa; 22 liverworts and 69 mosses); 4) North America, Europe and Asia (49 taxa of mosses); 5) North America and Europe (19 taxa of mosses); 6) Europe, Macaronesia, Africa (14 taxa of liverworts); 7) Europe, Macaronesia, Asia (9 taxa of liverworts); 8) Europe and Macaronesia (55 taxa; 13 liverworts and 42 mosses); 9) South America and Africa (11 taxa of liverworts); 10) others. The African, Endemic, and Global elements were the largest, comprising 11% of the total taxa (Table 3). Table 3 Phytogeographical patterns of the bryofloras of the five North Atlantic Ocean islands and archipelagos Patterns Liverworts Mosses Total Worldwide 23 107 130 Africa 117 77 194 N America, Europe and -- 49 49 N America and Europe -- 19 19 Europe, Macaronesia, Africa 14 -- 14 Europe, Macaronesia, Asia 9 -- 9 Europe and Macaronesia 13 42 55 South America and Africa 11 -- 11 Endemic 22 69 91 Others 217 352 569 Total 426 715 1141** ** This number is different because there are taxa identified only at genera level We found that the liverworts and mosses exhibited distinct phytogeographical patterns (Table 3). Liverworts showed low numbers of European and Macaronesia species (13 spp.), but the highest number of African elements (117), followed by global elements (23) and endemic species (22); mosses showed high numbers of global elements (107), followed by African and Endemic elements (77 and 69). The phytogeographical patterns of the South Atlantic Ocean islands and archipelagos were combined into eight categories for this analysis: 1) Global (73 taxa; 9 liverworts and 64 mosses); 2) Pantropical (24 taxa; 8 liverworts and 16 mosses); 3) Afro-America (30 taxa; 19 liverworts and 11 mosses); 4) Tropical America (27 taxa; 15 liverworts and 12 mosses); 5) Africa (26 taxa; 9 liverworts and 17 mosses); 6) South America (77 taxa; 66 liverworts and 10 mosses); 7) Endemic (100 taxa;43 liverworts and 57 mosses); 8) others (122 taxa; 72 liverworts and 50 mosses). Endemic elements was the largest category, comprising ca. 21% of all taxa (Table 4). Table 4 Phytogeographical patterns of the bryofloras of the five South Atlantic Ocean islands and archipelagos (Trindade, Fernando de Noronha, Ascension, St Helena, and Tristan da Cunha) Patterns Liverworts Mosses Total Worldwide 9 64 73 Pantropical 8 16 24 Afro-America 19 11 30 Tropical America 15 12 27 Africa 9 17 26 South America 66 10 77 Endemic 43 57 100 Others 72 50 122 Total 241 237 478** ** This number is different because there are taxa identified only at genera level The liverworts and mosses exhibited distinct phytogeographical patterns on the South Atlantic Ocean islands and archipelagos. Liverworts had high numbers of species with South American distributions (66 spp.), followed by endemic species (43), but low numbers of species with global distributions (9); mosses had high numbers of endemic species (57) and species having global distributions (64). The phytogeographical patterns of the islands and archipelagos of the North Atlantic Ocean were combined into ten categories for this analysis: 1) Global (130 taxa; 23 liverworts and 107 mosses); 2) Africa (194 taxa; 117 liverworts and 23 mosses); 3) Endemic (91 taxa; 22 liverworts and 69 mosses); 4) North America, Europe and Asia (49taxa of mosses); 5) North America and Europe (19 taxa of mosses); 6) Europe, Macaronesia, Africa (14 taxa of liverworts); 7) Europe, Macaronesia, Asia (9 taxa of liverworts); 8) Europe and Macaronesia (55 taxa; 13 liverworts and 42 mosses); 9) South America and Africa (11 taxa of liverworts); 10) others (569 taxa; 217 liverworts and 352 mosses). African, Endemic, and Global elements were the largest categories, comprising 11% of the total taxa (Table 5). Table 5 Phytogeographical patterns of the bryofloras on the five North Atlantic Ocean islands and archipelagos (Azores, Madeira, São Tomé and Príncipe, Cape Verde, and Canary Islands) Patterns Liverworts Mosses Total Worldwide 23 107 130 Africa 117 77 194 N America, Europe and -- 49 49 N America and Europe -- 19 19 Europe, Macaronesia, Africa 14 -- 14 Europe, Macaronesia, Asia 9 -- 9 Europe and Macaronesia 13 42 55 South America and Africa 11 -- 11 Endemic 22 69 91 Others 217 352 569 Total 426 715 1141** ** This number is different because there are taxa identified only at genera level The liverworts and mosses were found to exhibit distinct phytogeographical patterns on the South Atlantic Ocean islands and archipelagos (Table 5). Liverworts there had high numbers of African elements (117 spp.), followed by global elements (23) and endemic species (22), but low numbers of Europe and Macaronesia species (13); mosses there showed high numbers of global species (107), followed by African elements (77), and Endemic species (69). A striking feature of the bryofloras of the archipelagos in the North Atlantic Ocean is that species richness was concentrated in just a few families (Table 5, Fig. 3), with mosses, Brachytheciaceae (44 species), Bryaceae (47), Fissidentaceae (58), and Pottiaceae (135) standing out and totaling together 284 species (fully 31% of the total of 916 moss species recorded for the five archipelagos). In terms of the liverworts, just three families (Lejeuneaceae [121 species], Plagiochilaceae [33], and Ricciaceae [34]) concentrated 32% of the total number of liverwort species (593) recorded for the five archipelagos. The species richness of bryophytes was therefore concentrated in seven families that together represented 40% of the total number of species (1161). The species richness of the bryofloras of the South Atlantic Ocean islands and archipelagos was concentrated in six families (Table 6, Fig. 3), Lejeuneaceae (38 species), Lophocoleaceae (40), Bryaceae (25), Fissidentaceae (28), Grimmiaceae (11), and Pottiaceae (25), which together totaled 167 species, representing 27% of the total. Table 6 Total numbers of bryophyte families and genera, as well as the main families in each archipelago or island. Bold = families standing out in terms of their numbers of species Islands or Archipelagos Families Genera Main families Azores 90 128 LIVERWORTS : Aneuraceae, Calypogeiaceae, Cephaloziaceae, Cephaloziellaceae , Fossombroniaceae, Geocalycaceae, Lejeuneaceae , Lepidoziaceae, Lophocoleaceae , Plagiochilaceae, Radulaceae, Ricciaceae , Scapaniaceae. MOSSES : Bartramiaceae, Brachytheciaceae , Bryaceae, Ditrichaceae, Fissidentaceae, Grimmiaceae , Hypnaceae, Leucobryaceae , Neckeraceae, Polytrichaceae, Pottiaceae , Sphagnaceae Madeira 92 197 HORNWORTS : Anthocerotaceae LIVERWORTS : Cephaloziaceae , Cephaloziellaceae, Fossombroniaceae, Frullaniaceae, Lejeuneaceae , Marchantiaceae, Plagiochilaceae , Ricciaceae , Scapaniaceae MOSSES : Amblystegiaceae, Bartramiaceae, Brachytheciaceae , Bryaceae , Ditrichaceae, Fissidentaceae , Funariaceae, Grimmiaceae , Leucobryaceae, Mniaceae, Neckeraceae, Orthotrichaceae , Polytrichaceae, Pottiaceae , Sphagnaceae São Tomé and Príncipe 62 104 HORNWORTS : Dendrocerotaceae LIVERWORTS : Frullaniaceae, Lejeunaceae , Metzgeriaceae, Plagiochilaceae , Radulaceae, Ricciaceae MOSSES : Bryaceae, Calymperaceae, Fissidentaceae , Hypnaceae, Orthostichellaceae, Pilotrichaceae, Pottiaceae Cape Verde 49 105 LIVERWORTS : Frullaniaceae, Lejeuneaceae, Ricciaceae MOSSES : Bartramiaceae, Brachytheciaceae, Bryaceae, Fissidentaceae , Orthotrichaceae, Pottiaceae Canary 90 187 HORNWORTS : Anthocerotaceae LIVERWORTS : Cephaloziellaceae, Frullaniaceae, Lejeuneaceae, Lophocoleaceae, Plagiochilaceae, Radulaceae, Ricciaceae , Riellaceae, Sphaerocarpaceae MOSSES : Bartramiaceae, Brachytheciaceae, Bryaceae , Ditrichaceae, Fissidentaceae , Funariaceae, Grimmiaceae , Leucobryaceae, Neckeraceae, Orthotrichaceae, Polytrichaceae , Pottiaceae , Trindade 16 25 LIVERWORTS : Lejeuneaceae (12 spp.) Fernando de Noronha 14 18 HORNWORTS : Nothothyladaceae (2 spp.) MOSSES : Bryaceae (4 spp.), Fissidentaceae (12 spp.) Ascension 23 45 HORNWORTS : Nothothyladaceae (3 spp.) LIVERWORTS : Lejeuneaceae (5 spp.) MOSSES : Bryaceae (8 spp.), Fissidentaceae (4 spp.), Leucobryaceae (7 spp.), Pottiaceae (12 spp.) St Helena 40 63 LIVERWORTS : Lejeuneaceae (15 spp.) MOSSES : Bryaceae (7 spp.), Fissidentaceae (11 spp.), Leucobryaceae (4 spp.), Pottiaceae (9 spp.) Tristan da Cunha 68 142 LIVERWORTS : Adelanthaceae (10 spp.), Anastrophyllaceae (9 spp.), Aneuraceae (13 spp.), Cephaloziellaceae (10 spp.), Lejeuneaceae (14 spp.), Lepidoziaceae (14 spp.), Lophocoleaceae (36 spp.), Plagiochilaceae (9 spp.) MOSSES : Andreaeaceae (7 spp.), Bartramiaceae (10 spp.),Bryaceae (10 spp.), Ditrichaceae (8 spp.), Grimmiaceae (11 spp.), Polytrichaceae (6 spp.), Pottiaceae (6 spp.) There were differences among the islands and archipelagos with unique families, or among those which had significantly different numbers of species ( beta diversity ), Table 7. The Brazilian archipelagos In the South Atlantic Ocean (Trindade and Fernando de Noronha) are examples of how habit destruction by humans can influence bryophyte species diversity, as those archipelagoes have only 35 and 32 species respectively. Those numbers represent the lowest diversity of bryophyte species among the ten islands and archipelagos studied here, and neither has an endemic species. That situation is in contrast to the two English islands and one archipelago (Ascension, St. Helena, and Tristan da Cunha) which have between 81 and 312 species of bryophytes, including between 16 to 86 endemic species – demonstrating their degrees of preservation. The African archipelagos of Cape Verde and São Tomé and Príncipe in the North Atlantic Ocean have between 215 and 314 bryophyte species, including 5 to 16 endemic species. The Portuguese archipelagos of Azores and Madeira, and the Spanish Canary Islands archipelago evidenced the highest diversity and shared more species, with the Canary Islands being the most preserved (with a total of 501 bryophyte species, of which 54 are endemic). AZORES – Three bryophyte families occur only on this archipelago; Jamesoniellaceae is represented there by a species that occurs nowhere else outside of the Neotropics ( Syzygiella rubricaulis (Nees) Steph.). The Sphagnaceae family also stands out in this archipelago, with 17 species (Hodgetts and Lockhart 2020). This archipelago shares its highest number of species with the Madeira archipelago (379 species, including 37 endemic species). MADEIRA – Three families occur only on this archipelago: Blasiaceae (Liverworts), Aongstroemiaceae, and Flexitrichaceae (Mosses). The families Brachytheciaceae (34 species) and Grimmiaceae (27) also stand out (Hodgetts and Lockhart 2020). This archipelago shares the highest number of species with the Azores , as noted above. SÃO TOMÉ and PRÍNCIPE – Has the highest number of restricted families. This is the only archipelago with Dendrocerotaceae, a hornwort family, as well as moisture-indicating families such as Hypopterygiaceae and Meteoriaceae. It shares 23 species with Cape Verde , 20 with the Canary Islands, 21 with the Azores , and 22 with the Madeira archipelago. CAPE VERDE – Two bryophyte families occur only in this archipelago; 69 exclusive species are found there, with four being endemic. This is the most unique archipelago among the five studied in the North Atlantic Ocean, having the smallest total number of species and endemic species. It also shares a high number of taxa with the Canary Islands (118 species). CANARY ISLANDS – Three bryophyte families occur only on this archipelago. It shares the most species with the Azores and Madeira (283 species, with 27 being endemic) and with Cape Verde (118 taxa). This archipelago, together with Madeira , has the best-known bryofloras, which have been presented in many different publications. TRINDADE – No bryophyte family occurs exclusively on this island. Its native vegetation has been almost completely destroyed through the grazing of introduced animals. E. Knight visited the island in 1881 and 1889 and reported its already degraded vegetation and a large waterfall on the west coast. Vegetation restoration efforts were initiated in 1990 through the elimination of goats. The vegetation has slowly recovered since then, and its creeks have more water. The vegetation above 400 m is considered a “Giant Fern Forest" and is dominated by a single species, Cyathea delgadii Sternb.; it is the most diverse region for bryophytes. Most of the species are Neotropical, followed by cosmopolitan species; there are no endemic species (Costa and Rezende 2022). The growth of Campylopus introflexus was probably facilitated by human impacts on the original vegetation. This archipelago shares six species with Ascension Island and five species with the Fernando de Noronha archipelago. FERNANDO DE NORONHA – Only one moss family, Stereophyllaceae, occurs on this island. Charles Darwin visited the island and made the first botanical collections there. A later expedition by Ridley (1890) reported the predominance of herbaceous vegetation due to the intensive cutting of tall trees that could be used to build rafts and aid prisoner escapes. Fissidens , with 12 species, is the richest genus. Cosmopolitan species compose approximately 43% of vegetation; there are no endemic species (Costa and Rezende 2022). This archipelago shares five species with Ascension Island. ASCENSION – No bryophyte family occurs conclusively on this island. The island harbors over 200 introduced species ranging from large trees to shrubs and herbs; many native plants, such as the endemic fern Pitsana purpurascens (de Vriese) Murdock, are becoming severely outcompeted. Campylopus, with seven species, is the richest genus. There is a notable absence of cosmopolitan species ( Ceratodon purpureus (Hedw.) Brid. , Tortula muralis Hedw. , Octoblepharum albidum Hedw., etc.). There are no representatives of Grimmiales, despite the highly suitable volcanic rock substrate there. The island has fifteen endemic bryophyte species. A handful of taxa are probable human introductions, but despite the small size of the island and the destruction of its natural habitats, relatively few bryophytes are threatened with extinction. This island shares 21 species with St Helena Island, three of which are endemic. ST HELENA – No bryophyte family occurs only on this island, but there is a notable absence of cosmopolitan species, Ceratodon purpureus, Tortula muralis, Octoblepharum albidum , etc. There are no representatives of Grimmiales despite the highly suitable volcanic rock substrate there. Fissidens, with 11 species, is the richest genus (Wigginton 2013). The destruction of the natural vegetation began with the introduction of goats, and the establishment of permanent settlements there introduced exotic vegetation. Native trees were cut for different purposes (house building, fuel, etc.), and many endemic species are now extinct or critically endangered. This archipelago shares 16 species with the Tristan da Cunha archipelago, three of which are endemic. TRISTAN DA CUNHA – Eleven bryophyte families occur only on this archipelago. There are currently very few introduced species on the archipelago. The family Orthotrichaceae is represented by several endemic species, and Dicranoloma (Renauld) Renauld by three endemic species. This archipelago shares 28 species with the Azores archipelago and 23 with Madeira , none are endemic. There are strong indications of a close connection with South America, as mosses, liverworts, and hornworts can be dispersed by the western winds – ‘West Wind Drift’ – and connections with South America are conspicuous in all groups of non-vascular plants, except the algae (Galloway 1996, Milius 2004, Muñoz et al. 2004). Sunding (1979) considered Cape Verde to be different from the other archipelagos of Macaronesia, which was subsequently corroborated by Vanderpoorten et al. (2007). According to these authors, Macaronesia (comprising the archipelagos of Azores, Madeira, Canary Islands, and Cape Verde) is considered a single biogeographical unit, with their floras being relics of a widely distributed tertiary subtropical flora. These authors rejected the concept of Macaronesia sensu lato , and viewed the Cape Verde archipelago as being more associated with tropical Africa. Liverworts in the other Macaronesian archipelagos support an Azores-Madeira-Canary clade (Macaronesia sensu stricto ), while mosses support the Canary Islands as being related to North Africa, rejecting the concept of Macaronesia s.s. for the group. The exchange of taxa with neighboring continental areas better explains the relationships between the cryptogamic flora of Cape Verde and the mossy flora of the Canary Islands. In contrast, the relic flora is consistent with a monophyletic Macaronesia s.s. group of liverworts. These congruent patterns, however, can hide a complex mix of relict distributions and more recent speciation and dispersal events. Similarities among the ten islands and archipelagos (beta diversity) The results of our study, when comparing the bryofloras of the ten Atlantic Ocean islands and archipelagos , demonstrated that their bryofloras were very dissimilar, with low floristic affinities (Table 7). Of the ten islands and archipelagos, those demonstrating the greatest similarity were the archipelagos of Macaronesia, Azores, Madeira, and the Canary Islands, which share 283 species (Fig. 4 A,B,C). Our results also indicated Macaronesia as being formed by the Azores, Madeira, and the Canary Islands archipelagos, with Cape Verde outside the clade when the islands and archipelagos of the North Atlantic Ocean were analyzed separately or together with the islands and archipelagos of the South Atlantic Ocean (Fig. 4 A,C). As expected, the Azores, Madeira, and Canary Islands archipelagos (Macaronesia s.s. ) shared several species and evidenced the highest similarity. The archipelago of São Tomé and Príncipe was found to be highly dissimilar to the others, sharing few species with any of the North Atlantic Ocean archipelagos (Fig. 4 A,C). The results for the South Atlantic Ocean also showed low similarity among almost all of the islands and archipelagos, although the islands of St Helena and Ascension show the highest similarity and form a separate clade (with Tristan da Cunha outside of it). The islands of St Helena and Ascension share several species. The archipelagos of Trindade and Fernando de Noronha also formed a separate clade, although they share few species; this can be attributed to the degree of devastation suffered by these two archipelagos since their discovery and occupation (Fig. 4 B,C). When we analyzed the ten Atlantic Ocean islands and archipelagos together, their similarities were very low, with the exception of Madeira, Azores, and the Canary Islands (Marcaronesia s.s .), each demonstrating an essentially unique bryoflora. Our results also corroborate the separation of Cape Verde from Macaronesia (Fig. 4 C). Table 7 Similarities among the ten islands and archipelagos studied. TRI FN ASC STH TRC AZO MAD STP CV CAN TRI 1 0,07936508 0,03703704 0,0229008 0,01183432 0,01616162 0,01470588 0,01557632 0,0170213 0,01444 FN 0,079365 1 0,04761905 0,0076336 0 0,00804829 0,01309329 0,03184713 0,0171674 0,01083 ASC 0,037037 0,04761905 1 0,1503268 0,03225807 0,0341556 0,03755869 0,04843305 0,0446097 0,030717 STH 0,022901 0,00763359 0,1503268 1 0,03571429 0,04805915 0,03945372 0,03723404 0,0557491 0,036424 TRC 0,011834 0 0,03225807 0,0357143 1 0,04306864 0,03476246 0,01355932 0,0240481 0,033457 AZO 0,016162 0,00804829 0,0341556 0,0480592 0,04306864 1 0,53868613 0,03265306 0,1666667 0,421429 MAD 0,014706 0,01309329 0,03755869 0,0394537 0,03476246 0,53868613 1 0,02934272 0,1755952 0,518417 STP 0,015576 0,03184713 0,04843305 0,037234 0,01355932 0,03265306 0,02934272 1 0,0576923 0,031526 CV 0,017021 0,01716738 0,04460967 0,0557491 0,0240481 0,16666667 0,17559524 0,05769231 1 0,196399 CAN 0,01444 0,01083033 0,03071672 0,0364238 0,03345725 0,42142857 0,51841746 0,03152585 0,1963994 1 Conservation The ten islands and archipelagos studied here proved to be very rich areas for bryophytes, with their numbers being equivalent to 43% of all of the species known to Africa (3800 species), 41% of all bryophyte species known to tropical America (4000 species), and 8% of all species known globally. Our study indicated that the vegetation on the islands and archipelagoes must be preserved and protected, as 91 species are only known from them. Additionally, more botanical collections should be undertaken on Tristan da Cunha and São Tomé and Príncipe. Our results reinforce the importance of the bryofloras of the Atlantic Ocean islands and archipelagos to bryophyte conservation, as they hold high percentages of the total bryophyte diversity of Africa and tropical America, as well as the total species diversity known for the Macaronesia region within their small areas. The main threats to the biodiversity on all ten islands and archipelagos studied here include species introductions, pollution (maritime traffic, tourism, fishing), the introduction of invasive species with their potential to alter native habitats and compete with native species (as occurs in Saint Helena – New Zealand flax), and rising ocean levels due to climate change. Five of the North Atlantic Ocean islands and archipelagos studied here had 240 taxa classified as being threatened to some degree (2 EX - extinct, 15 CR - critically endangered, 72 EN - endangered, 145 VU - vulnerable, 6 - least concern LC ) – fully 20% of the total number of species. This study confirms the importance of these islands and archipelagos as important sites of bryophyte diversity for Macaronesia and Africa. There is, however, a lack of adequate protection for endemic and/or endangered species on these islands and archipelagos, which should be addressed with priority, as global warming will accelerate the destruction of such sensitive environments around the planet. The bryophytes of the Macaronesian laurel forests ( Laurisilva ) are also at considerable risk due to climate change (Patiño et al. 2016, Patiño and Vanderpoorten 2018). Many areas are predicted to become water-deficient, and forest fires are becoming more frequent – with projections indicating significantly increased risks to important endemic species (Hodgetts et al. 2019). There are still many other threatened bryophyte species from a conservation perspective, with tropical species frequently referenced solely from type specimens or based on small private collections from Africa; approximately 45 bryophyte taxa reported for São Tomé and Príncipe, for example, have not been collected again since the 19 th century. The ecology, habitat specificity, and distribution patterns of a large number of bryophyte species are not well known, and threats to the forest habitats in São Tomé and Príncipe (including habitat destruction and competition from invasive species, for example) can affect the survival of many bryophyte species (Garcia et al. 2022). Conclusions and conservation implications The present study represents the first survey of bryophyte diversity on ten Atlantic Ocean Islands and archipelagos. The results have increased our knowledge of the bryofloras of island habitats, and have introduced a considerable number of novelties and new records. We consider this data to be relevant to the conservation of bryophyte diversity on oceanic islands and archipelagos in the Atlantic Ocean, and to subsidize Global Strategy for Plant Conservation (GSPC) targets. This survey of ten Atlantic Ocean islands and archipelagos provides a better understanding of their floristic compositions and increases our estimates of species richness and distributions – although this information is far from complete and additional inventories will still be needed in unexplored areas on the Tristan da Cunha and São Tomé and Príncipe archipelagos. Bryophyte richness and diversity differed significantly among study sites (alpha diversity), and their low similarities (beta diversity) illustrated their high degrees of heterogeneity and reflected the importance of habitat differences for high diversity. A better understanding of the factors related to species richness and endemism on oceanic islands will be important for conservation efforts and biodiversity management, especially in view of global climate change (Grabherr et al. 2010). Climate change is expected to result in strong impacts on oceanic islands and archipelagos, with habitat losses and declines, and extinction risks, as isolated bryophyte species in these regions have no access to refuges. The complexity of island ecosystems is associated with their high diversity and endemism, but researchers will need to generate primary data and integrate available geological and climatic information into diversity models. Further studies will be necessary to develop proactive conservation plans, for these environments hold a large fraction of the world’s biodiversity. The main threats to species diversity, forest structure, and habitats include urbanization, habitat destruction and fragmentation, the introduction of invasive species that can affect the survival of native species, uncontrolled tourism, and rising oceans (which will mainly affect islands smaller than 1,000 ha - Médail 2017). Our results established baselines of bryophyte diversity and endemism for ten Atlantic Ocean islands and archipelagos, which can be compared with future surveys and could help to determine whether species can migrate or will become extinct due to climate change. The unique features of those oceanic islands and archipelagos demand conservation strategies considering their biological and environmental diversities and their importance as protected areas. Declarations Acknowledgments The authors would like to thank CNPq for the PDE grant (no. 200858/2022-0), also to University of Lisbon, National Museum of Natural History and Science for provide all the infrastructure for the development of the work. Finally, the collaboration of the colleagues César A. Garcia, Fátima Sales, Ireneia Melo, Jorge Paiva, José Cardoso, and Maria Cristina Duarte, and that collected and sending samples from different islands. Author contributions Denise Pinheiro Costa developed the idea of this publication, both authors contributes to the writing and editing draft and gave approval for its publication. Conflict of interest The authors declare no competing interest.s Ethical approval Not applicable. References Corley MFV, Crundwell, AC, Düll, R, Hill, MO, Smith, AJ E (1981) Mosses of Europe and the Azores; an annotated list of species, with synonyms from the recent literature. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3389905","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":236437635,"identity":"dcc52b28-4f87-44ef-988e-962f63844608","order_by":0,"name":"Denise Pinheiro Costa","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABMElEQVRIie3PsUrDQBjA8TuyXto1xYKvkOAgUqQP4hIpmKWH6w2xXCgki8W1YKmvEJfMOQ6S5dA1YMGIUJcOZksHxUvM1DTiKJg/yXF85MflAGhr+4uF5QMQ0MqNUo2JfBWF/ppACkRBYCP5TivXikC3onvqxDPGcrLqHx9MGSdkcNj1vLd0u7w663qS5CTYJT3xYHIk1uhkEZlMCMuYC2E4syDGcy5PuxZPu0RPxjqHLkfFhjkuh742hlQNIkwlUaBbJ88bnW0/C3L5XpChr1kvzsciwndNJEF6qNLyFFCQc18zjalKbew3kJ6QP4aitSQXOqPCGhV3ue1HIb6XhO25SycWR1lur4Z6MnrNKBmc3nhemm3sCV4+cpbmpEaqwtqEN8x/IJPmj9va2tr+W18JfIr8ZQN9XAAAAABJRU5ErkJggg==","orcid":"","institution":"Instituto de Pesquisas Jardim Botânico do Rio de Janeiro","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Denise","middleName":"Pinheiro","lastName":"Costa","suffix":""},{"id":236437636,"identity":"916097f6-c1c3-44d4-a1e5-a0427aed6861","order_by":1,"name":"Cecília Sérgio","email":"","orcid":"","institution":"Universidade de Lisboa, Museu Nacional de História Natural e da Ciência","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Cecília","middleName":"","lastName":"Sérgio","suffix":""}],"badges":[],"createdAt":"2023-09-26 18:59:19","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3389905/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3389905/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":44093917,"identity":"e7daf08f-03d1-4b27-9b51-4c1b4102b513","added_by":"auto","created_at":"2023-10-04 16:50:38","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":163560,"visible":true,"origin":"","legend":"\u003cp\u003eLocation of islands and archipelagos studied in the Northern and Southern Atlantic Ocean\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-3389905/v1/38c11da4aa7391086353cfe2.png"},{"id":44093914,"identity":"20950eb1-7b2b-45a6-a91c-082c6cbfae47","added_by":"auto","created_at":"2023-10-04 16:50:37","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":23596,"visible":true,"origin":"","legend":"\u003cp\u003eContributions of the oceanic islands and archipelagos to bryophyte species richness of the Atlantic Ocean and the world as a whole. The numbers in brackets signify the percentage of the islands in relation to the region.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-3389905/v1/0735e23ed66f4635f73bbeb5.png"},{"id":44095250,"identity":"1ec4c690-d1cc-4e5d-bf5e-0c4661602379","added_by":"auto","created_at":"2023-10-04 16:58:38","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":22390,"visible":true,"origin":"","legend":"\u003cp\u003eMain families of liverworts and mosses encountered on the ten Atlantic Ocean islands and archipelagos studied (578 species = 35% of the total number)\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-3389905/v1/b5f371f5fac1475021519676.png"},{"id":44093915,"identity":"f7d5e819-9f96-439a-a078-65ac1a83ec39","added_by":"auto","created_at":"2023-10-04 16:50:38","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":42946,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eA. \u003c/strong\u003eSimilarities among the five North Atlantic Ocean islands and archipelagos. \u003cstrong\u003eB.\u003c/strong\u003e Similarities among the five South Atlantic Ocean islands and archipelagos. \u003cstrong\u003eC.\u003c/strong\u003e Similarities among the ten islands and archipelagos studied. FN = Fernando de Noronha, TRI = Trindade, ASC = Ascension, STH = Saint Helena, TRC = Tristan da Cunha, AZO = Azores, MAD = Madeira, STP = São Tomé and Príncipe, CV = Cape Verde, CAN = Canary Islands\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-3389905/v1/0fc3d2746f88ce15926476ce.png"},{"id":48495791,"identity":"5bc78735-cd86-4b17-b5ba-06e5fc853faf","added_by":"auto","created_at":"2023-12-19 22:07:27","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":998700,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3389905/v1/5b275015-ef83-4182-b186-029ccb6ce864.pdf"},{"id":44093918,"identity":"844f3874-6937-4187-ba34-3d3a125ab0cc","added_by":"auto","created_at":"2023-10-04 16:50:38","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":279004,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementarymaterialTable1.docx","url":"https://assets-eu.researchsquare.com/files/rs-3389905/v1/bb39bf5203d673a0fa23c0b6.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Tropical Atlantic Oceanic Islands and Archipelagos: Physical structures, plant diversity, and affinities of the bryofloras of northern and southern islands","fulltext":[{"header":"Introduction","content":"\u003cp\u003eScientists have long been fascinated by the unique nature of remote oceanic islands, especially after Charles Darwin\u0026rsquo;s study of the Galapagos Islands. Young oceanic islands of volcanic origin represent the summits of large volcanoes that rose from the bottom of the ocean. They are isolated from other landmasses by deep seas and have never been connected to continental landmasses so their environments are unique products of trans-oceanic dispersal and eventual speciation (Wace \u0026amp; Holdgate 1976; Cowie and Holland 2006). Islands therefore represent natural laboratories that simplify the complexities of the natural world (Whittaker and Fernandez-Palacios 2007). The introduction of non-native species, however, is a constant threat to an island\u0026rsquo;s biota, and it is generally known that warmer temperatures can improve the chances of alien invasive species settling, reproducing, and successfully outcompeting the native biota, and can lead to significant biodiversity losses.\u003c/p\u003e\n\u003cp\u003eThere are numerous islands and archipelagos in the Atlantic Ocean: including the archipelagos of the Azores and Madeira within the territorial waters of Portugal; the archipelago of the Canary Islands (Spain); the archipelagos of Cape Verde and S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe (Africa); Fernando de Noronha, Trindade and Martin Vaz (Brazil); and the islands of St Helena, Ascension and the archipelago of Tristan da Cunha (England) (Fig. 1).\u003c/p\u003e\n\u003cp\u003eThe bryofloras of the ten islands and archipelagos were inventoried: Azores, Madeira, S\u0026atilde;o Tom\u0026eacute; e Pr\u0026iacute;ncipe, Cape Verde, and Canary islands (Northern Atlantic) and Trindade, Fernando de Noronha, Ascension, St Helena and Tristan da Cunha (Southern Atlantic) to: (1) provide an overview of their bryofloras; (2) describe the species richness of mosses and liverworts and incorporate new reports of bryophyte taxa; (3) analyze the distribution patterns and endemism of those species; (4) analyze the similarities among northern and southern islands; and (5) establish a diversity baseline for comparisons. We describe here the diversity, endemism, and similarities of mosses and liverworts on ten islands and archipelagos located in the Atlantic Ocean.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eStudied Islands and Archipelagos (physical structures and bryofloras)\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe \u003cstrong\u003ePortuguese archipelagos\u003c/strong\u003e are of volcanic origin and represent important geological sites. The Azores archipelago comprises nine islands. It is located ca. 1600 km from the European coast and represents a chain of submarine volcanic islands with sources of fresh water. The archipelago harbors a total of 436 species of bryophytes, with the diversity among the different islands varying greatly, from 104-324 species; only 5% of the archipelago is covered by forest vegetation. The Madeira archipelago is located 978 km from Portugal and ca. 700 km from Africa. It is also of volcanic origin, with two main islands (Madeira and Porto Santo), as well as two groups of uninhabited islands (Desertas and Selvagens). The highest points there are the Pico Ruivo (1862 m), Pico das Torres (1851 m), and Pico do Arieiro (1818 m). The mountainous landscape and their exposure to constant winds mean that the small islands have different microclimates and higher precipitation rates on the northern coast; few places have original intact vegetation, and they have no sources of fresh water. The Madeira archipelago has 577 species of bryophytes (Table 1).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 1\u003c/strong\u003e General data on the ten islands and archipelagos studied.\u0026nbsp;AZO = Azores, MAD = Madeira, STP = S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe, CV = Cape Verde, CAN = Canary Islands, FN = Fernando de Noronha, TRI = Trindade, ASC = Ascension, STH = St Helena, TRC = Tristan da Cunha, END = Endemic\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"945\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eAREA\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eORIGIN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eCLIMATE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eHABITAT\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eMOSSES\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eLIVERWORT\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eHORNWORT\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTOTAL\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eAZO\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (9 islands), 0.3-0.8 mya, 2,351 Km\u003csup\u003e2\u003c/sup\u003e, population: 236,440 permanent residents, highest point is Monte do Pico (2,351 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eSubtropical oceanic, 17\u0026deg;C, 1,026 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eLaurissilva forest\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e270 spp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e160 spp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e6 spp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e436 spp.\u003c/p\u003e\n \u003cp\u003e(40 END \u0026ndash; 10%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eMAD\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (4 islands), 5 mya, 724 Km\u003csup\u003e2\u003c/sup\u003e, population: 262,456 permanent residents (Madeira Island), highest peak is Pico Ruivo (1,862 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eMediterranean and Temperate in the highest parts, 20\u0026deg;C, 2,000 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eLaurissilva forest (best preserved evergreen forests in Macaronesia), with a great diversity of bryophytes covering the trunks and branches of trees and shrubs\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e389 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e182 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e6 spp.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e577 spp.\u003c/p\u003e\n \u003cp\u003e(68 END \u0026ndash; 12%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSTP\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (3 islands), 31 mya, 991 \u0026nbsp;Km\u003csup\u003e2\u003c/sup\u003e, population: 213.000 permanent residents, highest peak is Pico de S\u0026atilde;o Tom\u0026eacute; (2024 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eEquatorial oceanic (hot and wet), 22-30\u0026deg;C, 214 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eTropical moist broadleaf forest\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e133 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e170 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e7 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e310 spp.\u003c/p\u003e\n \u003cp\u003e(16 END \u0026ndash; 7%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eCV\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (10 islands, and one active vulcan in the Fogo Island), 8-20 mya, 4,033 Km\u003csup\u003e2\u003c/sup\u003e, population: 561.901 (nine islands have permanent residents), highest peak is Pico do Fogo (2829 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eArid to semi-arid, 22-27\u0026deg;C, 214 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eSavanna or steppe, and tropical climate depending on elevation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e168 spp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e54 spp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e2 spp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e224 spp.\u003c/p\u003e\n \u003cp\u003e(5 END \u0026ndash; 2%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eCAN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (7 islands), 8-20 mya, 7,493 Km\u003csup\u003e2\u003c/sup\u003e, population: 2.2 million permanent residents, highest peak is Monte Teide (3715 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eWarm subtropical and semi-desertic\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eSubtropical, with xerophytic, humid forest, the laurel forest, fayal-brezal, the pine forest and high mountain vegetation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e352 spp.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e143 spp.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e6 spp.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e501 spp.\u003c/p\u003e\n \u003cp\u003e(54 END \u0026ndash; 11%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eFN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (21 islands), 30 mya, 26 Km\u003csup\u003e2\u003c/sup\u003e, population: 3,100 permanent residents, highest peak is Desejado (323 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eTropical wet and dry, 26,5\u0026deg;C, 1,350 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eSeasonal deciduous forest, subxerophytic (species typical of the northeastern \u003cem\u003eagreste\u003c/em\u003e). Dry Forest (Ponta da Sapata), 25% of shrubs and trees on the main island. Mangrove (only occurrence of an insular mangrove in the South Atlantic Ocean) is located in the Sueste Bay. Creepers covering native bushes and trees during the rainy season.\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e23 spp.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e7 spp.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e2 spp.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e32 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTRI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (5 islands), 3 mya, 10.4 Km\u003csup\u003e2\u003c/sup\u003e, population: 32 soldiers + 8 researches (no permanent residents), highest peak is Pico do Desejado (620 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eTropical oceanic, 25\u0026deg;C, 923 mm precipitation\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eExotic plants (100-150 m), lowland (up to 400 m) grass fields, Giant Fern Forest or Giant Ferns Nebular Forest (\u0026gt; 400 m). Less than 5% of the island is covered by forest and ca. 60% by herbaceous vegetation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e14 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e20 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e1 sp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e35 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e(NO END)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eASC\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic, 1 mya, 91 Km\u003csup\u003e2\u003c/sup\u003e, population: 800 inhabitants (no permanent residents), highest peak is Green Mountain (859 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eDry tropical oceanic, 20-31\u0026deg;C, 130-680 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eDryland (0-300 m), mid-altitude area on Green Mountain (330-600 m), high-altitude with cloud forest area top of Green Mountain (\u0026gt; 660 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e51 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e26 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e4 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e81 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e(16 END \u0026ndash; 20%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSTH\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (one island), 7 mya, 122 Km\u003csup\u003e2\u003c/sup\u003e, population: \u0026nbsp; \u0026nbsp; 4,500 permanent residents, highest peak is Diana\u0026rsquo;s Peak (818 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eTropical, 17-28\u0026deg;C, 750-1,000 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eTree fern forest, pastures and coastal zones, dry and eroded, and middle elevations now destroyed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e62 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e35 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e2 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e99 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e(26 END \u0026ndash; 28 %)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"8.24524312896406%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTRC\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.799154334038056%\" valign=\"top\"\u003e\n \u003cp\u003eVolcanic (6 small islands -Nightingale \u0026ndash; \u0026gt;18 mya, Inaccessible \u0026ndash; 3-4 mya, Tristan da Cunha \u0026ndash; 200,000 mya; Gough \u0026ndash; 3-5 mya), 207 Km\u003csup\u003e2\u003c/sup\u003e, population: 264 permanent residents (only in Tristan da Cunha), highest peak is Queen Mary\u0026apos;s Peak (2,062 m)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.16490486257928%\" valign=\"top\"\u003e\n \u003cp\u003eCool temperate, 16-25\u0026deg;C, 1670 mm precipitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.78435517970402%\" valign=\"top\"\u003e\n \u003cp\u003eTussock grassland, pastures, fern bush, wet heath, feldmark and alpine, bogs and other wetland, and lava field\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.985200845665961%\" valign=\"top\"\u003e\n \u003cp\u003e158 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e142 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.993657505285412%\" valign=\"top\"\u003e\n \u003cp\u003e4 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"8.033826638477802%\" valign=\"top\"\u003e\n \u003cp\u003e305 spp.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e(86 END \u0026ndash; 22%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eThe \u003cstrong\u003eAfrican archipelagos\u003c/strong\u003e are likewise of volcanic origin and include two islands (S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe) and some islets. S\u0026atilde;o Tom\u0026eacute; is the largest island in the archipelago and consists of two islands formed during the tertiary era. The archipelago is located off the coast of central Africa (ca. 250 km distant), close to the equator. The island of S\u0026atilde;o Tom\u0026eacute; is the capital of the archipelago, with an area of ca. 1001 km\u0026sup2;. The climate there is equatorial, hot, and humid, with temperatures varying according to altitude and exposure (22-30\u0026deg;C). The island is steep, with a plain zone, high mountains to the south and west, a flat landscape to the north, and relatively fertile soil. It holds a total of 310 species of bryophytes. Cape Verde is an island country that is part of an archipelago formed by 10 islands covering ca. 4033 km\u0026sup2;, located off the coast of West Africa. A total of 224 species of bryophytes have been recorded there, with the islands of Santo Ant\u0026atilde;o, Santiago, and Fogo having the highest numbers of plant species. Santo Ant\u0026atilde;o has the highest species richness of bryophytes, with 111 species (73% of the archipelago\u0026apos;s total bryoflora), (Table 1).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe \u003cstrong\u003eSpanish archipelago\u003c/strong\u003e of the Canary Islands is of volcanic origin that emerged 8-12 million years ago and covers an area of 7747 km\u003csup\u003e2\u003c/sup\u003e. \u0026nbsp;It composes part of Macaronesia along with Madeira, Azores, and Cape Verde. A total of 501 species of bryophytes are recognized there (Table 1).\u003c/p\u003e\n\u003cp\u003eThe \u003cstrong\u003eBrazilian archipelagos\u003c/strong\u003e are of volcanic origin and represent important national and international geological sites. The \u003cstrong\u003eTrindade archipelago\u003c/strong\u003e is located ca. 1100 km from the Brazilian coast. Its mountainous islands belong to a chain of submarine volcanoes, with a source of fresh water. Most of the vegetation on the archipelago is herbaceous, with only 5% of the vegetation being forest; it harbors 35 species of bryophytes. The \u003cstrong\u003earchipelago of Fernando de Noronha\u003c/strong\u003e is formed by 21 small islands of volcanic origin, located ca. 350 km from the Brazilian coast. There are few places with original vegetation, and no source of fresh water; most of the vegetation is dry forest similar to Caatinga, with small trees, shrubs, and grasses. There are 32 species of bryophytes growing there, but neither Brazilian archipelago has any endemic species (Table 1).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe \u003cstrong\u003etwo islands\u0026nbsp;\u003c/strong\u003eand the\u003cstrong\u003e\u0026nbsp;English archipelago\u003c/strong\u003e are also of volcanic origin. \u003cstrong\u003eAscension\u0026nbsp;\u003c/strong\u003eis a small island covering only 91 km\u0026sup2;, 1660 km from Africa and 2250 km from South America; it is approximately 1 million years old, and has a dry tropical oceanic climate with little seasonal variation; its highest peak is Green Mountain (859 m). The island has three vegetation zones related to altitude: a bare and rocky dry zone at 0-330 m; the base surroundings of the mountain at altitudes between 330-660 m, with few bryophytes species; and the top of Green Mountain, altitude \u0026gt; 660 m, with a cloud forest and vegetation dominated by ferns and bryophytes (81 species of bryophytes, 4 hornworts, and 22 liverworts and 55 mosses, with 12 endemic species). The \u003cstrong\u003eIsland of Santa Helena\u003c/strong\u003e is a volcanic island covering 122 km\u003csup\u003e2\u003c/sup\u003e, 4,000 km east of Rio de Janeiro (Brazil) and 1,950 km west of Africa; it is approximately 7 million years old, has a rugged terrain, and is one of the most remote regions in the world. The climate there is tropical, marine, and mild, with continuous winds, with an average annual temperature of 17-28\u0026deg; C. The highest point is Pico Diana (818 m), whose high altitude areas concentrate endemic species. It has three vegetation zones (tree fern forests; dry and eroded pastures and coastal zones; medium elevations with highly altered vegetation), and a total of 99 bryophyte species. \u003cstrong\u003eTristan da Cunha\u003c/strong\u003e is an archipelago of volcanic origin formed by three islands: Gough, Inaccessible, and Nightingale, with a total of 305 species of bryophytes (61 being endemic), (Table 1).\u0026nbsp;\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003e\u003cem\u003eData collection\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eA dataset was compiled (Excel spreadsheet) containing the information on species growing on the archipelagos of Azores, Madeira, S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe, Cape Verde, and the Canary Islands. The plant names were updated, with information concerning their distributions and endemism, and conservation data, with a total of 1520 recorded species. The data set also included information concerning the bryophyte species found on the islands and archipelagos of Trindade, Fernando de Noronha, Ascension, St Helena, and Tristan da Cunha, with a final total of 1647 species (675 hornworts plus liverworts, and 972 mosses).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHerbarium collections\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eSubsequently, ca. 600 specimens from the archipelago of S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe, ca. 80 from the archipelago of Cape Verde, and ca. 330 samples from the Azores archipelago were identified at the LISU herbarium. The nomenclature and synonyms for each species were checked using databases such as TROPICOS and some new literature.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eData analysis\u0026nbsp;\u003c/em\u003e(\u003cem\u003ediversity and floristic affinities)\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eTo determine bryophyte diversity on the ten islands and archipelagos, a floristic matrix was prepared using incidence data (presence/absence) to quantify diversity (Table 1 Supplementary Material). Rare species (occurring on only a single island or archipelago) were included. We used this matrix to calculate the Jaccard index, verify floristic similarities, and build a dendrogram of plant diversity among the ten islands and archipelagos. These statistical analyses were performed using PAST program version 4.12 (Hammer et al. 2001).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe phytogeographic patterns of the taxa were characterized based on their global distributions, according to information available in the scientific literature and databases (Wigginton 2004, 2018; O\u0026rsquo;Shea 2006; Pati\u0026ntilde;o-Llorente et al. 2005; TROPICOS database; and the general literature).\u003c/p\u003e\n\u003cp\u003eThese datasets included the more important literature and were used to analyze differences in the bryofloras of the ten islands and archipelagos in the Atlantic Ocean in relation to their total numbers of taxa, endemic species, exotic species, and possibly extinct species (Table 2), as well as the main families and any exclusive ones (Table 3).\u003c/p\u003e"},{"header":"Results And Discussion","content":"\u003cp\u003e\u003cem\u003eDiversity of the ten islands and archipelagos of the Atlantic Ocean (alpha, beta, and gama diversity)\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eA total of 1647 species (\u003cstrong\u003egama diversity\u003c/strong\u003e) were listed for the ten islands and archipelagos in the \u003cstrong\u003eNorthern\u0026nbsp;\u003c/strong\u003eand\u003cstrong\u003e\u0026nbsp;Southern Atlantic Ocean\u003c/strong\u003e, including 1152 species of mosses and 495 species of liverworts and hornworts. Of these, 197 species are endemic (63 species of liverworts and hornworts and 134 species of mosses), representing 12% of all species surveyed (Table 2); only 10 species were shared by all five archipelagos. The number of endemic species in the five \u003cstrong\u003eNorth Atlantic Ocean\u003c/strong\u003e archipelagos (197 spp.) was higher than the five islands and archipelagos of the \u003cstrong\u003eSouth Atlantic Ocean\u003c/strong\u003e, demonstrating their importance to the conservation of bryophyte biodiversity on Atlantic Ocean islands. This is especially critical because they will soon suffer the consequences of climate change \u0026ndash; added to continuous pressure from tourism, pollution, population growth, the introduction of alien species, and habitat destruction.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2\u003c/strong\u003e Species diversity data for each studied island and archipelago. Spp = Total number of species, Ext = Extinct species, End = Endemic species, Exo = Exotic species\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"623\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eIslands and archipelagos\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSpp.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eExt\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eEnd\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eExo\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eMain references\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eAzores\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e436\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e40\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003eCorley et al. (1981), Grolle (1983), and results of this study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eMadeira\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e577\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e68\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003ePersson (1939), Sj\u0026ouml;gren (1975, 2001), S\u0026eacute;rgio et al. (2006), plus results of this study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eS\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e310\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003eS\u0026eacute;rgio and Garcia (2011), Garcia et al.\u0026nbsp;(2012), and results of this study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eCape Verde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e224\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003eLeyens and Lobin (1996); Arechavaleta et al. (2005); Medina \u0026amp; Gomes (2015), and results of this study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eCanary Islands\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e501\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e54\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003ePersson (1939), Eggers (1982), Dirkse et al. (1993), Gonz\u0026aacute;les et al. (2008)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eTrindade\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e35\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003eSerafin et al. (2010), Faria et al.\u0026nbsp;(2012), Costa and Rezende (2022)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eFernando de Noronha\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003eVital et al. (1991), Serafin et al.\u0026nbsp;(2010), Pereira and C\u0026acirc;mara (2015), Costa et al. (2021)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eSt Helena Island\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e99\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003eWigginton (2012)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eAscension Island\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e81\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003ePressel et al. (2014)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.2%\" valign=\"top\"\u003e\n \u003cp\u003eTristan da Cunha\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e305\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e86\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.12%\" valign=\"top\"\u003e\n \u003cp\u003e?\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.32%\" valign=\"top\"\u003e\n \u003cp\u003eDixon (1960), Wace (1961), Wace and Dickson (1965), V\u0026aacute;\u0026ntilde;a and Engell (2013)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFinally, with the inclusion of species from the South Atlantic Ocean islands and archipelagos (Fernando de Noronha, Trindade, St Helena, Ascension, and Tristan da Cunha) to analyze and compare them with those of the North Atlantic Ocean (\u003cstrong\u003egama diversity\u003c/strong\u003e), the total number of species increased to 1647 species (11 hornworts, 664 liverworts, and 972 mosses).\u003c/p\u003e\n\u003cp\u003eSpecies richness varied strongly among the ten southern Atlantic islands and archipelagos, ranging from 32 species on Fernando de Noronha, to 577 on Madeira (Table 2). The bryophyte diversity on the ten islands and archipelagos studied here (\u003cstrong\u003egama diversity)\u003c/strong\u003e was high when compared to Africa, Tropical America, and the world (Gradstein et al. 2001; Shaw and Goffinet 2000; O\u0026rsquo;Shea 2006; Wigginton 2018), (Fig. 2).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003ePhytogeographical patterns and endemism\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe analyzed the islands and archipelagos of the North and South Atlantic Oceans separately, and then together; the results are presented in Tables 3, 4.\u003c/p\u003e\n\u003cp\u003eThe phytogeographical patterns of the \u003cstrong\u003eNorth Atlantic Ocean\u003c/strong\u003e islands and archipelagos were combined into \u003cstrong\u003eten categories\u003c/strong\u003e for this analysis: 1) Global (130 taxa; 23 liverworts and 107 mosses); 2) Africa (194 taxa; 117 liverworts and 23 mosses); 3) Endemic (91 taxa; 22 liverworts and 69 mosses); 4) North America, Europe and Asia (49 taxa of mosses); 5) North America and Europe (19 taxa of mosses); 6) Europe, Macaronesia, Africa (14 taxa of liverworts); 7) Europe, Macaronesia, Asia (9 taxa of liverworts); 8) Europe and Macaronesia (55 taxa; 13 liverworts and 42 mosses); 9) South America and Africa (11 taxa of liverworts); 10) others. The \u003cstrong\u003eAfrican, Endemic,\u0026nbsp;\u003c/strong\u003eand\u003cstrong\u003e\u0026nbsp;Global\u0026nbsp;\u003c/strong\u003eelements were the largest, comprising 11% of the total taxa (Table 3).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 3\u003c/strong\u003e Phytogeographical patterns of the bryofloras of the five North Atlantic Ocean islands and archipelagos\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003ePatterns\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003eLiverworts\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003eMosses\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eWorldwide\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e107\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e130\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eAfrica\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e117\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e77\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e194\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eN America, Europe and\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e49\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e49\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eN America and Europe\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEurope, Macaronesia, Africa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEurope, Macaronesia, Asia\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e9\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEurope and Macaronesia\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e55\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eSouth America and Africa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEndemic\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e69\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e91\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eOthers\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e217\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e352\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e569\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e426\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e715\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e1141**\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e** This number is different because there are taxa identified only at genera level\u003c/p\u003e\n\u003cp\u003eWe found that the liverworts and mosses exhibited distinct phytogeographical patterns (Table 3). Liverworts showed low numbers of European and Macaronesia species (13 spp.), but the highest number of African elements (117), followed by global elements (23) and endemic species (22); mosses showed high numbers of global elements (107), followed by African and Endemic elements (77 and 69).\u003c/p\u003e\n\u003cp\u003eThe phytogeographical patterns of the \u003cstrong\u003eSouth Atlantic Ocean\u003c/strong\u003e islands and archipelagos were combined into eight categories for this analysis: 1) Global (73 taxa; 9 liverworts and 64 mosses); 2) Pantropical (24 taxa; 8 liverworts and 16 mosses); 3) Afro-America (30 taxa; 19 liverworts and 11 mosses); 4) Tropical America (27 taxa; 15 liverworts and 12 mosses); 5) Africa (26 taxa; 9 liverworts and 17 mosses); 6) South America (77 taxa; 66 liverworts and 10 mosses); 7) Endemic (100 taxa;43 liverworts and 57 mosses); 8) others (122 taxa; 72 liverworts and 50 mosses). Endemic elements was the largest category, comprising ca. 21% of all taxa (Table 4).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 4\u003c/strong\u003e Phytogeographical patterns of the bryofloras of the five South Atlantic Ocean islands and archipelagos (Trindade, Fernando de Noronha, Ascension, St Helena, and Tristan da Cunha)\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003ePatterns\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003eLiverworts\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003eMosses\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eWorldwide\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e64\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e73\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003ePantropical\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e24\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eAfro-America\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e30\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eTropical America\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e27\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eAfrica\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e26\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eSouth America\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e77\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eEndemic\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e43\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e57\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eOthers\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e122\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"31.47482014388489%\" valign=\"top\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e241\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e237\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.841726618705035%\" valign=\"top\"\u003e\n \u003cp\u003e478**\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e** This number is different because there are taxa identified only at genera level\u003c/p\u003e\n\u003cp\u003eThe liverworts and mosses exhibited distinct phytogeographical patterns on the South Atlantic Ocean islands and archipelagos. Liverworts had high numbers of species with South American distributions (66 spp.), followed by endemic species (43), but low numbers of species with global distributions (9); mosses had high numbers of endemic species (57) and species having global distributions (64).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe phytogeographical patterns of the islands and archipelagos of the \u003cstrong\u003eNorth Atlantic Ocean\u003c/strong\u003e were combined into ten categories for this analysis: 1) Global (130 taxa; 23 liverworts and 107 mosses); 2) Africa (194 taxa; 117 liverworts and 23 mosses); 3) Endemic (91 taxa; 22 liverworts and 69 mosses); 4) North America, Europe and Asia (49taxa of mosses); 5) North America and Europe (19 taxa of mosses); 6) Europe, Macaronesia, Africa (14 taxa of liverworts); 7) Europe, Macaronesia, Asia (9 taxa of liverworts); 8) Europe and Macaronesia (55 taxa; 13 liverworts and 42 mosses); 9) South America and Africa (11 taxa of liverworts); 10) others (569 taxa; 217 liverworts and 352 mosses). African, Endemic, and Global elements were the largest categories, comprising 11% of the total taxa (Table 5).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 5\u003c/strong\u003e Phytogeographical patterns of the bryofloras on the five North Atlantic Ocean islands and archipelagos (Azores, Madeira, S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe, Cape Verde, and Canary Islands)\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003ePatterns\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003eLiverworts\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003eMosses\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eWorldwide\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e107\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e130\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eAfrica\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e117\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e77\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e194\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eN America, Europe and\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e49\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e49\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eN America and Europe\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEurope, Macaronesia, Africa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEurope, Macaronesia, Asia\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEurope and Macaronesia\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e55\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eSouth America and Africa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e--\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eEndemic\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e69\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e91\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eOthers\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e217\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e352\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e569\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.983842010771994%\" valign=\"top\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.414721723518852%\" valign=\"top\"\u003e\n \u003cp\u003e426\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e715\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.800718132854577%\" valign=\"top\"\u003e\n \u003cp\u003e1141**\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e** This number is different because there are taxa identified only at genera level\u003c/p\u003e\n\u003cp\u003eThe liverworts and mosses were found to exhibit distinct phytogeographical patterns on the South Atlantic Ocean islands and archipelagos (Table 5). Liverworts there had high numbers of African elements (117 spp.), followed by global elements (23) and endemic species (22), but low numbers of Europe and Macaronesia species (13); mosses there showed high numbers of global species (107), followed by African elements (77), and Endemic species (69).\u003c/p\u003e\n\u003cp\u003eA striking feature of the bryofloras of the archipelagos in the \u003cstrong\u003eNorth Atlantic Ocean\u003c/strong\u003e is that species richness was concentrated in just a few families (Table 5, Fig. 3), with mosses, Brachytheciaceae (44 species), Bryaceae (47), Fissidentaceae (58), and Pottiaceae (135) standing out and totaling together 284 species (fully 31% of the total of 916 moss species recorded for the five archipelagos). In terms of the liverworts, just three families (Lejeuneaceae [121 species], Plagiochilaceae [33], and Ricciaceae [34]) concentrated 32% of the total number of liverwort species (593) recorded for the five archipelagos. The species richness of bryophytes was therefore concentrated in seven families that together represented 40% of the total number of species (1161).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe species richness of the bryofloras of the \u003cstrong\u003eSouth Atlantic Ocean\u003c/strong\u003e islands and archipelagos was concentrated in six families (Table 6, Fig. 3), Lejeuneaceae (38 species), Lophocoleaceae (40), Bryaceae (25), Fissidentaceae (28), Grimmiaceae (11), and Pottiaceae (25), which together totaled 167 species, representing 27% of the total.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 6\u003c/strong\u003e Total numbers of bryophyte families and genera, as well as the main families in each archipelago or island. \u003cstrong\u003eBold\u003c/strong\u003e = families standing out in terms of their numbers of species\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"633\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eIslands or Archipelagos\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eFamilies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eGenera\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eMain families\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eAzores\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e90\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e128\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: Aneuraceae, Calypogeiaceae, Cephaloziaceae, \u003cstrong\u003eCephaloziellaceae\u003c/strong\u003e, Fossombroniaceae, Geocalycaceae, \u003cstrong\u003eLejeuneaceae\u003c/strong\u003e, Lepidoziaceae, \u003cstrong\u003eLophocoleaceae\u003c/strong\u003e, Plagiochilaceae, Radulaceae, \u003cstrong\u003eRicciaceae\u003c/strong\u003e, Scapaniaceae.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: Bartramiaceae, \u003cstrong\u003eBrachytheciaceae\u003c/strong\u003e, Bryaceae, Ditrichaceae, \u003cstrong\u003eFissidentaceae, Grimmiaceae\u003c/strong\u003e, Hypnaceae, \u003cstrong\u003eLeucobryaceae\u003c/strong\u003e, Neckeraceae, Polytrichaceae, \u003cstrong\u003ePottiaceae\u003c/strong\u003e, \u003cstrong\u003eSphagnaceae\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eMadeira\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e92\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e197\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eHORNWORTS\u003c/u\u003e: Anthocerotaceae\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: \u003cstrong\u003eCephaloziaceae\u003c/strong\u003e, Cephaloziellaceae, Fossombroniaceae, Frullaniaceae, \u003cstrong\u003eLejeuneaceae\u003c/strong\u003e, Marchantiaceae, \u003cstrong\u003ePlagiochilaceae\u003c/strong\u003e, \u003cstrong\u003eRicciaceae\u003c/strong\u003e, Scapaniaceae\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: Amblystegiaceae, Bartramiaceae, \u003cstrong\u003eBrachytheciaceae\u003c/strong\u003e, \u003cstrong\u003eBryaceae\u003c/strong\u003e, Ditrichaceae, \u003cstrong\u003eFissidentaceae\u003c/strong\u003e, Funariaceae, \u003cstrong\u003eGrimmiaceae\u003c/strong\u003e, Leucobryaceae, Mniaceae, Neckeraceae, \u003cstrong\u003eOrthotrichaceae\u003c/strong\u003e, Polytrichaceae, \u003cstrong\u003ePottiaceae\u003c/strong\u003e, Sphagnaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eS\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e62\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e104\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eHORNWORTS\u003c/u\u003e: Dendrocerotaceae\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: \u003cstrong\u003eFrullaniaceae, Lejeunaceae\u003c/strong\u003e, Metzgeriaceae, \u003cstrong\u003ePlagiochilaceae\u003c/strong\u003e, Radulaceae, Ricciaceae\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: \u003cstrong\u003eBryaceae, Calymperaceae, Fissidentaceae\u003c/strong\u003e, Hypnaceae, Orthostichellaceae, Pilotrichaceae, Pottiaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eCape Verde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e49\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e105\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: Frullaniaceae, \u003cstrong\u003eLejeuneaceae, Ricciaceae\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: Bartramiaceae, \u003cstrong\u003eBrachytheciaceae, Bryaceae, Fissidentaceae\u003c/strong\u003e, Orthotrichaceae, \u003cstrong\u003ePottiaceae\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eCanary\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e90\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e187\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eHORNWORTS\u003c/u\u003e: Anthocerotaceae\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: Cephaloziellaceae, \u003cstrong\u003eFrullaniaceae, Lejeuneaceae,\u003c/strong\u003e Lophocoleaceae, Plagiochilaceae, Radulaceae, \u003cstrong\u003eRicciaceae\u003c/strong\u003e, Riellaceae, Sphaerocarpaceae\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: Bartramiaceae, \u003cstrong\u003eBrachytheciaceae, Bryaceae\u003c/strong\u003e, Ditrichaceae, \u003cstrong\u003eFissidentaceae\u003c/strong\u003e, Funariaceae, \u003cstrong\u003eGrimmiaceae\u003c/strong\u003e, Leucobryaceae, Neckeraceae, Orthotrichaceae, \u003cstrong\u003ePolytrichaceae\u003c/strong\u003e, \u003cstrong\u003ePottiaceae\u003c/strong\u003e,\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eTrindade\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e25\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: \u003cstrong\u003eLejeuneaceae\u0026nbsp;\u003c/strong\u003e(12 spp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eFernando de Noronha\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eHORNWORTS\u003c/u\u003e: Nothothyladaceae (2 spp.)\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: Bryaceae (4 spp.), \u003cstrong\u003eFissidentaceae\u003c/strong\u003e (12 spp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eAscension\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eHORNWORTS\u003c/u\u003e: Nothothyladaceae (3 spp.)\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: Lejeuneaceae (5 spp.)\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: \u003cstrong\u003eBryaceae\u0026nbsp;\u003c/strong\u003e(8 spp.), Fissidentaceae (4 spp.), \u003cstrong\u003eLeucobryaceae\u003c/strong\u003e (7 spp.), \u003cstrong\u003ePottiaceae\u0026nbsp;\u003c/strong\u003e(12 spp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eSt Helena\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e40\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: \u003cstrong\u003eLejeuneaceae\u003c/strong\u003e (15 spp.)\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: Bryaceae (7 spp.), \u003cstrong\u003eFissidentaceae\u003c/strong\u003e (11 spp.), Leucobryaceae (4 spp.), \u003cstrong\u003ePottiaceae\u003c/strong\u003e (9 spp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"25.31645569620253%\" valign=\"top\"\u003e\n \u003cp\u003eTristan da Cunha\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.443037974683545%\" valign=\"top\"\u003e\n \u003cp\u003e68\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.126582278481013%\" valign=\"top\"\u003e\n \u003cp\u003e142\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"54.11392405063291%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cu\u003eLIVERWORTS\u003c/u\u003e: \u003cstrong\u003eAdelanthaceae\u003c/strong\u003e (10 spp.), Anastrophyllaceae (9 spp.), Aneuraceae (13 spp.), Cephaloziellaceae (10 spp.), Lejeuneaceae (14 spp.), Lepidoziaceae (14 spp.), \u003cstrong\u003eLophocoleaceae\u003c/strong\u003e (36 spp.), Plagiochilaceae (9 spp.)\u003c/p\u003e\n \u003cp\u003e\u003cu\u003eMOSSES\u003c/u\u003e: \u003cstrong\u003eAndreaeaceae\u003c/strong\u003e (7 spp.), Bartramiaceae (10 spp.),Bryaceae (10 spp.), Ditrichaceae (8 spp.), \u003cstrong\u003eGrimmiaceae\u003c/strong\u003e (11 spp.), \u003cstrong\u003ePolytrichaceae\u003c/strong\u003e (6 spp.), Pottiaceae (6 spp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003eThere were differences among the islands and archipelagos with unique families, or among those which had significantly different numbers of species (\u003cstrong\u003ebeta diversity\u003c/strong\u003e), Table 7. The Brazilian archipelagos In the \u003cstrong\u003eSouth Atlantic Ocean\u003c/strong\u003e (Trindade and Fernando de Noronha) are examples of how habit destruction by humans can influence bryophyte species diversity, as those archipelagoes have only 35 and 32 species respectively. Those numbers represent the lowest diversity of bryophyte species among the ten islands and archipelagos studied here, and neither has an endemic species. That situation is in contrast to the two English islands and one archipelago (Ascension, St. Helena, and Tristan da Cunha) which have between 81 and 312 species of bryophytes, including between 16 to 86 endemic species \u0026ndash; demonstrating their degrees of preservation. The African archipelagos of Cape Verde and S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe in the \u003cstrong\u003eNorth Atlantic Ocean\u003c/strong\u003e have between 215 and 314 bryophyte species, including 5 to 16 endemic species. The Portuguese archipelagos of Azores and Madeira, and the Spanish Canary Islands archipelago evidenced the highest diversity and shared more species, with the Canary Islands being the most preserved (with a total of 501 bryophyte species, of which 54 are endemic).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAZORES\u0026nbsp;\u003c/strong\u003e\u0026ndash; Three bryophyte families occur only on this archipelago; Jamesoniellaceae is represented there by a species that occurs nowhere else outside of the Neotropics (\u003cem\u003eSyzygiella rubricaulis\u003c/em\u003e (Nees) Steph.). The Sphagnaceae family also stands out in this archipelago, with 17 species (Hodgetts and Lockhart 2020). This archipelago shares its highest number of species with the \u003cstrong\u003eMadeira\u003c/strong\u003e archipelago (379 species, including 37 endemic species).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMADEIRA\u0026nbsp;\u003c/strong\u003e\u0026ndash; Three families occur only on this archipelago: Blasiaceae (Liverworts), Aongstroemiaceae, and Flexitrichaceae (Mosses). The families Brachytheciaceae (34 species) and Grimmiaceae (27) also stand out (Hodgetts and Lockhart 2020). This archipelago shares the highest number of species with the \u003cstrong\u003eAzores\u003c/strong\u003e, as noted above.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eS\u0026Atilde;O TOM\u0026Eacute; and PR\u0026Iacute;NCIPE\u0026nbsp;\u003c/strong\u003e\u0026ndash; Has the highest number of restricted families. This is the only archipelago with Dendrocerotaceae, a hornwort family, as well as moisture-indicating families such as Hypopterygiaceae and Meteoriaceae. It shares 23 species with \u003cstrong\u003eCape Verde\u003c/strong\u003e, 20 with the \u003cstrong\u003eCanary\u003c/strong\u003e Islands, 21 with the \u003cstrong\u003eAzores\u003c/strong\u003e, and 22 with the \u003cstrong\u003eMadeira\u003c/strong\u003e archipelago.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCAPE VERDE\u0026nbsp;\u003c/strong\u003e\u0026ndash; Two bryophyte families occur only in this archipelago; 69 exclusive species are found there, with four being endemic. This is the most unique archipelago among the five studied in the North Atlantic Ocean, having the smallest total number of species and endemic species. It also shares a high number of taxa with the \u003cstrong\u003eCanary\u003c/strong\u003e Islands (118 species).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCANARY ISLANDS\u0026nbsp;\u003c/strong\u003e\u0026ndash; Three bryophyte families occur only on this archipelago. It shares the most species with the \u003cstrong\u003eAzores\u0026nbsp;\u003c/strong\u003eand\u003cstrong\u003e\u0026nbsp;Madeira\u0026nbsp;\u003c/strong\u003e(283 species, with 27 being endemic) and with \u003cstrong\u003eCape Verde\u003c/strong\u003e (118 taxa). This archipelago, together with \u003cstrong\u003eMadeira\u003c/strong\u003e, has the best-known bryofloras, which have been presented in many different publications.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTRINDADE\u003c/strong\u003e \u0026ndash; No bryophyte family occurs exclusively on this island. Its native vegetation has been almost completely destroyed through the grazing of introduced animals. E. Knight visited the island in 1881 and 1889 and reported its already degraded vegetation and a large waterfall on the west coast. Vegetation restoration efforts were initiated in 1990 through the elimination of goats. The vegetation has slowly recovered since then, and its creeks have more water. The vegetation above 400 m is considered a \u0026ldquo;Giant Fern Forest\u0026quot; and is dominated by a single species, \u003cem\u003eCyathea delgadii\u0026nbsp;\u003c/em\u003eSternb.; it is the most diverse region for bryophytes. Most of the species are Neotropical, followed by cosmopolitan species; there are no endemic species (Costa and Rezende 2022). The growth of \u003cem\u003eCampylopus introflexus\u003c/em\u003e was probably facilitated by human impacts on the original vegetation. This archipelago shares six species with \u003cstrong\u003eAscension\u003c/strong\u003e Island and five species with the \u003cstrong\u003eFernando de Noronha\u003c/strong\u003e archipelago.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFERNANDO DE NORONHA\u003c/strong\u003e \u0026ndash; Only one moss family, Stereophyllaceae, occurs on this island. Charles Darwin visited the island and made the first botanical collections there. A later expedition by Ridley (1890) reported the predominance of herbaceous vegetation due to the intensive cutting of tall trees that could be used to build rafts and aid prisoner escapes. \u003cem\u003eFissidens\u003c/em\u003e,\u003cem\u003e\u0026nbsp;\u003c/em\u003ewith 12 species, is the richest genus. Cosmopolitan species compose approximately 43% of vegetation; there are no\u0026nbsp;endemic species (Costa and Rezende 2022). This archipelago\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eshares\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003efive species with \u003cstrong\u003eAscension\u003c/strong\u003e Island.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eASCENSION\u003c/strong\u003e \u0026ndash; No bryophyte family occurs conclusively on this island. The island harbors over 200 introduced species ranging from large trees to shrubs and herbs; many native plants, such as the endemic fern \u003cem\u003ePitsana purpurascens\u0026nbsp;\u003c/em\u003e(de Vriese) Murdock, are becoming severely outcompeted. \u003cem\u003eCampylopus,\u003c/em\u003e with seven species, is the richest genus. There is a notable absence of cosmopolitan species (\u003cem\u003eCeratodon purpureus\u0026nbsp;\u003c/em\u003e(Hedw.) Brid.\u003cem\u003e, Tortula muralis\u0026nbsp;\u003c/em\u003eHedw.\u003cem\u003e, Octoblepharum albidum\u0026nbsp;\u003c/em\u003eHedw., etc.). There are no representatives of Grimmiales, despite the highly suitable volcanic rock substrate there. The island has\u0026nbsp;fifteen endemic bryophyte species. A\u0026nbsp;handful of taxa are probable human introductions, but despite the small size of the island and the destruction of its natural habitats, relatively few bryophytes are threatened with extinction.\u0026nbsp;This island\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eshares\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e21 species with \u003cstrong\u003eSt Helena\u003c/strong\u003e Island, three of which are endemic.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eST HELENA\u003c/strong\u003e \u0026ndash; No bryophyte family occurs only on this island, but there is a notable absence of cosmopolitan species, \u003cem\u003eCeratodon purpureus, Tortula muralis, Octoblepharum albidum\u003c/em\u003e, etc. There are no representatives of Grimmiales despite the highly suitable volcanic rock substrate there. \u003cem\u003eFissidens,\u003c/em\u003e with 11 species, is the richest genus (Wigginton 2013). The destruction of the natural vegetation began with the introduction of goats, and the establishment of permanent settlements there introduced exotic vegetation. Native trees were cut for different purposes (house building, fuel, etc.), and many endemic species are now extinct or critically endangered. This archipelago\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eshares\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e16 species with the \u003cstrong\u003eTristan da Cunha\u003c/strong\u003e archipelago, three of which are endemic.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTRISTAN DA CUNHA\u003c/strong\u003e \u0026ndash; Eleven bryophyte families occur only on this archipelago. There are currently very few introduced species on the archipelago. The family Orthotrichaceae is represented by several endemic species, and \u003cem\u003eDicranoloma\u003c/em\u003e (Renauld) Renauld by three endemic species. This archipelago\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eshares\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e28 species with the \u003cstrong\u003eAzores\u003c/strong\u003e archipelago and 23 with \u003cstrong\u003eMadeira\u003c/strong\u003e, none are endemic. There are strong indications of a close connection with South America, as mosses, liverworts, and hornworts can be dispersed by the western winds \u0026ndash; \u0026lsquo;West Wind Drift\u0026rsquo; \u0026ndash; and connections with South America are conspicuous in all groups of non-vascular plants, except the algae (Galloway 1996, Milius 2004, Mu\u0026ntilde;oz et al. 2004).\u003c/p\u003e\n\u003cp\u003eSunding (1979) considered Cape Verde to be different from the other archipelagos of Macaronesia, which was subsequently corroborated by Vanderpoorten et al. (2007). According to these authors, Macaronesia (comprising the archipelagos of Azores, Madeira, Canary Islands, and Cape Verde) is considered a single biogeographical unit, with their floras being relics of a widely distributed tertiary subtropical flora. These authors rejected the concept of Macaronesia \u003cem\u003esensu lato\u003c/em\u003e, and viewed the Cape Verde archipelago as being more associated with tropical Africa. Liverworts in the other Macaronesian archipelagos support an Azores-Madeira-Canary clade (Macaronesia \u003cem\u003esensu stricto\u003c/em\u003e), while mosses support the Canary Islands as being related to North Africa, rejecting the concept of Macaronesia \u003cem\u003es.s.\u003c/em\u003e for the group. The exchange of taxa with neighboring continental areas better explains the relationships between the cryptogamic flora of Cape Verde and the mossy flora of the Canary Islands. In contrast, the relic flora is consistent with a monophyletic Macaronesia \u003cem\u003es.s.\u003c/em\u003e group of liverworts. These congruent patterns, however, can hide a complex mix of relict distributions and more recent speciation and dispersal events.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eSimilarities among the ten islands and archipelagos (beta diversity)\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe results of our study, when comparing the bryofloras of the ten \u003cstrong\u003eAtlantic Ocean\u003c/strong\u003e islands and archipelagos\u003cstrong\u003e,\u0026nbsp;\u003c/strong\u003edemonstrated that their bryofloras were very dissimilar, with low floristic affinities (Table 7). Of the ten islands and archipelagos, those demonstrating the greatest similarity were the archipelagos of Macaronesia, Azores, Madeira, and the Canary Islands, which share 283 species (Fig. 4 A,B,C). Our results also indicated Macaronesia as being formed by the Azores, Madeira, and the Canary Islands archipelagos, with Cape Verde outside the clade when the islands and archipelagos of the \u003cstrong\u003eNorth Atlantic Ocean\u003c/strong\u003e were analyzed separately or together with the islands and archipelagos of the South Atlantic Ocean (Fig. 4 A,C). As expected, the Azores, Madeira, and Canary Islands archipelagos (Macaronesia \u003cem\u003es.s.\u003c/em\u003e) shared several species and evidenced the highest similarity. The archipelago of S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe was found to be highly dissimilar to the others, sharing few species with any of the \u003cstrong\u003eNorth Atlantic Ocean\u003c/strong\u003e archipelagos (Fig. 4 A,C).\u003c/p\u003e\n\u003cp\u003eThe results for the \u003cstrong\u003eSouth Atlantic Ocean\u003c/strong\u003e also showed low similarity among almost all of the islands and archipelagos, although the islands of St Helena and Ascension show the highest similarity and form a separate clade (with Tristan da Cunha outside of it). The islands of St Helena and Ascension share several species. The archipelagos of Trindade and Fernando de Noronha also formed a separate clade, although they share few species; this can be attributed to the degree of devastation suffered by these two archipelagos since their discovery and occupation (Fig. 4 B,C).\u003c/p\u003e\n\u003cp\u003eWhen we analyzed the ten \u003cstrong\u003eAtlantic Ocean\u0026nbsp;\u003c/strong\u003eislands and archipelagos together, their similarities were very low, with the exception of Madeira, Azores, and the Canary Islands (Marcaronesia \u003cem\u003es.s\u003c/em\u003e.), each demonstrating an essentially unique bryoflora. Our results also corroborate the separation of Cape Verde from Macaronesia (Fig. 4 C).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 7\u0026nbsp;\u003c/strong\u003eSimilarities among the ten islands and archipelagos studied.\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"716\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.43175487465181%\" valign=\"bottom\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"7.66016713091922%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eTRI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eFN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.30640668523677%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eASC\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eSTH\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eTRC\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eAZO\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eMAD\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.445682451253482%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eSTP\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eCV\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.938718662952646%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eCAN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.43175487465181%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eTRI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.66016713091922%\" valign=\"bottom\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,07936508\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.30640668523677%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,03703704\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,0229008\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01183432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01616162\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01470588\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.445682451253482%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01557632\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,0170213\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.938718662952646%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01444\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.43175487465181%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eFN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.66016713091922%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e0,079365\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.30640668523677%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,04761905\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,0076336\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,00804829\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01309329\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.445682451253482%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,03184713\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,0171674\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.938718662952646%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01083\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.43175487465181%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eASC\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.66016713091922%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,037037\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,04761905\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.30640668523677%\" valign=\"bottom\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,1503268\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,03225807\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,0341556\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,03755869\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.445682451253482%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,04843305\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,0446097\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.938718662952646%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,030717\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.43175487465181%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eSTH\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.66016713091922%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,022901\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,00763359\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.30640668523677%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e0,1503268\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n 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width=\"7.938718662952646%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e0,196399\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.43175487465181%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eCAN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.66016713091922%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01444\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,01083033\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.30640668523677%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,03071672\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,0364238\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,03345725\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.584958217270195%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e0,42142857\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.331476323119777%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e0,51841746\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.445682451253482%\" valign=\"bottom\"\u003e\n \u003cp\u003e0,03152585\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.192200557103064%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e0,1963994\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"7.938718662952646%\" valign=\"bottom\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eConservation\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe ten islands and archipelagos studied here proved to be very rich areas for bryophytes, with their numbers being equivalent to 43% of all of the species known to Africa (3800 species), 41% of all bryophyte species known to tropical America (4000 species), and 8% of all species known globally. Our study indicated that the vegetation on the islands and archipelagoes must be preserved and protected, as 91 species are only known from them. Additionally, more botanical collections should be undertaken on Tristan da Cunha and S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe. Our results reinforce the importance of the bryofloras of the Atlantic Ocean islands and archipelagos to bryophyte conservation, as they hold high percentages of the total bryophyte diversity of Africa and tropical America, as well as the total species diversity known for the Macaronesia region within their small areas.\u003c/p\u003e\n\u003cp\u003eThe main threats to the biodiversity on all ten islands and archipelagos studied here include species introductions, pollution (maritime traffic, tourism, fishing), the introduction of invasive species with their potential to alter native habitats and compete with native species (as occurs in Saint Helena \u0026ndash; New Zealand flax), and rising ocean levels due to climate change.\u003c/p\u003e\n\u003cp\u003eFive of the North Atlantic Ocean islands and archipelagos studied here had 240 taxa classified as being threatened to some degree (2 EX - extinct, 15 CR - critically endangered, 72 EN - endangered, 145 VU - vulnerable, 6 - least concern LC ) \u0026ndash; fully 20% of the total number of species. This study confirms the importance of these islands and archipelagos as important sites of bryophyte diversity for Macaronesia and Africa. There is, however, a lack of adequate protection for endemic and/or endangered species on these islands and archipelagos, which should be addressed with priority, as global warming will accelerate the destruction of such sensitive environments around the planet.\u003c/p\u003e\n\u003cp\u003eThe bryophytes of the Macaronesian laurel forests (\u003cem\u003eLaurisilva\u003c/em\u003e) are also at considerable risk due to climate change (Pati\u0026ntilde;o et al. 2016, Pati\u0026ntilde;o and Vanderpoorten 2018). Many areas are predicted to become water-deficient, and forest fires are becoming more frequent \u0026ndash; with projections indicating significantly increased risks to important endemic species (Hodgetts et al. 2019).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThere are still many other threatened bryophyte species from a conservation perspective, with tropical species frequently referenced solely from type specimens or based on small private collections from Africa; approximately 45 bryophyte taxa reported for S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe, for example, have not been collected again since the 19\u003csup\u003eth\u003c/sup\u003e century. The ecology, habitat specificity, and distribution patterns of a large number of bryophyte species are not well known, and threats to the forest habitats in S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe (including habitat destruction and competition from invasive species, for example) can affect the survival of many bryophyte species (Garcia et al. 2022).\u003c/p\u003e"},{"header":"Conclusions and conservation implications ","content":"\u003cp\u003eThe present study represents the first survey of bryophyte diversity on ten Atlantic Ocean Islands and archipelagos. The results have increased our knowledge of the bryofloras of island habitats, and have introduced a considerable number of novelties and new records. We consider this data to be relevant to the conservation of bryophyte diversity on oceanic islands and archipelagos in the Atlantic Ocean, and to subsidize Global Strategy for Plant Conservation (GSPC) targets. This survey of ten Atlantic Ocean islands and archipelagos provides a better understanding of their floristic compositions and increases our estimates of species richness and distributions \u0026ndash; although this information is far from complete and additional inventories will still be needed in unexplored areas on the Tristan da Cunha and S\u0026atilde;o Tom\u0026eacute; and Pr\u0026iacute;ncipe archipelagos. Bryophyte richness and diversity differed significantly among study sites (alpha diversity), and their low similarities (beta diversity) illustrated their high degrees of heterogeneity and reflected the importance of habitat differences for high diversity. A better understanding of the factors related to species richness and endemism on oceanic islands will be important for conservation efforts and biodiversity management, especially in view of global climate change (Grabherr et al. 2010). Climate change is expected to result in strong impacts on oceanic islands and archipelagos, with habitat losses and declines, and extinction risks, as isolated bryophyte species in these regions have no access to refuges. The complexity of island ecosystems is associated with their high diversity and endemism, but researchers will need to generate primary data and integrate available geological and climatic information into diversity models. Further studies will be necessary to develop proactive conservation plans, for these environments hold a large fraction of the world\u0026rsquo;s biodiversity. The main threats to species diversity, forest structure, and habitats include urbanization, habitat destruction and fragmentation, the introduction of invasive species that can affect the survival of native species, uncontrolled tourism, and rising oceans (which will mainly affect islands smaller than 1,000 ha - M\u0026eacute;dail 2017). Our results established baselines of bryophyte diversity and endemism for ten Atlantic Ocean islands and archipelagos, which can be compared with future surveys and could help to determine whether species can migrate or will become extinct due to climate change. The unique features of those oceanic islands and archipelagos demand conservation strategies considering their biological and environmental diversities and their importance as protected areas.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u0026nbsp;\u003c/strong\u003eThe authors would like to thank CNPq for the PDE grant (no. 200858/2022-0), also to University of Lisbon, National Museum of Natural History and Science for provide all the infrastructure for the development of the work. Finally, the collaboration of the colleagues C\u0026eacute;sar A. Garcia, F\u0026aacute;tima Sales, Ireneia Melo, Jorge Paiva, Jos\u0026eacute; Cardoso, and Maria Cristina Duarte, and that collected and sending samples from different islands.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003eDenise Pinheiro Costa developed the idea of this publication, both authors contributes to the writing and editing draft and gave approval for its publication.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e The authors declare no competing interest.s\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical approval\u0026nbsp;\u003c/strong\u003eNot applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eCorley MFV, Crundwell, AC, D\u0026uuml;ll, R, Hill, MO, Smith, AJ E (1981) Mosses of Europe and the Azores; an annotated list of species, with synonyms from the recent literature. 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Ser B, Biol Sci 249: 273\u0026ndash;360\u003c/li\u003e\n\u003cli\u003eWigginton MJ (2012) Mosses and liverworts of St Helena. Pisces Publications for St Helena Nature Conservation Group, Newbury, England\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Atlantic Oceanic, Islands, Bryophytes, Distribution, Endemism, Similarity","lastPublishedDoi":"10.21203/rs.3.rs-3389905/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3389905/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"We investigated the species richness, endemism, and similarities of the bryofloras on ten islands and archipelagos in the Atlantic Ocean to determine the floristic relationships among them using presence/absence. Cluster analysis was used to determine similarities among the bryofloras of ten islands and archipelagos. We examined the bryophyte species to address the following questions: (1) How many bryophyte species are there in the total area and on each island and archipelago? (2) Do mosses and liverworts share patterns of diversity? (3) What are their species distribution patterns? (4) How many endemic species are there on the islands? (5) Is there a high similarity among those bryophyte floras? We encountered 1,647 species, 325 genera, and 204 families, with the richest families (Lejeuneaceae, Lophocoleaceae, Plagiochilaceae, Brachytheciaceae, Bryaceae, Fissidentaceae, and Pottiaceae), representing 35% of all species. Over 12% of the bryophytes are African elements, 8% have global distributions, approximately 3% are Macaronesia elements, 5% are endemic, and 34% are included in other distributions. We present a checklist of 1,647 bryophyte species, of which 91 are endemic. Bryophyte richness and diversity differed significantly among the ten islands and archipelagos studied, and their low similarities illustrate their high degrees of heterogeneity. This paper is the first evaluation of bryophyte species on ten islands and archipelagos of the Atlantic Ocean, giving insight into their species richness, endemism and similarities, and providing information for analyzing species turnover, migration, invasion, etc., representing an increase in our general knowledge of the bryofloras of oceanic islands and archipelagos, contributing to the targets of the Global Strategy for Plant Conservation.","manuscriptTitle":"Tropical Atlantic Oceanic Islands and Archipelagos: Physical structures, plant diversity, and affinities of the bryofloras of northern and southern islands","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2023-10-04 16:50:33","doi":"10.21203/rs.3.rs-3389905/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"f8244f23-c4d9-45dd-97ac-b2df21432ad0","owner":[],"postedDate":"October 4th, 2023","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2023-12-19T21:59:20+00:00","versionOfRecord":[],"versionCreatedAt":"2023-10-04 16:50:33","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3389905","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3389905","identity":"rs-3389905","version":["v1"]},"buildId":"cBFmMYwuxLRRLfASyISRj","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}