1 Macrofungal abundance and distribution during the wet season in Monteverde, Costa Rica Jason Prior Matthew Prior Department of Biology, Central Michigan University ABSTRACT Fungi are a diverse group of organisms that play a myriad of important roles in all ecosystems. From providing medication for humans to interacting mutu alistically with tropical plants, this kingdom offe rs benefits to many organisms. However, there is much to be learned from this highly understudied collec tion of species. This study focused on expanding a data base of fungi in and around the Estacin biolgica de Monteverde. Each fungus collected was photographed and data such as date and time, weather conditions microhabitat conditions, and morphological characte ristics were recorded and entered into the database Milagro Mata identified 25 of the collected species to the family, genus, or species level. By expandi ng this database we hope to increase knowledge and interest in the local macrofungal communities. RESUMEN Los hongos son un grupo diverso de organismos que j uegan una mirada de papeles importantes en todos los ecosistemas. Desde proveer medicinas a los ser es humanos hasta interactuar mutualsticamente con plantas tropicales, este reino ofrece beneficios a muchos organismos. No obstante, hay mucho que aprender de esta coleccin de especies que ha sido muy poco estudiada. Este estudio se enfoc en expa ndir la base de datos de los hongos en, y alrededor de l a Estacin biolgica de Monteverde. Cada hongo colectado fue fotografiado y otros datos como la fe cha, la hora, las condiciones del tiempo, del microhabitat, y las caractersticas morfolgicas, f ueron registrados en una base de datos. Milagro Mat a identific 25 de las especies colectadas a nivel de familia, gnero, o especie. Al expandir esta base de datos, se espera aumentar el conocimiento y el inters en las comunidades de los macrohongos locales. INTRODUCTION Fungi are a group of living organisms that are dive rse, both in their methods of obtaining nutrients as well as their taxonomic nomenclature. This kingdom contains many species ranging from unicellular to multicellular organisms Although fungi are often thought to be plants, members of this kingdom are taxonomicall y more closely related to animals. Unlike plants, fungi lack light-harvesting organell es, such as chloroplasts, that are necessary for photosynthesis; thus they must obtain nutrients and energy from other organisms, living or dead, through absorption (Mata 1999). Most fungi are saprophytic and obtain energy by attacking organisms that have died from some other cause. Other fungi are parasitic and acquire nourishment from li ving organisms, but do not kill their host. Some fungi are necrophytic and attack a dyin g organism with toxins and enzymes, killing it in the process. Some symbiotic fungi in teract with living organisms mutualistically, usually exchanging water and nutri ents (fungi) for carbohydrates (plants), increasing the fitness of both organisms (Alexopoul os et al. 1996). In Costa Rica, approximately 2,000 species of fungi have been docu mented, and it is estimated that a total of 40,000 to 70,000 species of fungi inhabit this country (Mata 1999). As of 1991,
2 approximately 100,000 species of fungi were describ ed worldwide, and as many as 1.5 million were estimated to exist. Next only to inse cts, this kingdom is thought to have the second highest species richness (Hawksworth 1991). Fungi play a wide range of roles within all of the ecosystems distributed around the world (Mata 1999). Fungi cycle nutrients resu lting from dead plant and animal tissue, thus allowing the reuse of limited biotic a nd abiotic resources (Rossman et al. 1998). Mycorrhizal fungi interact mutualistically with many tropical plants. Since tropical soils tend to be rather infertile, nutrien t uptake can be difficult for many plants. Fungi help these tropical plants obtain the nutrien ts necessary to grow and reproduce. In return, the mycorrhizal fungi absorb carbohydrates from the plant (Mata 1999). Fungi also provide food for many organisms. For instance leaf-cutting ants bring leaves to their nests where they farm fungi to use as a prima ry food source (Stevens 1983). Humans also use fungi to aid in the preparation of foods such as cheese, beer, wine and bread. Conversely, fungi may have a negative impac t on an environment. Some fungal species are known to act as pests by devastating cr ops in agricultural communities around the world (Griffith et al. 2000). Within the realm of human beings, many fungi are used medicinally. Penicillin and many other heavily use d antibiotics are synthesized from fungal compounds and aid humans in the recovery fro m many bacterial infections (Rossman et al. 1998). With such a low estimated n umber of species discovered, and such a high medicinal potential, it is imperative t hat fungal research continues. We believe that it is important to make an inventor y and catalogue the abundance and distributions of fungal species located in Mont everde for many reasons. In order to learn about an organism, one must have an organized system of naming and classifying organisms of a similar taxa (e.g. fungi). With an organized database, information about the fungal diversity of Monteverde will be accessib le to the public and will increase interest in this area of study. This will result i n more research being conducted, which will lead to a better overall understanding of fung i. Similarly, it is important to know the fungal diversity of this area in order to determine if there is any medicinal or scientific significance associated with the local fungi. If a certain species of fungus is endemic to Monteverde it could be beneficial to conserve that area. On the other hand, if a species of fungus is threatening the survival of another endan gered plant or animal species, it would be advantageous to take the necessary precautions t o prevent further destruction. Not all fungi produce fruiting bodies at the same t ime throughout the year; thus it is important to create an inventory of fungi spanni ng all seasons. In Monteverde, seasons are characterized by differences in precipitation a nd cloud types (Clark et al. 2000). As precipitation increases, soil moisture levels also increase. All species of fungi require water, however, some require more than others. In unfavorable conditions (e.g. water reduction), fungi produce fruiting bodies that rele ase spores to increase an individualÂ’s fitness. Thus, precipitation levels can affect the abundance and growth of fungi. Cloud cover can also play a role in fungal species compos ition. Increased cloud cover will reduce the amount of solar heat that warms the soil thus creating a difference in soil conditions throughout the seasons (Pace 1998). Beca use precipitation and cloud types vary across seasons, this may result in the presenc e of certain species in one season and different fungi in another season. We hypothesized that the fungal species found between July and August in Monteverde are affected by these abiotic parameters. Therefore, we predicted that the community composition observed d uring the wet season (in which we
3 conducted our studies) would differ from the compos ition of species catalogued at other times of the year by previous CIEE students. MATERIALS AND METHODS Study Sites This study was conducted along the trails around th e Estacin biolgica de Monteverde. Established trails acted as transects with the samp le collection being restricted to ten meters on either side of the trail (as in Rogers 20 05). Our study focused on the areas of the trails that were previously studied by CIEE stu dents. Of these areas, the trails closest to the Station were the primary study sites. Data were collected from July 15 to August 2, 2006. Collection Methods Each species was photographed with a digital camera and specific field data were recorded. Microhabitat conditions such as elevatio n, relative humidity, and percent canopy coverage were noted, using an altimeter, rel ative humidity meter, and a spherical densiometer, respectively. Other data, such as the abundance, growth habit, and the observed substrate were also recorded on a specimen data sheet that was created by Rogers (2005). After specimen collection, addition al morphological characteristics were recorded, such as the shape of pileus, presence and shape of stipe, color of sample, and texture of the margin were documented on the specim en data sheet (as in Rogers 2005). Once all data were gathered and organized, our resu lts were statistically analyzed and compared to the results produced by Rogers (2005) a nd Williams (2006). The Sorenson Qualitative Index was used to look for similarity i n community composition between our results and the results found by Rogers (2005) and Williams (2006), individually. Identification While in the field, we observed fungi that appeared to be similar to species previously documented by Rogers (2005) and Williams (2006). Th ese specimens were compared to the photographs provided in the database for identi fication. When a species was determined to be part of the database, its presence was noted and new pictures were taken. No other data were recorded for previously d ocumented species. After each collection, an attempt was made to ident ify each species using Macrohongos de Costa Rica (Mata 1999; Mata et al. 2003). To aid in the iden tification, spore prints were successfully produced for some sp ecimens. Once all data collection were complete, two or three photographs of each spe cimen were compiled and sent to Milagro Mata, a mycologist at the National Biodiver sity Institute, for identification. Database Expansion In an effort to expand the current fungal database and create a more complete collection of the macrofungi of Monteverde, we followed the pr otocol set by Rogers (2005). Since
4 the fruiting bodies found in the wet season were no t catalogued, we thought it was necessary to include this part of the wet season in the database. All data collected were compiled into specimen data sheets for each species (as in Rogers 2005). In order to expand the database, slides were made that included the same data parameters used by Rogers (as in 2005) in the original database. Phot ographs and taxonomic nomenclature were also included in the slides. RESULTS Altogether there were 35 fungal species gathered be tween July 15, 2006 and August 2, 2006. However, five of the species were determined to be part of the database already. Therefore, we recorded data for 30 new species. We collected one fewer specimen than did Williams (2006) and five more specimens than di d Rogers (2005). Out of these 35 species, four were also identified and described by Rogers (2005) and five were identified and described by Williams (2006). Using the Sorenso n Qualitative Index, overlap between the species that we gathered and the specie s collected by Rogers (2005) and Williams (2006) was calculated to be Cs = 0.12 and Cs = 0.14, respectively. After sending the photos to Milagro Mata for identi fication, we received a list of all the available taxonomic information on the spec imens that we collected. There were 25 species that we received taxonomic information f or. Eight out of those 25 were classified at the family level. Eleven were identif ied at the genus level, while six were identified to the species name. Of the identified s pecies, three were in common with Rogers (2006) and one with Williams (2006). Howeve r, in the field, we identified one other species in common with Rogers (2005) and four other species with Williams (2006). These five specimens that were determined t o be previously part of the database were not sent for identification. Five species were unidentifiable. The most common substrate that fruiting bodies wer e found on was dead or rotting wood (19 out of 35; 54%). Twelve out of 35 (34%) were found growing on soil, three out of 35 (9%) were found growing on dead lea ves, and one species (3%) was found to have fruiting bodies on both a decaying lo g and the soil (Figure 1). The most common morphological form of fruiting bodies collec ted was a toadstool (23 out of 35). Six species of shelf fungi, two species of each, pu ffballs and coral fungi, and one species of slime fungus were collected (Figure 2). Most of the toadstool species that we collected had a pileus of less than two centimeters (15 out o f 23; 65.2%).
5 n rr rn rrrn Figure 1. The percentage of fruiting bodies that we re found growing on the four different substrates observed (dead leaf, dead wood, soil, de ad wood and soil). n rrrn n nn rn Figure 2. The different morphological forms of fung al fruiting bodies observed and the relative abundances of each gathered from July 15, 2006 to August 2, 2006.
6 DISCUSSION As we predicted, species composition was different in July and early August than at any other catalogued time (Rogers 2005; Williams 2006). However, there was overlap in species present during this study with species pres ent both in the late wet season (Rogers 2005) and the late dry season (Williams 2006). As our Sorenson Qualitative Indices showed, our collection of species was slightly more similar to the catalogued species from Williams (2006) than from Rogers (2005). Acco rding to Clark et al. (2000), as elevation increases, the amount of precipitation al so increases. Although WilliamsÂ’ study took place in the dry season, the higher elevations that were studied may have had a similar amount of precipitation to our lower elevat ion study during the wet season. According to Mata (1999), toadstools are the most w ell known morphological form of fungi. This is supported by our results, w hich show that 24 of the 35 fungi that we found were in the form of a toadstool. Shelf fu ngi comprised the second largest morphological form, accounting for six of our 35 sp ecimens. According to Alexopoulos (1996), many shelf fungi produce fruiting bodies th at are visible for a long period of time. Since many live for a long time, we may have seen s ome of the same specimens that were found by Williams in May 2006, only a couple months prior to our study. As shown in our results, the most common substrate fungi were found on was decaying wood. This coincides with Rossman et al. (1998) who found that fungi are the major decomposer of woody tissues in tropical ecosy stems. Since tropical soils tend to be infertile, nutrient rich decaying wood may provi de a better substrate for the growth and fitness of fungi. Having the taxonomic nomenclature available, we com pared our results to that of Williams (2006) and that of Rogers (2005). There we re five species not identifiable, so there may have been more overlap with previous stud ies. The specimens that were not classified taxonomically appeared nothing like any of the species previously described. Therefore, after receiving this information none of our data changed. This was not expected, but promotes new ideas about the differen ce of fungal diversity throughout the seasons in Monteverde. It suggests that there may be a lot more macrofungal species occurring in Monteverde than previously thought. T hese new findings also suggest that the seasonal abiotic differences truly do harbor a profound impact on fungal diversity. ACKNOWLEDGMENTS Throughout our independent project, there were many individuals who helped us complete our study. Fir st of all, we would like to thank Karen Masters for he lping us organize our project and giving us advice along the way. We would also like to thank Carmen Rojas for helping us communicate with Milagro Mata and helping us translate our abstract. Milagro Mata, t hank you for volunteering your time to help us iden tify the fungi that we collected. Camryn Pennington and Tom McFarland, thanks for answering all of our questions and obtaining all the supplies and equipm ent that we needed. Finally, we would like to than k the Estacin biolgica de Monteverde staff for keeping clean sheets on our beds and hot meals on the table LITERATURE CITED
7 Alexopoulos, C.J., C.W. Mims, and M. Blackwell. 199 6. Introductory Mycology ed. 4. John Wiley & Sons, Inc., U.S.A., pp. 29,39, 563-574 Clark, K.L., R.O. Lawton, and P.R. Butler. 2000. The Physical Environment In: Monteverde: Ecology and Conservation of a Tropical Cloud Forest N. M. Nadkarni and N. T. Wheelwright, eds. Oxford Univers ity Press, New York, NY, pp. 15-20. Griffith, K., D.C. Peck, and J. Stuckey. 2000. Agriculture in Monteverde: Moving Toward Sustainability In: Monteverde: Ecology and Conservation of a Tro pical Cloud Forest N. M. Nadkarni and N. T. Wheelwright, eds. Oxford University Press, New York, NY, pg. 402. Hawksworth, D.L., P.M. Kirk, B.C. Sutton, and D.N. Pegler. 1995. Ainsworth and BisbyÂ’s Dictionary of the Fungi (Including the Lich ens) 8th ed. CAB International, Wallingford, pg. 616. Mata, M. 1999. Macrohongos de Costa Rica Vol. 1. Instituto Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, Costa Rica, pp. 11-244. Mata, M., R. Halling, and G.M. Mueller. 2003. Macro hongos de Costa Rica Vol. 2. Instituto Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, Costa Rica, pp. 9-231. Pace, G. 1998. Mushrooms of the World Firefly Books Ltd., Buffalo, NY, pg. 10. Rogers, C. 2005. Database of the Macrofungi of the Monteverde Reserve. CIEE, Monteverde, Costa Rica, pp. 260-70. Rossman, A.Y., R.E. Tulloss, T.E. OÂ’Dell, and R.G. Thorn. 1998. Protocols for an All Taxa Biodiversity Inventory of Fungi in a Costa Ric an Conservation Area. Parkway Publishers, Boone, NC, pg. 1. Stevens, G.C. 1983. Atta cephalotes (Leaf-cutting Ants). In: Costa Rican Natural Histor y D. H. Janzen, ed. The University of Chicago Press, Chicago, IL, pp. 688-91. Williams, S. 2006. Dry Season Macrofungi of the Mon teverde Reserve. CIEE, Monteverde, Costa Rica, pp. 1-8.
8 Hygrocybe miniataÂ• Family: Tricholomataceae Â• Collection date: July 18, 2006 Â• Collection Time: 12.10 pm Â• Weather Conditions: Mostly sunny, few clouds. Â• Microhabitat conditions at collection site:-Type of substrate : decaying woodElevation: 1580 mObserved abundance: 10-15Observed growth habit: clumpedRelative Humidity: 75 % Â• Morphological Characteristics (of collectedsamples): Â–Basic form: ToadstoolÂ–Shape of pileus: HemisphericalÂ–Diameter of pileus:17 mmÂ–Surface texture of pileus: velvety and smoothÂ–Presence of stipe: yes; 50 mmÂ–Shape of stipe: cylindrical to equalÂ–Color of mature sample: redÂ–Color of young specimen: slightly orangerÂ–Shape of margin (using longitudinal section): decurved Â–Texture of margin: plicate-striateÂ– Hymenium or fertile surfacecharacteristics: Â•Color: orangish pinkÂ•Texture: gilledÂ•Type of juncture with stipe: decurrent Â•Space between gills: subdistantÂ•Color of spore print: brown Figure 3. An example of PowerPoint slides made for Hygrocybe miniata This shows the data that were typically included in the database s lides and the general organization of the information and pictures. Information such as taxo nomic nomenclature, collection date and time, weather conditions, microhabitat conditio ns, and morphological characteristics, (e.g. size and color, etc.) are included.
9 1. Polyporus tenuiculus 16. Morpho 3 2. Morpho 1 17. Morpho 4 3. Coprinus disseminatus 18. Tricholomataceae 4. Tricholomataceae 19. Pluteus sp. 5. Morganella fuliginea 20. Pluteus sp. 6. Morpho 2 21. Tricholomataceae 7. Oudemansiella cannarii 22. Tricholomataceae 8. Crepidotus sp. 23. Tricholomataceae 9. Clavaria sp. 24. Lepiota sp. 10. Tricholomataceae 25. Scleroderma verrucosum 11. Hygrocybe miniata 26. Tricholomataceae 12. Psathyrella sp. 27. Coprinus sp. 13. Marasmius sp. 28. Gymnopus sp. 14. Mycena sp. 29. Morpho 5 15. Tremella sp. 30. Tricholomataceae Figure 4. A map of the trails around the Estacin biolgica de Monteverde with a numbered list of fungi collected. The speciesÂ’ num ber corresponds to the numbered location on the map which shows where each specimen was found. Monteverde, Costa Rica, July 15 Â– August 2, 2006. 1 2 5 6 7 8 9 10 11 12 13 16 17 21 22 23 24 26 27 30
10 APPENDIX A: Specimen data sheet for fungal inventor ies at Monteverde. Specimen number: Date: Time: Weather conditions: Microhabitat Conditions: Type of substrate: Relative humidity: Canopy cover (%): Elevation: Abundance: Observed growth habit: Morphological Characteristics: Basic form: Shape of pileus: Diameter of pileus: Presence of stipe: Shape of stipe: If no stipe, length and width of pileus: Color of mature sample: Color of young sample: Shape of margin (using longitudinal section): Texture of margin: Surface texture of pileus: Color of Context: Hymenium or fertile surface characteristics: Color: Texture: Type of juncture with stipe (for those with l amellae): Space between gills: Stipe characteristics: Size: Color: Position: Presence of annulus: Location of annulus: Color of annulus: Presence of volva: Shape of volva: Color of volva: Texture of volva: Color of spore print: Color of bruising: Notes:
11 APPENDIX B: Morphological Terminology Keys (Adapted from Mata 1999) Pileus shapes: Textures of the pileus margin: Lamellae spacing: Stipe positions: Shapes of pileus margins: Stipe shapes: Lamellae attachments:
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Prior, Jason Prior, Matthew
Abundancia y distribucin de los hongos durante la estacin lluviosa en Monteverde, Costa Rica
Macrofungal abundance and distribution during the wet season in Monteverde, Costa Rica
Fungi are a diverse group of organisms that play a myriad of important roles in all ecosystems. From providing medication for humans to interacting mutualistically with tropical plants, this kingdom offers benefits to many organisms. However, there is much to be learned from this highly understudied collection of species. This study focused on expanding a database of fungi in and around the Estacin biolgica de Monteverde. Each fungus collected was photographed and data such as date and time, weather conditions, microhabitat conditions, and morphological characteristics were recorded and entered into the database. Milagro Mata identified 25 of the collected species to the family, genus, or species level. By expanding this database we hope to increase knowledge and interest in the local macrofungal communities.
Los hongos son un grupo diverso de organismos que desempean una gran variedad de funciones importantes en todos los ecosistemas. Desde proveer medicinas a los seres humanos hasta interactuar mutualisticamente con las plantas tropicales, este reino ofrece beneficios a muchos organismos. No obstante, hay mucho que aprender de esta coleccin de especies que ha sido poco estudiada. Este estudio se enfoco en expandir la base de datos de los hongos en y alrededor de la Estacin Biolgica de Monteverde.
Text in English.
Macrofungi--Costa Rica--Estacin Biolgica de Monteverde
Fungi--Taxonomy and ecology
Microhabitats--Costa Rica--Puntarenas--Monteverde Zone
Macrohongos--Costa Rica--Estacin Biolgica de Monteverde
Hongos --Morfologa --Costa Rica
Tropical Ecology 2006
Ecologa Tropical 2006
t Monteverde Institute : Tropical Ecology
M39-00066--[Macrofungal abundance and distribution during the wet season in Monteverde, Costa Rica--supporting materials--images][ppt]; M39-00067-- [Macrofungal abundance and distribution during the wet season in Monteverde, Costa Rica--supporting materials--additions to the database of fungi at the Estacion biologica de Monteverde (Summer 2006)]
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