AÇOREANA, 2011, Suplemento 7: 149-206
AZOREAN BRYOPHYTES: A PRELIMINARY REVIEW OF RARITY PATTERNS
Rosalina Gabriel1, Nídia Homem1, Adalberto Couto1, Silvia Calvo Aranda1,2 & Paulo A.V. Borges1
1Azorean Biodiversity Group – CITA-A, Dep. Ciências Agrárias, Universidade dos Açores, 9700-042 Angra do Heroísmo, Portugal e-mail: [email protected] 2Dep. Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales (CSIC). C/ José Gutiérrez Abascal, 2. 28006, Madrid, Spain
ABSTRACT Bryophytes are not exempt of rarity and threat, although their small size, mute colours and difficult field identification may mask their true conservation status. Actually, it is known that a quarter of all European bryophytes are under actual or potential threat. The first Red Data Book for European Bryophytes was produced in 1995, largely based on national red lists and on the work of a vast team of bryologists who assessed the conservation status of each European species. The red listing of bryophytes has undoubtedly contributed to increase the awareness of planners to this group of organisms, and several efforts have been made, through Europe, to preserve sites based on their bryological interest. Accordingly, a specific Red List for the Azorean Bryophytes may help regional managers to identify particularly endangered species, thus allowing for the creation of measures to improve their preservation. In this paper we have used an adaptation of the works of Deborah Rabinowitz (1981), who created a typology to access different forms of rarity, using three variables: Geographical Distribution, Abundance and Habitat Specificity. All the 480 species and subspecies known to occur in the Azores were surveyed; of these, 215 species lacked sufficient data to be analyzed (data deficient), 121 were not considered rare and 144 (1 hornworts, 56 liverworts and 87 mosses) were considered rare, at least in one of the three parameters considered. The benefits and limitations of the methodology are briefly discussed. Several practical suggestions are proposed in order to enhance the conservation of selected bryophyte species.
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RESUMO Os briófi tos podem ser tão raros e estar tão ameaçados como os demais organismos do planeta, apesar de o seu pequeno tamanho, cores discretas e difícil identifi cação no campo poderem mascarar o seu verdadeiro estatuto de conservação. De facto, é reconhecido que cerca de um quarto de todos os briófi tos da Europa estão efectiva ou potencialmente ameaçados. O primeiro “Livro Vermelho dos Briófi tos da Europa” foi produzido em 1995, amplamente baseado em listas vermelhas nacionais e no trabalho de uma vasta equipa de briólogos que avaliaram o estatuto de conservação para as espécies Europeias. A classifi cação de briófi tos em listas vermelhas tem contribuído para aumentar a sensibilidade dos gestores para este grupo de organismos e alguns esforços têm sido desenvolvidos na Europa, para preservar locais tendo como característica o seu interesse briológico. Consequentemente, uma lista vermelha para os briófi tos dos Açores pode auxiliar os gestores regionais a identifi car espécies particularmente ameaçadas, tornando-se o primeiro passo para assegurar a sua protecção. Neste artigo usamos uma adaptação dos trabalhos de Deborah Rabinowitz (1981), que criou uma tipologia para desocultar e avaliar várias formas de raridade, utilizando três variáveis: Distribuição Geográfi ca, Abundância e Especifi cidade do Habitat. Todas as 480 espécies e subespécies dos Açores foram investigadas: 215 taxa não tinham informação suficiente para ser analisados (deficientes em dados), 121 não foram consideradas raros e 144 briófitos (1 antocerota, 56 hepáticas e 87 musgos) foram considerados raros pelo menos num dos parâmetros considerados. Os benefícios e limitações desta metodologia são brevemente discutidos. São propostas algumas sugestões práticas para melhorar a estratégia de conservação dos briófitos seleccionados.
INTRODUCTION chipelagos (González-Mancebo et al., 2008; Sérgio et al., 2008), ne of the most interesting a feature unparalleled in other Ocharacteristics of the Azores groups of Azorean organisms is their extraordinary wealth (Izquierdo et al., 2004; Borges of bryophytes (480 species and et al., 2008, 2010a). In addition, subspecies, Gabriel et al., 2010), Azorean islands host a high pro- comparable to the diversity pre- portion of European bryophyte sent in other Macaronesian ar- species (Homem & Gabriel,
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2008) and also many endemic plantation forests dominate the species of vascular plants, mol- islands these days (Borges et al., luscs and arthropods (Borges et 2009). al., 2010b), many of which are in Most ecological studies in is- danger as a consequence of his- lands, and in particular in the torical human occupation and Azores, are limited in their time land-use changes (Borges et al., span and a detailed understand- 2000, 2009; Cardoso et al., 2010; ing of the long-term responses Martín et al., 2010; Triantis et of island bryophyte communi- al., 2010). In fact, the conserva- ties to global change drivers is tion of island biota was always not known. Bryophytes have considered a true priority since long been considered indicator most of the recorded extinctions groups for habitat change, as have occurred in islands (Sax & their lack of roots makes them Gaines, 2008). totally dependent on the at- In spite of more than four mospheric (or aquatic) inputs centuries of Human occupa- of nutrients (eg. Frego, 2007; tion, the Azores and other Gignac, 2010). Besides, bryo- Macaronesian archipelagos, phytes are a characteristic part still possess natural habitats of the Azorean native forests, (Borges et al., 2009; Gaspar et covering all kinds of substrata, al., 2011), and these islands are including leaves of vascular some of the very few places in species, with luxuriant com- Europe where the ‘biodiversity munities (Homem & Gabriel, crisis’ is particularly critical and 2008), and are generally con- a proper conservation strategy sidered remnants of the sub- may effectively contribute to tropical flora that endured the preserve unique pristine com- Quaternary glaciations (but see munities. Presently, about a Aigoin et al., 2009, who recently fifth of the Azorean islands area questioned the relictual origin is under some legal protection of Macaronesian bryophytes). status (Monteiro & Furtado, Thus, assessing the conserva- 2010), and a few remnants of tion status for bryophytes may native forests have persisted couple with policies for native since the Portuguese occupa- habitats protection. tion in the early 15th century, A red list ranks taxa accord- although grasslands and exotic ing to their threat level and
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extinction risk, and assess- Brummitt et al., 2008). In prac- ments for the red list compile tice, not many species have been current knowledge of conser- studied in any of these ways and vation status and threats to in- the information necessary to do dividual species (ex. Knapp & so is impressive. Nevertheless, Monterrosa Salomón, 2010). the need to better understand Few vascular plants and even the rarity of species is pressing fewer bryophytes (only 101 and simple methods of ranking of the ca. 18000 species!) have should at least be essayed for all been formally assessed using groups of organisms. the IUCN system (IUCN, 2010). The pioneer work of Deborah However, one of the targets of Rabinowitz (1981, 1986) has en- the Global Strategy for Plant lightened the rarity concept, ac- Conservation is “a preliminary knowledging that assessment of the conservation “There are many ways in which status of all known plant spe- a species can become rare and this cies, at national, regional and path has profound evolution- international levels” (UNEP, ary and ecological consequences” 2002). There are various ap- (Rabinowitz, 1981: 205). proaches to achieve this goal, To define rarity, she used a three including the use of expert’s dimensional system including opinions (ex. Sérgio et al., 1992; distribution, abundance and Schumacker, 2001; Sjögren, habitat specificity. Each one of 1995), the use of herbarium la- these dimensions was further bels information (ex. Krupnick subdivided into two qualita- et al., 2008), the creation of tive categories (wide or nar- specific software to create red row, large or small, generalist lists accommodating the IUCN or specialist, respectively), re- criteria (ex. RAMAS, 2007), but sulting in an eight cells table, lately it has been advised that a from which, only one cell in- thorough use of all available in- cludes common species, those formation, including georefer- with wide distributions, large enced herbarium specimens and populations and generalist. other parameters such as popu- All other combinations suffer lation size and local abundance, from at least one form of rar- would be a good way forward ity. Knowing the rarity status to stimulate conservation (ex. of species is critical to evaluate
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both their extinction proneness Azorean bryophytes, as in- and their roles in the ecosys- serted in ATLANTIS database, tems (Gaston, 1994, 2010). supplemented with literature One of the most useful re- and herbaria ecological fea- sources to study biodiversity tures, to analyze the rarity pat- in the Azores is the regional terns of the species and provide species database, ATLANTIS, a preliminary conservation as- where grid-based spatial inci- sessment, at the regional lev- dence information, allied to tem- el, of this important indicator poral data, has been gathered group. It is expected that it may for several groups of organisms be the basis of an Azorean Red (lichens, bryophytes, vascular List for Azorean bryophytes. plants, marine invertebrates, terrestrial molluscs, arthro- METHODS pods and vertebrates) (Borges et al., 2010c; Martín et al., 2010). Study Area Parallel to this work, the bio- The nine islands composing logical and ecological features the archipelago of the Azores, of bryophytes have also been are situated in the North Atlantic noted by RG and co-workers. Ocean, extending along a west- The information thus gathered northwest to east-southeast ori- may therefore serve as a start- entation (between 36° 30’ - 40° ing point for an assessment of North latitudes and 24° 30’ - 31° the rarity of bryophytes, using 30’ West longitudes). The joined Rabinowitz’ approach (consider- area of the islands is 2,323 km2 ing range, abundance and dis- (larger island, São Miguel, 745 tribution). Such a characteriza- km2; smaller island, Corvo, tion has been applied to vascular 17 km2) (Forjaz, 2004). The ar- plant species (ex. Rabinowitz et chipelago’s highest altitude is al., 1986; McIntyre, 1992), verte- reached in Pico Island, at 2,350 brates (ex. Kattan, 1992; Goerck, m, but the second highest is- 1997), insects (ex. Fattorini, land (São Miguel) is just 1,105 2011) and was essayed for m altitude. The Azores were European liverworts (Weibull & uninhabited when Portuguese Söderström, 1995). navigators arrived in the early In this study we used all 15th century, and the earlier de- the information available to scriptions of the islands por-
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tray them as intensely forested also examined. All informa- (Frutuoso, 1963). Nowadays tion was fed to the ATLANTIS the Azorean population in- database (Borges, 2005). This cludes about 241,800 people, database includes 29,323 spe- at a density of 104 inhabitants cies citations coming from lit- per square kilometre (Forjaz, erature (most of them, ca. 80%, 2004). It is estimated that laurel with recognizable locations and forests occupied about 85% of indication of date of collection) the present area of Azores; un- and 11,237 citations coming fortunately, most of this natural from herbarium records (most habitat has been disturbed, re- of them placed at the bryophyte maining only around 6,000 ha collection of the University of (Fernández-Palacios et al., 2011; the Azores). One of the authors Gaspar et al., 2011). (RG) has further compiled in- formation on fruiting, ecology Data Sources and substrate preferences for A tentative categorization of each bryophyte record; in some rarity was essayed for all the 480 occasions the habitat type was bryophyte species and subspe- inferred from other published cies included in the most recent sources or direct knowledge of check-list of the Azorean Islands local experts. Although gaps (Gabriel et al., 2010). The ana- of information are inevitable, lyzed data came from literature and have indeed been demon- and herbarium records. The strated (see Aranda et al., 2010), first step included a thorough this database is deemed to be as analysis and interpretation of complete as possible and a good the available literature, dating starting point to analyze rarity back to the 19th century (167 issues. A webpage, the Azorean sources; see Appendix 1); this Biodiversity Portal (http://www. list includes books, chapters and azoresbioportal.angra.uac.pt/), papers as well as some grey lit- with data on the taxonomy, de- erature such as academic thesis, tailed distribution of the species letters and fieldwork reports on the Azorean Islands (grid (see Borges et al., 2010c for de- of 500 m x 500 m), European tails). Secondly, the collection conservation status and some of bryophytes deposited at the pictures and common names University of the Azores was (whenever possible) has been
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available to the general public Vanderpoorten & Hallingback, since 2008 (Borges et al., 2010c). 2008). In order to reach an es- timation of abundance for bryo- Rarity dimensions phytes, we have taken advantage Geographical distribution of a recurrent pattern in ecologi- Due to the high disper- cal communities, i.e. the posi- sal ability characteristic of the tive intraspecific or interspecifc group (see revision in Rydin, relationship between mean lo- 2008), bryophytes occurring cal abundance and regional dis- in the Azores were considered tribution (Gaston, 1994, 1996), to have a narrow geographical which assumes that distribution when their pres- “Within a taxonomic assem- ence was known only from blage, locally abundant species the Macaronesian Islands (i.e. tend to be widespread and locally Macaronesian endemic species rare species tend to be restricted and subspecies) and a wide dis- in their distribution.” (Gaston, tribution, whenever they also 1996: 211). occurred elsewhere. This data The key issue here is the use of the was obtained from the recent small-scale distribution as a proxy checklist for Azorean bryo- of abundance. The importance of phytes (Gabriel et al., 2010). scaling, rarity and risk, has been highlighted by Hartley & Kunin Abundance (2003), working with two plant Abundance was the most dif- species (Dianthus armeria L. and ficult parameter to quantify, as it Silene otites (L.) Wibel) at a distri- refers to the size of the popula- bution resolution of 1-km in Great tions, which is not immediate in Britain. Bearing this in mind, and bryophyte studies (Hallingbäck using the ATLANTIS database, we et al., 1998; Hallingbäck, 2007). have calculated for each species the Different authors have used dif- number of geographical cells (500 ferent approaches to estimate m x 500 m) allocated with the high- abundance, such as the examina- est precision values (precision 1 – tion of museum specimens (see very precise locations, usually Fattorini, 2011 for arthropods), point UTM data; 2 – localities and others have not considered never exceeding 25 km2) in all this parameter for bryophytes Azorean Islands (see Borges et al., (ex. Söderström & Séneca, 2008; 2010c), and subsequently divided
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that value by the total number of Miguel is the worst. Likewise, 500 m2 cells of the archipelago the percentage of records to (10044 cells), thus reaching an which it was not possible to as- estimate of the area of occupan- sign a habitat varied according to cy (AOO) for each bryophyte. the islands (highest in Faial and This ratio of relative area of oc- São Jorge and lowest in Terceira cupancy was then considered a and Corvo) but, in average it did predictor of the local abundance not reach one tenth (9.7%) of the for each species. All the species 34976 records considered. were ranked by this index, and To appreciate the ecological those which fell below the me- range of a species, all the re- dian value were considered of cords where this was possible, low abundance while the others were allocated to one of the 12 were considered as abundant. habitat types. Then, the number of records present in one habitat Ecological tolerance was divided by the total number Habitat specificity was used of locations of that habitat (nor- as a proxy of ecological toler- malizing the records per habi- ance. RG’s species database on tat). Finally, for those species ecological traits was categorized that had 12 or more described in 12 different habitat types occurrences, the Lloyds Index (Coastal habitats, Mesic areas, of Patchiness (L) was applied: 2 2 Native forests, Semi-natural L = Sx – x / x + 1 (Basset, 1999), 2 grasslands, Mountainous ar- where Sx and x are respectively eas, Aquatic habitats, Peat the variance and mean of the bogs, Urban habitats, Parks and samples in the 12 different habi- Gardens, Intensive pastures, tat types. A specialist species Exotic forest plantations and in the present context is a spe- Cave entrances). Table 1 in- cies that showed preference for cludes a summary description of a particular habitat, the value each of the habitats considered. of the index increasing for more The islands survey is not eq- specialized species. According uitable (Table 2); for example, if to the interpretation of differ- the number of records per km2 is ent authors (eg. Basset, 1999; considered, Corvo, Terceira and Gabriel & Bates, 2005) those spe- Graciosa are the best inspected cies with an L value larger or of the Azorean Islands while São equal to three, were considered
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TABLE 1. Brief description of the habitats considered in this paper and an indication of the number of independent locations where bryophytes were collected in the Azores. Number of Habitat types Description locations Coastal habitats are situated at the lowest altitudes, near the Coastal sea, mostly up to an altitude of 50 m, which may be higher, 124 habitats depending on the Island. Mesic areas occur above the coastal habitats and receive intermediate amounts of precipitation. These areas are Mesic areas 204 presently dominated by fi elds (mostly corn fi elds), intensive pastures and exotic plantations. Native forests are the remnants of the former dominant ecosystem types, found by the fi rst se lers. They include Native forests 522 evergreen tree species such as Laurus azorica, Erica azorica, Ilex perado subsp. azorica and Juniperus brevifolia. Semi-natural These are open areas, mostly located among native forest 42 grasslands fragments, including several herbaceous plant species. Mountainous This habitat (high mountain) is restricted to Pico Island, above 57 areas 1200 m altitude. This habitat includes lagoon margins, temporary and Aquatic permanent rivulets, cascades and other interior waters 212 habitats habitats.
Peat bogs Large, open areas dominated by Sphagnum spp. 115
Habitats that may be found in cities and villages, including Urban habitats 70 buildings and other human constructions. Parks and Areas covered with exotic species, organized to appreciate 48 Gardens nature. Intensive Areas dominated by Holcus, Bromus or Lolium species, used by 129 pastures grazing ca le. Exotic Areas dominated by Eucalyptus spp., Cryptomeria japonica or plantation 163 Pi osporum undulatum. forests Specifi c habitat, including all the rocky walls of caves (lava Cave entrances tubes) and volcano entrances (pits and pit caves), where 81 light penetrates.
with restricted habitat require- normalized for the number of ments. Before proceeding to the total occurrences in that habitat. calculus of L, the number of oc- For instance, while there were currences in a given habitat was 522 locations inside native for-
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TABLE 2. General characteristics of the Azorean Islands, including the total number of bryophyte’ records made in the archipelago and the absolute and relative frequencies of records to which no habitat could be attributed. (1. Forjaz, 2004). Highest Number of Records without Area1 Inhabitants1 Azorean point1 records habitat information Island 2 (km ) (m) (Censos 2001) (NT)(NH) (%)
Santa Maria 97 587 5578 942 98 10,4
São Miguel 745 1105 131609 3897 224 5,7
Terceira 400 1021 55833 13104 433 3,3
Graciosa 61 405 4780 1576 32 2,0
São Jorge 244 1053 9674 4054 744 18,4
Pico 445 2350 14806 6501 780 12,0
Faial 173 1043 15063 2076 404 19,5
Flores 141 911 3995 1551 126 8,1
Corvo 17 720 425 1275 103 8,1
ests, there were only 163 loca- ing categorization: 1. Species tions placed in exotic plantation that are not rare; 2. Scarce spe- forests. cies (rare in abundance); 3. Species with narrow ecological Vulnerability index tolerance; 4. Restricted spe- Species considered rare on dis- cies (species rare by geographi- tribution, abundance and ecologi- cal range); 5. Scarce species cal tolerance, tend to be the most with narrow ecological toler- prone to extinction (Ka an, 1992; ance; 6. Scarce and restricted Manne & Pimm, 2001). The con- species; 7. Restricted species sequent application of the three with narrow ecological range criteria, with their binomial mea- and 8. Restricted and scarce surements: Distribution (large/ species with narrow ecological small), Abundance (common/ range. Similar categories may rare) and Ecological tolerance be appreciated for other groups (wide/narrow), led to the follow- such as vertebrates (ex. Kattan,
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1992) and arthropods (Fattorini, that all measures were not inde- 2011) and also for bryophytes pendent (p<0.05). Söderström (1995). Twenty four species, nine liv- erworts and 15 mosses, previ- RESULTS ously classifi ed in the European Red List of Bryophytes (ECCB, Of the 480 species referred 1995; Dierssen, 2001), four of to the Azores, only 265 (55.2%) which (Acanthocoleus aberrans, could be analyzed following Jamesoniella rubricaulis, Fissidens the combination of criteria used azoricus and Neckera cephalonica) (Appendix 2). From the evalu- also suggested by Sjögren (1995) ated species, about half (121; to become protected species in 45.7%) were not considered rare the Azores, could not be evaluat- (1. Species that are not rare) but ed in this analysis. All of these 24 six of the seven types of rarity species are scarce (rare by abun- proposed by Rabinowitz (1981) dance) and none had the neces- were found within the Azorean sary number of collections to al- bryophytes’ dataset (absolute low a full assessment of their eco- and relative frequencies of the logical tolerance. Among them eight categories may be seen in there are fi ve restricted species, Figure 1). If one considers sin- two Azorean endemics (Fissidens gle categories of rarity by them- azoricus and Trematodon persso- selves, less than half (112; 42.3%) niorum) and three Macaronesian of the evaluated species pre- endemics (Leucodon canariensis, sented narrow ecological tol- Neckera cephalonica and Tortula bo- erance, more than one fi h (56; gosica). Trematodon perssoniorum 21.1%) were considered scarce and which, so far, was only found only 17 evaluated species (6.8%) in São Miguel Island seems to had restricted distributions. prefer aquatic habitats, and was The results of the Chi-square collected mostly around Lagoa test show that the hypothesis das Furnas and Ribeira Quente of overall independence of the (seven records at diff erent times), three factors may be rejected while Riccia ligula was only re- (X2=47.36; df=2; p < 0.05), indi- corded in intensive pastures (six cating that these factors are not records) and Jamesoniella rubri- independent. Separate analysis caulis was only collected above of the 2 x 2 tables also indicated 1000 m (five records).
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DISCUSSION O’Malley, 2007) and is being considered for larger organisms Only about half (265) of all with microscopic dispersing Azorean bryophytes species and stages (e.g. spores), such as ferns subspecies (480) could be classi- or bryophytes (ex. Fontaneto, fied using the three rarity cate- 2011). An indirect evidence of gories proposed by Rabinowitz this wide distribution ability is (1981). In itself, this exposes the low endemism value found a serious lack of information, among Azorean bryophytes regarding mostly abundance (n=7; 1,5%), much lower those and ecological tolerance, which found among native vascular thwarts the design of a compre- species or arthropods (Borges et hensive conservation policy for al., 2010b). Moreover, accord- bryophytes. Without appropri- ing to the study of Söderström & ate knowledge of the biology of Séneca (2008), the liverwort flora the species, it is not possible to of Europe and Macaronesia con- understand why a bryophyte is sists of mainly widespread spe- rare or threatened and it is very cies, and, unlike what happens difficult to propose measures with vascular species, the rarest that would induce its recovery. species occur in oceanic areas The data presented in Figure (and not in the Mediterranean 1 and Appendix 2, shows that region). most of the analyzed bryophytes Eight of the 17 Macaronesian that may be considered rare and Azorean endemic bryo- have wide range distributions phytes evaluated, exhibited re- (247 species), which is not sur- stricted distributions (Appendix prising, considering that bryo- 2, “vulnerability index 4”), while phytes successfully disperse by not appearing to be scarce or re- spores. Actually some authors stricted in their habitat require- such as Medina, Draper & Lara ments. Interestingly enough, (2011), have argued that due to all of these eight species have their high dispersal ability, bryo- been reported to the three geo- phytes would tend to ubiquity. graphical groups of islands The hypothesis “Everything is and are presently known of six everywhere, but the environment (Breutelia azorica) or more, of selects” (EiE) has generally been the nine Azorean islands (other accepted by microbiologists (ex. seven species). Although the
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DISTRIBUTION (d) wide range (1) restricted range (0)
dense (1) ABUNDANCE (a) scarce (0)
generalist (1) specialist (0) generalist (1) specialist (0)
HABITAT (h)
FIGURE 1. Distribution of rarity types within the evaluated bryophyte species (n=265) in the Azores. Numbers indicate number of species per category; dark areas of pie charts indicate the percent of the dataset each rarity type represents. 1, Species that are not rare; 2, Scarce species (rare by abundance); 3, Species with narrow ecological tolerance; 4, Restricted species (rare by range); 5, Scarce species with narrow ecological tolerance; 6, Restricted and scarce species; 7, Restricted species with narrow ecological range; 8, Restricted and scarce species with narrow ecological tolerance.
Chi-square tests indicated a sig- abundantly collected in different nificant association among dis- types of habitats and all islands tribution, rarity and abundance, of the Azores (eg. González- endemism is not always asso- Mancebo et al., 2009). If these ciated with narrow ecological species evolved in Macaronesia tolerance or with scarcity; spe- (neoendemics), or survived cies such as Andoa berthelotiana only in Macaronesia (paleoen- and Leucodon treleasei have been demics), they should indeed
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be well adapted to the Azorean Forty liverworts and 24 mosses, ecosystems. Notwithstanding, more than half (!) of the special- Echinodium renauldii, which was ist bryophyte species evaluated recently confirmed as a true en- in this study (n=112) and about demic species (Stech et al., 2008), a quarter of all evaluated species is considered vulnerable by the show preference for natural for- IUCN (BSG, 2010), on account of ests (n=64; 24.2%). This is not its decreasing population trend surprising in view of what we and occurrence in less than ten know about the original plant localities in fi ve islands of the cover of the islands – a dense Azores; this view is shared by forest ecosystem (ex. Frutuoso, Sjögren (1995). Fortunately, the 1963) that is lavishly covered number of places where this spe- with bryophytes in all occurring cies has been collected is now substrata. In spite of its obvi- known to be higher (more than ous decrease in area (Silveira, 40 locations) and its presence was 2007), the diversity and luxuri- confi rmed in two more islands ance of the communities that (Corvo and Terceira) since 1995. may be observed in the remain- Almost half (n=112; 42,3%) ing native forest fragments (ex. of all the analyzed bryophytes Gabriel & Bates, 2005; Homem, were considered specialists in 2005) is still staggering; thus, it their habitat requirements, as re- is understandable, that this is ferred by their high Lloyd index the single most important habi- values, achieved when a high tat for bryophyte conservation proportion of the total number in the Azores. Recently it was of collections are grouped into also demonstrated that Azorean one, or mostly two, habitats. native forests are a unique habi- Man-made habitats, such as ex- tat for the conservation of most otic forests, grasslands or urban endemic arthropods (Triantis et habitats do not seem to harbour al., 2010), and a high proportion specialist bryophyte species. of those species are now under This in itself has sobering impli- threat of extinction due to its re- cations for conservation, because duction. Bryophytes depending of the historical decrease and on native fragments are prob- fragmentation of native habitats ably under the same pressures (Triantis et al., 2010; Gaspar et and would greatly benefit from al., 2011). an increase in the areas devoted
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to natural forests and from a longispina, Cyclodictyon laete- careful control of the quality of virens, Plagiothecium nemorale, remaining fragments, such as Tetrastichium virens and others, the removal of invasive species. may be found both in native Peat bogs are structurally forest and cave entrances. This very different from forests, in ability to colonize cave rocks is their openness and permanent likely to expand their altitudi- access to water and eleven spe- nal range, as many of the lowest cies were considered specialists altitude records were from cave from this habitat, taking ad- entrances. It is clear that caves vantage of these special condi- are acting as refugia for some of tions. Obviously, Sphagnum and these species. Due to the particu- Polytrichum species (the green lar way abundance was inferred and brown makers of peatlands) from the data, this is the least are prone to be found in these independent rarity dimension habitats, but the persistent pres- studied. The main issue is the ence of the rare Isopterygium lack of standardized data from tenerum in Furnas do Enxofre where to take sound information (Terceira Island), should also be (but see Gabriel & Bates, 2005 noted. and Homem, 2005). However as Surprisingly, or not (see showed by Couto (2010), stan- Gabriel et al., 2006; Jennings, dardized data on abundance 2009), caves (lava tube and pit obtained for several sites, was caves entrances) are an equally well correlated with distribution interesting habitat for the spe- at the scale of Terceira Island. cialist group of bryophytes. Bearing this in mind, additional Beyond Asterella africana, that efforts to get standardized in- has been collected specifical- formation for different habitats ly in such habitats, two other and islands should be made. A liverworts and eight mosses relatively high number of the find refuge in these harsh en- evaluated species (n=56; 21.1%), vironments, where competi- showed low abundance values, tion from vascular species is and were thus considered scarse. certainly lower. Besides the 11 Scarse species include represen- species that have mostly been tatives from the three taxonomic collected at cave entrances, groups (hornworts, liverworts some species such as Plagiochila and mosses); some examples in-
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clude species that have been col- tion, generally following IUCN lected in a few places and were criteria for protected areas (ex. considered specialists such as Dudley, 2008). Nevertheless, Asterella africana (cave entrances), there is a quarter of all conser- Leptoscyphus azoricus (native vation concern species (n=35 forests) or Isopterygium tenerum species, 22 liverworts and 13 (peat bogs) but also species such mosses; 24.5%), including as Cephalozia lunulifolia, Fissidens Acrobolbus wilsonii, Adelanthus coacervatus or Campylopus bre- decipiens, Aphanolejeunea ma- vipilus. deirensis, Leptoscyphus azoricus, It is important to recog- Pallavicinia lyellii, Campylopus nize that among the species shawii or Cyclodictyon laetevirens that could not be evaluated are that exist in less than fi ve Island Azorean rare bryophytes seem to Parks. While some of these spe- be found mostly in three impor- cies have a restricted range in tant habitats: native forests, peat the archipelago, occurring also in bogs and cave entrances. While few islands (ex. Kurzia paucifl ora, a commendatory effort has been Cheilolejeunea cedercreutzii), oth- made in order to create natural ers, such as Plagiochila punctata (7 parks in all islands including Islands – 3 Island Parks), Calypogeia most native forests fragments, azorica (6 – 3), Cyclodictyon laetevi- no such effort has been made to rens (6 – 3), Pallavicinia lyellii (6 – 2) encompass lava tubes (Pereira et or Trichocolea tomentella (4 – 1) are al., 2011), which are largely un- not adequately protected by the der private land and require ade- current design of the Island Parks. quate legislation to protect them, This work illustrates that and peat bogs are presently very even among relatively well stud- disturbed habitats. ied groups of organisms – bryo- About a third (n=43, 26 liv- phytes, in a very confined region erworts and 17 mosses; 30.1%) – the Azores, where a continuing of all conservation dependent collection, identification and re- bryophytes (n=143) exist in five porting effort has been made or more Island Parks. These through time, it was not pos- Parks (one for each Island) have sible to have a clear picture of recently been created in the the general rarity patterns of all Azores and incorporate areas species, and only about half of using different levels of protec- the reported taxa (n=265; 55.2%)
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could be assessed using a simple also mentioned by the ex- method of categorization. This perts Erik Sjögren (1995) hinders conservation efforts, as and / or René Schumacker only a fraction of knowledge is (2001): Acanthocoleus aber- available to managers and deci- rans, Jamesoniella rubricaulis, sion makers, while enlightening Fissidens azoricus and Neckera the way forward. It is clear that cephalonica. better floristic knowledge and 3. One liverwort (Aphanolejeunea expertise on bryophytes is nec- madeirensis) and three essary in the Azores, if we are to mosses (Fissidens coacerva- preserve the wealth of species tus, Sphagnum nitidulum, and the natural communities Thamnobryum rudolphianum) where they occur. As Knapp & have come out as restricted, Monterrosa Salomón have stated: scarce and with a narrow “[this] method is not a substitute ecological tolerance, which for a quantitative conservation as- means they were considered sessment…” (2010: 527), however rare in the three dimensions it is a way of setting priorities considered. While it is obvi- for further study or monitoring. ous that their conservation Some suggestions follow: in the Azores should be care- 1. The 143 species selected at fully planned, the taxonomic least by one of Rabinowitz’s status of S. nitidulum and F. dimension of rarity should coacervatus, should be clari- be followed and all efforts fied. should be made to adequate- 4. Island Parks are acting as ly conserve their habitats. “safe areas” for a number of 2. The 24 species previously bryophyte species however, selected by IUCN criteria other conservation concern (ECCB, 1995; Dierssen, 2001), species would benefit from that could not be evaluated a reshape, sometimes quite in this study for lack of col- straightforward, of those pro- lection records, should be tected areas. very carefully prospected 5. Some species that are not in the field and their evolu- routinely included in red tion monitored, especially lists have nonetheless come the four species that were up as rare in one or two
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dimensions, an aspect al- LITERATURE CITED ready discussed for mosses by Sjögren (2006). This en- AIGOIN, D.A., N. DEVOS, S. lightens the scale problem HUTTUNEN, M.S. IGNATOV, of conservation: it is impor- J.M. GONZALEZ-MANCEBO tant to acknowledge that & A. VANDERPOORTEN, 2009. regional, as well as global, And if Engler was not completely conservation plans should wrong? Evidence for multiple evo- be enforced. lutionary origins in the moss fl ora of Macaronesia. Evolution, 63: 3248- ACKNOWLEDGEMENTS 3257. ARANDA, S.C., R. GABRIEL, P.A.V. We are grateful for the careful BORGES & J.M. LOBO, 2010. text revision and suggestions Assessing the completeness of given by Simone Fattorini. We bryophytes inventories: an oceanic wish to thank Eva Sousa Borges, island as a case study (Terceira, Berta Martins for their careful Azorean archipelago). Biodiversity feeding of the ATLANTIS data- and Conservation, 19: 2469-2484. base and Enésima Mendonça for BASSET, Y., 1999. Diversity and abun- her help throughout the project dance of insect herbivores forag- and Fernando Pereira for his ing on seedlings in a rainforest in help with old names of locali- Guyana. Ecological Entomology, 24: ties and general information 245-259. on their habitats. RG and PAVB BORGES, P.A.V., 2005. Introduction. worked on this project under In: BORGES, P.A.V., R. CUNHA, the EU projects INTERREGIII B R. GABRIEL, A.M.F. MARTINS, “ATLÂNTICO” (2004-2006) and L. SILVA & V. VIEIRA (eds.), A list BIONATURA (2006-2008). NH of the terrestrial fauna (Mollusca and has a short grant from CITA-A. Arthropoda) and flora (Bryophyta, SCA was funded by the Azorean Pteridophyta and Spermatophyta) Fundo Regional da Ciência e from the Azores, pp. 11-20. Direcção Tecnologia (M311/I009A/2005). Regional de Ambiente and Field work was partly funded by Universidade dos Açores, Horta, FCT - PTDC/AMB/70801/2006 - Angra do Heroísmo and Ponta Understanding Underground Delgada. Biodiversity: Studies of Azo- BORGES, P.A.V., A.R.M., SERRANO & rean Lava Tubes. J.A. QUARTAU, 2000. Ranking the
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APPENDIX 1. List of references used in this bryophyte survey. Nº Citation
ADE, A., & F. KOPPE, 1942. Beiträge zur Kenntnis der Moosfl ora der atlantis en Inseln und 1 der pyrenais en Halbinsel. Hedwigia, 81: 1-34.
ALLORGE, P., & V. ALLORGE, 1946. Les étages de la végétation muscinale aux îles Açores et 2 leurs elements. Mémoires de la Societé de Biogéographie, 8: 369-386.
ALLORGE, P., & V. ALLORGE, 1950. Hépatiques recoltées par P. et V. Allorge aux îles Açores 3 en 1937. Révue Bryologique et Lichénologique, 19: 90-118.
ALLORGE, P., & V. ALLORGE, 1952. Mousses recoltées par P. et V. Allorge aux îles Açores en 4 1937. Révue Bryologique et Lichénologique, 21: 50-95. ALLORGE, P., & H. PERSSON, 1938a. Contribution à la fl ore hepaticologique des îles Açores. 5 Annales Bryologici, 11: 6-14. ALLORGE, P., & H. PERSSON, 1938b. Mousses nouvelles pour les Açores. Le Monde des 6 Plantes, 39 (232): 25-26. ALLORGE, V., & P. ALLORGE, 1938. Sur la répartition et l’écologie des hépatiques epiphylles 7 aux Açores. Boletim da Sociedade Broteriana, 2ª série, 13: 211-236. ALLORGE, V., & P. ALLORGE, 1944. Le Telaranea nematodes (Go sche) Howe dans le 8 domaine ibero-atlantique. Compte Rendu Sommaire des Séances de la Société de Biogéographie, 182-184: 58-60. ALLORGE, V., & P. ALLORGE, 1948. Végétation bryologique de l’île de Flores (Açores). 9 Révue Bryologique et Lichénologique, 17: 126-164. ALLORGE, V., & S. JOVET-AST, 1950. Aphanolejeunea teotonii nov.sp., hépatique des Açores. 10 Révue Bryologique et Lichénologique, 19: 19-24.
ALLORGE, V., & S. JOVET-AST, 1955. Cololejeunea azorica V.A. et S.J.-A., Lejeunéacée nouvelle 11 de l´Îlle San Miguel. Mi eilungen der Thüringischen Botanischen Gesellscha , 1 (2/3): 17-22.
ALLORGE, V., & S. JOVET-AST, 1956. Targionia lorbeeriana K.M. dans la Péninsule Ibérique, 12 aux Açores et aux Canaries. Révue Bryologique et Lichénologique, 25: 134-135.
ALLORGE, V., 1951. Trematodon perssonorum Allorge et Theriot espèce nouvelle des Açores. 13 Révue Bryologique et Lichénologique, 20: 179-181. ANDO, H., 1973. Révision des espèces africaines de Gollania (Hypnaceae). Révue Bryologique 14 et Lichénologique, 39: 529-538. 15 ARMITAGE, E., 1931. Some bryophytes of the Açores. Journal of Botany, 69: 75-76. ARTS, T., 1989. Rhamphidium purpuratum Mi .: its vegetative propagation and distribution. 16 Lindbergia, 15: 106-108. 17 BARROS, G., 1942. Notas briológicas, II. Agronomia Lusitana, 4(1): 155-166. BATES, J., 2000. Introduction to the Azores and its Bryophytes. Bulletin of the British Bryological 18 Society, 76: 21-23. BATES, J.W., & R. GABRIEL, 1997. Sphagnum cuspidatum and S. imbricatum ssp. affi ne new 19 to Macaronesia, and other new island records for Terceira, Azores. Journal of Bryology, 19(3): 645-648. BISCHLER- CAUSSE, H., 1993. Marchantia L. The European and African taxa. Bryophytorum 20 Bibliotheca, 45: 1-129. BISCHLER, H., 1970. Les espèces du genre Calypogeia sur le continent africain et les îles 21 africaines. Revue Bryologique et Lichénologique, 37: 63-134.
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Nº Citation
BISCHLER, H., 1976. Exormotheca pustulosa Mi en. Distribution, écologie, caryotype, spores, 22 parois sporales, germination. Revue Bryologique et Lichénologique, 42(3): 769-783. BISCHLER-CAU E, H., 1993. Mar antia L. The European and African Taxa. J. Cramer, 23 Berlin. BOUMAN, A.C., & G.M. DIRKSE, 1990. The genus Radula in Macaronesia. Lindbergia, 16: 24 119-127. BROWN, C.E., & E.V. WATSON, 1963. A note on a small collection of bryophytes from São 25 Miguel, Azores. Révue Bryologique et Lichénologique, 32: 181-182. BRUGGEMAN-NANNENGA, M.A., 1982. The section Pachylomidium (genus Fissidens). III. The F. crassipes-subcomplex (F. bryoides-complex), F. sublineaefolius (Pot. Varde) Brugg.- Nann. 26 and F. fl uitans (Pot. Varde) Brugg.- Nann. Proceedings of the Koninklij ke Nederlandse Akademie van Wetenschappen, Series C, 85(1): 59-104. BRUGGEMAN-NANNENGA, M.A., 1985. The section Pachylomidium (genus Fissidens). IV. 27 Further species from Europe, the Mediterranean and the Atlantic African islands. Proceedings of the Koninklij ke Nederlandse Akademie van Wetenschappen, Series C, 88(2): 183-207. BUCH, H., & H., PER ON, 1941. Bryophyten von den Azoren und Madeira. Societas 28 Scientiarum Fennica. Commentationes Biologicae, 8(7): 1-15. CARDOT, J., 1897. The mosses of the Azores. Annual Report of the Missouri Botanical Garden, 29 8: 51-72. CARDOT, J., 1905. Nouvelle contribution a la fl ore bryologique des Îles Atlantiques. Bulletin 30 du Herbier Boissier, Série 2, 5: 201-215. CASAS, C., M. BRUGUES, R.M. CROS & C. SÉRGIO, 1985. Cartografi a de Briòfi tos. Península 31 Ibèrica i les Illes Baleares, Canàrias, Açores i Madeira. I. Institut d’estudis Catalans, Barcelona.
CASAS, C., M. BRUGUES, R.M. CROS & C. SÉRGIO, 1989. Cartografi a de Briòfi tos. Península 32 Ibèrica i les Illes Baleares, Canàrias, Açores i Madeira. II. Institut d’estudis Catalans, Barcelona.
CASAS, C., M. BRUGUES, R.M. CROS & C. SÉRGIO, 1992. Cartografi a de Briòfi tos. Península 33 Ibèrica i les Illes Baleares, Canàrias, Açores i Madeira. III. Institut d’estudis Catalans, Barcelona.
CASAS, C., M. BRUGUES, R.M. CROS & C. SÉRGIO, 1996. Cartografi a de Briòfi tos. Península 34 Ibèrica i les Illes Baleares, Canàrias, Açores i Madeira. IV. Institut d’estudis Catalans, Barcelona. CHURCHILL, S.P., 1986. A revision of Echinodium Jur. (Echinodiaceae: Hypnobryales). 35 Journal of Bryology, 14: 117-133. CHURCHILL, S.P., 1989. Transfer of Lepidopilum virens Cardot to Tetrastichium (Leucomiaceae). 36 Journal of Bryology, 15: 537-541. CLARO, D., C. SÉRGIO & R. SCHUMACKER, 2009. Bryophytes of S. Jorge Island (Azores, 37 Portugal). Conservation and biogeographic characterization. Portugaliae Acta Biologica, 23: 147-223. CLARO, G.D., 2008. Briófi tos da Ilha de S. Jorge: Conservação e caracterização biogeográfi ca. 38 Mestrado em Biologia da Conservação. Faculdade de Ciências da Universidade de Lisboa. Lisboa. CRUNDWELL, A.C., H.C. GREVEN & R.C. STERN, 1994. Some additions to the bryophyte 39 fl ora of the Azores. Journal of Bryology, 18: 329-337. CRUNDWELL, A.C., 1981. Reproduction in Myurium hochste eri. Journal of Bryology, 11: 715- 40 717. CUNHA, A.G., & G. BARROS, 1942. Algumas espécies de Musgos da Terceira novas para os 41 Açores ou para a ilha. Boletim da Sociedade Portuguesa de Ciências Naturais, 13: 156-157.
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Nº Citation DIAS, E., & R. GABRIEL, 1994. Distribuição das Comunidades vegetais no Algar do Carvão (Terceira, Açores). In: Actas do 3º Congresso Nacional de Espeleologia e do 1º Encontro Internacional 42 de Vulcanoespeleologia das Ilhas Atlânticas, 214-226 pp. (30 de Setembro a 4 de Outubro de 1992). Angra do Heroísmo.
DIAS, E., & C. MENDES, 2007. Characterisation of a basin mire in the Azores archipelago. 43 Mires and Peat, 2: 1-11. h p://www.mires-and-peat.net/ .
DIAS, E., 1986. Estudo Bio-Ecológico da Bacia da Lagoa do Negro. Relatórios e Comunicações 44 do Departamento de Biologia, 16: 1-131. Ponta Delgada.
DIAS, E., 1989. Métodos de estudo e análise da vegetação. Comunidades herbáceas. M. Sc. thesis. 45 Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo.
DIAS, E., 1996. Vegetação natural dos Açores. Ecologia e sintaxonomia das fl orestas naturais. Ph.D. 46 thesis. Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo.
DIAS, E., C. MENDES & A.J. SHAW, 2009. Sphagnum recurvum P. Beauv. on Terceira, Azores, 47 new to Macaronesia-Europe. Journal of Bryology, 31: 1999-201. DIXON, H.N., 1909. Contributions to the moss fl ora of the Atlantic islands. Journal of Botany, 48 47: 365-374. DUELL-HERMANNS, I., 1986. Taxonomy of Homalia lusitanica Schimp. and H. subrecta 49 (Mi .) Jaeg: is H. subrecta a species or a variety? Bryologische Beiträge, 6: 67-84. EDWARDS, S.R., 1997. Key to the Fissidens species from the Azores, 6 pp. Manuscript (last 50 edited 2 August 2002)... EGGERS, J., 1982. Artenliste der Moose Makaronesiens. Cryptogamie, Bryologie et Lichénologie, 51 3: 283-335. ENGEL, J.J., & G.L.S. MERRILL, 2004. Austral Hepaticae. 35. A Taxonomic and Phylogenetic 52 Study of Telaranea (Lepidoziaceae), with a Monograph of the Genus in Temperate Australasia and Commentary on Extra-Australasian Taxa. Fieldiana, 44: 149-174. FELDBERG, K., H. GROTH, R. WILSON, A. SCHÄFER-VERWIMP & J. HEINRICHS, 2004. 53 Cryptic speciation in Herbertus (Herbertaceae, Jungermanniopsida): Range and morphology of Herbertus sendtneri inferred from nrITS sequences. Plant Systematics Evolution, 249: 247-261. FONTINHA, S., & C. SÉRGIO, 1995. 11. Eucladium verticillatum (Brid.) B.S.G. novo musgo 54 para briofl ora da ilha Terceira (Açores). In: Notulae Bryofl orae Macaronesicae III. Revista de Biologia (Lisboa) “1994”, 15: 189. FRAHM, J.-P., 1975. Taxonomis e Notizen zur Ga ung Campylopus. Révue Bryologique et 55 Lichénologique, 41(3): 321-332. FRAHM, J.-P., 1982. Taxonomis e Notizen zur Ga ung Campylopus. XII. Cryptogamie, 56 Bryologie et Lichénologie, 3(1): 59-65. 57 FRAHM, J.-P., 2004. A Guide to Bryological Hotspots in Europe. Archive for Bryology, 3: 4-14. FRAHM, J.-P., 2005. Briófi tos colhidos por Jan-Peter Frahm e colaboradores nas ilhas Terceira, Pico e 58 Faial, em Agosto de 2004. Manuscrito. FRAHM, J.-P., 2005. New or interesting records of bryophytes from the Azores. Tropical 59 Bryology, 26: 45-48. GABRIEL, R., & J.W. BATES, 2003. Reponses of photosynthesis to irradiance in bryophytes of 60 the Azores laurel forest. Journal of Bryology, 25: 101-105.
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Nº Citation GABRIEL, R., & E. DIAS, 1994. First approach to the study of the Algar do Carvão fl ora (Terceira, Azores). In: Actas do 3º Congresso Nacional de Espeleologia e do 1º Encontro Internacional 61 de Vulcanoespeleologia das Ilhas Atlânticas (30 de Setembro a 4 de Outubro de 1992), 206-213 pp. Angra do Heroísmo. GABRIEL, R., & C. SÉRGIO, 1991. Notas acerca dos endemismos da fl ora briológica açoreana: 62 Sphagnum nitidulum Warnst. IX Simposio Nacional de Botánica Criptogámica, Poster. Libro de Resúmens. Salamanca. GABRIEL, R., & C. SÉRGIO, 1995. Bryophyte survey for a fi rst planning of conservation areas 63 in Terceira (Açores). Criptogamica Helvetica, 18: 35-41.
GABRIEL, R., 1994a. Briófi tos de pastagem. Algumas noções de ecologia. M.Sc. thesis. 64 Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo. GABRIEL, R., 1994b. Briófi tos da Ilha Terceira (Açores). Ecologia, distribuição e vulnerabilidade 65 de espécies seleccionadas. M.Sc. thesis. Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo. GAUTHIER, R., & M. BRUGUÉS, 1997. Note sur la présence de Sphagnum affi ne Ren. & Card., 66 Sphagnum centrale C. Jens. et Sphagnum papillosum Lindb. aux Açores. Cryptogamie, Bryologie et Lichénologie, 18(2): 121-125. GEHEEB, A., 1910. Bryologia atlantica - Die Laubmoose der atlantis en Inseln. E. 67 Schweizerbat´sche Verlagsbuchhandlung, Stu gart. GIESE, M., & J.-P. FRAHM, 1985. A revision of Microcampylopus (C.Müll.) Fleisch. Lindbergia, 68 11: 114-124. GONZÁLEZ-MANCEBO, J.M., A. LOSADA-LIMA & C.D. HÉRNANDEZ-GARCIA, 1991. 69 A contribution to the fl oristic knowledge of caves on the Azores. Mémoires de Biospéologie, 17: 219-226. GROLLE, R., & H. PER ON, 1966. Die Ga ung Tylimanthus auf den atlantis en Inseln. 70 Svensk Botanisk Tidskri , 60(1): 164-174. GROLLE, R., & R. SCHUMACKER, 1982. Zur synonymik und vebreitung von Plagio ila 71 spinulosa (Di s) Dum. und P. killarniensis Pears. Journal of Bryology, 12: 215-225. GROLLE, R., 1962. Monographie der Lebermoosga ung Leptoscyphus Mi . Nova Acta 72 Leopoldina, 25(161): 1-143. GROLLE, R., 1966. Dicranolejeunea auf den Atlantis en Inseln. Transactions of the British 73 Bryological Society, 5: 95-99. GROLLE, R., 1968. Monographie der Ga ung Nowellia. Journal of the Ha ori Botanical 74 Laboratory, 31: 20-49. GROLLE, R., 1970. Zur Kenntnis der Frullanien in Europa und Makaronesien. Wiss. Ztschr. 75 Friedrich-Schiller-Univ., Jena, Math.-Naturwiss. Reihe, 19 (3): 307-319. GROLLE, R., 1972. Bazzania in Europa und Makaronesien. Zur Taxonomic und Verbreitung. 76 Lindbergia, 1: 193-204. GUERKE, W.R., 1978. A monograph of the genus Jubula Dumortier. Bryophytorum Bibliotheca, 77 17: 1-119. HE, S., 1997. A Revision of Homalia (Musci: Neckeraceae). Journal of the Ha ori Botanical 78 Laboratory, 81: 1-52. HEDENÄS, L., 1992. Notes on Madeiran Pseudotaxiphyllum, Brachythecium and Rhynchostegiella 79 species (Bryopsida). Nova Hedwigia, 54(3-4): 447-457. 80 HEDENÄS, L., 2006. Bryoxiphium norvegicum in Azores.
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Nº Citation HEIRCHS, J., D.S. RYCROFT, H. GROTH & W.J. COLE, 2002. Morphological and 81 phytochemical studies of Plagiochila papillifolia Steph., a Neotropical liverwort new to Europe. Journal of Bryology, 24: 119-126. HEINRICHS, J., H. ANTON, S.R. GRADSTEIN & R. MUES, 2000. Systematics of 82 Plagiochila sect. Glaucescentes Carl (Hepaticae) from tropical America: a morphological and chemataxonomical approach. Plant Systematics and Evolution, 220: 115-138. HENTSCHEL, J., H.-J. ZÜNDORF, F.H. HELLWIG, A. SCHÄFER-VERWIMP & J. HEINRICHS, 83 2006. Taxonomic studies in Chiloscyphus corda (Jungermanniales: Lophocoleaceae) based on nrITS sequences and morphology. Plant Systematics and Evolution, 262: 125-137. HÜBSCHMANN, A. von., 1974. Bryologis e Studien auf der Azoreninsel São Miguel. 84 Revista da Faculdade de Ciências de Lisboa, Série 2. C, 17: 627-702. JOVET-AST, S., 1948. Bazzania tricrenata (Wahl.) Trevis. aux Açores. Révue Bryologique et 85 Lichénologique, 17(1-4): 174. LONG, D.G., & M.O. HILL, 1982. Tortula solmsii (Schimp.) Limpr. in Devon and Cornwall, 86 newly recorded in the British Isles. Journal of Bryology, 12: 159-169. LUISIER, A., 1937. Recherches bryologiques récentes à Madère (Deuxième Série). Brotéria, 87 Série de Ciências Naturais, 6: 88-95. 88 LUISIER, A., 1938. Mousses des Açores. Brotéria, Série de Ciências Naturais, 7: 96-98. 89 LUISIER, A., 1938. Hepáticas dos Açores. Brotéria, Série de Ciências Naturais, 7: 187-189. LUISIER, A., 1945. A família das Hookeriáceas na Península Ibérica e nas Ilhas da Madeira, 90 Açores e Canárias. Las Ciencias, 1: 1-8. LÜPNITZ, D. von, 1975. Geobotanis e Studien zur natürli en Vegetation der Azoren unter 91 Berü si tigung der Corologie innerhalb Makaronesiens. Beiträge Biological Pfl anzen, 51: 149-319. MASCHKE, J., 1976. Taxonomische Revision der Laubmoosga ung Myurium (Pterobryaceae). 92 Bryophytorum Bibliotheca, 6: 1-219. MASTRACCI, M., 2004. Thamnobryum rudolphianum (Neckeraceae, Musci), a new species 93 from the Azores. Lindbergia, 29: 143-147. 94 MAY, R., 1986. Notes on some Macaronesian Tortella species. Bryologische Beiträge, 6: 58-66. MENDES, C., 1998. Contributo para a caracterização de turfeiras de Sphagnum spp. na ilha 95 Terceira. Relatório de Estágio de Licenciatura em Engenharia Agrícola. Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo. MITTEN, W., 1870. Musci, Hepaticae. In: GODMAN, F. (ed.), Natural history of the Azores on 96 Western Islands, 286-328 pp. John Van Voorst, London. OCHYRA, R., 2005. Racomitrium aquaticum colhido por Frahm em 2004 no Pico, Açores. 97 Manuscrito. PERSSON, H., 1937. Briófi tos colhidos por H. Persson em São Miguel em 1937, 1 pp. Registo de 98 Herbário. Manuscrito. PERSSON, H., 1938. Account of the botanical travel to the Açores in 1937 by Dr. Herman Persson 99 and his wife, 11 pp. Manuscrito. 100 PERSSON, H., 1939. Bryophytes from Madeira. Botaniska Notiser, 566-590 PERSSON, H., 1973. The Azorean bryophytes collected by P. Dansereau and A. R. Pinto da 101 Silva in 1964. Agronomia Lusitana, 35: 5-19 RICHARDS, P.W., 1937. A collection of bryophytes from the Azores. Annales Bryologici, 9: 102 131-138.
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Nº Citation RUSSEL, J.L., 1862. Some notes on the cryptogamic vegetation of Fayal, Azores. Proccedings 103 of the Essex Institute, 2: 134-137. RYCROFT, D.S., W.J. COLE, N. ASLAM, Y.M. LAMONT & R. GABRIEL, 1999. Killarniensolide, 104 methyl orsellinates and 9,10-dihydrophenanthrenes from the liverwort Plagiochila killarniensis from Scotland and the Azores. Phytochemistry, 50: 1167-1173. RYCROFT, D.S., J. HEINRICHS, W.J. COLE & H. ANTON, 2001. A phytochemical and 105 morphological study of the liverwort Plagiochila retrorsa Go sche, new to Europe. Journal of Bryology, 23: 23-34. SANTOS, N., 2005. Elaboração de uma ferramenta promocional da ZPEPVRG (Zona de Protecção Especial do Pico da Vara e Ribeira do Guilherme) numa perspectiva de educação ambiental. Relatório 106 de Estágio de Licenciatura em Engenharia do Ambiente. Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo. SCHUMACKER, R., & R. GABRIEL, 2002. Gymnocolea infl ata (Huds.) Dumort., on Terceira 107 island (Azores, Portugal). Portugaliae Acta Biologica, 20: 101-104. SCHUMACKER, R., & J. VÁNA, 1999. Two new liverworts for Europe in Macaronesia: 108 Odontoschisma prostratum (Sw.) Trevis. on the Azores and Jungermannia callithrix Lindenb. & Go sche on the Azores and Madeira. Tropical Bryology, 17: 115-127. SCHUMACKER, R., 2001. The hepatic fl ora of the Azores: brief historical outline, present 109 knowledge, endemics and phytogeographical aspects. Belgium Journal of Botany, 134(1): 51-63.
SCHUMACKER, R., 2002/2003. Lista de espécies de briófi tos da Serra do Labaçal observados por R. 110 Schumacker, 1 p. Verão de 2000 e 2003. Manuscrito.
SCHUMACKER, R., 2003. New national and regional bryophyte records, 8; 2. Cephalozialla 111 dentata (Raddi) Steph. Journal of Bryology, 25: 217-221. 112 SCHUMACKER, R., 2005. Jungermannia hyalina Lyell, 1 p. Manuscrito. 113 SCHUMACKER, R., 2005. Lepidozia stuhlmanii Steph, 1 p. Azores. Manuscrito. 114 SCHWAB, G., 1981. Azoren. Herbar Go fried S wab, 19 pp. Manuscrito SÉRGIO, C., & R. GABRIEL, 1995. 7. Novos dados para os Açores sobre o género Riccia. In: 115 Notulae Bryofl orae Macaronesicae III. Revista de Biologia (Lisboa) “1994”, 15: 184-185.
SÉRGIO, C., & E. DIAS, 1991. 4 - Algumas Ricciaceae e Anthocerotaceae novas para ilha 116 Terceira. In: Notulae Briofl orae Macaronesicae. II. Portugaliae Acta Biologica, Série B, 15: 421-423.
SÉRGIO, C., & R. GABRIEL, 1997. 2. Fissidens papillosus Lac. um novo musgo para a briofl ora 117 Açoreana. In: Notulae Bryofl orae Macaronesicae IV. Portugaliae Acta Biologica, Série B, 17: 267.
SÉRGIO, C., & J.P. HERBRARD, 1982. Orthothecium duriaei (Mont.) Besch., étude systématique, 118 écologique et phytogéographique. Collectanea Botanica, 13(1): 247-255. SÉRGIO, C., 1974. Le genre Fossombronia au Portugal, à Madère et aux Açores. Bulletin de la 119 Societé Botanique de France, 121: 319-326. SÉRGIO, C., 1976. Deux nouveautés pour la fl ore bryologique des Açores: Kiaeria bly ii (B. 120 S. G.) Broth. et Orthodicranum fl agellare (Hedw.). Boletim da Sociedade Broteriana, 2ª série, 50: 99-105. SÉRGIO, C., 1978. Lejeunea eckloniana Lindenb. (hépatique) dans la Macaronesie. Boletim da 121 Sociedade Portuguesa de Ciências Naturais, 18: 39-41. 122 SÉRGIO, C., 1983. Do liverworts really like beer? The Bryological Times, 19: 4.
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Nº Citation SÉRGIO, C., 1985. 1. Considerações sobre a presença de Frullania muscicola Steph. e Frullania 123 ericoides (Nees) Mont. nos Açores e Madeira. Notulae Bryofl orae Macaronesicae. I . Portugaliae Acta Biologica, Série B, 14: 161-168. SÉRGIO, C., 1987. Contribuição para o estudo taxonómico e fi togeográfi co de Anthoceros caucasicus 124 Steph. na Península Ibérica e Macaronésia. Actas del VI Simpósio Nacional de Botánica Criptogámica (1987), pp. 605-614. SÉRGIO, C., 1991. 1. Sphaerocarpus texanus Aust. nova hepática para a fl ora da Macaronésia. 125 In: Notulae Briofl orae Macaronesicae II. Portugaliae Acta Biologica, Série B, 15: 419.
SÉRGIO, C., 1995. 5. Duas novas espécies de Fossombronia para a briofl ora dos Açores. In: 126 Notulae Bryofl orae Macaronesicae III. Revista de Biologia (Lisboa) “1994”, 15: 183. SÉRGIO, C., 1995. 6. Petalophyllum ralfsii (Wils.) Nees et Go ex Lehm., espécie nova para 127 os Açores e para a Macaronésia. In: Notulae Bryofl orae Macaronesicae III. Revista de Biologia (Lisboa) “1994”, 15: 184. SÉRGIO, C., 1995. 8. Notas sobre o género Sphagnum no arquipélago dos Açores. In: Notulae 128 Bryofl orae Macaronesicae III. Revista de Biologia (Lisboa) “1994”, 15: 185.
SÉRGIO, C., 1997. 3. Grimmia pulvinata (Hedw.) Sm. um novo musgo para a ilha Terceira 129 (Açores). Notulae Bryofl orae Macaronesicae IV. Portugaliae Acta Biologica, Série B, 17: 268. 130 SÉRGIO, C., 2005. Comunicação pessoal em 3 de Junho de 2005. SÉRGIO, C., & R. SCHUMACKER, 1999. Notulae Bryofl orae Macaronesicae V. 3. Àcerca da 131 presença de Campylopus introfl exus (Hedw.) Brid. Nos Açores. Anu. Soc. Brot., 65: 90-91. SÉRGIO, C., R. GABRIEL & E. DIAS, 1995. 10. Novos musgos para a fl ora da ilha Terceira 132 (Açores). In: Notulae Bryofl orae Macaronesicae III. Revista de Biologia (Lisboa) “1994”, 15: 187- 188. SÉRGIO, C., R., GABRIEL & E., DIAS, 1995. 9. Hepáticas novas para a fl ora da ilha Terceira 133 (Açores). In: Notulae Bryofl orae Macaronesicae III. Revista de Biologia (Lisboa) “1994”, 15: 186-187. SÉRGIO, C., Z. IWATSUKI & A. EDERRA, 1997. Fissidens luisierii P. Varde (Fissidentaceae, 134 Musci), a neglected species from Macaronesia. Journal of the Ha ori Botanical Laboratory, 83: 237-249. 135 SILVEIRA, T., 1937. Flora briológica. Espécies novas para os Açores. Açoreana, 1: 240-247. SIM-SIM, M., L. LUÍS & M. STECH, 2009. Hygroamblystegium fl uviatile. In: BLOCKEEL, T. et 136 al., New national and regional bryophyte records. Journal of Bryology, 31: 135.
SIM-SIM, M., M. STECH & G. ESQUÍVEL, 2009. Southbya tophacea. In: BLOCKEEL, T. et al., 137 New national and regional bryophyte records. Journal of Bryology, 31: 137. SIM-SIM, M., C. SÉRGIO, R. MUES & L. KRAUT, 1995. A new Frullania species (Trachycolea) 138 from Portugal and Macaronesia. Frullania azorica sp. nov. Cryptogamie, Bryologie-Lichénologie, 16(2): 111-123. SJÖGREN, E., 1973. Plant communities of the Natural vegetation of Madeira and the Azores. 140 Monographiae Biologicae Canariensis. Las Palmas, 4: 107-111.
SJÖGREN, E., 1973. Recent changes in the vascular fl ora and vegetation of the Azores Islands. 141 Memórias da Sociedade Broteriana, 22: 1-451. SJÖGREN, E., 1978. Bryophyte vegetation in the Azores Islands. Memórias da Sociedade 143 Broteriana, 26: 1-273.
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Nº Citation SJÖGREN, E., 1990. Bryophyte fl ora and vegetation on the island of Graciosa (Azores), with 142 remarks on fl oristic diversity of the Azorean islands. Arquipélago. Life and Earth Sciences, 8: 63-96. SJÖGREN, E., 1993. Bryophyte fl ora and vegetation on the island of Corvo (Azores). 143 Arquipélago. Life and Marine Sciences, 11A: 17-48. SJÖGREN, E., 1995. Report on investigations of the bryofl ora and bryovegetation in 1995 on the 144 Azorean islands of Faial, S. Jorge, Pico and Flores, 30 pp. LIFE-project. (mimeogr.) Angra do Heroísmo. SJÖGREN, E., 1996. Report on investigations of the bryofl ora and bryovegetation on the Azorean 145 island of Santa Maria, 24 pp. LIFE-project. (mimeogr.) Angra do Heroísmo. SJÖGREN, E., 1997. Report on investigations of the bryofl ora and bryovegetation in 1997 on 146 the Azorean island of Terceira. LIFE project. Departamento de Ciências Agrárias, Angra do Heroísmo. SJÖGREN, E., 2001. Distribution of Azorean bryophytes up to 1999, their island distribution 147 and information on their presence elsewhere, including Madeira and the Canary Islands. Boletim do Museu Municipal do Funchal, Sup. Nº 7: 1-89. SJÖGREN, E., 2003. Azorean Bryophyte Communities - A Revision of Diferencial Species. 148 Arquipélago. Life and Marine Sciences 20A: 1-29. SJÖGREN, E., 2004. List of species in herbario UPSV collected in the Azores Islands, 12 pp. 149 Manuscrito. SJÖGREN, E., 2005. List of species in herbario UPSV collected in the Azores Islands, 50 pp. 150 Manuscrito. SJÖGREN, E., 2005. Revisão da colecção de briófi tos recolhidos pelo casal Allorge nos Açores em 151 1937 e por outros autores do fi nal do século XIX e início do século XX e depositados no Museu Carlos Machado (Ponta Delgada, São Miguel), 1 p. Manuscrito. SJÖGREN, E., 2006. Bryophytes (Musci) unexpectedly rare or absent in the Azores. 152 Arquipélago, 23A: 1-17. SMOOKLER, M. M., 1967. Bryophyte report. Chelsea College Azores Expedition, 1965. Chelsea 153 College of Science and Technology, University of London, London. SOARES, P.C.M.G., 2003. Avaliação de elementos vestigiais atmosféricos na Ilha de Santa Maria, 154 por biomonitorização. Relatório de Estágio de Licenciatura em Engenharia do Ambiente. Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo. STECH, M., M. SIM-SIM, S. MARTINS & C. SÉRGIO, 2009. Plagiomnium undulatum var. 155 madeirense. In: BLOCKEEL, T. et al., New national and regional bryophyte records. Journal of Bryology, 31: 136. SMNH - Swedish Museum of Natural History 2006. Moss Register. (h p://andor.nrm.se/ 156 fmi/xsl/kbo/publFinditems.xsl?-token.nav=items&-view&-db=kbo_mossregister&-token. languagecode=en-GB). TAVARES, C.N., 1956. Teotónio da Silveira Moniz: 13.X.1891-5.V.1953. Revue Bryologique et 157 Lichénologique, n.s. 25: 188. The New York Botanical Garden, 2007. International Plant Science Center. State/Province: 158 Azores. (h p://sciweb.nybg.org/Science2/vii2.asp) THERIOT, J., 1938. Campylopodioidees des Îles Açores recoltées par P. Allorge et H. Persson. 159 Révue Bryologique et Lichénologique, 11: 100-109. TRELEASE, W., 1897. Botanical observations on the Azores. Annual Report of the Missouri 160 Botanical Garden, 8: 77-220 + Plates 12 to 66. UBC - University of British Columbia 2007. Herbarium Databases. (h p://herbarium.botany. 161 ubc.ca/index.html).
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Nº Citation VARDE, R.P., 1945. Formes atlantiques de trois mousses des Açores. Révue Bryologique et 162 Lichénologique, 15: 40-45. VARDE, R.P., 1955. Noveau Fissidens aux Açores. Mi eilungen der Thüringis en Botanis en 163 Gesells a , 1(2/3): 15-16. VIEIRA, B.J.H., 2002. Avaliação de elementos vestigiais atmosféricos na Ilha Terceira, por 164 biomonitorização. Relatório de Estágio de Licenciatura em Engenharia do Ambiente. Departamento de Ciências Agrárias, Universidade dos Açores, Angra do Heroísmo. WATSON, H.C., 1844. Notes on the botany of the Azores. The London Journal of Botany, 3: 165 582-617.
PPaginação_21.inddaginação_21.indd 181181 119-01-20129-01-2012 11:51:5711:51:57
182AÇOREANA 2011, Suplemento 7: 149-206 Vulnerability Index Vulnerability tolerance tolerance ecological tolerance tolerance with narrow ecological tolerance abundance)
2. Scarce species (rare by 3. Species with narrow ecological 3. Species with narrow ecological 5. Scarce species with narrow 3. Species with narrow ecological 8. Restricted and scarce species
Experts’ opinion Experts’
Old IUCN criteria criteria IUCN Old
Records >1975 Records
Records <1976 Records Maximum altitude (m) altitude Maximum
ed in the Rabinowitz (1981) rarity criteria
fi
cient; na, not applicable; es, SJÖGREN, 1995; cient; na, not applicable; es, SJÖGREN, 1995; (m) altitude Mean
fi Minimum altitude (m) altitude Minimum
50 701,8 1100 468 185 150 549,8 975 85 195 v 1. Species that are not rare
number of records of number Habitat with highest highest with Habitat
(180) (570) NaFor NaFor
records)
Lloyd index (>11 (>11 index Lloyd
m cells m
Number of 500 x 500 500 x 500 of Number
Parks
Presence in Island Island in Presence
Number of Islands of Number Groups of Islands of Groups
33 91 9 63 4 7 146 8 1,8 1 447 ExFor (18) 1,3 5 200 3 NaFor (24) 520,5 100 70 dd 825 372,3 0,6 14 700 na (1) NaFor (6) 45 49 175 350 51 r 440,0 350,0 700 350 3 0 1. Species that are not rare 12 4 1. Species that are not rare 3 41 1 3 67 dd 2 na (2) 20 dd dd NaFor (4) dd 500 690,0 dd 750 4 4 3 1 r es; rs 2 4 3 117 6,1 NaFor (47) 500 864,3 1050 20 32 v es; rs 3 6 4 179 6,4 NaFor (92) 500 856,8 1500 34 65 es 3 8 5 137 1,6 NaFor (26) 250 692,7 2300 15 33 1. Species that are not rare 3 7 5 79 7,9 (21) Caves 10 387,8 875 13 25 v rs 3 9 7 198 2,0 NaFor (76) 150 592,9 1050 75 65 v 1. Species that are not rare 3 5 4 92 dd NaFor (5) 500 616,7 850 6 5 3 9 8 390 2,4 3 8 6 624 4,2 Endemic grouping Endemic MAC 2 5 4 83 4,6 NaFor (36) 250 658,8 925 36 6 r rs Ast)
et n.) et ff . reb.) R. M. (Taylor) A. (Taylor) (Schi (Brot.) Stotler, (Brot.) Stotler, (S Steph. (Michx.) Prosk. (V. Allorge (V. (Lindenb. Steph. L. (Hook.) Mi Nees (L.) Prosk. (Mont.) A. Evans t ij Crand.-Stotl. (L.) Dumort. et e) Kru e) s