Additions , corrections and comments to the Red List of bryophytes from mainland Spain and the Balearic Islands

This article is a review of the conservation status of several taxa of bryophytes that were included in the Red List of Mainland Spain and the Balearic Islands, and those that were omitted, but should have been featured on the basis of most recent data. Of the 33 studied species that were regarded as threatened in the Red List, 31 have been downlisted upon reevaluation, mostly as a result of better knowledge of species and their distribution. On the other hand, this study highlights the most urgent need to review all taxa assigned to the “Deficient Data” category in the Red List since most of these taxa are precisely the most likely to be truly threatened.


Introduction
In recent years there has been an increase in the interest for conservation of bryophytes in Spain, in the form of both, compiling red lists (Brugués et al., 2014), and legislation at various administrative levels that provide protection to certain taxa (Infante et al., 2017).Significant progress on IUCN Red List assessments has been made for bryophytes nationaly and regionaly despite difficulties when applying IUCN Red List Criteria to regional assessments (Garilleti & Albertos, 2012).The establishment of detailed IUCN Red List assessments for plants is a crucial step in conservation planning.Red lists, in addition to being a useful tool for selecting areas of botanical interest for threatened species and habitats (Anderson, 2002), allow to identify which taxa within a specific territory are threatened.IUCN (2012) criteria are widely used for classifying taxa into risk categories.Evaluating the risk status of bryophyte taxa according to IUCN (2012) criteria demands a detailed knowledge of the species' demographics and population dynamics, which is very seldom feaseable.This limiting factor affects not only bryophytes but also vascular plants (Sáez et al., 2010) although for the former the dearth of available data is comparatively much worse.
The aim of this article is to update the red list of Mainland Spain and the Balearic Islands (Brugués et al., 2014) following IUCN (2012) criteria, through: i) the addition of new taxa not considered threatened in Brugués et al. (2014) and ii) the reevaluation of the conservation status of some taxa included in the red list.

Methods
The extinction risk of several bryophyte species included in the red list of Mainland Spain and the Balearic Islands (Brugués et al., 2014) was assessed using the IUCN Red List Categories and Criteria version 3.1 (IUCN, 2012).This is not, however, an assessment of the entire red list, but only of those taxa occurring in the northeastern Iberian Peninsula and in the Balearic Islands with sufficiently detailed and current information that begs for correcting the criteria used by Brugués et al. (2014) and thus assignment to a different IUCN category.A preliminary documentation of distribution and historical information has been carried out by verifying herbarium material (mainly BCB) and data either published or recorded in databases such as Banc de Dades de Biodiversitat de Catalunya (BDBC, http://biodiver.bio.ub.es/biocat/, accessed in April 2018).In some cases, data from field surveys were also used.Attribution to a risk category follows prescriptions and recommendations of IUCN (2012IUCN ( , 2017) ) with a justification, in each case, for assigning a taxon to a specific category.For those species assessed as threatened or near threatened (IUCN, 2012), the extent of occurrence (EOO) and the area of occupancy (AOO) are provided.EOO and AOO were determined using the Geospatial Conservation Assessment Tool (Bachman et al., 2011) with a default cell width of 2 km (IUCN, 2017).A potential rescue effect from populations located in adjacent territories has not been considered since, although it may exist in some cases, there is no evidence of its occurrence.Many of such species rarely develop sporophytes or their populations are extremely small, so rescue effect from adjacent areas (France, Andorra, etc.) through obvious geographic barriers (mostly the Pyrenees range) is regarded here as negligible.The herbaria codes follow Thiers (2018, continuously updated).

Results
Liverwort and moss taxa are listed below separately, arranged alphabetically.Acronyms for province names follow Flora Briofítica Ibérica (Guerra et al., 2006).
New category: VU B1ab(iii)+2ab(iii) The presence of this circumboreal species is confirmed in, at least, 6 areas from central Pyrenees (L province): Vall de Mulleres, Güells del Joeu, Liat and Mata de València (data from BCB), Aigüestortes (Casas Sicart, 1986) and the Aiguamog area (Carrillo et al., 2008).These localities (most of them correspond to recent collections) are found in 7 UTM 1x1 km squares (AOO = 24 km 2 , EOO = c.510 km 2 ).C. suecica is also known from nearby areas in France: Midi-Pyrénées region (Infante Sánchez, 2015) and Pyrénées-Orientales department [6 localities] (Hugonnot et al., 2018), although the possibility of any rescue from French populations is unknown.Brugués et al. (2014) assigned C. suecica to "EN" since a population decline is inferred due to natural causes and disturbances induced by human activity.The available information suggests that the species' range, in spite of being relatively restricted, is not severely fragmented (Figure 1) and exceeds the number of localities (sub-criterion "Ba") for "EN".Nevertheless, C. suecica may fulfill this subcriterion for category "VU", as proposed here.

Gymnomitrion corallioides Nees
Previous category: CR B2a(ii,iv) (Brugués et al., 2014).New category: excluded from the Spanish Red List According to the most recent data (Cros et al., 2015), this bipolar species (circumpolar, arctic-alpine in the Northern Hemisphere) is only known from a single locality in Spain, which corresponds to the Pica d'Estats (L province).Previously , Casas Sicart (1986) reported it from another Pyrenean locality (L province, Amitges, 31TCH31) but the voucher specimen (BCB 10550) corresponds to G. concinnatum (Lightf.)Corda.A record from "Aigüestortes i Sant Maurici" (Infante & Heras, 2012) is based on an erroneous report by Casas Sicart (1986).However, the locality where G. corallioides was collected in Pica d'Estats corresponds to its northern slope ("cara N, 3130 m", BCB 52776), entirely located within France (Ariège department), so, despite its proximity, this species should be excluded altogether from the Spanish bryophyte flora and thus its Red List.

New category: VU D2
This circumboreal species is reported from many European countries but generally regarded as rare (Söderström et al., 2002).It is found in central Pyrenees (Hu and L provinces) where it grows on decaying wood.Information on its occurrence in Spain was published by Casas & Infante (1998) for localities in Artiga de Lin and Goelhs deth Joeu (Aran Valley, L province), where several subpopulations are found between 1,400 and 1,600 m asl.In addition, other localities are known: Ordesa (BCB 54051), Marimanya, (BCB 30679), Mata de València (BCB 53706) and Portilló de Bossost (BCB 59293) (AOO=20 km 2 , EOO=c.800 km 2 ).L. ascendens is also known from nearby areas: Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez et al., 2015) and Pyrénées-Orientales department [9 localities] (Hugonnot et al., 2018).In absence of evidence regarding a continuous decrease in its Spanish populations (the subpopulation of Portilló de Bossost, which includes several stands, has been verified by us recently in July 2017 and there are no short-term threats), the "Bb" subcriterion ["B": small range (EOO and/or AOO), "b": continuing decline, observed, inferred or projected (IUCN, 2012)] cannot be applied.However, in view of the very restricted range and specificity for a fragile habitat, this species is here assessed as VU under criterion D2.

Nardia insecta Lindb.
Previous category: DD-n (Brugués et al., 2014).New category: CR B1ab(iii)+2ab(iii); D This species is known only from a single location in the northern slope of the Roca Blanca peak (L province), where it is extremely rare (Sáez et al., 2011).In the French central Pyrenees (Hautes Pyrénées) it is a very rare species, which has been assigned to the CR category in the Midi-Pyrénées region (Infante Sánchez et al., 2015).In Roca Blanca it has a very restricted population size (AOO and EOO=4 km 2 ).A single patch measuring about 25 cm 2 was found.In this study a single discrete patch of N. insecta is regarded as an individual.It therefore meets the IUCN (2012) criteria for Critically Endangered, based on the number of individuals (fewer than 50) and a decline due to natural plant competition is inferred.The attribution of this species to CR is consistent with that same assignment for its conspecific, N. breidleri (Limpr.)Lindb., with a similar conservation problem (Brugués et al., 2014).Few minor changes related to the subcriteria [CR B1ab(iii)+ 2ab(iii)] are proposed here.

Petalophyllum ralfsii (Wilson) Nees & Gottsche
Previous category: VU B2ab(iii,iv) (Brugués et al., 2014).New category: NT This species is listed as Vulnerable in the Red Data Book of European Bryophytes (Steward et al., 1995) and it has a scattered distribution in the Mediterranean region.Since its first discovery in the Balearic Islands (Casas de Puig, 1956), our knowledge of the distribution of P. ralfsii in the Balearic archipelago has improved significantly.Moreover, a recent record has expanded the limits of its distribution in eastern Spain (Segarra-Moragues & Puche, 2016).In Spain it is currently present in all the main islands of the Balearic archipelago (Figure 2) and a single locality in V province.The values of AOO and EOO are 84 km 2 and c. 4,630 km 2 , respectively.Habitats listed for this species in mainland Spain and the Balearic Islands are diverse, e.g.: clayish soil in a stream mouth, streambanks, roadsides, among pine litter in a shaded north-facing gully, on flat clayish surfaces, temporary freshwater pools on siliceous or calcareous soil, and a rushing stream.Although there is no doubt that if mainland Spain was to be considered in isolation, the species well merits assessment as CR on the basis of D criterion, the assessment is very different when mainland Spain and the Balearic archipelago are taken together.There is no evidence of a continuous decline in the number of Balearic localities or AOO, and several populations currently contain hundreds of plants.Even if some of the less recently prospected populations were lost (possibly the case of a single coastal Majorcan locality: Port de Sóller towards Muleta), this circumstance cannot be considered as a continuous decline.To the contrary, some revisited Balearic populations do persist without reasonable conservation concerns.AOO value for P. ralfsii in the Balearic Islands and Mainland Spain has increased in recent decades (Figure 3).In fact Blockeel (2003) indicated that this species may be more common than presumed in some Mediterranean regions.Since Brugués et al. (2014) assessment, the species has increased its AOO by 16 km 2 (Pericàs et al., 2016;Segarra-Moragues & Puche, 2016) and it seems that previous considerations about P. ralfsii being threatened were based on inaccurate knowledge of its actual distribution and status.

Plagiochasma appendiculatum Lehm. & Lindenb.
Previous category: CR B2ab(ii,iii,iv) ["B2a (ii,iii,iv)"] (Brugués et al., 2014).New category: VU D2 The only European population of this species is located in northern Majorca, where it was initially known only from a karstic ravine: torrent de Coma Freda o des Guix (Cros et al., 2005).Due to its extreme rarity and the existence of anthropic impacts on the population, it was attributed to the CR category.In the locality of Coma Freda there are small stands in a section of this ravine, between 235 and 320 m asl, and there is a risk of anthropogenic disturbances due to canyoning.Recently, J.A.W. Nieuwkoop (in Ellis et al., 2016) has found a second and more abundant population of this species, which also corresponds to another karstic ravine (Torrent de Pareis) also located in northern Majorca.According to J.A.W. Nieuwkoop (l.c.), P. appendiculatum is found in several points of this ravine, between 20 and 100 m altitude.Considering all these new data, the values of AOO and EOO are 12 km 2 .P. appendiculatum does not have an extremely restricted distribution in the territory considered as initially assumed (Cros et al., 2005), so the category assigned by Brugués et al. (2014) must be reconsidered.Since there is no evidence of decline in the population or in quality, extent or area of habitat in order to qualify under subcriteria b of criteria B, P. appendiculatum is assessed as VU D2 on the basis of its restricted range and number of known locations locations, which makes it potentially vulnerable to stochastic events, as well as human activities.Extensive surveying in northern Majorca would also help to determine if further populations exist.
Scapania cuspiduligera (Nees) Müll.Frib.Previous category: DD (Brugués et al., 2014).New category: VU D2 This is a very rare species in Spain, where it is currently known from central and eastern Pyrenees: Vall d'Àssua (Manobens i Rigol & Casas Sicart 1985) and Serra de Cadí (B and L provinces) (Ruiz et al., 2018a) (AOO = 20 km 2 , EOO = c.70 km 2 ).In addition, S. cuspiduligera was reported from Valle de Oro (Casares, 1911) (Lu province) where its presence has not been confirmed recently.S. cuspiduligera was assessed as DD (Brugués et al., 2014) since it was excluded from the Spanish bryoflora by Brugués et al. (2011).Both Pyrenaean populations grow on shady calcareous rocky places (between 1,700 and 2,700 m asl) and are found within protected areas.It is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez, 2015) and Pyrénées-Orientales department (Hugonnot et al., 2018) although its rarity would prevent an eventual rescue effect (IUCN, 2003).There could be a decline in EOO and AOO if we accept that the population from Lugo province was lost (which would require confirmation), but this is not considered a continuous decline.The species is assessed as VU D2 given its rarity in our area (small area of occupancy, small number of known locations) which makes it potentially vulnerable to stochastic events.
Tritomaria scitula (Taylor) Jörg.Previous category: DD-n (Brugués et al., 2014).New category: CR B1ab(iii)+2ab(iii) It is known from the north slope of the Pic del Pinetó in L province (Sáez et al., 2011) where there are two small stands located in the same UTM 1x1 km square (31TCH3918) (AOO and EOO = 4 km 2 ), with the coverage of c. 4 m 2 .These stands are found near the base of some cliffs where there are several equipped climbing routes, so there is a risk of anthropogenic disturbances on the population and its habitat.In fact, since its discovery, it has maintained and expanded the climbing routes in the area, so a population decline is inferred due to disturbances induced by human activity.In the Iberian Peninsula, T. scitula was previously only known from Andorra (Sotiaux & Schumacker, 2002), where it is currently considered rare (Sotiaux & Vanderpoorten, 2017), while its presence in the French slope of the Pyrenees (Hautes Pyrénées) is uncertain (Infante Sánchez, 2015), so it seems that the isolation of the population is important or, at least, rule out an eventual rescue effect (IUCN, 2003).

Mosses
Acaulon fontiquerianum Casas & Sérgio Previous category: VU D2 (Brugués et al., 2014).New category: NT This species has a relatively wide but scattered distribution in eastern Iberian Peninsula.Until a decade ago this species was only known from Almería province (Guerra, 2006a), but in recent years its Spanish distribution has been significantly expanded.Brugués et al. (2016a) listed A. fontiquerianum from 6 provinces (Ab, Al, CR, Ge, Mu and V) where it is distributed in 13 UTM 10x10 km squares.Assuming that A. fontiquerianum exists only in 1 UTM 1x1 km square for each UTM 10x10 km squares, the values of AOO (c.52 km 2 ), EOO (> 20,000 km 2 ) and the number of localities greatly exceed the thresholds established for criterion D2.Although there is no clear evidence of recent population declines or significant population fluctuations, in some localities its habitat is vulnerable to anthropogenic disturbances.Therefore, it is assessed as Near Threatened.

Andreaea nivalis Hook.
Previous category: VU D2 (Brugués et al., 2014);EN B2ab(ii,iv,v) (Luceño et al., 2017).New category: NT This artic-alpine species is distributed through the Iberian Central System, Cantabrian mountain range and central Pyrenees (Av, Hu, L, O, S and Sa provinces).Luceño et al. (2017) provide a detailed distribution of this species in the Central Iberian System, where there are at least 4 main well delimited populations groups exist.In Spain at least 17 localities are known (AOO = 68 km 2 , EOO = c.115,600 km 2 ), what precludes the assignation of this species to VU D2 (IUCN, 2012), as Luceño et al. (2017) rightly suggested.However, according to the latter authors this species could be considered as EN B2ab(ii,iv,v) in Spain, since i) the threshold for AOO in the criterion B2 is not exceeded in EN category, ii) its distribution is fragmented (subcriterion a) and iii) the observed climate warming and fluctuations in precipitations are probably reducing the effective number of localities and individuals (subcriterion b).
According to IUCN (2012), 'severely fragmented' refers to the situation in which increased extinction risk to the taxon results from the fact that most of its individuals are found in small and relatively isolated populations and these small populations may go extinct, with a reduced probability of recolonization.However, this situation does not seem to be the case of A. nivalis in mainland Spain, since this species can be locally "abundant at high elevations" in the Iberian Central System (Luceño et al., 2017) [11 localities, 8 UTM 1x1 km squares, AOO=32 km 2 ].This regional population is self-sustaining, and it is not dependent on immigration for long-term survival.On the other hand, the connection and continuity of the populations from the Spanish Pyrenees with those located in France (Infante-Sánchez et al., 2015;Hugonnot et al., 2018) and Andorra (Sotiaux & Vanderpoorten, 2017) has not been considered.Certainly, the climate warming is probably affecting the viability of some populations (subcriterion Bb), but at present there are no empirical data to support the supposed continue demographic decline.For taxa with very restricted distribution, classification as VU D2 is only permissible if the effects of climate change are such that the taxon is capable of becoming CR or extinct in a very short period of time after the effects of the threat become apparent (IUCN, 2017: 87-88).If the species has a AOO<20 km 2 but is not declining or under specific threat or experiencing extreme fluctuations, although it may be affected by the climate change but the effects are expected to be gradual and slow (which will not trigger any criteria for CR or cause extintion within three generations) it does not meet VU D2.The case of the chionophile species Polytrichastrum sexangulare (see below) is especially interesting, because although it is only known from a single locality in Iberian Central System (Luceño et al., 2017) where it grows together with A. nivalis, in this case the effects of climate warming have not been considered (although AOO in the criterion B2 is not exceeded neither in EN nor in VU categories), and it was assessed as LC in Spain.In our opinion, considering the Spanish populations, it does not seem possible to infer or project a continuous decline in the AOO or in the number of subpopulations, as established by the criterion "Bb".This species is listed here as NT as it is close to qualifying for VU under criterion D2.
Anoectangium aestivum (Hedw.)Mitt.Previous category: DD (Brugués et al., 2014).New category: VU D2 This species was considered as Regionally Extinct (RE) for the Iberian Peninsula and Balearic Islands by Brugués & González-Mancebo (2012), as only a few confirmed records for Hu, Le and Po provinces in the late nineteenth and early twentieth century were known (see Cano et al., 2016).Recently, Guerra & Cano (2013) and Cano et al. (2016) have found this species in the central Pyrenees (La Renclusa, Hu province) and in the Cantabrian range (El Angliru, O province).The values of AOO and EOO are 8 km 2 .In the Midi-Pyrénées region it is also a rare species (Infante-Sánchez, 2015, Infante-Sánchez et al., 2015) while there are no records from eastern Pyrenees (Hugonnot et al., 2018), so it seems unlikely an eventual rescue effect (IUCN, 2003) from French populations.If ancient records were considered, the apparent decline of this species in Spain may not be real or severe, it may be overlooked or the concrete areas from where A. aestivum was found have not been explored in detail subsequently.
Although the category DD highlights taxa lacking sufficient information for a sound status assessment, if the taxon range is suspected to be narrow, or if a considerably period of time has elapsed since the last record of the taxon, considering it as threatened may be justified (IUCN, 2017: 10).Since A. aestivum is a very rare species in Spain, it is assessed as VU D2 based on its geographical restriction, which makes it vulnerable to stochastic events, as well as human activities.
Anomodon longifolius (Schleich.ex Brid.)C.Hartm.Previous category: VU D2 (Brugués et al., 2014).New category: NT In the Iberian Peninsula this species is found in northern mountainous areas (B, Ge, Hu, Na, SS and Vi provinces), where it grows on calcareous rocky places.Based on bibliographic references (Allorge, 1955;Granzow de la Cerda, 1988;Heras et al., 2002), herbarium data and field surveys values of AOO and EOO are 36 km 2 and c. 1,810 km 2 , respectively.It is also known from nearby areas in France: Midi-Pyrénées region (Infante Sánchez, 2015).The number of localities and AOO value preclude the assignation of this species to VU D2 (IUCN, 2012).Therefore, in absence of evidence regarding a continuous decrease in AOO or the disappearance of a significant number of subpopulations, it is assessed as NT.
Anomodon rostratus (Hedw.)Schimp.Previous category: VU B2ab(ii,iv) ["B2a(ii,iv)"] (Brugués et al., 2014).New category: NT This species is found in Spain in Ge, Hu, Na, SS, Te and Vi provinces, in 15 localities (AOO = 52 km 2 , EOO = c.47,290 km 2 ).It is also known from nearby areas in France: Midi-Pyrénées region (Infante Sánchez et al., 2015) -where it was regarded as VU and Pyrénées-Orientales department [6 localities] (Hugonnot et al., 2018).The assignment to the VU category (Brugués et al., 2014) was based on the AOO, severe fragmentation of the area (or known less than 10 localities) and that there is a continuous decrease in the area of occupation and the number of locations or subpopulations.It is unclear if the fragmentation can be classified as severe (it does not seem attributable to destruction of its habitat), because only in the area of Vidrà (Ge province) 7 subpopulations are close to each other and spread over 4 UTM of 1x1 km squares.It has not disappeared from a significant number of subpopulations, nor can a continuous decline in the occupation area be inferred or projected.On the basis of this reinterpretation of the available information, A. rostratus is listed as NT.
Aschisma carniolicum (F.Weber & D.Mohr) Lindb.Previous category: VU B2ab(ii,iii,iv) ["B2a(ii,iii,iv)"] (Brugués et al., 2014).New category: NT It is a widely distributed species in peninsular Spain and the Balearic Islands according to Brugués et al. (2016c) in A, Al, Ab, CR, Ge, H, Mu, PM, V and Vi provinces (AOO = 84 km 2 , EOO = c.300,000 km 2 ).This species grows on bare soil in open grassland and scrublands.The fragmentation of its area is not severe (in southern Spain it is rather relative) and the number of known localities is clearly higher than 10.On the other hand, in relation to the sub-criteria of criterion B, it is possible that some populations have been exposed (or may be exposed in the near future) to anthropogenic disturbances, but this can not be extrapolated to the whole Spanish population of A. carniolicum, since most of the populations are in well-conserved areas.The probability of immigration from southern Portugal is unknown, thus the category is not up-or downlisted.Aschisma carniolicum in Spain no longer meets the criteria for being globally threatened and it is hence listed as NT.

Atrichum flavisetum Mitt.
Previous category: DD-n (Brugués et al., 2014).New category: VU D2 Two confirmed Iberian populations of this species are known, corresponding to Baga de Queràs (BCB 22767) and Valle de Pineta (BCB 52091) (G and Hu provinces, respectively).The values of AOO and EOO are 8 km 2 .Although no detailed population data are available, both populations were found in subalpine forests located in protected areas (National and Natural Parks).Its presence is not known in Andorra (Sotiaux & Vanderpoorten, 2017) or in the Midi-Pyrénées region (Infante-Sánchez et al., 2015).Due to its strong geographical restriction and apparently without probability of immigration from neighbouring countries, it is advisable to assign this species, at least temporarily, as VU D2, according to IUCN (2017) recommendations (see comments under Anoectangium aestivum).

Brachythecium erythrorrhizon Schimp.
Previous category: Not listed by Brugués et al. (2014).New category: VU D2 Only two Iberian locations of this species are known (Orgaz et al., 2012, Guerra et al., 2015;Orgaz, 2018), corresponding to recent collections: Tavascan, estany de Romedo Baix (MUB 38468, MUB 38469) and Ansó, valle de Las Cabretas, entre Las Cabretas y el ibón de Estanés (VIT 29624) (Hu and L provinces).The values of AOO and EOO are 8 km 2 .The species is not listed for nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France (Infante Sánchez, 2015;Hugonnot et al., 2018).In the absence of more or less concrete demographic data, it is provisionally assessed as VU D2 based on low AOO value and the small number of localities, which makes it potentially vulnerable to stochastic events, as well as human activities.
Brachythecium mildeanum (Schimp.)Schimp.Previous category: VU D2 (Brugués et al., 2014).New category: NT According to the data provided by Orgaz et al. (2009) and Orgaz (2018) the distribution of B. mildeanum is relatively broad in the N and CW of Spain (B, Ge, Hu, Na, Sg, SS, To Vi and Za provinces) where there are, at least, 13 localities, which mostly correspond to mountain areas (between 580 and 1,900 m asl) in which no significant disturbances are predictable.The values of AOO and EOO are 52 km 2 and c. 116,000 km 2 , respectively.The species is also known from nearby areas in France: Midi-Pyrénées region (Infante Sánchez, 2015).It is classified as Near Threatened because it is close to qualify as threatened under criterion D2 due to its relatively small AOO and number of localities.
Brachythecium turgidum (C.Hartmann) Kindb.Previous category: VU D2 (Brugués et al., 2014).New category: NT In the Iberian Peninsula this species is found in seven Pyrenaean localities (B and L provinces) where it grows on humid calcareous rocky places, between 1,900 and 2,600 m asl (Cros et al., 2010;Sáez et al., 2011;Orgaz et al., 2012;Ruiz et al., 2018a;BCB).It is also known from nearby areas in France (Midi-Pyrénées region), where B. turgidum has been assessed as NT (Infante Sánchez et al., 2015).Values of AOO and EOO are 28 km 2 and c. 1,530 km 2 , respectively.All the Spanish localities are not subject to any problem to their conservation.All of them correspond to protected spaces.In a locality recently discovered by one of us (LS) in the Pedraforca massif (B province) B. turgidum is not particularly scarce, it appears in the form of several stands in almost inaccessible areas.Available data (number of localities and AOO value) in addition to the fact that the Spanish population of B. turgidum is not prone to the effects of human activities or stochastic events within a very short time period in an uncertain future that could drive the species to CR or EX preclude the assignation of this species to VU D2 (IUCN, 2012).However, since the species has a relatively restricted range in mainland Spain, it is assessed as NT.
Buxbaumia viridis (Moug.ex Lam.& DC.) Brid.ex Moug.& Nestl.Previous category: VU B2ab(ii,iii,iv) (Brugués et al., 2014).New category: NT This species, which is generally restricted to decaying wood, is currently known from 24 UTM 10x10 km squares in Spain (B, Ge, Hu, L, Na and Vi provinces).The values of AOO and EOO are 136 km 2 and c. 24,900 km 2 , respectively.In recent years new populations have been found even in Pyrenean sectors where B. viridis was unknown, such as the eastern Pre-Pyrenees (Carnicero & Unzeta, 2016;Ruiz et al., 2018a) and Andorra (Ellis et al., 2017).It is also known from nearby areas in France: Pyrénées-Orientales department [15 localities] (Hugonnot et al., 2018) and Midi-Pyrénées region (Infante Sánchez, 2015).Some Spanish populations that were regarded extinct at the regional level have been relocated, as for example in their southernmost Iberian population (Sáez et al., 2018).Most populations are found within protected areas: National Parks (Ordesa-Monte Perdido and Aigüestortes and Estany de Sant Maurici), Natural Parks (Posets-Maladeta, Alt Pallars, Cadí-Moixeró, Capçaleres del Ter and Freser, Montseny, etc.) and no reductions in their habitat or anthropic threats are predicted in the short or medium term (at least in Catalonia), although sporophyte consumption has been detected in central and western Pyrenees (Infante & Heras, 2018).Our preliminary results of population monitoring in three Pyrenaean locations (L province) and the Montseny massif (B province), over 5-7 years, do not indicate, for the moment, a decline in the sporophytes production.Population management actions have even been developed to favor subpopulations in some protected areas (Ordesa-Monte Perdido National Park and Montseny Natural Park).On the basis of all this information, it is most reasonable to assign B. viridis to NT.We have here assigned this species to NT because their populations are not strong from a demographic point of view (although this is an intrinsic characteristic of the species) and are found at the limit of its range, but cannot be ruled out the rescue effect (IUCN, 2003) from the populations of the northern slope of the Pyrenees.IUCN (2012) recommends the assimilation to the LC category to those abundant and widely distributed taxa, which does not conform to B. viridis, at least within our area.Our assessment of not considering this species as threatened at regionally level agrees with other studies conducted in other southern European peninsulas (Spitale & Mair, 2017).
Conostomum tetragonum (Hedw.)Lindb.Previous category: Not listed by Brugués et al. (2014).New category: VU D2 It is a very rare species in Spain (Heras & Infante, 2016) where only 5 localities are known: one from the Cantabrian mountain range (Pico Curavacas, Pa province) and the rest from the central Pyrenees (Hu and L provinces): Ibón de Paderna, Amitges, Certescan and Liat (AOO = 20 km 2 , EOO = c.5,470 km 2 ).Its presence in Sierra Nevada is doubtful (Rams et al., 2014).Conostomum tetragonum is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez et al., 2015) and Pyrénées-Orientales department (Hugonnot et al., 2018).The species is here assessed as VU D2 given its rarity in our territory (small area of occupancy, small number of known locations) which makes it potentially vulnerable to stochastic events, as well as human activities.

New category: VU D2
There are only 3 localities of this species known in our territory, all correspond to the Pyrenees: Vilallonga de Ter (Ge), Aigüestortes (L) and Monte Mendaur (Na).The values of AOO and EOO are 12 km 2 and c. 1,250 km 2 , respectively.The species grows in fissures of acidic rocks, between 1,100 and 1,800 m asl.It is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez et al., 2015) and Pyrénées-Orientales department (Hugonnot et al., 2018) although its rarity would prevent an eventual rescue effect (IUCN, 2003).Cynodontium polycarpon is categorized as VU in Spain according to IUCN (2017) recommendations (see comments under Anoectangium aestivum).
Cyrtomnium hymenophylloides (Huebener) T. Kop.Previous category: Not listed by Brugués et al. (2014).New category: VU D2 This circumpolar arctic-alpine species is known only from two distinct locations in the Pyrenees: Pineta and Serra de Cadí (Ruiz et al., 2018b), where it grows in calcareous rocky places.The values of AOO and EOO are 8 km 2 .These populations are relatively isolated, since the nearest populations are found in the Alps (Hofmann, 2013).In the population of Serra de Cadí, there are several stands so this species is not particularly scarce in the concrete locality in which it has been found.Its persistence in this area is favored by vegetative spread, since it was recorded sterile (sporophytes and male plants are rare in this species).Both Pyrenaean populations are within protected areas (National and Natural Parks).Since there is no evidence of decline in the population or in quality, extent or area of habitat in order to qualify under subcriteria b of criteria B, C. hymenophylloides is assessed as VU D2 on the basis of its restricted AOO and presence at two locations, which makes it potentially vulnerable to stochastic events.If there was any evidence of decline, this species would qualify as EN under criterion B1ab+2ab.Cyrtomnium hymenophylloides is not redlisted in Europe (ECCB, 1995), but is threatened at a regional level in some countries (Sabovljevic et al., 2009).

New category: RE
The only known locality of this species in the Iberian Peninsula corresponds to a beech forest in the region of La Garrotxa (Ge province), where this species was collected growing on rocks by C. Casas in 1952.Although this area has been subject of bryological studies (see Brugués et al., 2007) and that several specific surveys have been conducted to relocate this species, the presence of D. muehlenbeckii in the area has not been confirmed, so it is assigned to the RE category.Dicranum spadiceum J.E.Zetterst.Previous category: EN B2ab(ii,iv) (Brugués et al., 2014).New category: NT The Spanish populations are found in the Pyrenees and Cantabrian range, between Pico Tresmares and the valley of Setcases (Ge, Hu L and S provinces) (Brugués & Ruiz, 2015).The values of AOO and EOO are 32 km 2 and c. 3,570 km 2 , respectively.It grows on rocky places and subalpine scrubs, between 2,000 and 2,600 m asl.In the Midi-Pyrénées region, the occurrence data of D. spadiceum dates back to the end of the 19th century (Infante-Sánchez, 2015), while in eastern Pyrenees there are at least 12 localities of this species (Hugonnot et al., 2018), most of which are very close to the Núria and Setcases populations (Gerona province).In Catalonia there are 4 main areas in which the presence of this species is known: Restanca, Portarró d'Espot, Núria and Setcases (in these last two areas with two subpopulations in each of them).These Catalan localities are found in little frequented protected areas, and without remarkable threats for the habitat of D. spadiceum.Considering all the Spanish populations, it does not seem possible to infer or project a continuous decline in the area of occupation or the number of subpopulations, as established by the criterion "B".This species is listed as NT as it is close to qualifying for VU under criterion D2.

Encalypta affinis Hedw.
Previous category: VU D2 (Brugués et al., 2014).New category: NT It is known from few sites in the central and eastern Pyrenees in Catalonia (B, Ge and L provinces: La Molina, Comabella, Meranges, Muntanyó de Llacs, Taüll, Pla de Beret and Tuc de Crabera (Casas Sicart, 1986;Canalís & Casas, 1992;Casas et al., 2006;Ruiz et al., 2018a) and in a single Aragonese locality (Hu province): Pico de Paderna, Maladeta massif (Casas Sicart, 1986).This species grows on calcareous rocky places between 1,800 and 2,600 m asl.The values of AOO and EOO are 32 km 2 and c. 1,810 km 2 , respectively.The species has been reported from nearby areas in France: Midi-Pyrénées region (Infante Sánchez, 2015) although the possibility of any rescue from French populations is unknown.Most of the Spanish localities correspond to protected areas and without remarkable threats for the habitat of E. affinis.The number of localities (8) and AOO value preclude the assignation of this species to VU D2 (IUCN, 2012).
Eurhynchium angustirete (Broth.)T.J.Kop.Previous category: VU D2 (Brugués et al., 2014).New category: NT This species is currently known from central and eastern Pyrenees in Spain (Ge, Hu and L provinces), where it grows in humus-rich-soils and rocks, in montane or subalpine areas (between 1,100 and 1,800 m asl).At least 10 concrete localities are known, mostly Catalan except two Aragonese: Benasque (BCB 30261) and Sierra de Chía (Heras et al., 2004).The values of AOO and EOO are 40 km 2 and c. 2,490 km 2 , respectively.Eurhynchium angustirete also has populations in nearby areas: Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez, 2015) and Pyrénées-Orientales department [5 localities] (Hugonnot et al., 2018).In Spain there is no indication of a continuous population decline, and most of its range lies within protected areas.The AOO value and the number of localities in Spain greatly exceed the thresholds established for criterion D2.However, since E. angustirete has a restricted range in Spain limited to few locations on mountain areas, it is assessed as NT.
The species is assessed as VU D2 because of the small area of occupancy, small number of known locations, which makes it potentially vulnerable to stochastic events, as well as human activities (anthropic impacts can not be ruled out in Portalet locality).Moreover, G. elongata has been assessed as VU D2 in the Midi-Pyrénées region (Infante Sánchez et al., 2015) which would prevent an eventual rescue effect (IUCN, 2003), and species of the same genus and similar ecological requirements, with a similar number of known localities were regarded as VU D2 by Brugués et al. (2014).
Herzogiella striatella (Brid.)Z.Iwats.Previous category: EN B2ab(ii,iii,iv) (Brugués et al., 2014).New category: NT The Spanish populations are found in the Central Pyerenees (Hu and L provinces), between Maladeta massif (ibón Alto de Vallimuerta, MUB 27378) and Tavascan (estany Romedo Baix, MUB 45092).The values of AOO and EOO are 32 km 2 and c. 410 km 2 , respectively.It grows in wet places in subalpine forests, sometimes in rocky places, between 975 and 2,050 m asl (Guerra, 2018).It is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and French Pyrenees (Luchon, Gorges du Lys, BCB 1855) where we have verified its presence.All Spanish localities correspond to protected areas and without remarkable threats for the habitat of H. striatella.Considering all these populations, it does not seem possible to infer or project a continuous decline in the area of occupation or the number of subpopulations, as established by the criterion "Bb".This species is listed as NT as it is close to qualifying for VU under criterion D2.
Previous category: VU B2ab(ii,iv) (Brugués et al., 2014).New category: NT It is a relatively rare species in Spain.It is known from several localities in the central Pyrenees (Hu and L provinces) where it grows on the rocks in or beside streams, between 1,850 and 2,490 m asl (Oliván & Fuertes, 2018).The values of AOO and EOO are 52 km 2 and c. 1,310 km 2 , respectively.It is also known from nearby areas in France: Midi-Pyrénées region (Infante Sánchez et al., 2015) and Pyrénées-Orientales department [6 localities] (Hugonnot et al., 2018).Most of the Spanish localities correspond to protected areas (a National Park and two Natural Parks).The available information suggests that the area of the species, in spite of being relatively restricted, is not severely fragmented and far exceeds the number of 10 localities (subcriterion "a").On the other hand, a decline in the area of occupation or the number of subpopulations cannot be inferred.According to Oliván et al. (2007), H. smithii cannot be considered as a vulnerable species in the Iberian Peninsula since it only agrees with the first part of criterion B as defined by Hallingbäck et al. (1998).Moreover, Oliván et al. (2007) observed that Pyrenaean populations are large and usually with sporophytes, which suggests a healthy condition.Although rare in Spain, H. smithii, no longer meets the criteria for being regionally threatened and is hence listed as NT.

Orthotrichum philibertii Venturi
Previous category: VU D2 (Brugués et al., 2014).New category: LC This montane Mediterranean-oceanic species exists in 14 provinces according to Lara & Garilleti (2014).Neither data related to its distribution in Spain, nor those related to its ecology (it grows on a remarkable number of phorophytes, in a wide altitudinal range, colonizing even artificial substrates such as fiber-cement siding) suggest that this species can accommodate VU on the basis of criterion D2.We assign it to LC since it does not even seem to be considered as NT according to IUCN criteria (2012).

Orthotrichum rogeri Brid.
Previous category: Not listed by Brugués et al. (2014).New category: VU D2 Surprisingly, this rare epiphyte was not included in the Spanish Red List (Brugués et al., 2014) although it is worthy of being assigned to a threat category IUCN (2012) in our territory, and it is also included in European legislation (Annex II of the Habitats Directive).At least 4 localities of this species in the Iberian Peninsula are known, all relatively recent and corresponding to the axial Pyrenees (Hu and L provinces): Bujaruelo, port de la Bonaigua, Estany de Sant Maurici y Parque Natural de Posets-Maladeta (Garilleti et al., 2002;Infante & Heras 2012: 237).A report from Montgarri (Venturi, 1887) is excluded, since it is referable to O. stramineum Brid.(Garilleti et al., 2002).There is a recent record (year 2005) of O. rogeri from Sierra del Cadí-Moixeró (BDBC), and in case it is confirmed it would expand its Iberian distribution to the eastern Pyrenees.According to Infante Sánchez (2015), this species is found in the Ariège and Hautes-Pyrénées departments (recent observations, 2004 and 2014).On the other hand, Infante Sánchez et al. (2015) do not consider O. rogeri a threatened species, and assessed it as NT.Similarly, O. rogeri has a significant number of populations (around 20) in Pyrénées-Orientales department, where it is in a clearly satisfactory situation (Hugonnot et al., 2018).There is currently no evidence of decline in Spain, although its geographic restriction (AOO=16 km 2 , EOO=c.395 km 2 ) and low number of localities makes it vulnerable to stochastic events.

Orthotrichum scanicum Gronvall
Previous category: VU D2 (Brugués et al., 2014).New category: LC It is a mainly sub-Mediterranean species, present in the Pyrenees, Galicia, Iberian System, reaching even the Sierra de la Alhamilla (Medina et al., 2008).Similarly to O. philibertii, it has a wide distribution, since it is found at least in 11 provinces (Lara & Garilleti, 2014).Moreover, there are no indications that the subpopulations are especially vulnerable to the effects of human activity, at least in the medium or short term, such as to consider CR or disappear (at the regional level), as established by IUCN (2012).By not satisfying criteria that allow this species to be included in a risk category, it must be considered LC.
Pohlia andalusica (Höhn.)Broth.Previous category: VU D2 (Brugués et al., 2014).New category: NT According to Guerra (2010), this species is found in our area in the Pyrenees and Sierra Nevada (Gr, Hu, L and Na provinces).Its presence has been documented of at least 16 UTM 1x1 km squares (AOO = 60 km 2 , EOO = c.97,450 km 2 ).At least 5 populations are found in Sierra Nevada located in high altitude areas, above 2,860 m (Rams et al., 2014) and in the Pyrenees, where the altitudinal gradient is much wider, and correspond to well conserved areas.Both, the number of localities and the value of AOO far exceed the geographical restriction threshold established for VU D2.In the absence of evidence regarding a continuous decrease in AOO or the disappearance of a significant number of subpopulations, P. andalusica is assigned to NT.
Polytrichastrum sexangulare (Brid.)G.L.Sm.Previous category: VU D2 (Brugués et al., 2014); LC (Luceño et al., 2017).New category: LC The Spanish populations of this chionophile species are found in the Pyrenees and the Central System (Gredos) (Av, Ge, Hu and L provinces).Excluding the Andorran localities, this species has a total of 30 localities (AOO = 104 km 2 , EOO = 40,300 km 2 ).In addition, P. sexangulare grows in a wide altitudinal range, between 2,100 and 3,100 m asl, so the effects of an eventual loss of potential habitat related to changes in the climate would not be immediate.The available data far exceed the thresholds established for D2 criterion, so this species is assigned to LC.
Pseudoleskeella rupestris (Berggr.)Hedenäs & L. Söderstr.Previous category: DD (Brugués et al., 2014).New category: VU D2 This saxicolous species is extremely rare in the Iberian Peninsula.It seems to be restricted to the central Pyrenees: Ordesa and Escarrilla (Hu province) and Loseron forest, Valle de Arán (L province), where it grows in calcareous rocky places, between 1,120 and 1,600 m asl (Brugués & Ruiz, 2018).The collections are relatively recent (end of the last century) except in the case of the collected by Allorge in Escarrilla (PC 002592).In addition, there is a small population near the top of Puig Major in Majorca (cover c. 10 cm 2 ) (Sáez & Brugués, 2018).The values of AOO and EOO are 16 km 2 and c. 11,620 km 2 , respectively.On the basis of its strong geographical restriction in our territory which makes it potentially vulnerable to stochastic events, as well as human activities and encroatchment of habitat by other bryophytes (in the case of the Balearic subpopulation), it is advisable to assimilate P. rupestris to VU D2.Pseudostereodon procerrimus (Molendo) M.Fleisch.Previous category: VU D2 (Brugués et al., 2014, sub Hypnum procerrimum Molendo).New category: NT According to Ríos et al. (2018), this species is distributed through the Pyrenees, the Cantabrian range and mountains of Guipúzcoa (Aitzgorri) in the provinces Hu, L, Lo, O, S and SS.It grows in forests and meadows in a wide altitudinal range (530-2,500 m asl).Ríos et al. (2018) also listed this species for Cu province based on a bibliographic reference.At least 14 localities are known (AOO = 56 km 2 , EOO = c.33,500 km 2 ), which greatly exceeds the threshold established for VU D2.Considering the available corological data and the fact that there is no evidence of a continuous decline in the occupation area of P. procerrimus or disappearance of a significant number of subpopulations, it is assessed as NT.
Ptilium crista-castrensis (Hedw.)De Not.Previous category: EN D ["D2"] (Brugués et al., 2014).New category: VU D2 It is a rare species in Spain (Ge, L and Na provinces), known from 6 Pyrenean localities (AOO = 20 km 2 , EOO = c.2,560 km 2 ).These populations are found between the Port of Eraize (Montserrat et al., 1988) and Ter valley (Lloret, 1989) and mostly correspond to protected areas.The subpopulations are found in humid places in montane forests, between 1,100 and 1,600 m asl (Gallego, 2018).The supposed isolation of Spanish populations is scarce: at least one Aranese population (Portilló de Bossost) is connected with those of the French Pyrenees (populations located in Val d'Astau, L. Sáez pers.obs.) and with the adjacent Bois Neuf (L.Sáez pers.obs.).P. crista-castrensis is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez et al., 2015) -where it was assessed as LC-and Pyrénées-Orientales department [3 localities] (Hugonnot et al., 2018).The attribution to EN based on criterion D (population size less than 250 individuals) is difficult to maintain since in some of the localities P. crista-castrensis occurs in the form of scattered stands.In fact, this criterion is rarely used solely to assign a species of bryophyte to a specific IUCN (2012) category.There is currently no evidence of decline in Spain, although its geographic restriction and low number of localities makes it vulnerable to stochastic events as well as forest management activities.
Sarmentypnum sarmentosum (Wahlenb.)Tuom.& T.J. Kop.Previous category: DD (Brugués et al., 2014).New category: EN B2ab(iii) It is a very rare species in Spain, known from 4 Cantabrian-Pyrenean localities (Ge, Hu, Na and P provinces) (AOO = 16 km 2 , EOO = c.10,130 km 2 ) where it grows in permanently wet and oligotrophic places, between 1,000 and 1,850 m asl (Brugués & Ruiz, 2003;Fuertes et al., 2006Fuertes et al., , 2010)).The localities that are based on collections after 1970 correspond to the Curavacas area (P province) and the Estanys de la Pera (Ge province).The locality from Huesca province is based on a collection of Fourcade in 1866 (Brugués & Ruiz, 2003), whereas the label of a specimen from Roncesvalles (MA 12657) does not include date or collector name.The species is also very rare in the Pyrénées-Orientales department (Hugonnot et al., 2018) and the Midi-Pyrénées region where it has been assessed as EN (Infante Sánchez et al., 2015).Sarmentypnum sarmentosum has an extremely restricted range in our area and it is especially sensitive to environmental changes.The species is assessed as EN because of the small area of occupancy, small number of known locations.A population decline is inferred due to modifications of the water regime induced by human activity and natural causes.

Schistidium papillosum Culm.
Previous category: DD (Brugués et al., 2014).New category: LC This saxicoulous species is found in mountain areas in northern Spain (Ge, Gu, Hu, L, Le, O and S provinces;Suárez & Muñoz, 2015).It is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez, 2015) and Pyrénées-Orientales department [5 localities] (Hugonnot et al., 2018).Almost twenty localities (AOO = 64 km 2 ) are known solely in Ge and L provinces that mostly correspond to protected areas (one National park and several Regional natural parks), none subject to any problem to its conservation.Since it is not a rare species and that most populations are located in protected areas it is assessed as LC.
Sciuro-hypnum curtum (Lindb.)Ignatov Previous category: DD (Brugués et al., 2014).New category: VU D2 In the Iberian Peninsula this species is known from four localities that correspond to the Cantabrian Range and the central Pyrenees (Hu, L, P and S provinces) (Orgaz et al., 2012;Orgaz 2018).The values of AOO and EOO are 16 km 2 and c. 2,600 km 2 , respectively.Apparently the isolation of these Iberian populations is important, since the presence of this species in the French Pyrenees and in Andorra is unknown (Orgaz et al., 2011;Infante Sánchez, 2015;Hugonnot et al., 2018;Sotiaux & Vanderpoorten, 2017).The collections are recent except in the case of the locality of La Renclusa, dating from 1966, where we can not rule out direct anthropic impacts, related to works carried out (after 1966) in this area.On the other hand, trampling by cattle and habitat changes can not be ruled out in these Spanish localities.With the data available, it is somewhat risky to apply criterion B2ab(iii,iv), both for EN and for VU, since it is not possible to infer a continuous population decrease in view of the habitats it occupies (subalpine and alpine meadows and snowbeds, between 1,900 and 2,100 m).The species is here assessed as VU D2 given its rarity in our territory (small area of occupancy, small number of known locations) which makes it potentially vulnerable to stochastic events, as well as human activities.Sciuro-hypnum glaciale (Schimp.)Ignatov & Huttunen Previous category: VU D2 (Brugués et al., 2014).New category: NT According to Orgaz et al. (2011) and Orgaz (2018), this species is distributed in Spain along the Pyrenean-Cantabrian ranges, with one population in Sierra Nevada (Ge, Gr, Hu, L, S and P provinces).It grows in humid rocky places and snowbeds between 1,600 and 3,000 m asl.At least 14 localities are known (AOO = 56 km 2 , EOO = c.192,770 km 2 ).The number of localities and the AOO value far exceed the threshold established for VU D2.However, S. glaciale has a restricted range, limited to few locations on mountain ridges, it is here assessed as NT.

Sphagnum warnstorfii Russow
Previous category: VU D1 (Brugués et al., 2014).New category: NT According to Brugués et al. (2016b) and BDBC, there are c.20 localities of this species are known in Spain which are found in the Pyrenees, Ge, Hu and L provinces (AOO = 64 km 2 , EOO = c.2,700 km 2 ).In our area there is no indication of a continuous population decline, and most of its range lies within protected areas.Sphagnum warnstorfii was assigned to VU D1, which establishes that the estimated population is less than 1,000 mature individuals.The delimitation of individuals is complex in Sphagnum, but considering the remarkable number of localities and subpopulations, it is unlikely that less than 1,000 individuals exist.Until further data based on field research can be gathered, it is assessed as NT.
Stegonia latifolia (Schwägr.)Venturi ex Broth.Previous category: VU B2ab(ii,iv) (Brugués et al., 2014).New category: NT This is a rare species in Spain.It is known from few sites in the central Pyrenees (L province: Montsent de Pallars, Pla de Baqueria, Coll de Cabús and Roca Blanca) and the Cantabrian Mountains (Bu and O provinces: macizo de Castro Valnera, Peña Ubiña and alto de la Farrapona) (Cano et al., 2017), where it occurs on calcareous rocky places across a wide altitudinal range (between 1,445 and 2,647 m asl).The values of AOO and EOO are 32 km 2 and c. 15,995 km 2 , respectively.This species is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez et al., 2015) and Pyrénées-Orientales department [5 localities] (Hugonnot et al., 2018).The available data do not suggest a continuous decline.On the other hand, the number of localities and AOO value preclude the assignation of this species to VU D2 (IUCN, 2012), thus it is assessed as NT.
Stereodon callichrous (Brid.)Braithw.Previous category: VU B2ab(ii,iv) (Brugués et al., 2014, sub Hypnum callichroum Brid.).New category: NT The Spanish populations of this species are found in the central and eastern Pyrenees (Ge, Hu and L provinces), between the Pico Crabioules and Puigmal (Ruiz & Brugués, 2011).It grows on rocks, soil and bark between 1,500 and 2,200 m (Ríos et al., 2018).In total, 13 localities are known (AOO = 52 km 2 , EOO = c.1,510 km 2 ).Some records (especially those corresponding to Aragon region) are old and correspond to high mountain areas that are in good condition with very little anthropogenic disturbances and that in some cases have not been prospected in recent times.The Catalan subpopulations are found in 9 UTM squares 1x1 km (EOO= c. 1,330 km 2 ; AOO = 36 km 2 ) and all of them correspond to protected areas (a National park and two Natural parks).This species also exists in nearby localities of Andorra (Ruiz & Brugués, 2011;Sotiaux & Vanderpoorten, 2017) and the French Pyrenees (Ariège, Haute Garonne, Hautes-Pyrénées) (Infante Sánchez, 2015).This species is assigned to NT instead of LC following the same reasoning as exposed for Hygrohypnum smithii.
Syntrichia sinensis (Müll.Hal.)Ochyra Previous category: VU D2 (Brugués et al., 2014).New category: NT In the Iberian Peninsula this species is known from two main areas in Eastern Pyrenees (B and Ge provinces): Castellar de n'Hug and Alta Vall del Ter (Lloret Maya, 1986;Lloret, 1989;Gallego, 2005;Ros & Werner, 2006).The values of AOO and EOO are 28 km 2 and c. 80 km 2 , respectively.The known populations are found on shaded siliceous rocky places, between 1,100 and 2,300 m asl.At least two populations are also known from nearby areas in the Pyrénées-Orientales department (Hugonnot et al., 2018).Although S. sinensis has a relatively restricted range in Spain, limited to a few locations on mountain areas, it is unlikely to be declining markedly.This species is therefore here listed as NT, although it is close to qualifying for VU under criterion D2.

Timmia austriaca Hedw.
Previous category: VU D2 (Brugués et al., 2014).New category: LC It is distributed through the Cantabrian mountain range and Pyrenees (B, Ge, Hu, L and O provinces) where it grows in rocky places, slopes and woods (Álvaro, 2010;Ruiz et al., 2018a) throughout a wide altitudinal range (120-2,495 m).At least 24 localities are currently known (AOO = 92 km 2 , EOO = c.15,000 km 2 ) [data from Manobens i Rigol (1984, 1985), Cros et al. (2010), Ruiz et al. (2018a), BCB and BCN], most of them located in protected areas (National and Natural Parks).In the Pedraforca massif and Serra de Cadí (B and L provinces) T. austriaca is not particularly scarce in almost inaccessible areas.In eastern French Pyrenees there are at least 19 localities for the species (Hugonnot et al., 2018), whereas in the Midi-Pyrénées region it was regarded as "DD" (Infante-Sánchez et al., 2015).The value of AOO and the number of localities greatly exceed the threshold established for criterion D2.
On the other hand, there is no evidence to indicate that T. austriaca is especially vulnerable to anthropogenic disturbances or to fortuitous events in a short period of time in an uncertain future to be regarded as CR or EX categories.It is thus here assigned to LC.
Tortula viridifolia (Mitt.)Blockeel & A.J.E.Sm.Previous category: VU D2 (Brugués et al., 2014).New category: NT According to Ros et al. (2011), this species is restricted in Spain to a few localities in the sourheastern Iberian Peninsula (Al, Gr and Mu provinces, corresponding to 4 UTM squares 10x10 km).Based on this restricted range, it was assigned to VU D2 by Brugués et al. (2014).The species also exists in 3 localities in the northeastern Iberian Peninsula (Ge province) and also in the Balearic Islands (2 localities in Minorca; Brugués & Ros, 2017).A Majorcan report (Sáez et al., 2002) accepted by Brugués & Ros (2017) is due to confusion with T. pallida (Lindb.)R.H. Zander, whereas reports from M and Po provinces require verification.The confirmed Spanish populations of T. viridifolia are found across a wide altitudinal range, from sea level to 1500 m.The values of AOO and EOO are 36 km 2 and c. 102,400 km 2 , respectively.The number of localities and AOO value precludes assigning this species to VU D2 (IUCN, 2012).It has some potential threats (see Ros et al., 2011), but these are not thought to be severe at the moment, so the species is unlikely to be declining markedly.It is to be therefore listed as NT.
Weissia wimmeriana (Sendtn.)Bruch & Schimp.Previous category: VU D2 (Brugués et al., 2014).New category: NT It is a relatively rare species restricted to high mountains or regions with cold climates (Werner et al., 2004).In mainland Spain W. wimmeriana is known from around 20 mountain localities from the Pyrenees and Sierra Nevada (Ge, Gr, Hu and L provinces) where it grows in rock crevices and slopes (Guerra, 2006c).AOO and EOO values are 48 km 2 and c. 57,000 km 2 , respectively (data from BCB and Rams et al., 2014).It is also known from nearby areas in Andorra (Sotiaux & Vanderpoorten, 2017) and France: Midi-Pyrénées region (Infante Sánchez et al., 2015) and Pyrénées-Orientales department [5 localities] (Hugonnot et al., 2018).In Spain there is no indication of a continuous population decline, and most of its range lies within protected areas.The number of localities and AOO value precludes assigning this species to VU D2 (IUCN, 2012).However, since the species has a relatively restricted range in mainland Spain, it is assessed as NT.

Discussion
The review of the information available for the species studied, together with new data on their distribution and relative frequency in the territory considered, led to establish new IUCN categories for 48 taxa.Four species that were not included in Brugués et al. (2014) red list have been identified and subsequently assessed as VU, while an additional species has been assessed as regionally extinct.Given that this study is mainly focused on species present in northeastern Iberian Peninsula, it is advisable to review the Spanish list of bryophytes in order to extend the current red list further as to correct for possible absences.
Regarding category changes -taxa assessed by Brugués et al. (2014) as threatened [CR, EN and VU]our results indicate that downlisting (moving a species to a category of lower threat) is more frequent than uplisting.The vast majority of the taxa studied (31 of a total of 33, excluding Gymnomitrium coralloides, to be excluded from the Spanish bryoflora) see an improvement in their situation by qualifying for a lower IUCN (2012) category.Specifically, of the 31 taxa downlisted, 25 no longer qualify for risk categories (that is, they pass to NT or LC).This is in accordance to a general trend of downlisting species when information about its distribution improves.In other cases (i.e., Orthotrichum philibertii, O. scanicum) the fact that the taxa can no longer be assigned to a risk category is not due to better corological data available, but rather to their initial assignment to a risk category that is questionable and needed to be reconsidered.
In contrast, of the 10 taxa presented here that were regarded as DD by Brugués et al. (2014), all but one (Schistidium papillosum) have been found to be threatened species according to IUCN (2012) criteria: 6 VU, 1 EN and 2 CR.This highlights the need to review in greater detail any taxon before it is designated as DD, which may have been the problem with some in Brugués et al. (2014), especially since DD makes up the largest group (82 taxa) in that red list.Species assessed as DD are very likely to be threatened, like Scorpidium revolvens (Sw.ex anon.)Rubers and Warnstorfia fluitans (Hedw.)Loeske.Both mosses are extremely rare in Spain (Canalís et al., 1984;Fuertes et al., 2006Fuertes et al., , 2010)), restricted to few localities and are especially sensitive to anthropic disturbances.They can confidently be assessed as CR or EN under criteria B2ab.On the other hand, included within the DD group are also species of obvious wide distribution in Spain, such as Gyroweissia tenuis (Hedw.)Schimp., wich is listed from 14 Spanish provinces (Guerra, 2006b).
The classification protocol provided by IUCN (2012, 2017) is a simple and flexible method for determining species' vulnerability to extinction.Accurate and standardized conservation status assessments for the IUCN Red List are limited by a lack of adequate information, so there is a need for consistent and unbiased interpretation of that information (Hayward et al., 2015).Although the IUCN has developed a set of detailed guidelines for using categories and criteria, biases and incorrect applications of the protocols (or mistakes interpreting thresholds and criteria) are not uncommon (Regan et al., 2005, Hayward et al., 2015).The result of these evaluations usually translates into an overrepresentation of taxa in red lists, but also in a higher attribution of the risk status, unrealistically superior to what a taxon may actually has.Although data referring to a species' population dynamics are seldom available for applying criterion B correctly (although IUCN precedures allow for establishing inferences), it is nonetheless widely used despite no clear evidence of population decline.Yet precision of assessments remains a non-trivial problem because, among other things, the criteria provide no real guidance for the use of inference (Regan et al., 2005).
Another source of inaccurate assessments is the inadequate use of the VU D2 criterion.IUCN (2017: 69) emphasizes that the restricted AOO (typically < 20 km 2 ) -or five or less locations-is to be applied to species whose populations are prone to effects of human activities or to stochastic events in an uncertain future, and the species is thus subject to become Critically Endangered or even Extinct in a very short time (e.g., within one or two generations after the threating event occurs).Much of category changes proposed in this study correspond to improper use of the criterion D2.
Finally, another aspect that may cause deviations is to assign risk categories to taxa that, strictly, do not meet the minimum objective criteria for being granted any risk category according to IUCN (2012).Such inaccurate assessments may be explained because these taxa are under some kind of legal protection.This could be the case with species such as Petalophyllum ralfsii and Buxbaumia viridis, which have become protected by various European and Spanish regulations.Reviewing available data for these species shows that the "Bb" subcriterion [small range (EOO and/or AOO) and continuing decline] is not applicable in these and several other cases, at least not in the short-medium term.Perhaps this legal protection does not favor a critical re-evaluation of the available data that will likely lead to their removal from red lists.Higher attribution of the risk situation than a species merits could actually be very detrimental to conservation because it distracts from more serious conservation problems.Attention is drawn in the wrong direction and resources are diverted away from species that are genuinely at risk of extinction, while favouring species that are not.Unwarranted uplisting of species is considered for these reasons counterproductive to the general goals of biodiversity conservation (Webb, 2000(Webb, , 2008)).

Figure 2 .
Figure 2. Distribution area of Petalophyllum ralfsii in the Balearic Islands by UTM 2 x 2 km squares.

Figure 3 .
Figure 3.Total cumulative area of occupancy (AOO) by decade for Petalophyllum ralfsii in mainland Spain and Balearic Islands.