Mycosphere Doi 10.5943/mycosphere/2/5/8/ New additions to the myxobiota of Peru 1 2* 3 4 Rojas C , Stephenson SL , Pavlich M 1 Instituto de Investigaciones en Ingeniería, Universidad de Costa Rica, San Pedro de Montes de Oca 11501-2060, Costa Rica – corresponding author crojas@fungica.com 2 Escuela de Biologia, Universidad de Costa Rica, San Pedro de Montes de Oca 11501-2060, Costa Rica 3 Department of Biological Sciences, University of Arkansas, Fayetteville AR 72701-1201, United States of America – slsteph@uark.edu 4 Departamento de Ciencias Biológicas y Fisiológicas, Universidad Peruana Cayetano Heredia, Lima, Peru – magdalena.pavlich@upch.pe Rojas C, Stephenson SL, Pavlich M 2011 – New additions to the myxobiota of Peru. Mycosphere 2 (5), 583-592, Doi 10.5943/mycosphere/2/5/8/ Recent efforts to study myxomycetes in many areas of the world have resulted in a considerable increase in the number and quality of regional species lists, especially during the last decade. However, in South America certain countries such as Peru continue to be highly understudied with respect to this group of organisms. During a series of surveys carried out in 2007 in several different areas of Peru, 49 species of myxomycetes not previously known for this country were recorded. Basic ecological information was also collected and analyzed in the context of available data on myxomycetes for the Neotropics. With the new species reported herein, the myxobiota of Peru is increased to 80 species, which is still a low number when the natural and biogeographical characteristics of Peruvian landscapes are considered. Key words – biodiversity, biogeography, microbial ecology, mycetozoans, myxogastrids, South America. Article Received 27 October 2011 Accepted 2 November 2011 Published online 9 November 2011 *Corresponding author: Carlos Rojas – e-mail – crojas@fungica.com Introduction ecosystems (Stephenson 2003), there are still The mycetozoans make up a group of large gaps in the distributional information protists within the supergroup Amebozoa (see available for most species. Adl et al. 2005). The myxomycetes (or In spite of the former constraint, in myxogastrids), one of the subgroups of recent years several authors have provided mycetozoans (Pawlowsky & Burki 2008), are evidence to explain the distribution of primarily characterized by the capacity to form myxomycetes at large scales under a type of a vegetative, multinucleate but essentially moderate endemicity model (Stephenson et al. single-celled structure known as a plasmodium. 2008, Rojas et al. 2011b). However, before The myxomycetes also exhibit the capacity of meaningful distributional patterns can be producing spores within relatively large (for a constructed using the techniques of group of microorganism) fruiting bodies, computational biology, still more field-based showing an ecological strategy for dispersion information on the occurrence of species in that seems to be analogous to the one used by different parts of the world is needed. most macrofungi. Even though they are now Moreover, the importance of the collections known to occur in practically all terrestrial obtained from the surveys that have been 583 carried out is largely justified if plans are made that is outside the scope of this paper. to study various aspects of their biology such However, in an effort to close the gap on as population genetic makeup, gene flow and research projects and availability of regional the prevalence of apomixis in more detail in the data within the Neotropics, the present study future. For this reason, basic distributional was carried out as part of a larger research studies that permit researchers to extend effort directed towards an investigation of the species ranges and describe new species are dynamics of myxomycetes in different forest still essential if the determination of species types of Peru. The primary objective of this diversity, one of the key questions for science paper is to provide an updated overview of the to answer during of the first part of the current myxomycete biota of the country that can serve century (see Pennisi 2005), is to be addressed. as a source of baseline information for future Since the publication of the only analyses and studies on this group of monograph devoted specifically to the organisms, both locally and for the entire myxomycetes of the Neotropics (Farr 1976), Neotropics. there have been more than 250 articles on these organisms published for the region (updated Methods from Lado & Wrigley de Basanta 2008). A The study described herein was carried number of understudied ecosystems and out in 2007 in Peru. All species names follow particular microhabitats have been examined the nomenclatural treatment of Lado (2005–- during this time (e.g., Schnittler & Stephenson 2011) except for Stemonitis smithii, for which 2002, Estrada-Torres et al. 2009). Interestingly, the original protologue is provided. Due to its the research effort has not been homogeneous ecological similarity to the myxomycetes, the throughout the Neotropics, and published genus Ceratiomyxa was considered in the studies from some countries, like Mexico, present study. The morphological concept of represent almost 50% of the total, whereas species was used for all of the myxomycetes investigations in understudied areas like Peru reported herein. represent only about 1% of the research effort carried out over the entire region (Lado & Study areas Wrigley de Basanta 2008). Five study areas representing different Such disparity in scientific productivity ecological situations in the central-southern among Latin American countries has been portion of Peru were selected for this study. thoroughly analyzed from the socio-economic These areas represent an east-west transect that perspective (e.g., Romero et al. 2009). extends from the southwestern section of the However, in the case of myxomycete studies, it Amazon basin to the Pacific coast, crossing the is also a partial product of the presence of entire Andean region (Fig. 1). Each area specialists in only a handful of countries in the encompassed a series of two to five collecting region and the low availability of funding for sites selected a priori using GIS-based research on less well-known and generally information in order to increase the probability obscure (i.e., cryptic) groups of organisms. For of including different microenvironments. In instance, in the case of Peru, the last published this way, all collecting sites within one study article specifically on myxomycetes, which area differed in terms of the dominant appeared in 1994, did not involve the vegetation, structural characteristics of that participation of local researchers (see vegetation and the extent of human influence. Stephenson & Mitchell 1994). That is also the Located in the Department of Madre de case of two other recent studies that report Dios, the first study area (1) was Centro de myxomycetes from Peru (i.e., Wrigley de Investigación y Capacitación Rio Los Amigos Basanta & Lado 2005, Wrigley de Basanta et (hereafter referred to as Los Amigos, Figs 2a al. 2008). and 2b). This area corresponds to a land Unfortunately, this type of situation is concession administered by the Amazon the result of the circumstances related to the Conservation Association, intended to protect social, academic and scientific conditions of 453 hectares of lowland Amazon moist forest particular countries in the Neotropics, a subject along the Madre de Dios river watershed. The 584 Mycosphere collecting sites in this area were located For field collections, both the opportunistic between 12˚ 33’–35’ S latitude and 70˚ 5’–6’ method (Cannon & Sutton 2004) and a survey W longitude at elevations between 234–282 m on decaying wood were used to look for above sea level (m asl). Two of the more specimens. In the one instance, fruiting bodies distinctive habitats present in this area are are searched for in the field in a random seasonally flooded forests and Mauritia fashion without any limitations imposed by a flexuosa-dominated swamps. particular sampling scheme. This method is The first two study areas on the Andean very useful for recovering singletons and section of the transect are part of the Peruvian doubletons represented in the assemblage being Yungas ecosystem, a lower montane studied, especially in the case of organisms transitional area between lowland Amazon such as the myxomycetes, which are forests and high-elevation wet Puna grasslands characterized by rather heterogeneous fruiting on the eastern side of the Andes. These were patterns. For the survey of decaying wood, (2) the Wayqecha Cloud Forest Biological only used in Los Amigos, a series of dead logs Station (Wayqecha); collecting sites between on the forest floor was selected at random 13˚ 10’–11’ S latitude and 71˚ 35’–36’ W during each of the collecting periods, and longitude at elevations between 2805–2966 m fruiting bodies of myxomycetes were searched asl; and (3) the town of Aguas Calientes or for in an intensive manner along each log. This Macchu Picchu (Macchu Picchu, Fig 2d); technique is useful in forest environments, due collecting sites at 13˚ 9’–10’ S latitude and 72˚ to the relatively high moisture levels observed 31’–32’ W longitude at elevations between within forested zones and the relationship of 2050–2150 m asl. Also on the Andes, but evapotranspiration, moisture retention in wood representing the high-elevation wet Puna and the phenology of some species of grassland, a dry ecosystem found between the myxomycetes. cloud forests and the permanent snow line, the For the moist chamber technique, fourth study area corresponds to (4) the samples of dead plant material were collected Saqsaywaman archeological park (Saqsay- in each of the study sites, taken back to the waman, Fig 2c); collecting sites between 13˚ laboratory and then used to set up moist 30’–31’ S latitude and 71˚ 58’–59’ W chamber cultures for isolation of myxomycetes. longitude at elevations between 3560–3600 m With this technique, Petri dishes were lined asl. with pieces of filter paper before the samples The last study area is located on the were placed in them. Distilled water was then western side of the country, in the Sechura added to plates and the excess water poured off Desert ecosystem and corresponds to (5) the after 24 hr. Before the latter occurred, the value city of Lima (Lima). The collecting sites were of pH was determined for each culture. All located between 12˚ 4’–5’ S latitude and 76˚ cultures were maintained at room temperature 56’–77˚ 1’ W longitude at elevations between and checked at regular intervals for a period of 100–250 m asl. The collections in this study at least 10 weeks. Distilled water was added area were carried out in recreational areas and when needed to keep cultures from drying out. the Botanical Gardens of the Universidad In the case of myxomycetes, this technique is Nacional Agraria La Molina. useful for recording and quantifying the relative abundance of the most common species at a given locality and to record rare Sampling species not encountered in the field. All study areas were visited during both In all cases, specimens were placed in the dry and the wet season in 2007. In all cases, pasteboard boxes after being collected or a combination of techniques was used to record removed from moist chamber cultures. myxomycetes. Both field occurrences and Vouchers for all species recorded during this moist chamber records were considered for study were deposited in the mycological biodiversity purposes. In the case of the latter herbarium of the University of Arkansas technique, the standard protocol described by (UARKM). Duplicates of collections also have Stephenson & Stempen (1994) was used. been also deposited in the herbarium of the 585 Universidad Peruana Cayetano Heredia separate forthcoming publication (Rojas et al., (HUPCH), as required by the Peruvian unpublished data). However, for the purpose of legislation. the present study, simple examinations of the Only those species not previously complete dataset have been included in the recorded from Peru are included in the list of results. A calculation of the taxonomic species reported herein. As such, this list does diversity index (TDI) sensu Stephenson et al. not correspond to the complete catalogue of (1993) was carried out. myxomycetes for the entire country. In order to compile the present list, the records of species Results reported by Lado & Wrigley de Basanta (2008) A total of 557 specimens representing were used as a starting point. All species were 66 species were documented during the present arranged alphabetically by genus and then study. Approximately 62% of the records were specific epithet, and author names were added. obtained from moist chamber cultures and 39% The latter followed the nomenclatural in field conditions. Excluding the species treatment mentioned earlier. In all cases, a already known for Peru, 49 taxa not previously series of annotations corresponding to basic reported from the country were recorded. The ecological data associated with particular most commonly encountered species were records have been included after the species Arcyria cinerea, Ceratiomyxa fruticulosa and names. For practical purposes, this information Stemonitis fusca. For the complete dataset has been coded using abbreviations for the obtained, the value calculated for the TDI different ecological factors that collectively index was 3.3. represent the microhabitat of each species. The most productive forest types were An indication of the source of the old growth and old successional habitats, with record follows the author name. Those species about 24% and 22% of the total number of that were collected in the field are marked as records, respectively. Substrates with the field collections (FC), whereas those obtained highest number of records were decaying wood from moist chamber cultures in the laboratory and aerial litter, accounting for 33% and 23% are indicated as moist chamber collections of the total dataset, respectively. For field (MC). Following this, the number of records collections, about 62% of records were for a particular species is included before the encountered during the dry season. In contrast, name of the study area where collections of 58% of the specimens recorded from moist that species were recorded. The forest type was chamber cultures appeared on substrates then indicated, using a one letter code for old collected during the wet season. For the growth forest sections (A), old (B) and young complete dataset, pH values ranged between (C) successional patches, former plantations 2.8 and 7.9. (D), seasonally flooded (E), very disturbed An annotated list of the new records open (F) habitats, and palm swamps (G). In a documented during the present study areas is similar manner, substrates upon which fruiting provided below. bodies of each species were recorded, used a two letter code, and were abbreviated as GL for ground litter, AL for aerial litter, TW for twigs, Annotated list of new records for Peru DW for decaying wood, FS for fruits and seeds and LP for living plants. Finally, an indication Arcyria afroalpina Rammeloo: MC; 4 of the pH value or range associated with collections; Los Amigos, in B, C and E; species occurrences is provided at the end of on GL and AL; pH range = 4.4–5.7. the annotations. Ceratiomyxa morchella A.L. Welden: FC; 17 collections; Los Amigos; in A and B; Data analysis on DW; pH range = 3.0–4.6. Due to the heterogeneity of protocols Ceratiomyxa sphaerosperma Boedijn: FC; 8 used for collecting data during this study, a collections; Los Amigos; in A, B and F; separate analysis, carried out only for the on DW, FS and LP; pH range = 2.8– subset of the Amazon data, is the subject of a 5.9. 586 Mycosphere Clastoderma debaryanum A. Blytt: MC; 4 Didymium flexuosum Yamash.: MC; 1 collections; Los Amigos; in A and G; collection; Los Amigos; in G; on GL; on TW; pH range = 4.1–6.0. pH = 5.6. Collaria arcyrionema (Rostaf.) Nann.-Bremek. Didymium iridis (Ditmar) Fr.: MC; 17 ex Lado: MC and FC; 2 collections; collections; Los Amigos; Wayqecha Los Amigos; in D; on TW and DW; pH and Macchu Picchu; in A, B, D, E, G range = 5.0–5.5. and F; on GL, AL, TW and DW; pH Comatricha elegans (Racib.) G. Lister: MC; 1 range = 5.1–7.9. collection; Los Amigos; in G; on TW; Didymium ochroideum G. Lister: MC; 1 pH = 6.6. collection; Los Amigos; in D; on GL; Comatricha laxa Rostaf.: MC; 2 collections; pH = 7.0. Los Amigos; in A; on AL and TW; pH Didymium pertusum Berk.: MC; 1 collection; range = 4.2–4.9. Los Amigos; in A; on GL; pH = 6.0. Comatricha pulchella (C. Bab.) Rostaf.: MC; 8 Didymium trachysporum G. Lister: MC; 2 collections; Los Amigos; in A, B, D collections; Saqsaywaman; in C; on and G; on GL, AL and TW; pH range = TW; pH range = 6.3–6.5. 4. 4–6.4. Didymium vaccinum (Durieu & Mont.) Buchet: Craterium leucophaeum Fr.: FC; 1 collection; MC; 1 collection; Saqsaywaman; in C; Los Amigos; in A; on DW; pH = 4.36. on TW; pH = 6.5. Cribraria intricata Schrad.: FC; 4 collections; Fuligo septica (L.) F.H. Wigg.: FC; 4 Los Amigos; in A; on DW; pH range = collections; Los Amigos; in B; on DW; 4.3–5.4. pH range = 3.7–5.7. Cribraria languescens Rex: FC; 4 collections; Hemitrichia leiocarpa (Cooke) Lister: MC; 5 Los Amigos; in A; on DW; pH range = collections; Los Amigos; in A, B and E; 4.3–6.0. on GL and AL; pH range = 4.8–5.6. Cribraria microcarpa (Schrad.) Pers.: MC and Hemitrichia minor G. Lister: MC; 2 FC; 9 collections; Los Amigos; in B collections; Los Amigos; in A; on TW; and G; on TW and DW; pH range = pH = 6.1. 4.2–5.5. Hemitrichia pardina (Minakata) Ing: MC; 7 Cribraria splendens (Schrad.) Pers.: FC; 1 collections; Los Amigos; in A, D, E; on collection; Los Amigos; in A; on DW; AL and GL; pH range = 5.1–6.3. pH = 4.9. Lamproderma scintillans (Berk. & Broome) Cribraria violacea Rex: MC; 2 collections; Los Morgan: MC; 20 collections; Los Amigos; in A; on GL; pH range = 5.6– Amigos; in A, D and E; on GL and AL; 6.1. pH range = 5.4–7.9. Diachea leucopodia (Bull.) Rostaf.: MC; 2 Lycogala conicum Pers.: FC; 1 collection; Los collections; Los Amigos; in E; on GL; Amigos; in B; on DW; pH = 5.8. pH range = 5.3–7.8. Lycogala epidendrum (L.) Fr.: FC; 12 Diderma effusum (Schwein.) Morgan: MC and collections; Los Amigos; in A, E and F; FC; 14 collections; Los Amigos; in A, on DW; pH range = 3.5–7.1. B, D and E; on GL and AL; pH range = Lycogala exiguum Morgan: FC; 1 collection; 3.9–7.4. Los Amigos; in F; on DW; pH = 5.6. Didymium clavus (Alb. & Schwein.) Rabenh.: Perichaena chrysosperma (Curr.) Lister: MC; FC; 1 collection; Los Amigos; in G; on 1 collection; Los Amigos; in E; on AL; DW; pH = 3.9. pH = 5.9. Didymium difforme (Pers.) Gray: MC; 6 Perichaena corticalis (Batsch) Rostaf.: FC; 1 collections; Los Amigos and collections; Saqsaywaman; in B; on Saqsaywaman; in A, B, C and D; on TW; pH = 6.5. GL, AL, TW and DW; pH range = 5.5– 7.7. 587 Perichaena depressa Lib.: MC; 27 collections; Physarum tenerum Rex: FC; 1 collection; Los Los Amigos, Wayqecha, Macchu Amigos; in B; on DW; pH = 5.5. Picchu and Lima; in A, D, E, F, G; on Stemonaria longa (Peck) Nann.-Bremek., R. GL, AL and TW; pH range = 5.4–7.6. Sharma & Y. Yamam.: FC; 1 Physarum album (Bull.) Chevall.: MC and FC; collection; Los Amigos; in F; on DW; 10 collections; Los Amigos; in A, B, D pH range = 5.3. and E; on AL, TW and DW; pH range = Stemonitis flavogenita E. Jahn: FC; 2 3.5–6.2. collections; Los Amigos; in A and F; on Physarum didermoides (Pers.) Rostaf.: MC; 4 DW; pH range = 4.9–7.2. collections; Macchu Picchu and Lima; Stemonitis smithii T. Macbr.: FC; 1 collections; in F; on GL and AL; pH range = 6.2– Los Amigos; in A; on DW; pH = 4.5. 6.8. Stemonitopsis hyperopta (Meyl.) Nann.- Physarum flavicomum Berk.: FC; 2 collections; Bremek.: MC; 1 collection; Los Los Amigos; in F; on DW; pH range = Amigos; in D; on TW; pH = 6.4. 3.5–4.0. Stemonitopsis microspora (Lister) Nann.- Physarum galbeum Wingate: MC; 1 collection; Bremek.: MC; 1 collection; Los Los Amigos; in D; on AL; pH = 5.3. Amigos; in D; on TW; pH = 5.6. Physarum nucleatum Rex: FC; 1 collection; Tubifera bombarda (Berk. & Broome) G.W. Los Amigos; in B; on DW; pH = 7.6. Martin: FC; 3 collections; Los Amigos; in B; Physarum polycephalum Schwein.: FC; 2 on DW; pH range = 2.8 – 3.1. collections; Los Amigos; in A and B; on DW; pH range = 4.8–5.4. Physarum stellatum (Massee) G.W. Martin: FC; 1 collection; Los Amigos; in F; on DW; pH = 7.6. Fig 1 – Map of Peru showing the location of the areas sampled in the central-southern portion of the country. For location names, please refer to the section on materials and methods. Discussion now almost two decades old. With the The last publication on myxomycetes important biogeographical and ecological from Peru (i.e., Stephenson & Mitchell 1994) is contribution provided by a series of studies in 588 Mycosphere South America during the last decade (e.g., Neotropical countries do not reflect the true Stephenson et al. 2004, Lado et al. 2007, diversity of myxomycetes. For example, in the Cavalcanti et al. 2009), the number of species Neotropical review of Lado & Wrigley de of myxomycetes known for Peru prior to the Basanta (2008), the number of myxomycetes present study was only 31, which seems to be a known for Costa Rica was 143 species. clear example of undersampling, especially However, a more recent publication based on since at least 100 species have been reported the results of detailed local studies in that from neighboring countries such as Brazil, country reported a total diversity of 208 species Chile and Ecuador (Lado & Wrigley de (see Rojas et al. 2010), a number similar to the Basanta 2008). In that sense, it is exceedingly published total number of species for Brazil. likely that the number of species of Under the current concepts of myxomycete myxomycetes for Peru will continue to increase biogeography, it is very unlikely that Costa with additional studies. In the meantime, Rica actually supports a similar number of increasing the number of Peruvian species as Brazil, which is characterized by a myxomycetes to 80 species represents a higher diversity of ecosystem types and a total significant step towards developing a more land area 166 times the size of Costa Rica. complete understanding of the myxobiota of Therefore, it seems probable that a country like this country. Brazil has been just moderately studied in In fact, it is very likely that the numbers comparison to Costa Rica. of species known for even better studied Fig 2 – Examples of the study areas surveyed as part of the present investigation: (a) old successional forest and (b) disturbed open area in the lowland rain forest at Los Amigos; (c) shrub- dominated open area in Cusco at Saqsaywaman and (d) lower montane wet forest vegetation in an open area near the town of Macchu Picchu. From that perspective, the 49 new number of species of myxomycete reported records of myxomycetes for Peru generated in from Peru prior to the updated list provided the present study may represent just a partial herein, the present study is a significant view of the true biodiversity of these organisms contribution to our knowledge of Peruvian and in that country. However, given the low South American myxomycetes. This is borne 589 out when these data are considered within the seems apparent that it is the structure of the context of previously available data. For forest itself and not particular habitats or instance, approximately 74% of the species substrates within that forest that represents the recorded during the present study represented major factor determining the abundance of new records for Peru, and about 55% of all myxomycetes. A more detailed examination of species previously known for the country also this situation will be provided in a separate were recorded. The inference from this is that analysis. However, it is interesting to notice Peruvian forests are still poorly studied and that aerial litter is important in Peruvian that the research effort involved in the present forests, since it has been said that this is an study simply substantiates the lack of important substrate for myxomycetes in generalized knowledge on the diversity and tropical areas (see Stephenson 2003). In any distribution of Neotropical myxomycetes. case, similar observations regarding forest In spite of the latter, it is not surprising structure and particular substrates have already that the three most commonly encountered been documented in other areas of the species in this study were Arcyria cinerea, Neotropics (Rojas et al. 2011a). Ceratiomyxa fruticulosa and Stemonitis fusca. The present study increased the number These three species have been considered as of myxomycetes known from Peru. For cosmopolitan for a long time (see Martin & ecologically fragile areas of the world such as Alexopoulos 1969). Even for Peru, they had the Andes and the Amazon Basin, it is been previously recorded in other studies (Farr important to have information on biodiversity 1976, Stephenson & Mitchell 1994). Moreover, before management plans for natural areas, the calculated value for the taxonomic diversity including those related to conservation plans, index seems to be comparable to values can be adequately developed. Therefore, this obtained in other studies that have involved a study, contributes directly to the overall moderate research effort in tropical areas. For knowledge of one of the most biodiverse areas instance, Stephenson et al. (1993) reported in the world and also to our knowledge of values of 3.04 and 4.13 for tropical areas in myxomycete distribution worldwide. India and Tran et al. (2006) a value of 3.44 for a tropical area in Thailand. In Neotropical Acknowledgements areas, Rojas et al. (2010) reported a value of Appreciation is extended to the Amazon 5.77 for Costa Rica and Stephenson et al. Conservation Association for providing funds (1999) a value of 1.76 for Puerto Rico. for this project. We are grateful to Daniella However, the last two cases are exceptional in Biffi, Pedro Romero and Rina Ramirez for that the first encompassed a collecting effort their help in the field; to Alicia Tuggle, Robin that has extended over more than a century and Doss and Amanda Bates for their help in the the second represents an island. In any case, the laboratory, to Gabriel Moreno for the calculated TDI in the present study supports confirmation of the identification of some taxa the idea that myxomycete assemblages in the and to the University of Arkansas for logistic tropics are characterized by a high intrageneric support. Finally, we would like to express our diversity, probably due to the large availability gratitude to INRENA, for granting us of microhabitats (see Schnittler & Stephenson permission to study the myxomycete diversity 2002). of Peru. When forest type associations were analyzed, both old growth and old successional References forest patches were associated with the highest diversity of myxomycetes. Interestingly, the Adl S, Simpson A, Famer M, Andersen R, two most productive substrates were decaying Anderson R, Barta J, Bowser S, wood and aerial litter. The presence of these Brugerolle G, Fensome R, Fredericq S, substrates requires a particular vegetation James T, Karpov S, Kugrens P, Krug J, structure, since both of them are related to Lane C, Lewis L, Lodge J, Lynn D, forest patch age (see DeWalt et al. 2003). 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