Differences in Diversity of Corticolous Lichens Between Interior and Edge

Differences in diversity of corticolous lichens between interior and edge of the Monte Barranca semi-deciduous forest, Santiago de Cuba Author(s): Dania Rosabal, Ana Rosa Burgaz, Adeline Altamirano, and Gregorio Aragón Source: The Bryologist, 115(2):333-340. 2012. Published By: The American Bryological and Lichenological Society, Inc. DOI: http://dx.doi.org/10.1639/0007-2745-115.2.333 URL: http://www.bioone.org/doi/full/10.1639/0007-2745-115.2.333

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Differences in diversity of corticolous lichens between interior and edge of the Monte Barranca semi-deciduous forest, Santiago de Cuba

Dania Rosabal1,4, Ana Rosa Burgaz2, Adeline Altamirano1, and Gregorio Arago´n3

1 Departamento de Biologıá, Universidad de Oriente, Patricio Lumumba s/n, CP. 90500, Cuba; 2 Departamento de Biologıá Vegetal I, Facultad de Biologıá, Jose´ Antonio Novais 2, Universidad Complutense de Madrid, 28040 Madrid, Spain; 3 Departamento de Biologıá y Geologıá, Universidad Rey Juan Carlos, c/Tulipań s/n, 28933-Mo´stoles, Madrid, Spain

ABSTRACT. The epiphytic lichen flora in the semi-deciduous forest of Monte Barranca, Santiago de Cuba differs between the edge and interior due to differences in light levels. Four transects of 50 m were establish: two on the edge and two on the interior. Sixty randomly-chosen trees were surveyed, fifteen trees in each transect. The trees were sampled from the ground to two meters in height. Seventy-four corticolous lichen species were recorded, including 51 newly recorded for Cuba. Reproductive strategies were as varied for those lichens on the edge of the forest as for those in the interior, but most of the species were reproducing sexually. Porina distans, a generalist species, is highly adapted to the Monte Barranca semi-deciduous forest.

KEYWORDS. Semi-deciduous forest, edge effect, Neotropic, Caribbean lichens, epiphytes.

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Edge effect enhances the microclimatic variability in Renhorn 1996, 1998; Hilmo & Holien 2002; Jansson an environmental gradient from the outside toward et al. 2009; Kivisto¨ & Kuusinen 2000; Renhorn et al. the interior of a forest (Bustamante & Grez 1995), 1997; Rheault et al. 2003; Will-Wolf et al. 2006). affecting changes in population size and interspecific Little is, however, known about how this effect relationships (Fahrig 2003). Its influence can influences their reproductive strategies (Dettki et al. penetrate from ten to 100 meters into a stand and can 2000; Sipman & Harris 1989). have a particularly negative influence on small forest The Monte Barranca semi-deciduous forest is fragments (Feinsinger 2004). Lichens are generally considered the only relic of the ancient forests of the sensitive to environmental change, and several Cauto basin in Cuba (Borhidi & Mun˜iz 1980). This studies have focused on assessing the edge effect on small and isolated patch of forest today lies adjacent their distribution (Belincho´n et al. 2007; Dettki & to pastures, sugar cane and corn fields. Monte Essen 1998; Dettki et al. 1998; Esseen 2006; Esseen & Barranca semi-deciduous forest was designated a Floristic Managed Reserve protected area in 2006. 4 Corresponding author’s e-mail: [email protected] Before that, the stand underwent intensive logging, DOI: 10.1639/0007-2745-115.2.333 and a number of heliophilous species occupied the

The Bryologist 115(2), pp. 333–340 0007-2745/12/$0.95/0 Copyright E2012 by The American Bryological and Lichenological Society, Inc. 334 The Bryologist 115(2): 2012 area. It is our hypothesis that corticolous lichen the trunk of trees on edge and interior of the stand composition, richness, frequency and reproductive were measured at breast height (DBH 5 1.30 m). strategies at the edge will be different from those at The roughness of the bark was estimated by the interior, due to differences in light levels, means of a roughness coefficient. Firstly, a line of roughness of the bark and trunk diameter. three cm length was marked on the trunk, and a thread was fixed at one end. The thread was then

MATERIALS AND METHODS pushed into cracks and fissures, following the trajectory of the line until the other end. The length Study area. Field studies were carried out in of the thread was then measured and this number February 2008, in the semi-deciduous forest of Monte Barranca of 311.4 ha defined by a perimeter of was divided by three (to normalize the values), 10.32 km, located in Palma Soriano-San Luis obtaining the roughness coefficient (Feinsinger, pers. flatland, Santiago de Cuba, at about 239 m above sea comm. October 2007). level. This forest lays between two important The incoming light was recorded from four mountain systems: the Sierra Maestra (44.3 km points marking the cardinal directions around the toward south) and Sierra Cristal (46.6 km toward tree. It was determined using a 6 cm diameter north). Two small forest patches occur 7.3 and cylinder. The interior of the disk was divided within 13.8 km westwards from the Monte Barranca semi- four quadrants, therefore each quadrants represents deciduous forest. The mean annual rainfall varies 25% (James & Shugart 1970). The cylinder was between 1000 to 1200 mm, the mean annual Tu placed at 2 m in height and it was looked through the oscillates between 24 to 26 uC and the mean annual cylinder, the number of the quadrants partial or atmospheric humidity is 80% (Montenegro 1991). totally cover by vegetation of the canopy in each of The Monte Barranca semi-deciduous forest is four points was counted. The percentage of incoming composed of two strata: the first with trees 12 m in light was calculated for each point as 100% minus the height and the second with trees up to 25 m. The sum of the quadrants covered with vegetation. Lastly, phorophytes surveyed in the edge of the forest were the average of percentage of incoming light was Oxandra lanceolata (Sw.) Baill. (Anonaceae), calculated for each tree as the sum of the percentage Guazuma ulmifolia Lam. (Sterculiaceae), Swietenia of the incoming light for each point divided by four. mahagonii (L.) Jacq. (Meliaceae), Cedrela odorata L. Reproductive structures (symbiotic propagules or (Meliaceae), Eugenia floribunda West ex Willd. ascocarps producing sexual ascospores) developed by (Myrtaceae), Trema lamarkiana (R. & Sch.) Blume the lichens were recorded. (Ulmaceae), Chrysophyllum oliviforme L. Specimens were identified using specialized keys (Sapotaceae), Spondias mombim L. (Anacardiaceae), (Aptroot et al. 2007; Brodo et al. 2001; Gunderley 1999; Gerascanthus gerascanthoides (Kunth) Borhidi Moberg 1990; Wirth & Hale 1978). A Novel XSZ-N207 (Boraginaceae) and Cecropia peltata L. Biological Microscope and a Novel NTB-2B (Cecropiaceae). The phorophytes sampled in the Stereoscope were used for to determine the specimens. interior of the forest were Oxandra lanceolata, The nomenclature follows Kirk (2008). The vouchers Cupania glauca Sw. (Sapindaceae) and Ocotea were deposited in the Charles Ramsdem Herbarium of coriacea (Sw.) Brito (Lauraceae). Universidad de Oriente, Santiago de Cuba. Sampling design. Four transects of 50 m were We follow the standard terms and definitions established: two on the edge and two on the interior, proposed by Harper et al. (2005) for commons located more than 25 m from each other. The edge of concepts used in research on forest edge. the forest is considered to begin one meter into the Data analysis. To determine differences in stand and extends for five meters, and the interior of species composition and reproductive strategies the stand begins 15 meters inward. We sampled 60 between the edge and interior of the forest along a randomly-chosen trees, 15 along each of the four light, diameter of the trunk and roughness of the transects and recorded the lichens from ground to two bark gradient, the data were analyzed using a meters up. The roughness of the bark and diameter of Canonical Correspondence Analysis (CCA) Rosabal et al.: Edge effect on epiphytic lichens 335 ordination method from ‘PC-ORD for Windows’ richness and frequency (richness: z 5 3.70; p 5 (McCune & Mefford 1999). The presence/absence 0.0002, frequency: z 5 2.95; p 5 0.003), with a data on each tree were used for the main matrix and higher diversity of epiphytic lichens at the edge of the all rare species were included. The reproductive forest. The b-diversity between both zones was 0.48, strategies of the taxa were recorded in the field, and which mean a 48% of dissimilarity between them. included in the second matrix. A Mann-Whitney U-test (P 5 0.05), according to DISCUSSION (Siegel & Castellań 1995) was performed to test for Leopold (1933) recognized that edge habitat differences in species richness and frequency between supported a high diversity of game species and this the edge and interior stands. The b-diversity between phenomenon was termed edge effect. Harper et al. edge and interior of the forest was calculated using the (2005) suggested that landscapes characterized by Proportional Dissimilarity Index (PD) (Feinsinger patchy habitats may also have many edge-adapted 2004). The formula is: PD 5 1-PS, where: PS 5 S min species. The epiphytic lichen communities at the edge (Pie;Pii), being Pie 5 nie/n, ni: frequency of the species and in the interior of the semi-deciduous forest of in the edge, n: total frequency of species in this zone, Monte Barranca differ markedly in their Pii 5 nii/n, ni: frequency of the species in the interior, composition. The edge of the forest was more diverse n: total frequency of species in this zone, min (Pie;Pii): in corticolous lichens and included species such as the minimum value resulting of comparison between Glyphis cicatricosa, Arthonia cinnabarina, Physcia the Pie and Pii. sorediosa, Trypethelium eluteriae, Chrysotrix candelaris, Graphis leptocarpa and Bacidia RESULTS circumspecta, which are considered tolerant of high Seventy-four corticolous lichen species were light (Brodo et al. 2001). Porina distans was a found, 51 of which are newly recorded for Cuba dominant species in both the interior and along the (Table 1). Ninety-three percent of the species were edge of this forest, suggesting that this species may crustose, only few were foliose-appressed (7%) and have a broad physiological tolerance, unlike typical none were fruticose. Porinaceae, a family considered by Rivas Plata et al. The Pyrenulaceae were the most diverse family (2007) to be more characteristic of undisturbed and with 23 species, followed by the Graphidaceae (9) old growth secondary forests. and Porinaceae (7). The families best represented These differences best explained by changes in both in the edge as in interior stands were the light conditions and the pattern is congruent with Pyrenulaceae and Porinaceae. Fifty-six species were that observed in different tropical and boreal forests found on the edge and 43 on the interior of the (Kivisto¨ & Kuusinen 2000; Pe´rez et al. 2005; Rosabal forest, 25 of which are shared by both zones. Porina & Aragoń 2010; Rosabal et al. 2010). By contrast, we distans was the most frequent species in both the did not find any trend in species richness with edge and the interior parts of the forest. Thirty-five changes in tree diameter or bark roughness, as would species with the minimum value of frequency (rare be expected if lichen communities diversified during species) were found. Lichens with sexual successions on aging trees (Barkman 1958), as their reproduction were more frequent than species with diameter increases and their bark becomes rougher symbiotic propagules (Table 1). (Belinchoń et al. 2007; Broad 1989; Brodo 1973; The CCA analysis showed a strong pattern in Burgaz et al. 1994; Fuertes et al. 1996; Kantvilas & species composition against light conditions between Jarman 2004). the edge and interior of the forest (Fig. 1). Diameters Most epiphytic lichens in this forest developed of the trunk and roughness of the bark did not show crustose thalli, as is typical in lowland forests any discriminative pattern. The analysis also resolved (Aptroot & Sipman 1997). Foliose and fruticose a correlation between lichens species with perithecia lichen diversity increases with altitude (Sipman & and trees on the edge of the forest. The edge and Harris 1989) and atmospheric humidity (Bu¨del & interior of the forest exhibit differences in species Scheidegger 1996), which could explain the low 336 The Bryologist 115(2): 2012

Table 1. List of the species, distribution and reproductive strategies of corticolous lichens in Monte Barranca semi-deciduous forest.

Frequency Mode of Family Species Abbreviation Edge Interior reproduction

Arthoniaceae *Arthonia aciniformis Stirt. Arth-aci 1 - Apothecia Arthonia cinnabarina (DC.) Wallr. Arth-cin 11 3 Apothecia Bacidiaceae Bacidia circumspecta (Norrl. & Nyl.) Malme Bac-circ 4 - Apothecia *Bacidia hostheleoides (Nyl.) Zahlbr. Bac-host 1 4 Apothecia Caliciales Calicium lenticulare Ko¨rb. Cal-lent 1 1 Apothecia Celotheliaceae *Celothelium aciculiferum (Nyl.) Vain. Cel-acic 1 - Perithecia Chrysothricaceae Chrysothrix candelaris (L.) J. R. Laundon Chry-can 2 - Soredia Coenogoniaceae *Coenogonium lutescens (Veˇzda & Malcolm) Malcolm Coen-lut - 1 Apothecia *Coenogonium roumeguerianum (Mu¨ll. Arg.) Kalb Coen-rou 1 4 Apothecia Collemataceae *Collema subflaccidum Degel. Col-subf - 1 Isidia Leptogium austroamericanum (Malme) Dodge Lep-aust - 1 Isidia Graphidaceae Glyphis cicatricosa Ach. Glyp-cic 6 1 Apothecia Graphina incrustans (Feé) Mu¨ll. Arg. Grapn-in 1 - Apothecia *Graphina suberythrella M. Wirth & Hale Grapn-su 1 - Apothecia *Graphis adpressa Vain. Grap-adp 2 2 Apothecia *Graphis descuamescens Feé Grap-des 1 - Apothecia *Graphis humilis Vain. Grap-hum 1 - Apothecia Graphis hyphosa Staiger Grap-hyp - 4 Apothecia Graphis leptocarpa Feé Grap-lep 2 - Apothecia Graphis librata C. Knight Grap-lib 1 1 Apothecia Lecanoraceae Lecanora flavidofusca Mu¨ll. Arg. Lec-flav 1 - Apothecia *Lecanora leproplaca Zahlbr. Lec-lepr 3 6 Apothecia Lecanora praeferenda (Nyl.) Nyl Lec-prae - 1 Apothecia Lecideaceae *Lecidea nylanderi (Anzi) Th. Fr. Leci-ny 8 11 Apothecia Letrouitiaceae *Letrouitia domingensis (Pers.) Hafellner & Bellem. Let-domi 2 7 Apothecia Monoblastiaceae *Anisomeridium ambiguum (Zahlbr.) R. C. Harris Anis-amb - 1 Perithecia *Anisomeridium leucochlorum (Mu¨ll. Arg.) R. C. Harris Anis-leu - 1 Perithecia Mycoporaceae Mycoporum lacteum (Ach.) R. C. Harris Myco-lac - 1 Perithecia *Mycoporum sparsellum Nyl. Myco-spa - 1 Perithecia Pertusariaceae *Pertusaria albescens (Huds.) M. Choisy & Werner Pert-alb 1 2 Apothecia *Pertusaria mesotropa Mu¨ll. Arg. Pert-mes 1 - Apothecia Physciaceae *Amandinea punctata (Hoffm.) Coppins & Scheid. Ama-punc - 1 Apothecia *Physcia atrostriata Moberg Phy-atro 1 - Soredia *Physcia biziana (A. Massal.) Zahlbr. Phy–biz 1 - Soredia Physcia sorediosa Lynge Phy-sor 5 2 Soredia Porinaceae *Porina curtula Malme Pori-cur 1 - Perithecia *Porina distans Malme Pori-dis 17 16 Isidia *Porina imitatrix Mu¨ll. Arg. Pori-imi 1 - Perithecia Porina nucula Ach. Pori-ncu 5 8 Perithecia *Porina nuculastrum (Mu¨ll. Arg.) R. C. Harris Pori-nuc 1 1 Perithecia *Porina subnucula Lumbsch, Lu¨cking & Veˇzda Pori-sub - 1 Perithecia Porina tetracerae (Afzel.) Mu¨ll. Arg. Pori-tet 1 - Perithecia Pyrenulaceae *Anthracothecium prasinum (Eschw.) R. C. Harris Anth-pra - 2 Perithecia *Pyrenula acutispora Kalb & Hafellner Pyr-acu 1 - Perithecia *Pyrenula andina Aptroot Pyr-andi 2 - Perithecia *Pyrenula anomala (Ach.) A. Massal. Pyr-anom 5 4 Perithecia *Pyrenula aspistea (Afzel.) Ach. Pyr-aspi 1 2 Perithecia *Pyrenula cocoe¨s Mu¨ll. Arg. Pyr-coco 1 1 Perithecia Rosabal et al.: Edge effect on epiphytic lichens 337

Table 1. Continued.

Frequency Mode of Family Species Abbreviation Edge Interior reproduction

Pyrenula costaricensis Mu¨ll. Arg. Pyr-cost - 1 Perithecia *Pyrenula dermatodes (Borrer) Schaer. Pyr-derm 1 2 Perithecia Pyrenula laii Aptroot Pyr-laii 1 - Perithecia Pyrenula leucostoma Ach. Pyr-leuc 5 4 Perithecia *Pyrenula luteopruinosa Etayo & Aptroot Pyr-lute - 1 Perithecia Pyrenula mamillana (Ach.) Trevis. Pyr-mami 1 4 Perithecia *Pyrenula massariospora (Starba¨ck) R. C. Harris Pyr-mass 2 - Perithecia *Pyrenula mastophoroides (Nyl.) Zahlbr. Pyr-mast 1 - Perithecia *Pyrenula papilligera (Leight.) Mu¨ll. Arg. Pyr-papi 1 1 Perithecia Pyrenula pyrenuloides (Mont.) R. C. Harris Pyr-pyr 1 - Perithecia *Pyrenula pyrgillospora Aptroot Pyr-pyrg 2 - Perithecia *Pyrenula rubroanomala Aptroot & Lu¨cking Pyr-rubr 2 - Perithecia *Pyrenula subcongruens Mu¨ll. Arg. Pyr-subc - 2 Perithecia *Pyrenula subferruginea (Malme) R. C. Harris Pyr-subf 1 - Perithecia *Pyrenula subgregantula Mu¨ll. Arg. Pyr-subg 1 - Perithecia *Pyrenula subsoluta (Mu¨ll. Arg.) Aptroot Pyr-subs - 1 Perithecia *Pyrenula tenuisepta R. C. Harris Pyr-tenu 2 - Perithecia Thelotremataceae *Myriotrema microporellum (Nyl.) Hale Myr-micr - 1 Apothecia *Myriotrema myriocarpum (Nyl.) Hale Myr-myr 2 - Apothecia *Thelotrema neei (Hale) Hale The-neei - 1 Apothecia Thelotrema porinoides Mont. The-pori 2 - Apothecia Trypetheliaceae *Polymeridium quinqueseptatum (Nyl.) R. C. Harris Poly-quin 1 - Perithecia *Polymeridium subcinereum (Nyl.) R. C. Harris Poly-sub 2 - Perithecia *Pseudopyrenula subnudata Mu¨ll. Arg. Pseu-sub 2 2 Perithecia Trypethelium eluteriae Spreng. Try-elut 3 1 Perithecia *Trypethelium subcatervarium Malme Try-subc 1 2 Perithecia Total frequency 130 115

* identify new records for Cuba. number of foliose lichens and none fruticose species high light levels. Porina distans, the most frequent lichen, in this forest. reproduce asexually by isidia. Corticolous snails such as Most epiphytic lichens in the Monte Barranca semi- Polymita venusta and Liguus fasciatus (species highly deciduous forest reproduce sexually. The dispersal agents abundant), others invertebrates and small vertebrates, as of the sexual diaspores, particularly wind, could not be a well as, the rainfall could be the most probable agents for limiting factor inside the forest, determining that species the dispersion of this lichen species in Monte Barranca with sexual propagules prevailed. Lichens that usually semi-deciduous forest. reproducing via meiotic spores are thought to be more The lichen communities in a semi-deciduous forest frequently found in areas with high illumination levels, in Cuba differ in composition between the forest margin whereas species relying on vegetative propagules occur and interior as would be predicted by the edge effect. The primarily under closed canopy (Stofer et al. 2006). This Monte Barranca semi-deciduous forest has been could explain the correlation found between lichens adjoined to pastures and cultures for decades and has species with perithecia and trees on the edge of the forest. low trees diversity, which increase the patch contrast and On the other hand, light conditions in the Monte enhance the distance of edge influence. Indeed, Harper Barranca forest vary seasonally given the deciduous et al. (2005) hypothesized that the edge effect is more nature of the trees, and hence the prevalence of sexually pronounced in homogenous forest stands that edged reproducing lichens may be related with such periodic highly contrasted habitats for prolonged times. 338 The Bryologist 115(2): 2012

Figure 1. CCA ordination of the species composition with regard to light and perithecia as reproductive strategy. Filled triangles correspond to trees on the interior (I); unfilled triangles correspond to trees on the edge (B).

ACKNOWLEDGMENTS inventory in Costa Rica: Pyrenocarpous Lichens with This study was partly supported by Oficina Territorial para la bitunicate asci. Bibliotheca Lichenologica 97: 1–162. Conservacioń de la Flora y la Fauna, Santiago de Cuba. We Barkman, J. J. 1958. On the ecology of cryptogamic epiphytes want to thank to Dr. Gerardo Hechavarrıá for the access to the with special reference to the Netherlands. Van Gorcum, area. We are grateful to two anonymous reviewers for valuable Assen, The Netherlands. critical comments on the manuscript. Belinchoń, R., I. Martıńez, A. Escudero, G. Aragoń& F. Valladares. 2007. Edge on epiphytic communities in a Mediterranean Quercus pyrenaica forest. Journal of LITERATURE CITED Vegetation Science 18: 81–90. Aptroot, A. & H. J. M. Sipman. 1997. Diversity of lichenized fungi Borhidi, A. & O. Mun˜iz. 1980. Mapa de la vegetacioń potencial in the tropics. Pages 93–106. In K. D. Hyde (ed.), Biodiversity of de Cuba. Acta Botanica Hungarica 28: 25–33. tropical microfungi. Hong Kong University Press, Hong Kong. Broad, K. 1989. Lichens in southern woodland. Forestry ———, R. Lu¨cking, H. Sipman, L. Umanã & J. Cha´vez. 2007. Commission. Handbook 4. Her Majesty’ Stationery Office, The first assessment of the Ticolichen Biodiversity London, United Kingdom. Rosabal et al.: Edge effect on epiphytic lichens 339

Brodo, I. M. 1973. Substrate ecology. Pages 401–441. In Hilmo, O. & H. Holien. 2002. Epiphytic lichen response to the V. Ahmadjian & M. E. Hale (eds.), The lichens. Academic edge environment in a Boreal Picea abies forest in central Press, New York. Norway. The Bryologist 105: 48–56. ———, S. D. Sharnoff & S. Sharnof. 2001. Lichens of James, F. C. & H. H. Shugart. 1970. A quantitative method of North America. Yale University Press, New Haven and habitat description. Audubon Field Notes 24: 727–736. London. Jansson, K. U., K. Palmqvist & P. A. Esseen. 2009. Growth of Bu¨del, B. & C. Scheidegger. 1996. Thallus morphology and the old forest lichen Usnea longissima at forest edges. anatomy. Pages 37–64. In T. Nash (ed.), Lichen Biology. Lichenologist 41: 663–672. Cambridge University Press, Cambridge, United Kingdom. Kantvilas, G. & S. J. Jarman. 2004. Lichens and bryophytes on Burgaz, A. R., E. Fuertes & A. Escudero. 1994. Climax epiphytic Eucalyptus obliqua in Tasmania: management implications communities in Mediterranean Spain. Botanical Journal of in production forest. Biological Conservation 117: 359–373. the Linnean Society 115: 35–47. Kirk, P. M. 2008. Species Fungorum. http://www. Bustamante, R. & A. Grez. 1995. Consecuencias ecolo´gicas de la speciesfungorum.org. fragmentacioń de los bosques nativos. Ambiente y Kivisto¨, L. & M. Kuusinen. 2000. Edge effects on the epiphytic Desarrollo XI (2): 5–63. lichen flora of Picea abies in middle boreal Finland. The Dettki, H. & P. A. Esseen. 1998. Epiphytic macrolichens in Lichenologist 32: 387–398. managed and natural forest landscapes: a comparison at Leopold, A. 1933. Game management. The University of two spatial scales. Ecography 21: 613–624. Wisconsin Press, Madison. ———, M. Edman, P. A. Esseen, H. Hedena˚s, B. G. Jonsson, McCune, B. & M. J. Mefford. 1999. PC-ORD. Multivariate N. Kruys, J. Moen & K. E. Renhorn. 1998. Screening for analysis of ecological data. Version 5.0, MjM Software, species potentially sensitive to habitat fragmentation. Gleneden Beach, Oregon, U.S.A.. Ecography 21: 649–652. Moberg, R. 1990. The lichen genus Physcia in Central and ———, P. Klintberg & P. A. Esseen. 2000. Are epiphytic lichens South America. Nordic Journal of Botany 10: 319–342. in young forests limited by local dispersal? Ecoscience 7: Montenegro, U. 1991. Clima. Atlas de Santiago de Cuba. 317–325. BIOECO. MEGACEM. Esseen, P. A. 2006. Edge influence on the old-growth forest Pe´rez, R., N. Miramontes, J. Rosales & H. Quiroz. 2008. indicator lichen Alectoria sarmentosa in natural ecotones. Macrolı´quenes cortıćolas en dos especies de conı´feras del Journal of Vegetation Science 17: 185–194. Parque Nacional Laguna de Zempoalas. Acta Universitaria ——— & K. E. Renhorn. 1996. Epiphytic lichen biomass in 18: 33–39. managed and old-growth boreal forests: effect of branch Renhorn, K. E., P. A. Esseen, K. Palmqvist & B. Sundberg. 1997. quality. Ecological Applications 6: 228–238. Growth and vitality of epiphytic lichens. I. Responses to ——— & ———. 1998. Edge effects on an epiphytic lichen in microclimatic along a forest edge-interior gradient. fragmented forests. Conservation Biology 12: 1307–1317. Oecologia 109: 1–9. Fahring, L. 2003. Effects of habitat fragmentation on Reyes, J., N. Vinã & O. Mun˜iz. 1991. Endemismos y rutas biodiversity. Annuual Review of Ecology, Evolution, and migratorias. Atlas de Santiago de Cuba. BIOECO. MEGACEM. Systematics 34: 487–515. Rheault, H., P. Drapeau, Y. Bergeron & P. A. Esseen. 2003. Edge Feinsinger, P. 2004. El disenõ de estudios de campo para la effects on epiphytic lichens in managed black spruce forests conservacioń de la biodiversidad. Editorial FAN, Santa Cruz of eastern North America. Canadian Journal of Forestry de Sierra, Bolivia. Research 33: 23–32. Fuertes, E., Burgaz, A. R. & A. Escudero. 1996. Pre-climax Rivas Plata, E., R. Lu¨cking & H. T. Lumbsch. 2007. When epiphyte communities of bryophytes and lichens in family matters: An analysis of Thelotremataceae Mediterranean forest from the Central Plateau (Spain). (Lichenized Ascomycota: Ostropales) as bioindicators of Vegetatio 123: 139–151. ecological continuity in tropical forests. Biodiversity and Gunderley, R. 1999. Die Lecanora subfusca-gruppe in Su¨d- und Conservation 17: 1319–1351. Mittelamerika. Journal of the Hattori Botanical Laboratory Rosabal, D. & G. Aragoń. 2010. Lı´quenes epı´fitos en el matorral 87: 131–257. costero de la Reserva Ecolo´gica Siboney-Juticı´ (Cuba). Halffter, G., J. Soberoń, P. Koleff & A. Melic (eds.). 2005. Sobre Botanica Complutensis 34: 21–25. diversidad biolo´gica: el significado de las diversidades Alfa, ———, A. R. Burgaz & R. De la Masa. 2010. Diversity and Beta y Gamma. Monografıás Tercer Milenio. Vol: 4. distribution of epiphytic macrolichens on tree trunks in two CONABIO, DIVERSITAS & S.E.A., Zaragoza, Spain. slopes of the montane rainforest of Gran Piedra, Santiago Harper, K., E. MacDonald, P. Burton, J. Chen, K. Brosofske, de Cuba. The Bryologist 113: 313–321. S. Saunder, E. Euskirchen, D. Roberts, M. Jaiteh & P.-A. Esseen. Siegel, S. & N. J. Castellań. 1995. Estadı´stica no parame´trica: 2005. Edge influence on forest structure and composition in aplicada a las ciencias de la conducta. Editorial Trillas, fragmented landscapes. Conservation Biology 19: 768–782. Mexico D. F., Mexico. 340 The Bryologist 115(2): 2012

Sierk, H. A. 1964. The genus Leptogium in North America groups in relation to land use intensity. The Lichenologist North of Mexico. The Bryologist 67: 245–317. 38: 331–353. Sipman, H. J. M. & R. C. Harris. 1989. Lichens. Pages 303–309. In Will-Wolf, S., L. Geiser, P. Neitlich & A. Reis. 2006. Forest H. Lieth & M. J. A. Werger (eds.), Ecosystems of the World. lichens communities and environment – How consistent 14B. Tropical rainforest ecosystem. Biogeographical and are relationships across scales? Journal of Vegetation ecological Studies. Elsevier, Amsterdam, The Netherlands. Science 17: 171–184. Stofer, S., A. Bergamini, G. Arago´n, P. Carvalho, B. J. Coppins, Wirth,M.&M.E.Hale,Jr.1978.Morden-Smithsonian S. Davey, M. Dietrich, E. Farkas, K. Ka¨rkka¨inen, Ch. Keller, Expedition to Dominica: The Lichens (Graphidaceae). L. Lo¨ko¨s, S. Lommi, C. Ma´guas, R. Mitchell, P. Pinho, Smitsonian Contributions to Botany 40: 1–64. V. J. Rico, A. Truscott, P. A. Wolseley, A. Watt & C. Scheidegger. 2006. Species richness of lichen functional ms received May 28, 2011; accepted April 2, 2012.