Resumen En los bosques montanos tropicales los briófitos son importantes en términos de diversidad y funcionamiento; allí, los briófitos que crecen sobre hojas son particularmente abundantes. El objetivo fue evaluar la riqueza y composición de comunidades epífilas en tres zonas del dosel de un árbol hospedador en el Parque Internacional La Amistad, Panamá. En cada árbol fueron colectadas seis hojas al azar, dos por zona de la copa (baja, media y alta). Se registró en total 48 especies, 89 % pertenecen a Marchantiophyta y 11 % a Bryophyta. La familia Lejeuneaceae representó el 70 % de las especies. La riqueza de especies disminuye desde la zona baja del dosel hacia la parte alta; sin embargo, las comunidades son similares entre zonas. Condiciones microclimáticas específicas del sotobosque podrían favorecer una mayor riqueza en la zona baja del dosel. Realizar investigaciones teniendo en cuenta factores ambientales y estructurales del bosque permitirá una mejor comprensión de estos cambios en diversidad.

Abstract In tropical montane forests, bryophytes are important in terms of diversity and functioning, in which epiphyllous species grow on tree leaves of trees and shrubs, and they are particularly abundant. The objective of this article was to evaluate epiphyllous bryophyte species richness and community composition in three canopy zones (low, medium, high) of a host tree in “La Amistad” International Park, Panama. From each sampled tree, six random leaves were randomly collected, two per canopy zone. A total of 48 species were recorded, of which 89% belong to Marchantiophyta and 11% to Bryophyta. The Lejeuneaceae family accounted for 70% of the species. Species richness decreased from the lowest to the highest canopy zone, however, species communities were similar between zones. Microclimatic conditions characteristic of the forest understory might have favoured greater species richness in the lower canopy zone. Further research is needed, for taking into account environmental factors and forest structure may lead to a better understanding of the observed changes in diversity.

ABSTRACT Some biogeographical features of bryophytes associated with subtropical-temperate ecosystems in Chile are examined, including diversity, concentration of species richness, endemism, biogeographical composition and disjunctions. The historical-biogeographical interpretation of these characteristics considers the major evolutionary changes of the group in three different temporal scenarios, according to geological, paleobotanical and molecular evidences. First, simple- and complex-thalloids liverworts represented in Chile are examined, belonging to the Classes Haplomitriopsida and Marchantiopsida- Marchantiidae, archaic lineages linked to the pre-Cenozoic colonization of the Pangea mega-continent. Examples of simple-thalloids liverworts of Class Jungermanniopsida-Pelliidae, hornworts of the Class Anthocerotopsida and mosses of the Classes Bryopsida and Polytrichopsida illustrate the paleo-austral distribution patterns of bryophytes from the temperate region of Chile, related to the Gondwana breakup and expansion of Angiosperms during the Cretaceous/Paleogene. A second scenario, considers the final uplift of the Andes and the development of the Arid Diagonal of South America during the Neogene, processes that have played a major role in the biogeographical structure of the Chilean and South American floras. In this context, leafy liverworts associated with the relict forests of the Chilean semi- arid coast stand out, particularly epiphyllous liverworts of the two most diverse families of the Class Jungermanniopsida-Jungermanniidae, Lejeuneaceae and Plagiochilaceae, adapted to microclimates favored by coastal fogs. The last scenario discusses the significant role of the Pleistocene glacial cycles in the expansion of the sub-Antarctic temperate-cold element throughout central-southern Chile, and its effects on the concentration of bryophyte richness associated with Chile’s temperate-rain ecosystems.

RESUMEN Se examinan algunos rasgos biogeográficos de las briófitas asociadas a los ecosistemas subtropical- templados de Chile, como diversidad, concentración de la riqueza de especies, endemismo, composición biogeográfica y disyunciones. La interpretación histórico-biogeográfica de estas características considera los grandes cambios evolutivos del grupo en tres escenarios temporales distintos, de acuerdo con las evidencias geológicas, paleobotánicas y moleculares. Para un primer momento, se examinan hepáticas talosas simples y complejas de las Clases Haplomitriopsida y Marchantiopsida-Marchantiidae representadas en Chile y pertenecientes a linajes arcaicos vinculados al poblamiento pre-Cenozoico del megacontinente Pangea. Ejemplos de distintos taxa de hepáticas talosas simples de la Clase Jungermanniopsida-Pelliidae, antocerotes de la Clase Anthocerotopsida y musgos de las Clases Bryopsida y Polytrichopsida ilustran patrones de distribución paleo-austral de briófitas de la región templada de Chile, relacionados a la fragmentación del Gonwana y expansión de las Angiospermas, durante el Cretácico/Paleógeno. Un segundo escenario Neógeno considera el levantamiento final de los Andes y desarrollo de la Diagonal Árida de Sudamérica, procesos que han jugado un rol significativo en la estructura biogeográfica de la flora de Sudamérica y de Chile. En este contexto se destacan casos de hepáticas foliosas asociadas a los bosques relictos de la costa semiárida chilena, particularmente hepáticas epifilas de las dos familias más diversas de la Clase Jungermanniopsida-Jungermannidae, Lejeuneaceae y Plagiochilaceae, adaptadas a microclimas favorecidos por neblinas costeras. En un último escenario se discute el rol relevante que han jugado las repetidas glaciaciones del Pleistoceno en la expansión del elemento sub-antártico templado- frío a lo largo de Chile central-sur, y sus efectos en la concentración de la riqueza de briófitas asociada a los ecosistemas templado-lluviosos de Chile.

Abstract Introduction: Many land molluscs survival strategies are still poorly understood or have not been even reported, especially in the Neotropics. Methods: I collected 25 adult Tikoconus.Tikoconus. costarricanus from Reserva Forestal Río Macho, Costa Rica. I kept the specimens for 8 days in terrariums to film their behavior. Objective: To analyse the behavior of T. costarricanus, with emphasis on its strategies to survive drought and probably also predation. Results: This snail has at least three unusual behaviors that probably help them reduce dehydration and may be escape from enemies and avoid diseases: hanging upside down like bats, falling and grooming. During aestivation, they compress the body and hang upside down from leaves, like bats hang from perches. They attach to the underside of leaves with mucus from a caudal gland. Disengagement is done with vigorous shell rotations and foot twisting in contorting sequences, and can be done as reaction to direct sunlight, and probably to avoid predators and parasites. They groom their own shell, shell lappets and foot, an unusual behavior among land snails. This species feeds mainly on epiphyllous mosses, algae and lichens, occasionally adding arthropod eggs and carrion. Egg laying is similar to other euconulids and valloniid snails. Conclusions: Aestivating hanging upside down is a drought avoiding trait described here for the first time and is also a new function for the caudal gland mucus. Leaf detaching is done by a contortion sequence of shell rotations and foot twisting; its complexity and duration varies according to the leaf side where the snail is located.

Resumen Introducción: Muchas estrategias de sobrevivencia en moluscos terrestres todavía están poco estudiadas o no han sido descubiertas, especialmente en el Neotrópico. Métodos: Recolecté 25 adultos de Tikoconus .Tikoconus. costarricanus de la Reserva Forestal Río Macho, Costa Rica. Mantuve los especímenes 8 días en terrarios para grabar su comportamiento. Objetivo: Analizar el comportamiento de T. costarricanus, con énfasis en sus estrategias para sobrevivir a la desecación del ambiente y posiblemente a los depredadores. Resultados: Este caracol tiene al menos tres comportamientos inusuales que le ayudan a reducir la deshidratación y probablemente escapar de enemigos y evitar enfermedades: colgarse como murciélagos, tirarse y acicalarse. Durante la estivación, ellos contraen el cuerpo y se cuelgan cabeza abajo de hojas, como los murciélagos en sus perchas. Ellos se adhieren al envés de las hojas con mucus de su glándula caudal. Para poder tirarse de la vegetación desarrollan una serie de contorsiones compuestas de giros de la concha y torsiones del pie, lo hacen para evitar la incidencia directa de los rayos del sol, y posiblemente, depredadores y parásitos. Ellos acicalan su propia concha, lóbulos de la concha y pie. Esta especie se alimenta principalmente de musgos, algas y líquenes epífilos, ocasionalmente agregan huevos y cadáveres de artrópodos. Depositan sus huevos de manera similar a otros caracoles euconúlidos y vallónidos. Conclusiones: La estivación colgándose como murciélagos es una estrategia para evitar la desecación que se describe aquí por primera vez y representa una función nueva del mucus generado por la glándula caudal. Para separarse del sustrato hacen una secuencia de contorsiones con la concha y con el pie, cuya duración depende del lado de la hoja donde se encuentren.

Resumo O Parque Estadual da Serra do Mar (PESM) está localizado no sudeste do estado de São Paulo, Brasil sendo a maior unidade de conservação de Mata Atlântica do país. O parque é dividido em 10 núcleos, sendo o Núcleo Itutinga-Pilões (NIP) selecionado para este estudo, visto que não foi realizado nenhum estudo analisando a riqueza de espécies de hepáticas nessa área. As similaridades florísticas entre o NIP e diferentes áreas da Mata Atlântica no estado de São Paulo também foram avaliadas. Cento e oitenta espécies de hepáticas foram identificadas nos três tipos de vegetação (floresta ombrófila de terras baixas, submontana e montana) encontradas no NIP, distribuídas em 62 gêneros e 21 famílias. A família Lejeuneaceae apresentou a maior riqueza com 80 espécies. O elevado número de espécies epífilas encontradas na área de estudo (26% de todas as espécies) foi notável. Em termos de suas distribuições mundiais, 57% das espécies são Neotrópicais. A análise de agrupamento mostrou que as áreas do PESM e Estação Ecológica Jureia-Itatins foram agrupadas com os maiores valores de similaridade. A flora de hepática do NIP demonstra a importância desse núcleo para a conservação da diversidade de hepáticas na Mata Atlântica do estado de São Paulo, bem como no Brasil.

Abstract Serra do Mar State Park (PESM) is located in southeastern São Paulo state, Brazil, and is the largest Atlantic Rainforest conservation area in the country. The park is divided into 10 nuclei, and the Itutinga-Pilões nucleus (NIP) was selected for study since no survey of liverworts species richness had yet been conducted there. The floristic similarities between the NIP and different areas of Atlantic Rainforest in São Paulo state were also assessed. One hundred and eighty liverworts species were identified in the three vegetation types (lowland, submontane, and montane ombrophilous forests) encountered in the NIP, distributed among 62 genera and 21 families. Lejeuneaceae showed the highest species richness with 80 species. The high number of epiphyllous species found in the study area (26% of all species) was notable. In terms of their worldwide distributions, 57% of the species are Neotropical. Clustering analysis showed that the areas of the PESM and Jureia-Itatins Ecological Station were grouped together with the highest similarity values. The liverworts flora of the NIP demonstrates the importance of that nucleus for the conservation of liverworts diversity in the Atlantic Rainforest of the São Paulo state, as well as in Brazil.

ResumenSe conoce muy poco sobre la ecología y biología del caracol terrestre Drymaeus tripictus. Se trata de una especie extremadamente escasa y endémica de las zonas altas de Costa Rica. Estudié la ecología y biología reproductiva de D. tripictus de abril 2009 a junio 2010 en un bosque maduro, un bosque jóven y una plantación de Cupressus lusitanica en Costa Rica. Cada tres meses visité esos hábitats y recolecté especímenes en 20 puntos de muestreo (cada uno de 3x3 m2). Estudié actividad y preferencias de hábitat en la naturaleza. En el laboratorio grabé en alta definición el comportamiento de apareamiento y analicé la morfología reproductiva con un microscopio estereoscópico de luz. El caracol es mas abundante en el bosque maduro (0.017 ind/m2) y prefiere hojas con poca cobertura de epífilos (25% o menos de cobertura, chicuadrado, p < 0.0001). En la época seca, se activan entre las 8:00 pm y las 8:00 am principalmente (chi-cuadrado = 22.65, p <0.0001). En la tarde se inactivan (11:00 am to 16:59 pm). Los individuos activos se encuentran principalmente en el haz de las hojas (Chi-square =6.76, df =1, p = 0.0093), que es donde se alimentan. El apareamiento es unilateral por "montaje de concha", no hacen cambio de roles sexuales y la intromisión fálica es críptica. Su sistema reproductivo es morfológicamente similar al de Drymaeus costarricensis. El comportamiento de apareamiento es como se espera de un caracol de concha con espira alta, excepto por la ausencia de cambio de roles. La densidad de D. tripictus es mas baja que la de otros bulimulidos considerados en peligro.

Abstract Very little is known about the ecology and biology of Drymaeus tripictus, an extremely rare and endemic land snail species from Costa Rican highlands. I studied the ecology and reproductive biology of D. tripictus from April 2009 through June 2010 in an old forest, a young forest and a Cupressuslusitanica plantation in central Costa Rica. Every three months I visited each habitat and collected specimens in 20 random sampling plots (3x3 m2 each). I observed the snail's activity and microhabitat preference in the field, and in the laboratory I recorded high definition videos of its mating behavior and analyzed reproductive morphology with light microscopy. The snail is more abundant in the old forest (0.017 ind./m2) and prefers leaves with little epiphyllous cover (0-25 % cover, chi-square test, p <0.0001). During the dry season the snails become active between 20:00 pm and 8:00 am (chi-square = 22.65, df=3, p < 0.0001); they are inactive mainly during the afternoon (11:00 am to 16:59 pm). I found active individuals mostly on the upper side of leaves, where they feed (Chi-square =6.76, df=1, p = 0.0093). Mating is unilateral, by shell mounting, with cryptic phallus intromission and without role switching or multiple mating. Its reproductive system is morphologically similar to that of Drymaeus costaricensis. Mating behavior is as expected for snails with high-spired shells, except for the lack of role switching. The density of D. tripictus is low even when compared with other endangered bulimulids. Rev. Biol. Trop. 64 (1): 55-68. Epub 2016 March 01.

El presente estudio constituye la primera parte de un inventario de la brioflora epifila de diferentes localidades de la región central de la Cordillera de la Costa, Venezuela. Las muestras analizadas proceden de colecciones de uno de los autores (T. Morales) y de otras que, en su mayoría, permanecían sin analizar en los estantes del Herbario Nacional de Venezuela (VEN). Se determinaron taxonómicamente 47 especies de briofitos epifilos, agrupados en 4 familias, 22 géneros y 42 especies de hepáticas; así como 3 familias, 4 géneros y 5 especies de musgos. Se adicionan 10 registros a la brioflora venezolana. Se enfatiza la extrema necesidad de estudiar taxonómica y ecológicamente, las comunidades de briofitos epifilos y epífitos de la Cordillera de la Costa, dada la incesante devastación de la misma por actividades humanas de diversa índole.

The present paper is the first part of an epiphyllous bryophyte inventory of the central region of the Cordillera de la Costa, Venezuela. The records are based on the collections of one of the authors (T. Morales) and on unidentified specimens kept in the National Herbarium of Venezuela. 47 species of epiphyllous bryophytes were taxonomically identified, grouped into 4 families, 22 genera and 42 species of liverworts, and 3 families, 4 genera and 5 species of mosses. 10 species are added to the Venezuelan bryoflora. The critical need for taxonomic and ecological studies of the communities of epiphyllous bryophytes of the Cordillera de la Costa is emphasized, given the relentless devastation of the area.

Foi realizado um inventário da brioflora epífita e epífila da RPPN Frei Caneca (8º42'41"S e 35º50'30"W, 750 m alt., 630 ha), remanescente de Floresta Atlântica, localizado no município de Jaqueira, Pernambuco. Um total de 120 amostras de briófitas epífitas foi coletado desde a base até a primeira ramificação do tronco em forófitas arbóreas representativas na área de estudo. Para o inventário das epífilas foram coletadas 40 amostras compostas de três a cinco folhas de arbustos e árvores, desde a base até cerca de 25 m de altura. A brioflora é composta por 71 espécies, distribuídas em 46 espécies de hepáticas e 25 espécies de musgos. Das 21 famílias registradas, Lejeuneaceae (31 spp.) e Calymperaceae (7 spp.) apresentaram maior riqueza genérica e específica. Lejeunea (6 spp.) e Syrrhopodon (4 spp.) foram os gêneros mais representativos. A distribuição das espécies é predominantemente Neotropical (67%) e Pantropical (15%). Neste trabalho são apresentadas oito novas ocorrências para o Estado de Pernambuco: Colura tortifolia (Nees & Mont.) Steph., Cyclolejeunea convexistipa (Lehm. & Lindenb.) A. Evans, Diplasiolejeunea brunnea Steph., D. pellucida (Meissn.) Schiffn., Prionolejeunea denticulata (Weber) Schiffn., Radula mexicana Lindenb. & Gottsche, R. recubans Tayl.; e uma delas Calymperes smithii E.B. Bartram é ocorrência nova para a região Nordeste. São apresentados comentários taxonômicos e ecológicos, informações sobre tipos de substrato e distribuição geográfica no mundo e no Brasil para as espécies de ocorrência nova.

A floristic survey of epiphyllous and epiphytic bryophytes was carried out at the Frei Caneca RPPN (8°42'41"S, 35°50'30"W, 750 m alt., 630 ha), a remnant Atlantic forest site located in the state of Pernambuco. A total of 120 samples of epiphytic bryophytes were taken from the trunks (base to first ramification) of host trees commonly found at the site. For the epiphyllous epiphytes, 40 samples consisting of 3-5 mature angiosperm leaves were collected, from soil level to c. 25 m. The bryoflora checklist is composed of 71 species (46 liverworts; 25 mosses). Of the 21 families, Lejeuneaceae (31 spp.) and Calymperaceae (7 spp.) are richest in genera and species. Lejeunea (6 spp.) and Syrrhopodon (4 spp.) were the most representative genera. The species are predominantly Neotropical (67%) and Pantropical (15%). Eight new records for the state of Pernambuco are presented here: Colura tortifolia (Nees & Mont.) Steph., Cyclolejeunea convexistipa (Lehm. & Lindenb.) A. Evans, Diplasiolejeunea brunnea Steph., D. pellucida (Meissn.) Schiffn., Prionolejeunea denticulata (Weber) Schiffn., Radula mexicana Lindenb. & Gottsche, R. recubans Tayl. and Calymperes smithii E.B. Bartram. The last species is a new record for Northeast Brazil. Ecological and taxonomic comments, substrate types and geographical distribution in Brazil and worldwide are given for these newly recorded species.

Organismos epifílicos em C. macrophyllus foram encontrados em duas parcelas amostradas em julho de 1996, em duas áreas com diferentes graus de perturbação pelo fogo, no Parque Estadual do Rio Doce, MG. Quatorze espécies epifílicas foram encontradas incluindo: 11 líquens, uma alga, uma hepática e um fungo. Os líquens totalizaram 81,8% das espécies na área menos perturbada e 72,7% na mais perturbada. A espécie mais abundante foi Porina epiphylla, que ocorreu em 80,8% dos forófitos da primeira área e em 97% dos forófitos da segunda. O número de folhas cobertas em cada forófito, o grau de cobertura em cada folha e a altura dos forófitos foram significativamente diferentes entre as duas áreas, enquanto que a área foliar e o número de folhas por forófito não o foram. Embora diferenças significativas não tenham sido encontradas na composição específica do epifilo entre as duas áreas, a maior ocorrência de folhas cobertas pelo epifilo pode estar associada à perturbação. A penetração de luz, intensificada pelas clareiras, pode ter aumentado o crescimento em altura do forófito e a colonização das folhas pelos organismos epifílicos.

Epiphyllous organisms on C. macrophyllus were found in two plots examined in July 1996, in two areas with different fire disturbance intensity, in the Parque Estadual do Rio Doce, MG. Fourteen epiphyllous species were found including: 11 lichens, one alga, one hepatica and one fungus. Lichens accounted for 81.8% of the epiphyllous species found in the less disturbed area and 72.7% in the other one. The most abundant species was Porina epiphylla, found in 80.8% of phorophytes in the former area, and 97% in the latter. The number of epiphyllous covered leaves per phorophyte, the coverage in each leaf, and the height of the phorophytes were significantly different for both areas, whereas the leaf area and the number of leaves per phorophyte were not. Although no significant differences were found in the epiphyllous specific composition between both areas, the highest occurrence of epiphylls could be associated with disturbance events. The intensified light penetration caused by gaps in the disturbed forest may have enhanced the phorophyte height growth and the colonization of leaves by epiphylls.

Se comunica la presencia de 153 especies de briófitas y antocerotófitas en la Isla de Cocos (5°32’N, 87°04’W), Costa Rica. Estas son 98 especies de hepáticas en 43 géneros y 10 familias; 54 especies de musgos en 33 géneros y 17 familias y una especie de antocerotófita. Más del 60% de las briófitas tiene amplia distribución en el Neotrópico, 10.4% distribución pantropical, 9.7% pertenecen al elemento Caribe, 1.3% al elemento Centroamericano, 4.5% al Nor-sudamericano, 3.2% son endémicos, 1.3% subendémicos y 1.3% subcosmopolitas. En comparación con Galápagos, la Isla de Cocos muestra una brioflora más tropical, en su mayor parte compuesta por hepáticas foliosas (e. g. Lejeuneaceae, Lepidoziaceae) y una menor representación de hepáticas talosas y musgos. La mayor variedad de hábitats tanto húmedos como secos y expuestos y el area mayor del Archipiélago explican el mayor número de especies de briófitas encontradas en Galápagos. Un análisis del uso de sustratos revela que la mayor parte de las briófitas (46%) utilizan corteza como sustrato, 25% son epífilas, 23% epilíticas y 12% terrestres. Se registró la ocurrencia de briófitas en parcelas de 10 x 10 m con cuadrículas de 30 x 30 cm en distintos hábitats y elevaciones en la Isla. La distribución de briófitas en la Isla coincide con bosque de bajura (0-100 m), bosque secundario de bajura (0-200 m) y bosque montano (200-600 m). La excesiva humedad del ambiente provoca que el cambio altitudinal de la composición de especies en la isla se vea acortado y se presenten unidades montañosas que se encuentran a mayor altitud en areas continentales (efecto "Massenerhebung"). Physantholejeunea portoricensis (Hampe & Gott.) Schust. es un nuevo registro para Costa Rica.

A total of 98 liverwort species (43 genera, 10 families), 54 moss species (33 genera, 17 families) and one species of hornwort have been reported for Cocos Island (5°32’N, 87°04’W), Costa Rica. Over 60% of the bryophytes have a Neotropical or Pantropical distribution, about 10% are Caribbean, and less than 5% are endemic or subendemic. In comparison to the Galapagos Archipelago, Cocos Island harbors a more typical tropical bryoflora with foliose hepatics (e. g. Lejeuneaceae, Lepidoziaceae) constituting the bulk of diversity; fewer thallose liverworts and moss taxa as in the Archipelago were found. A richer habitat variety including wet and dry habitats, as well as its bigger area, seem to account for the higher number of bryophyte species in Galapagos Archipelago. Most bryophytes in Cocos Island are corticolous (46%), the remaining are epiphyllous (25%), saxicolous (23%) or terrestrial (12%). Bryophyte occurrence in eight plots (10 x 10 m) with 20 quadrates (30 x 30 cm) were recorded at different habitats and altitudes (0-600 m). Bryophyte distribution within the island coincides with lowland forest (0-100 m), secondary lowland forest (0-200 m) and montane forest (to 600 m). Physantholejeunea portoricensis (Hampe & Gott.) Schust. is reported as new to Costa Rica.