Greater Diversity of Fungi on Atlantic Slope of Costa Rican Cloud Forest Joshua Obermeyer Department of Biology, St. Cloud State University ABSTRACT Biodiversity in much of the Neotropics is impacted by windward/leeward effects. North East trade winds d ump moisture as they hit the mountains, creating a wetter Atlantic slope, and a drier more seasonal Pacific slope. These factors often favor higher biodiversity for a given taxon on the Atlantic slope. Fungi were collected between the two slopes of the Mon teverde Cloud Forest. A total of 130 fungi specimens were collected from 1550 meters on both slopes of the Continental Divide to 1750 m. As predicted, fungal species richness (Pacific = 24 species, Atlantic = 39 species), abundance (Pacific = 40, Atlantic 3.44) were significantly higher on the Atlantic slope than on the Pacific. It was also found that the two slopes, though in close proximity, have almost entirely different fungal communities. The resu lts indicate that the climatic factors of high precipitation and lower seasonality caused by windward/leeward effects greatly effect fungal species diversity and composition. RESUMEN La biodiversidad en la mayoria de los neotrpicos es afectada por los efectos de la direccin de los vientos. Los vientos del este depositan humedad en la montaas del atlntico, creando un atlntico mas hmedo y un pacfico mas estacional. Estos factores la mayoria del tiempo resultan en mayor diversidad de algunos taxones en la vertiente atlntica. Colect hongos en las dos vertientes del bosque nuboso de Monteverde, Costa Rica. Un total de 130 hongos fueron encontrados en transectos entre 1550 y 1750 msl. Como era de esperarse la riqueza de especies de hongos fue mayor en el atlntico (Pacfico=24 especies, Atlntico=39) al igual que la abundancia (Pcifico = 40, Atlntico pesar de la proximidad de los sitios, tienen dos comunidades muy di stintas de hongos. Los resultados indican que los factores climticos como precipitacin y estacionalidad afectan la composcin de las comunidades de hongos. INTRODUCTION Climate has a dramatic impact on biodiversity, including birds, plants, insects a nd mammals (Blake and Loiselle 2000, Heaney 2001, Samson et al 1997). Windward/leeward effects on precipitation and seasonality impact regional climate and
may have profound effects on species composition and abundance (Heaney 2001). Increased precipitati on often results in high productivity that increases biomass, therefore promoting species diversity (Pianka 1966, Smith 2009). Windward/leeward impacts correlate with many elevational breeding birds, along with climate change causing a shift in species pos itioning and richness in the Monteverde Cloud Forest (Pounds et al. 2006). As is true of their biology in general, little is known about windward/leeward effects and whether certain forest types have greater diversity than others on Neotropical fungal (Lut zoni et al 2004). Fungi are composed of numerous hyphal filaments that generally need w et conditions to live and grow (Alexopoulos et al. 1996, Mata et al. 2003). The direct impact of moisture on fungal growth, and consequently diversity, may be especia lly important in understanding the structure and function of ecosystems. While macro organisms are the least studied organisms on the planet, they play a significant role in nutrient cycling, making them an essential component of tropical communities growi ng on highly weathered soils (Alexopoulos et al 1996, Arora 1986). This study documents differences in the species richness and abundance of macrofungi on the Atlantic and Pacific slopes of Monteverde, Costa Rica. The Monteverde Cloud Forest cradles the C ontinental Divide, which occurs at 1800m. Windward/leeward effects create a mosaic of Holdridge Life Zones that are highly compressed (Haber 2000). On the Pacific slope, Tropical Montane Wet Forest with a 2 3 month dry season gives way to aseasonal Tropica l Montane Rain Forest on the Atlantic slope (Haber 2000). Orographic cloud formation between the two slopes is rising, which as an affect is increasing temperature and may be attributed to current taxon findings between the two slopes (Pounds et al. 2006). Thus, local differences in fungal diversity, as with other taxon, may be attributed to a combination of topographic climatic factors that influence seasonality, precipitation, and temperature (Blake and Loiselle 2000, Haber 2000, Heaney 2001, Samson et al 1997). Because fungi normally thrive in moist conditions, there may be higher species richness, abundance, and diversity of macrofungi on the Atlantic slope. In addition, I will be adding to a fungal database for Monteverde that Rogers (2005) created. M y research differs from Rogers in that I include the Atlantic slope in my study, along with a comparison of the two slopes. Given the importance of fungi to the function of ecosystems, including the biodiverse Tropics; their response to precipitation, seas onality and temperature as influenced by slope and altitude, is vital to our understanding of Cloud Forest function. MATERIALS AND METHODS Study Site This study took place in the Monteverde Cloud Forest located in the Cordillera de Tilaran mountain rang e, Costa Rica. The collections were performed during the end of the dry season in the month of April. The dry season brings slightly cooler temperatures and the trade winds carry moist air from the Caribbean up the Atlantic slope where it cools, and clouds form that dump mist on the Pacific slope (Haber and Zuchowski 2000). Recently, global warming has caused a growing number of dry days and reduced mist frequency, which is more prominent on the Pacific Slope (Pounds et al. 2006). In addition, there was
no rainfall on the Pacific slope, and little on the Atlantic during collection. Two sites were surveyed in this study: A Lower Montane Wet Forest on Pacific slope, and a Lower Montane Rain Forest on the opposite Atlantic slope of the mountain. Both of these s ites are amid a continuous forest, which is part of the 22,000 hectare Monteverde Cloud Forest Reserve system. Pacific Slope: Lower Montane Wet Forest Holdridge (1967) classifies the Pacific slope of the Cordillera de Tilara mountain range to be a Lower Montane Wet Forest. It is characterized by having more seasonality, a slightly higher canopy, and less rainfall (about 3 m) than the Atlantic (Haber 2000). The Tropical Lower Montane Wet Forest is an evergreen forest of intermediate canopy height reaching 25 35 m tall (Haber 2000, Hartshorn 1983). The understory is comparatively open to the Atlantic, with the shrub layer being quite dense, though less than the Lower Montane Rain Forest (Hartshorn 1983, Haber and Zuchowski 2000). The ground is covered mainly with ferns, Begonia and a thick layer of moist, rotting leaves (Hartshorn 1983). The mist that the dry season brings fosters a diverse epiphyte community and maintains the lush character of this forest through the distinct dry period (Haber 2000). Though this is an old pattern of precipitation, and the reduced mist frequency is associated with shifts in populations of birds, reptiles and amphibians (Pounds et al. 2006). Therefore, it is likely the changing climate is affecting fungi species diversity and composition between the Pacific and Atlantic slopes. Atlantic Slope: Lower Montane Rain Forest The Atlantic slope of the continental divide in Monteverde is a Lower Montane Rain Forest (Holdridge 1967). The Atlantic slope receives 4 8 m of rainfall annua lly, is constantly wet, almost aseasonal and has many wind borne clouds coming from the Atlantic Ocean (Haber and Zuchowski 2000). The Tropical Montane Rain Forest is evergreen as well, with a shorter canopy of 20 30m (Hartshorn 1983, Haber 2000). The unde rstory is open, with trees mostly 5 15 m tall, and common tree ferns (Haber 2000). The shrub layer is dense, the ground is open under bamboo, and the trees are densely covered with moss and an abundant epiphyte community mainly consisting of orchids, ferns and a few species of large bromeliads (Hartshorn 1983). Collection and Identification An equal amount of time was spent collecting along elevational bands between 1550 and 1750 m on the Pacific and Atlantic slopes. Daily collections were made within a 100 m band of elevation on a given slope on trails, off trails, and along streams. Attention was mostly focused on collecting the fungi on moist and soft dead wood, among moist leaf litter, and under exposed root systems. Upon spotting a fruiting body, sev eral pictures were taken along with recording the altitude. Next, the fruiting body was carefully removed from the substrate. Data were then taken of color, texture, and any distinctive characteristics of the fruiting body or where it was growing. Fungi we re then carefully wrapped in wax paper and carried in a basket to the lab. Once in the lab, additional pictures were taken to record the size of the fungi, followed by identifying the macrofungi using guides (Arora 1986, Lincoff 1989, Fallas 2005, Mata 200 3, Mata 1999). The unidentifiable mushrooms were kept in wax paper and placed in a refrigerator
for preservation. For further identification, I loaded many photographs onto the Internet for the viewing of fungi specialists (Daniela Lizano, Ignacio Arroyo T rejos, and Melissa Mardones Hidalgo). RESULTS The Atlantic coast had a considerably higher abundance of macrofungi compared to the Pacific (Fig. 1). A total of 90 specimens were collected on the Atlantic, and 40 on the Pacific, with the statistical analy sis being significant ( x 2 = 19.23, df = 1, p = < 0.001). As for species richness, a total of 39 species were collected on the Atlantic, and 24 on the Pacific (Fig. 1). Despite a difference of 15 species, or a more than 50% increase from Pacific to Atlantic richness, the species richness between the two communities was only marginally different statistically ( x 2 = 3.57, df = 1, p = 0.058). FIGURE 1. The abundance of macrofungi found on the Atlantic and Pacific slopes of the Monte verde Cloud Forest, Costa Rica (x 2 = 19.23, df = 1, p = < 0.001), along with the species richness to the left, given for the Atlantic and Pacific (x 2 = 3.57, df = 1, p = 0.058). The Shannon Weiner index and a modified t test were used to compare the diver sity between the Pacific and Atlantic slopes (ZAR 1996). The tests showed significantly higher diversity, and a more evenly distributed community on the Atlantic slope compared to the Pacific (Fig. 2) (t = 7.66, df = 46.08, p = < 0.001).
FIGURE 2. The species relative abundance and community evenness for macrofungi on the Atlantic and Pacific slopes of the Monteverde Cloud Forest, Costa Rica 0.001). Figure 3 shows the families conta ining the highest abundance among the 90 specimens on the Atlantic, Tricholomataceae (typical mushrooms, by far the most diverse family of pale spored agarics) with 22 specimens (24%), 12 in Plutaceae (small to medium sized mushrooms with a central stem, f ree gills, and smooth, pinkish to red spores) (13%), and eight in Xylariaceae (tough club like mushrooms, often looking like a the 18 families found on the Atlantic slop e.
FIGURE 3. The relative abundance of macrofungi specimens collected within each of the 18 families on the Atlantic slope of the Monteverde Cloud Forest, Costa Rica. The pie shows percentages of 90 specimens collected. Figur e 4 shows the families containing the highest abundance of the 40 specimens on the Pacific, Tricholomataceae with nine specimens (22%), nine in Xylariaceae (22%), and seven in Russulaceae (differ from most other gilled mushrooms, having nests of large roun dish cells that are interspersed with the filamentous hyphae, giving the mushroom a characteristic brittle, granular texture. Also, the spores are ornamented with warts, spines, and/or ridges) (17%). These three families with the highest abundance make up 61% of the total specimens found between the11 families on the Pacific.
FIGURE 4. The relative abundance of macrofungi specimens collected within each of the 11 families on the Pacific slope of the Monteverde Cloud Forest, Costa Rica. The pie shows percentages of 90 specimens collected. Trichlomotaceae had more than twice the amount of species as any other family on the Atlantic, totaling in 10 (25.6%) of the 39 different species identified (Fig. 5). Four species were id entified in Coprinaceae (large family of fragile mushrooms with deep brown to black spores and a cartilaginous stem, though characteristics largely vary between Genus) (10%), and three species in Xylariaceae (7.7%), Plutaceae (7.7%, and Polyporaceae (this large diverse group comes in mind boggling multiplicity of shapes and comprise 58.7% of the 39 species collected on the Atlantic.
FIGURE 5. The num ber of macrofungi species identified between the 18 families on the Atlantic slope of the Monteverde Cloud Forest, Costa Rica. The figure also shows the four families that shared species on both slopes: Tricholomataceae (red), Xylariaceae (bright green). R ussulaceae (orange), and Helveliaceae (yellow) (Cn = 0.046, 6% of the total species). Figure 6 shows the species collected on the Pacific slope were found mostly in four families: six in Tricholomataceae (25%), four in Russulaceae (16.7%), four in Xylari aceae (16.7%), and three in Polyporaceae (12.5%). 70.9% of the species collected on the Pacific were in four of the 11 families.
FIGURE 6. The number of macrofungi species identified between the 11 families on the Pacific slope of the Monteverde Cloud Forest, Costa Rica. The figure also shows the four families that shared species on both slopes: Tricholomataceae (red), Xylariaceae (bright green). Russulaceae (orange), and Helveliaceae (yellow) (Cn = 0.046, 6% of the total specie s). It was found that 6% of the species shared the two slopes, or 4 of the 60 species n = 0.046). The families of the four species are color matched in figure 5 and 6. The shared species being: Trcholomataceae sp., Xylaria sp., Russula sp., and a Hexigonia sp. in the family Heleveliaceae. DISCUSSION It was predicted that there would be higher species richness, abundance, and diversity of macrofungi on the Atlantic slope, f ollowing trends seen in other taxon. Data of macro fungi presented here clearly show that the Atlantic slope has higher species richness, a significantly greater amount of abundance, more diversity, a more even distribution, and a largely distinct fungal c ommunity, compared to the Pacific slope. The Atlantic slopes higher abundance and species richness reflect the windward/leeward climatic effect that brings twice the amount of rain, and less seasonality (Haber 2000). Majority of fungi need moisture to gr ow, and where there is more moisture, it is more likely for fungi to be present (Mueller et al. 2007). This was clearly shown on the Atlantic side where it is constantly wet and fungi were easily spotted. The constant rain also moistens dead wood and loose ns top layers of soil, making it an ideal place for fungal spores to establish their mycelium, hyphae and then produce
its fruiting bodies when the conditions are right (Arora (1986, Lutzoni 2004). Further, the Atlantic slope is likely to have higher produ ctivity, and this may promote greater fungal diversity, which in part is because of a more evenly distributed community. The most common families found between the two slopes are large groups and are commonly found in great abundances throughout the Ameri cas. The species found on the conditions to live and grow, along with finer niche partitioning that promotes the wide array of family type. This diversity would be expected to be even greater in the following rainy season months. The fungi found on the Pacific side appeared to be dry and plain, which suggests they can survive during lon g dry periods, as it did not rain throughout the collections. The fungal families on the Pacific are also likely to be different than in the past, due to less precipitation and more seasonality caused by climate change. Though, further examination and more time is needed to find a trend in the type of families found on either slopes. Many of the macrofungi found in this study were to be out of season, not found in Monteverde, or growing on different substrates according to identification guides. This shows the little information known about macrofungi and the climatic conditions that require them to grow. Also, each species develops different, and largely unknown tolerances for pH, substrate nutrients, precipitation, and temperature that will allow them to s urvive. differs from the Lower Montane Wet Forest, yet they share the same continuous forest with only opposing mountainsides. The high diversity compacted into a fairly small spac e suggests high fungal diversity on windward sloped Cloud forests. Windward sloped fungal communities are likely maintained by high and constant precipitation, along with increased biomass, and greater stability that allows for finer niche partitioning. Th e Pacific slope has been maintained by a pronounced wet season and mist from the Atlantic during the dry season. Climate change has decreased the amount and frequency of precipitation, and the forest experiences edge effects at lower altitudes, more light gaps, and the plant layer is less dense. Given these characteristics, the Pacific slope is more vulnerable to climate change that will disrupt the fungal community. Considering the lack of study on fungi and whether or not certain forest types have greater diversity than others, especially windward/leeward climatic effects on its diversity, the research presented offers information on the fungi community found between the Atlantic and Pacific slopes of Monteverde, Costa Rica. The constant precipitation of t he Atlantic slope shows a rich and complex fungal community, while the Pacific is experiencing local impacts that are likely due to global climate change. This experiment opens up opportunity for the study of climatic effects on fungi, taxonomic informatio n, and the overall importance that fungi have on the function of the Cloud Forest ecosystem. ACKNOWLEDGEMENTS I would like to thank Alan Masters for his advice and guidance in formulating and carrying out this study. I would like to thank Yemen Araya for his assistance in statistical analysis. I would like to thank Pablo Alan and Jose Calderon for their advice. I would like to thank Daniela Lizano, Ignacio Arroyo Trejos, and
Melissa Mardones Hidalgo for their help with identification. Lastly, I would like to thank the Estacion Biologica for the use of their forest and lab. LITERATURE CITED ALEXOPOULOS, C.J., C.W. Mims and M. Blackwell. 1996. Introductory Mycology, Fourth Edition, Wiley. ARORA, D. 1986. Mushrooms Demystified Ten Speed Press. BLAKE, J .G., and B.A. Loiselle. 2000, Diversity Of Birds Along an Elevation Gradient in the Cordillera Central, Costa Rica. The Auk, Vol. 117: 663 686. FALLAS, R.C. 2005. Hongo de Costa Rica UNA. HABER, W.A. 2000. Plants and Vegetation in Nadkarni, N.M and heekw right, N.T (eds) Monteverde : Ecology and Conservation of a Tropical Cloud Forest, Oxford Univ. Press. HABER, W.A., W. Zuchowski and E. Bello., 2000. An Introduction to Cloud Forest Trees Monteverde, Costa Rica Second Edition, Mountain Gem. HARTSHORN, G.S. 1983. Plants. Costa Rican Natural History Ed. Daniel Janzen. Chicago: University of Chicago Press. 118 157. HEANEY, L.R. 2001. Small Mammal Diversity Along Elevational Gradients in the Philippines: an Assesment of patterns and hypo theses. Global Ecology and Biogeography, Vol 10: 15 39. HOLDRIDGE, L.R. 1967. Life Zone Ecology San Jose: Tropical Science Center. LINCOFF, G.H. 1989. Mushrooms Fireside. LUTZONI, F., F. Kauff, C.K. Cox, D. Mclaughlin, G. Cel io, B. Dentinger, and Mahajabee. 2004. Assembling the Fungal Tree of Life: Progress, Classification, and Evolution of Subcellular Traits. American Journal of Botany. Vol. 91: 1446 1480. MATA, M, R., Halling and G.M. Mueller. 2003. Costa Rica Macrofungi Volume 2, INBio. MATA, M. 1999. Costa Rica Mushrooms Volume 1, INBio. MUELLER, G.M, J.P. Schmit, P.R. Leacock, B. Buyck, J. Cifuentes, D. E. Desjardin, R.E. Halling, K. Hjortstam, T. Iturriaga, K. Larsson, D.J. Lodge, T.W. May, D. Minter, M. Rajchenber g, S.A. Redhead, L. Ryvarden, J.M. Trappe, R. Watling and Q. Wu. 2007. Global diversity and distribution of macrofungi. Biodiversity and Conservation, vol 16: 37 48. PIANKA, ERIC R. 1996. Latitudinal gradients in species diversity: a Review of Conepts. T he American Naturalist 100.910: 33 46. POUNDS, A.J., M.R. Bustamante, L.A. Coloma, J.A. Consuegra, M.P.L. Fogden, P.N. Foster, E.La Marca, K.L. Masters, A. Merino Viteri, R. Puschendorf, S.R. Ron, G. A. Sanchez Azofeifa, C.J. Still and B.E. Young. 2006 Widespread amphibian extinctions from epidemic disease driven by global warmig. Nature, vol 439: 161 167. ROGERS, C. 2005 Database of the Macrofungi of the Monteverde Reserve. Fall 2005. CIEE Tropical Ecology Program. SAMSON, D.A. Rickar, E.A. Gonzales, P.C. 1997. Diversity and Abundance along an Elevational Gradient in the Philippines. Biotropica, Vol 29: 349 363. SMITH, JUSTINE. 2009. Rodent Diversity in Relation to Atlantic and Pacific Slopes in a Neotropical Cloud Forest. Fall 2009. CIEE Tropical Eco logy Program. ZAR, J.H. 1996. Biostatistical Analysis Third Edition. Prentice Hall.
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Una mayor diversidad de hongos en la vertiente atlntica del Bosque Nuboso de Costa Rica
Greater diversity of fungi on Atlantic slope of Costa Rican cloud forest
Biodiversity in much of the Neotropics is impacted by windward/leeward effects. North
East trade winds dump moisture as they hit the mountains, creating a wetter Atlantic
slope, and a drier more seasonal Pacific slope. These factors often favor higher
biodiversity for a given taxon on the Atlantic slope. Fungi were collected between the two slopes of the Monteverde Cloud Forest. A total of 130 fungi specimens were
collected from 1550 meters on both slopes of the Continental Divide to 1750 m. As predicted, fungal species richness (Pacific = 24 species, Atlantic = 39 species), abundance (Pacific = 40, Atlantic = 90), and diversity (H Pacific = 1.55, H Atlantic = 3.44) were significantly higher on the Atlantic slope than on the Pacific. It was also found that the two slopes, though in close proximity, have almost entirely different fungal communities. The results indicate that the climatic factors of high precipitation and lower seasonality caused by windward/leeward effects greatly effect fungal species diversity and composition.
La biodiversidad en la mayora de los neotrpicos es afectada por los efectos de la direccin de los vientos. Los vientos del este depositan la humedad en las montaas del atlntico, creando un atlntico ms hmedo y un pacfico estacional ms seco. Estos factores a menudo favorecen una mayor biodiversidad a un determinado taxn en la vertiente atlntica. Colect hongos en las dos vertientes del bosque nuboso de Monteverde, Costa Rica. Un total de 130 hongos fueron encontrados en los trayectos entre los 1550 y 1750 metros. Como era de esperarse la riqueza de especies de hongos fue mayor en el atlntico (Pacfico=24 especies, Atlntico=39) al igual que la abundancia (Pacfico = 40, Atlntico =90) y la diversidad (H Pacfico=1.55, H Atlntico=3.44). Tambin se encontr que a pesar de la proximidad de los sitios, tienen dos comunidades muy distintas de hongos. Los resultados indican que los factores climticos como la precipitacin y la estacionalidad afectan la composicin de las comunidades de hongos.
Text in English.
Costa Rica--Puntarenas--Monteverde Zone
Costa Rica--Puntarenas--Zona de Monteverde
Tropical Ecology Spring 2010
Ecologa Tropical Primavera 2010
t Monteverde Institute : Tropical Ecology