The Case of Ferns Flora of the Gibraltar Arc

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Botanica Complutensis ISSN-e 1988-2874

https://dx.doi.org/10.5209/bocm.75454

The historical biogeography and conservation value of taxonomic distinctness: The case of ferns flora of the Gibraltar Arc

Ángel Enrique Salvo Tierra1; José C. Báez2; Antonio Flores-Moya3

Abstract. The pteridofloras of nine locations in the Gibraltar Arc were analyzed using a taxonomic distinctness index. We found that the index could be a proxy of historical biogeography of the pteridofloras from this area. Moreover, the value of the taxonomic distinctness index of the different locations showed relevant relationships with certain geographic variables. Finally, we hypothesize about the value of the information derived from taxonomic distinctness index for conservation of the pteridoflora in the GibraltarArc. Keywords: Pteridophytes, Western Mediterranean, Historical biogeography, Conservation

[es] Biogeografía histórica y valor conservacionista de la diferenciación taxonómica: ejemplo de la pteriflora de Arco de Gibraltar

Resumen. La pteridoflora de nueve localidades singulares situadas en el Arco de Gibraltar ha sido analizada utilizando un índice de distinción taxonómica. Encontramos que el índice podría ser una aproximación a la historia biogeográfica de las pteridofloras de esta zona. Además, el valor del índice de distinción taxonómica de las diferentes localidades mostró relaciones relevantes con determinadas variables geográficas. Finalmente, discutimos el posible papel de la información derivada del índice de distinción taxonómica en la conservación de la pteridoflora del Arco de Gibraltar. Palabras clave: Pteridofitas, Mediterráneo Occidental, Biogeografía histórica, Conservación

Introduction rica, and eleven chorotypes. Finally, considering the diversity, age, singularity, and degree of threat to its The fern flora of the Gibraltar Arc, i.e., the geolog- pteridoflora, it was proposed that the Gibraltar Arc ical region corresponding to an arcuate orogen sur- could be considered as a ‘Sanctuary for ferns’, an in- rounding the Alboran Sea, extending from the Iberi- creasingly widespread conservation concept. an Peninsula to Africa, has been extensively studied Taxonomic structure (based on the number of taxa recently by Salvo Tierra et al. (2020). These authors included in higher levels of the taxonomic hierarchy) suggested that the factors influencing diversity in has been widely used in ecology to measure impacts fern floras of nine different mountain areas is so rele- on the diversity of ecosystems (for example, Clarke vant that they should be included in the management & Warwick 1998, 1999, 2001, Tucker et al. 2017, of the territory. They concluded that the altitudinal Pérez Hernández 2019). Taxonomic structure shows gradient and the amplitude of the dry period have de- a latitudinal gradient and is maximum at the Equator termined the distribution of pteridophytes in the Gi- (Krug et al. 2008). Among taxonomic richness esti- braltar Arc, a distribution in which the longitudinal mators, taxonomic distinctness (TD; Pérez Hernán- gradient predominates over the latitudinal gradient. dez 2019, and references therein) is particularly sig- In particular, the oldest fern floras (with greater di- nificant because it appears to be less influenced by versity) are located at the western ends of the arc on sample size than are species richness or Shannon spe- both continents. Five pterido-geographic units were cies diversity. Moreover, TD may be a more sensitive detected, two of them spanning S Europe and N Af- univariate index of community perturbation than spe-

1 Department of Botany and Plant Physiology, Faculty of Sciences, University of Málaga, Campus de Teatinos. Bulevar Louis Pasteur s/n 29010 Málaga. Spain. Corresponding author. Contributions: Conceptualization, Methodology, Formal analysis, Investigation, Writing – Original Draft. E-mail: [email protected] ORCID: https://orcid.org/0000-0002-1464-9770 2 Instituto Español de Oceanografía, Centro Oceanográfico de Málaga (sede Fuengirola), 29640 Fuengirola. España. Instituto Iberoamericano de Desarrollo Sostenible, Universidad Autónoma de Chile, Temuco, Chile. Contributions: Conceptualization, Methodology, Formal analysis, Investigation, Writing – Original Draft. E-mail: [email protected] ORCID: https://orcid.org/0000-0003-2049-0409 3 Department of Botany and Plant Physiology, Faculty of Sciences, University of Málaga. Campus de Teatinos. Bulevar Louis Pasteur s/n 29010 Málaga. Spain. Contributions: Conceptualization, Methodology, Formal analysis, Investigation, Writing – Original Draft. E-mail: [email protected] ORCID: https://orcid.org/0000-0002-0940-4541

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cies diversity (Pérez Hernández 2019, and referenc- In addition, in a second step, linkage cluster anal- es therein). Furthermore, the statistical signifi cance ysis of the ∆+ index per locality was performed using of the TD index can be tested (Clarke & Warwick a multivariate agglomerative method. Thus, we used 2001, Pérez Hernández 2019). Clarke & Warwick the Bray-Curtis distance to generate the similarity (1998) proposed the ∆+ index (i.e., Delta-plus) to es- matrix, using the free PAST version 4.02 software timate taxonomic distinctness. This index measures (Hammer et al. 2001). We chose Bray-Curtis distance the overall mean taxonomic path length between any to generate the similarity matrix, because it is espe- two randomly chosen taxa. Because the index has no cially recommended for standardized data (Yoshioka dependence on sampling effort, the ∆+ fi gures com- 2008) (Fig. 2). puted from different sources of data can be compared For each OGU, the values of latitude, longitude, across studies with differing and uncontrolled de- maximum altitude, TD and TD range values (the dif- grees of sampling effort. ference between the upper and lower limits), specifi c The main aim of the current note is to analyze the richness, index m/t (Ito 1972, 1978) and degree of TD of fern fl oras from the Gibraltar Arc. The goal is ploidy, as well as the percentages of the presence of to assess the differences in biogeographical composi- each of the generalized tracks detected in the study tion on opposite shores of the Alboran Sea. area, were included in the analysis (Salvo Tierra et al. 2020).

Material and Methods Results and Discussion Data origin The ∆+ index for each region (i.e., Tingitania, The determination of the OGUs (operative geograph- Chauen, Gurugú, Tlemcen, Aljibe, Ronda, Tejeda, ical units) was carried out according to criteria of ge- Nevada, and Gata) was estimated. We did not ob- ophysical homogeneity and in correspondence with serve signifi cant differences according to the criteria the biogeographic units, according to Salvo Tierra et of the index. However, we observed three different al. (2020). Nine mountain areas located in the Gibral- and coherent similar groups according their ∆+ va- tar Arc were selected, one for each biogeographical lues (Fig. 1). Because the signifi cance of the index sector; the areas include the entire range of thermo- was developed under other criteria (in relation to the and ombrotypes found in these regions (Rivas Mar- number of sampling stations, and the number of spe- tinez 2008): Gata, Nevada, Tejeda, Ronda, Aljibe, cies) (Clarke & Warwick 1998, 1999, 2001), we con- Tingitania, Chaouen, Gurugú and Tlemcen (Fig. 1). sider that they may not be useful here. The basic matrix of data is showed in the supplemen- From Table 1 we conclude: 1) There is a strong tary data (metadata available in Salvo Tierra et al., inverse correlation between the value of TD and the 2020). maximum altitude (the lower the altitude, the greater the distinctness), as well as with the geographical Data Analysis longitude (decreasing from west to east) and with the index m/t. 2) From this last analysis, it can be inferred The ∆+ index is computed according to the equation: that there is a correspondence between higher values of TD and the older pteridofl oras (the proportion of taxa with trilete spores = primitive character, is high compared to those with monolete spores = advanced character). 3) This same observation is corroborated on the one hand by analysing the correspondence between TD and the degree of ploidy (the lowest values of the degree of ploidy correspond to the most primitive pteridofl oras), and on the other by observing

where s is the number of species observed, and wij that the lowest values of TD are correlated with those is the weight or distinctiveness value given to each pteridofl oras in which the generalized tracks of wider taxonomic branch of the hierarchical classifi cation, distributions (heterochoric and circumboreal) have a from species i to the fi rst node in common with spe- greater weight. 4) Consequently, the highest values

cies j. Thus, wij = 0 if i and j are the same genus, wij = of TD correspond to the most relict pteridofl oras, 1 if they are the same family. although they arose from different historical events, We calculated the ∆+ index for each region (i.e., and are scarcely ‘contaminated’ by species with wid- Tingitania, Chauen, Gurugú, Tlemcen, Aljibe, Ron- er geographical ranges that were incorporated into da, Tejeda, Nevada, and Gata) using the free PAST the territory during subsequent events. 5) Finally, TD software (Hammer et al. 2001). Confi dence intervals has a complementary signifi cance in the defi nition of were calculated from 1000 random replicates taken priority conservation areas. In this sense, the lower from the pooled data set (Hammer 2020). values in the range of the upper and lower limits of TD for Aljibe and Nevada indicate a more ancestral

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and consolidated flora, which would support a high Krug, A.Z., Jablonski, D., & Valentin, J.W. (2008). Spe- priority as regions for the conservation of pterido- cies-genus ratios reflect a global history of diversifica- phytes. This is confirmed when applying the Vane- tion and range expansion in marine bivalves. Proceed- Wright et al. (1993) these two units are defined in the ings of the Royal Socociety B, 275, 1117-1123. https:// first cycle on the application method. doi.org/10.1098/rspb.2007.1729. Pérez Hernández, C.X. (2019). Distintividad taxonómi- ca: Evaluación de la diversidad en la estructura tax- Acknowledgements onómica en los ensambles. In C.E. Moreno (Ed) La biodiversidad en un mundo cambiante: Fundamentos Dr. Eric C. Henry kindly revised the English style teóricos y metodológicos para su estudio. Universidad and usage. Autónoma del Estado de Hidalgo/Libermex, Ciudad de México, pp. 285-306. Pielou, E.C. (1975). Ecological diversity. Wiley, New York. References 165 pp. https://doi.org/10.4319/lo.1977.22.1.0174b PPG I (2016). A community-derived classification for ex- Clarke, K. R. & Warwick, R.M. (1998). A taxonomic tant lycophytes and ferns. Journal of Systematics and distinctness index and its statistical properties. Jour- Evolution, 54, 563–603. doi: https://doi.org/10.1111/ nal of Applied Ecology 35 (4), 523-531. https://doi. jse.12229 org/10.1046/j.1365-2664.1998.3540523.x Rivas Martínez, S. (2004). Global Bioclimatics (Cla- Clarke, K. R. & Warwick, R.M. (1999). The taxonomic sificación Bioclimática de la Tierra). https://www. distinctness measure of biodiversity: weighting of step globalbioclimatics.org/book/bioc/global_bioclimat- lengths between hierarchical levels. Marine Ecology ics-2008_00.htm Progress Series 184, 21-29. https://www.jstor.org/sta- Salvo Tierra, Á.E., Melero-Jiménez, I.J., García-Sánchez, ble/24853226 J., Báez, J.C., & Flores-Moya, A. (2020). Biogeo- Clarke, K. R. & Warwick, R.M. (2001). A further biodiver- grafía analítica de la pteridoflora del arco de alborán: sity index applicable to species lists: variation in tax- consecuencias para su status de protección. Acta onomic distinctness. Marine Ecology Progress Series Botánica Malacitana, 43: 1-19 and Atlas. https://doi. 216, 265-278. doi:10.3354/meps216265 org/10.24310/Actabotanicaabmabm.v45i.7743. Given, D.R. (1993). Changing Aspects of Endemism and Tucker, C.M., Cadotte, M.W., Carvalho, S.B., Davies, J., Endangerment in Pteridophyta. Journal of Bioge- Ferrier, S., Fritz, S.A., Grenyer, R., Helmus, M.R., Jin, ography, 20 (3), 293-302. https://www.jstor.org/sta- L.S., Mooers, A.O., Pavoine, S., Purschke, O., Red- ble/2845638 ding, D.W., Rosauer, D.F., Winter, M. & Mazel, F. Hammer, Ø. (2020). PAST manual version 4.02. Available (2017). A guide to phylogenetic metrics for conserva- from website: https://folk.uio.no/ohammer/past/ tion, community ecology and macroecology. Biologi- Hammer, Ø., Harper, D.A.T. & Ryan, P.D. (2001). PAST: cal reviews of the Cambridge Philosophycal Society Paleontological statistics software package for edu- 92(2), 698- 715. DOI: 10.1111/brv.12252 cation and data analysis. Palaeontologia Electronica Vane-Wright, R. I., C. J. Humphries & P. H. Williams. 4(1), 1-9. (1991). What to protect?- Systematics and the agony of IOTC (2019). Report of the 21st Session of the IOTC choice. Biological Conservation 55, 235-254. https:// Working Party on Tropical Tunas. Seychelles, 21 - 26 doi.org/10.1016/0006-3207(91)90030-D October 2019. IOTC–2019–WPTT21–R[E]. Warwick, R.M. & Clarke, K.R. (1995). New ‘biodiversity’ Ito, H. (1972). Distribution of monolete and trilete ferns in measures reveal a decrease in taxonomic distinctness eastern Asia and Northern Oceania. The Journal Japa- with increasing stress. Marine Ecology Progress Series nese Botany 37,455-462. 129, 301-305. https://doi.org/10.3354/meps129301. Ito, H. (1978). Distribution of two spore patterns in ferns Yoshioka, P.M. (2008). Misidentification of the Bray-Cur- floras of the world (A preliminary survey).The Journal tis similarity index. Marine Ecology Progress Series Japanese Botany 47, 321-326. 368, 309–310.

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Figure 1.- Location of the study area and the operating geographic units (OGU) The ∆+ index plotted for each locality, ordered from lowest to highest according to their range of distinction (difference between the upper and lower limit) and identifi cation according to the Pteridogeographic Units of Salvo Tierra et al. (2020).

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Figure 2.- Dendrogram of the OGUs in the Alboran Arc carried out with UPGMA based on the similarity calculated with the similarity index of Bray & Curtis (Correlation coefficient 88.8%; Phenon line at 88%) (PU1:Aljíbica Unit, PU 2: Tingitania Unit, PU 3. Chauen-Ronda Unit, PU 4. Tejeda-Nevada Unit, PU 5. Gurugú-Gata-Tlemcen Unit)

CUARTAS_BotanicaComplutensis45.indd 5 25/5/21 21:01 6 Salvo Tierra, A. E.; Báez, José C.; Flores-Moya, A. Bot. complut. 45, 2021: 1-11 7,69 3,03 7,50 2,17 4,35 4,88 10,00 13,64 13,04 REPAME 7,69 3,03 5,00 4,35 4,55 4,35 4,35 10,00 12,20 ENMED 26,92 13,33 15,15 20,00 17,39 22,73 30,43 17,07 19,57 SUBMED 7,69 6,67 9,09 7,50 6,52 4,55 8,70 9,76 6,52 MROP Tracks generalized Tracks LAT 23,08 26,67 24,24 22,50 23,91 13,64 17,39 17,07 19,57 0,00 3,33 3,03 2,50 2,17 0,00 4,35 2,44 6,52 SUBTROP CIR 3,85 9,09 10,00 21,21 15,00 23,91 13,04 17,07 13,04 ploidy: ploidy grade; m/t: index monolete/trilete) and tracks generalized tracks and monolete/trilete) index m/t: grade; ploidy ploidy: o HET 23,08 20,00 21,21 20,00 17,39 31,82 17,39 19,51 15,22 m/t 1,00 1,00 1,75 1,22 1,71 1,00 0,64 1,28 1,09 3,08 2,53 3,21 3,05 3,22 3,09 2,87 3,07 2,78 ploidy o 26 30 33 40 46 22 23 41 46 Pteridofloristic index Specific richesse 890 562 1418 1362 2066 1919 3749 2159 1092 elevation Maximum 37,9 35,1 36,11 34,45 35,47 36,52 36,37 35,16 36,59 Latitude Geographical references 1,26 5,28 3,46 5,07 3,26 2,57 1,53 5,12 5,29 Longitude 1,30 1,22 1,13 0,98 0,93 1,35 1,37 0,96 0,92 Range (Dif Ul-Ll) TD 3,59 3,65 3,38 3,53 3,46 3,65 3,79 3,62 3,84 Value (CIR: Circumboreal; ENMED: Mediterranean endemisms; HET: Heterochoric; LAT: Latimediterranean; MROP: (CIR: Mediterranean Circumboreal; and ENMED: neighboring Mediterranean eastern Heterochoric; endemisms; LAT: regions; HET: REPAME: Paleomediterranean relicts; SUBMED: Submediterranean; SUBTROP: Subtropical) Table 1.- Values for each of the analyzed OGUs: TD, geographical references, pteridofloristic index ( index pteridofloristic references, geographical TD, OGUs: analyzed the of each for Values 1.- Table Tlemcen Tingitania Tejeda Ronda Nevada Gurugú Gata Chauen Aljibe

CUARTAS_BotanicaComplutensis45.indd 6 25/5/21 21:01 Salvo Tierra, A. E.; Báez, José C.; Flores-Moya, A. Bot. complut. 45, 2021: 1-11 7 0 0 0 0 0 0 1 1 1 1 1 0 0 1 0 0 Gata 0 0 1 0 0 1 1 1 1 1 1 1 0 0 0 0 Nevada 0 0 0 0 0 1 0 1 1 1 1 0 0 0 0 0 Tejeda 0 0 0 0 1 0 1 1 0 1 1 0 0 1 0 1 Ronda 1 1 0 1 1 0 1 1 0 1 1 1 0 1 1 1 Aljibe 0 0 0 0 0 0 1 0 0 1 1 0 0 1 0 1 Tlemcen 0 0 0 0 1 0 0 1 0 1 1 0 1 0 0 0 Gurugú 0 0 1 0 1 1 1 1 0 1 1 0 1 0 1 0 Chauen 0 0 0 1 0 0 1 1 0 1 1 0 0 0 1 0 Tingitania CODE ISO HIS ISO TIG ISO VEL PSI NUD SEL BAL SEL ISO DUR VAN SPE VAN SEL DEN SEL EQU TEL OPH LUS OPH VUL EQU ARV BOT LUN BOT OSM REG CUL MAC CUL EQU RAM (L.) - (A. Ehrh. (L.) L. Gennari Bory Bory A. Braun some authors) recognized as species by recognized SPECIEs (incl. subspecies C. Presl Culcita macrocarpa (L.) P. Psilotum nudum (L.) P. Beauv. L. Osmunda regalis specio Vandenboschia Kunkel sa (Willd.) Ophioglossum lusitanicum L. Ophioglossum vulgatum L. Botrychium lunaria Swatrz Equisetum ramosissimum Desf. Equisetum telmateia Equisetum arvense Selaginella balansae Braun) Heiron. Selaginella denticulata Spring. Isoëtes tiguliana Isoëtes velata Isoëtes histrix Isoëtes durieui GENUS Culcita Psilotum Osmunda Vandenboschia Ophioglossum Botrychium Equisetum Selaginella Isoëtes FAMILY Culcitaceae Psilotaceae Osmundaceae Hymenophyllaceae Ophioglossaceae Equisetaceae Selaginellaceae Isoëtaceae ORDER SUPPLEMENTARY DATA - Checklist of pteridophytes from the Gibraltar Arch ordered according to the PGP1 (2016) systematics and presence (1) / absence (0) for the OGUs the for (0) absence / (1) presence and systematics (2016) PGP1 the to according ordered Arch Gibraltar the from pteridophytes of Checklist - DATA SUPPLEMENTARY studied. Cyatheales Psilotales Osmundales Hymenophyllales Ophioglossales Equisetales Selaginellales Isoëtales

CUARTAS_BotanicaComplutensis45.indd 7 25/5/21 21:01 8 Salvo Tierra, A. E.; Báez, José C.; Flores-Moya, A. Bot. complut. 45, 2021: 1-11 0 0 0 1 0 0 0 1 1 0 1 1 1 0 0 1 0 0 0 1 0 0 1 0 1 1 1 1 1 1 0 1 1 0 1 0 1 0 1 0 1 1 0 1 0 1 0 1 0 1 1 0 1 1 0 1 0 0 0 1 0 1 0 1 1 0 1 0 1 0 1 1 1 1 1 0 1 0 0 0 0 0 1 1 1 1 1 1 0 1 0 1 0 1 1 1 0 0 1 0 0 0 1 1 1 0 0 0 1 0 1 0 0 0 1 1 1 0 1 0 0 1 0 0 1 1 0 0 0 1 0 0 0 1 0 0 1 1 0 0 1 0 1 0 0 1 0 1 0 0 1 0 1 1 1 1 1 1 1 1 1 0 0 0 1 0 1 1 1 1 0 1 0 1 0 0 1 0 1 1 1 0 1 0 1 0 0 1 1 BLE SPI ASP BIL ASP ASP TRI ASP PLE HIS ATH FIL ATH ASP SEP ASP ASP VIR ASP ASP ADI ASP ASP PET ASP ASP PAC ASP DIP CAU DIP CET OFF CET ASP FOR ASP ASP RUT ASP ASP FON ASP PHY SCO PHY ASP OBO ASP PHY SAG PHY ASP ONO ASP ASP HEM ASP ASP MAR ASP L. L.

(DC.) L. (L.) (L.) Viv. Viv. Viv. Viv. L. (L.) L. Hudson Kümmerle (Schultz) (Cav.) Diplazium caudatum (Cav.) Jermy Struthiopteris spicant Weiss F.W. Asplenium viride Athyrium filix-femina Roth Asplenium septentrionale (L.) Hoffm. Asplenium trichomanes Asplenium ruta-muraria L. Asplenium sagittatum Bange Asplenium scolopendrium Asplenium petrarchae Asplenium petrarchae (Guérin) DC. Asplenium onopteris Asplenium obovatum subsp. billotii Kerg. Asplenium marinum L. Asplenium obovatum subsp. obovatum Asplenium hemionitis Asplenium hispanicum (Coss.) Greuter & Burdet Asplenium fontanum Bernh. Mag - Asplenium foreziense nier Asplenium csikii András . & Asplenium adiantum-nigrum L. Asplenium ceterach Diplazium Struthiopteris Athyrium Asplenium Blechnaceae Athyriaceae Aspleniaceae Polypodiales

CUARTAS_BotanicaComplutensis45.indd 8 25/5/21 21:01 Salvo Tierra, A. E.; Báez, José C.; Flores-Moya, A. Bot. complut. 45, 2021: 1-11 9 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 1 1 0 1 1 1 0 1 0 0 1 0 1 0 1 1 0 1 1 0 0 0 1 1 1 1 1 1 0 1 1 1 0 1 1 1 1 1 0 0 1 0 0 0 0 1 0 1 1 0 0 1 0 0 0 0 1 1 1 1 0 0 1 0 0 0 1 1 1 1 1 0 0 1 0 0 0 0 1 1 0 0 0 0 POL INT POL CYS VIR CYS DIC THE PAL CYS FRA POL VUL POL PTE AQU CHR DEN POL CAM POL DAV CAN DAV GYM ROB L.

R. (L.) Schott (L.) (Desv.) (Desv.) (L.) Cyclosorus dentatus (Forssk.) Ching Pteridium aquilinum Kuhn Thelypteris palustris Polypodium vulgare L. Polypodium vulgare Polypodium interjectum Shivas Polypodium cambricum Cystopteris dickieana Sim Cystopteris fragilis Bernh. Cystopteris viridula Desv. Gymnocarpium robertianum Newman (Hoffm.) Davallia canariensis Sm. Cyclosorus Pteridium Thelypteris Polypodium Cystopteris Gymnocarpium Davallia Dennstaedtiaceae Thelypteridaceae Polypodiaceae Cystopteridaceae Davalliaceae Polypodiales

CUARTAS_BotanicaComplutensis45.indd 9 25/5/21 21:01 10 Salvo Tierra, A. E.; Báez, José C.; Flores-Moya, A. Bot. complut. 45, 2021: 1-11 0 1 0 0 1 1 0 1 1 0 1 1 0 1 1 0 1 1 0 1 1 1 1 1 0 1 0 0 1 1 0 1 1 0 1 1 0 1 0 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 0 1 1 0 1 1 0 0 0 0 1 1 0 1 1 1 1 1 0 0 0 0 1 1 0 0 1 0 1 1 0 1 0 1 1 1 0 1 1 1 1 1 1 1 0 0 1 1 0 0 0 1 0 1 PTE VIT PTE INC CHE HIS CRY CRI CRY CHE TIN ADI CAP COS VEL ANO LEP CHE ACR CHE GUA NOT MAR NOT CHE MAD (L.) (Aiton) Cav. L. Pteris vittata Paragymnopteris marantae (L.) K.H. Shing Pteris incompleta Cryptogramma crispa (L.) ex Hook. R. Br. Cosentinia vellea Tod. Anogramma leptophylla Link Oeosporangium guanchi - & cum (Bolle) Fraser-Jenk. Pariyar Oeosporangium hispani - & cum (Mett.) Fraser-Jenk. Pariyar Oeosporangium pteridioides & (Reichard) Fraser-Jenk. Pariyar subsp. pteridioides Oeosporangium tinaei Fraser-Jenk. (Tod.) Adiantum capillus-veneris L . Oeosporangium pteridioides & (Reichard) Fraser-Jenk. Pariyar subsp . acrosticum & (Balb.) Fraser-Jenk. Pariyar Paragymnop - teris Pteris Cryptogramma Cosentinia Anogramma Adiantum Oeosporangium Pteridaceae Polypodiales

CUARTAS_BotanicaComplutensis45.indd 10 25/5/21 21:01 Salvo Tierra, A. E.; Báez, José C.; Flores-Moya, A. Bot. complut. 45, 2021: 1-11 11 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 0 1 1 1 0 0 0 1 0 0 1 0 1 1 1 0 0 1 1 0 0 0 0 1 0 1 0 0 1 0 0 0 0 1 1 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 1 1 1 0 0 0 0 0 0 0 0 PIL MIN PIL DRY FIL DRY POS SET DRY AFF DRY POS LON POS ACU DRY SUB DRY DRY THY DRY DRY BOR DRY MAR STR DRY GUA DRY - (L.) Gibby (L.) Durieu Pilularia minuta Marsilea strigosa Willd. Polystichum lonchitis Roth Polystichum setiferum Woyn. (Forssk.) Polystichum aculeatum Roth Fras - Dryopteris tyrrhena & Reichst. er-Jenkins Dryopteris mindshelken sis N.Pavl. Dryopteris guanchica & Jermy Dryopteris filix-mas (L.) Schott (New - Dryopteris borreri Tavel man) Oberh. & Dryopteris affinis (Lowe) Fraser-Jenkins Pilularia Marsilea Polystichum Dryopteris Marsileaceae Dryopteridacae Salviniales Polypodiales

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