Anura distribution and richness in a tropical montane forest in Monteverde, Costa Rica


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Anura distribution and richness in a tropical montane forest in Monteverde, Costa Rica

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Title:
Anura distribution and richness in a tropical montane forest in Monteverde, Costa Rica
Translated Title:
Distribución y riqueza de anuros en Monteverde, Costa Rica
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McDonald, Elizabeth L.
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Tropical regions, especially tropical montane regions, are biodiversity hotspots for amphibians. In tropical montane forests, varying temperatures and precipitation along an elevational gradient create distinct Holdridge life-zones for the 51 species of frogs and toads in Monteverde, Costa Rica. However, changing climatic conditions, and other complex factors, have contributed to recent anuran declines in the Monteverde area and worldwide. Shifting abiotic conditions cause species to move to new areas with suitable conditions. Within this study, I establish a data set complimentary to previous baselines of anuran distributions in local areas. In doing so, current encountered distributions can be compared to expected distributions. I conducted visual and auditory surveys across 5 locations and between 4 Holdridge life-zones in the Monteverde area. In a comparison to past studies and literature, I encountered 4 species in zones that contrasted expected life-zones. However, I did not determine an upslope trend that parallels the upslope shift in abiotic conditions. Because changes are complex, it is imperative to monitor how vulnerable species react to changing environmental conditions over time. ( ,, )
Abstract:
Las regiones tropicales, especialmente las tropicales montañosas, son puntos calientes de biodiversidad para anfibios. En las zonas montañosas la variación de temperatura y precipitación a lo largo del gradiente de elevación crean distintas zonas de vida, para 51 especies de ranas y sapos en Monteverde, Costa Rica. Sin embargo, el cambio climático global, así como otros factores, han contribuido a recientes disminuciones en las poblaciones de anfibios en Monteverde y en el mundo. En este estudio establecí una serie de datos complementados con estudios previos sobre la distribución de ranas en el área. También realicé recorridos escuchando y encontrando ranas en cinco sitios que incluyeron cuatro zonas de vida en Monteverde. En comparación con estudios previos, encontré 4 especies en zonas donde no se habían registrado. Sin embargo, no encontré tendencias de cambios hacia elevaciones mayores. Debido a que los cambios son complejos, es imperativo monitorear qué tan vulnerables son las especies frente a los cambios en condiciones medioambientales en el tiempo.
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Student affiliation : University of California, Merced – School of Natural Sciences
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Monteverde Institute
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Monteverde Institute
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M39-00790 ( USFLDC DOI )
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Mc D onald 1 Anura d istribution and r ichness in a t ropical m ontane f orest in M onteverde, Costa Rica Elizabeth L. McDonald University of California, Merced School of Natural Sciences EAP Tropical Biology and Conservation 29 May 2019 A BSTRACT Tropical regions, espec ially tropical montane regions, are biodiversity hotspots for amphibians. In tropical montane forests, varying temperature s and precipitation along an elevational gradient create distinct Holdridge life zones for the 5 1 species of frogs and toads in Montev erde, Costa Rica. However, changing climatic conditions , and other complex factors, have contributed to recent anuran declines in the Monteverde area and worldwide . Shifting abiotic conditions cause species to move to n ew areas with suitable conditions. Wi thin this study, I establish a data set complimentary to previous baselines o f anura n distribution s in local areas . In doing so, current encountered distributions can be compared to expected distributions. I conducted v isual and auditory surveys across 5 l ocations and between 4 Holdridge life zones in the Monteverde area. In a comparison to past studies and literature , I encountered 4 species in zones that contrasted expected life zones. However, I did not determine an u pslope trend that parallels the upslo pe shift in abiotic conditions. Because changes are complex, it is imperative to m onitor how vulnerable species react to changing environmental conditions over time. Distribución y riqueza de anuros en Monteverde, Costa Rica RESUMEN Las regiones tropi cales, especialmente las tropicales montañosas, son puntos calientes de biodiversidad para anfibios. En las zonas montañosas la variación de temperatura y precipitación a lo largo del gradiente de elevació n crean distintas zonas de vida, para 51 especies d e ranas y sapos en Monteverde, Costa Rica. Sin embargo, el cambio climático global, así como otros factores, han contribuido a recientes disminuciones en las poblaciones de anfibios en Monteverde y en el m undo. En este estudio establecí una serie de datos complementados con estudios previos sobre la distribución de ranas en el área. También realicé recorridos escuchando y encontrando ranas en cinco sitios que incluyeron cuatro zonas de vida en Monteverde. E n comparación con estudios previos, encontré 4 espe cies en zonas donde no se habían registrado. Sin embargo, no encontré tendencias de cambios hacia elevaciones mayores. Debido a que los cambios son complejos, es imperativo monitorear qué tan vulnerables s on las especies frente a los cambios en condiciones medioambientales en el tiempo.

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Anura Distributions McDonald 2 The structure of an ecological community is determined by biotic and abiotic factors. Abiotic factors and available resources govern species composition, abundances, and distributions. Areas like the Neotropics, where tem perature and precipitation are high, have high relative floral and faunal diversity (Vredenburg and Wake, 2007). At the species level, niches are defined areas of conditions and resources in which organisms can survive and reproduce successfully. A specie etermines the geographical distribution of the species. As environmental factors change, through natural or anthropogenic causes, the distribution range of organisms ma y shift. Specifically, changes in temperature and moisture of an ar ea create new conditions for species to adapt to or migrate from. Currently there are over 6,600 recognized species of frogs and toads within the order Anura. Because amphibians have perme able skin used for fluid and gas exchange, they have adapted to cool er and more humid habitats to prevent dehydration. Especially in warm tropical environments, many species are active at night, live near water sources or in the microclimates provided by f orest leaf litter. During tropical dry seasons, many amphibians will retreat underground to remain cool, emerging again once heavy rains signal the rainy season (Leenders, 2016). Differences in climatic and environmental aspects, such as seasonal rainfall, dry season length, and consistent cloud cover determine species div ersity of a region. The greatest diversity and abundance of amphibians is expected between the end of the dry season and the start of the rainy season (Leenders, 2016). Higher elevation, humid mountain ridges are often home to endemic species that are isol ated by surrounding drier lowlands, such as the golden toad of Monteverde, Costa Rica. Golden toad populations decreased from an observed 1500 individuals in 1987 to none in 1989 (Pounds, 2004). Along with 19 other species, the diminishing abundances and e ntire disappearances of amphibians may be related to factors such as habitat loss, climatic influence (Pounds, 2004), and infectious disease (Vredenburg and Wake, 2007). As global tempera tures e nviro nmental conditions of specific areas are altered (Pounds, 2004 ) . In tropical montane regions, t hese changes impact the composition of species that can survive in specific elevational areas . As environmental conditions shift upward in elevation, a nura n popu lations often move upslope to find suitable habitats. When a species does not have the physical space to move upslope, disappearances and declines of species occur. Within this study, I establish ed a data set on the current distributions and abundances of a nura n species in areas surrounding Monteverde (Fi g. 1), based on individuals I detect ed through direct observation. I a ddress ed the question: What are the current altitudinal distributions of common anuran in the Monteverde area? Through the use of past d ata on amphibian d istributions in similar locations, I compare d changes in altitudinal and zonal distributions of Anura. Aiming to establish a data set complimentary to previous baselines, I note d changes in species distributions if observed. Because curre nt literature examines the role of shifting montane cloud cover on amphibian distributi ons (Pounds and Puschendorf, 2004), I hypothesize that certain species have moved to new locations, following suitable conditions over time.

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Anura Distributions McDonald 3 M ATERIALS AND METHODS Stud y Area: I studied frog distributions in the Cordillera de Tilaran, four mountain ranges. Rainforest San Gerardo Biological Station. Although I studied locations acr oss an elevational range, sites were not geographically connected. At elevations between 1200 1 7 00m, I classified the areas according to literature Holdridge l ife zones , which are based on mean annual temperature and precipitation ( Table 1, Holdridge 1967 ). I used areas on and near trail systems of the Santuario Ecológico Mo nteverde (Zone 2), Rachel and Dwight Cramdell Reserve (Zone 3), Estación Biológica de Monteverde (Zones 3/4), San Gerardo Biological Station (Zone 5), and the Continental Divide (Zone 5 ). Site elevations were recorded with an altimeter. Additionally, t he c limatic characteristics of this region include a dry and rainy season . is typically between November / December until April / May , I conducted surveys between 6 23 M ay 2019 (Leenders, 2016). Holdridge Life Zone Study Sites Zone 1 Tropical Moist Forest NA Zone 2 Tropical Premontane Wet Forest Santuario Ecológico Zone 3 Tropical Lower Montane Wet Forest Rachel and Dwight Cramdell Reserve , Estación Biológica Zone 4 Tropical Lower Montane Rain Forest Estación Biológica Zone 5 Tropical Premontane Rain Forest San Gerardo Biological Station, Continental divide Zone 6 Tropical Wet Forest NA Data Collection: I recorded individuals using Visual Encounter Surveys, a common practice for amphibian surveys (Heyer et al., 1994). Using this method, other researchers and I search ed a specific habitat for a given amount of time. We conducted v isual and auditory searches during evenings, between 7 :00 and 10:00 pm. We used existing trail systems and followed streams when conducting searches. To aid in the surveys, we disturbed leaf litter, turned over logs, and listened to acoustic signals. I recorded e levation and species identification for each am phibian encountered. I inc luded t he following components in measurements: date and time of encounter, elevation of site, species identification of individual, and substrate and position of frog (Appendix, raw data) . We photographed i ndividuals and returned each to the habitat in wh ich found.

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Anura Distributions McDonald 4 Data Analysis: I analy z ed the data using Microsoft Excel software. Through direct auditory and visual observation, I measured species richness and abundance. I organize d species encounters across study sites , comparing species richness over a n elevational gradient. Additionally, I compared encountered species and elevations to expected zonal distributions in literature . R ESULTS Amphibian Distribution Across an Elevational Gradient Throughout the study, I encountered 140 individual s , across 17 species and amongst seven families. I included photographs of individuals from each of the 17 species, as well as the taxonomic names I used to classify them (Appendix). I organized species encountered across study sites ( Table 1). I encountered the follow ing species at relatively high abundance s: Pristimantis ridens (16.4%), Rana (Lithobates) taylori (15%), Diasporus diastema (14.3%), Espadarana prosoblepon (14.3%), and Craugastor fitzingeri (11.4%). I encountered a group of six species ( H. talamancae , S. phaeota, D. rufioculus, P. cruentus, C. underwoodi, C. crassidigitus ), only in elevations and areas corresponding to the Holdridge life zone 5, Tropical P remontane Rain Forest (Table 1) . Across an altitudinal gradient of from 1134 1700 meters, elevation d id not significantly influence the species diversity of amphibians ( Fi g. 1, R ² =0.4136, However, c omparisons between species richness across low to mid elevations were significantly varied ( Table 2 , d.f. = 1, s were not ( Table 2 , d.f.= 1,

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Anura Distributions McDonald 5 Table 2 Species richness and abundances across a spatial gradient in the Monteverde area. Individuals were recorded using visual and auditory surveys between 7:00 10:00 pm throughout the month of May 2019. A tota l of 17 species comprising 7 families were encountered at sites which include: the Santuario Ecológico Monteverde, Rachel and Dwight Cramdell Res erve , Estación Biológica de Monteverde, San Gerardo Biological Station, and the Continental Divide. Rates of in dividuals encountered per day were calculated to reduce bias associated with sites visited more frequently. Species Name San Gerardo (1134 1215m) Santuario (1230m) Cramdell Reserve (1319 1396m) Estacion (1422 1617m) Divide (1700m) Family Ranidae Ra na (Lithobates) taylori 1 >10 0 >10 0 Rana (Lithobates) warszewitschii 1 >5 0 0 0 Family Hylidae Duellmanohyla rufioculis 6 0 0 0 0 Duellmanohyla uranochroa 0 0 0 0 1 Smilisca phaeota 2 0 0 0 0 Isthmohyla psuedopuma 3 0 0 5 0 Isthmohyla zeteki 0 0 0 3 0 Family Centrolenidae Espadarana prosoblepon 10 0 2 0 8 Hyalinobatrachium colymbiphyllum 0 0 0 0 7 Hyalinobatrachium talamancae >5 0 0 0 0 Family Bufonidae Rhinella marina 0 0 0 1 0 Family Strabomantidae Pristimantis ridens 20 1 0 2 0 Pristimantis cruentus 1 0 0 0 0 Family Craugastoridae Craugastor fitzingeri 8 0 6 2 0 Craugastor underwoodi 3 0 0 0 0 Craugastor crassidigitus 2 0 0 0 0 Family Eleutherodacylidae Diasporus diastema >10 0 0 >10 0 Total Individua ls 72 16 8 33 16 Rate: Individuals/day 14.4 8 2.667 11 16 Number of Species 13 3 3 7 3

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Anura Distributions McDonald 6 Figure 1 Species richness across an elevational gradient in the Monteverde area. Individuals were recorded using visual and auditory surveys between 7:00 10:00 pm d uring May 2019. I encountered a total of 17 species comprising 7 families across an elevational range of 1134 1700 meters. Zonal Distribution A comparison of expected distribution zones (Hayes 1989, Savage 2002) with currently observed distribution zones r eveals disparities. In total, five species belonging to the families Ranidae, Hylidae, Strabomantidae , Centrolenidae and Craugastoridae were encountered in zones outside of their expected Holdridge life zones. A group of 3 species ( R. taylori, H. talamanca e and C. crassidigitus ) were encountered in Zone 5 Tropical Premontane Rain Forest yet were not previously recorded in this zone (Table 3 ). R² = 0,4136 0 2 4 6 8 10 12 14 1000 1100 1200 1300 1400 1500 1600 1700 1800 Number of Species Elevation (m)

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Anura Distributions McDonald 7 Table 3 Comparison of expected and encountered zonal distributions based on past guides of amphibian distributions (Hayes and Pounds, 1989, Savage, 2002). Zones were distinguished based on the Holdridge life zones as follows: Zone 1 Tropical Moist Forest (690 1300 m); Zone 2 Tropical Premontane Wet Forest (1300 1470); Zone 3 Tropical Lower Montane Wet Forest (1470 1600 m); Zone 4 Tropical Lower Montane Rain Forest; Zone 5 Tropical Premontane Rain Forest (1150 1450m); Zone 6 Tropical Wet Forest (600 1150m). Species Expected Distribution Zone Encountered Zone Differences in Zone Family Ranidae Rana (Lithobates) taylo ri 2,3 2,3, 5 X Rana (Lithobates) warszewitschii 2,3,4,5,6 2,5 Family Hylidae Duellmanohyla rufioculis 1,2,5,6 5 Duellmanohyla uranochroa 2,3,4,5 4 Smilisca phaeota 5,6 5 Isthmohyla psuedopuma 2,3,4,5,6 3,5 Isthmohyla zeteki 5,6 3,4 X Family Centrolenidae Espadarana prosoblepon 1,2,3,4,5,6 3, 4, 5 Hyalinobatrachium colymbiphyllum 2,3,4 4 Hyalinobatrachium talamancae 2 5 X Family Bufonidae Rhinella marina 1,2,3 3 Family Strabomantidae Pristimantis ridens 4,5,6 2 , 3 ,5 X Pristim antis cruentus 3,4,5,6 5 Family Craugastoridae Craugastor fitzingeri 2,3,5,6 3,5 Craugastor underwoodi 3,5 5 Craugastor crassidigitus 4 5 X Family Eleutherodacylidae Diasporus diastema 3,4,5,6 3,4,5 D ISCUSSION Within this study, I recor ded a nura n species distribution across a spatial gradient to document species moving to new geographical areas over time. On the basis that abiotic conditions of areas are changing, a nura n species are expected to move to areas that currently exhibit suitab le conditi ons. In the Monteverde area, t emperatures have increased while cloud cover has decreased over the past few decades ( Pounds et al. 2006) . According to these abiotic changes , as well as recorded upslope movement in distribution patterns , I expected to find an increase in elevational distribution amongst Monteverde species ( Mattoon 2000, Schlimm 2007).

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Anura Distributions McDonald 8 In past distributional studies a long the trail systems of the Estacion Biologica de M onteverde , two species, C raugastor (Eleuthrodactylus) bransfordd ii and C raugastor (Eleuthrodactylus) fitzingeri were encountered in Zone 4 while expected in Zone 3 (Schlimm 2007). These trends parallel an upslope shift in abiotic conditions. I did not find similar shifts in distribution explicitly from lower to higher elevation zones (Table 3). However, I did record varying distributional zone shifts among five species. Contrary to expected zonal distributions, three of the five species ( R. ta ylori, H. talamancae and C. crassidigitus ), were encountered in Zone 5, Tropic al Premontane Rain Forest on the Caribbean slope. Additionally, two of these three species ( H. talamancae and C. crassidigitus ), were encountered only in Zone 5, despite expected distributions in other zones. A possible explanation for this could be a pref erence for the conditions of Zone 5, as these species were not encountered in literature distribution zones (2,4). D iscrepan cies in expected and encountered distributions were notable due to changes in slope orientation. Specific e ncounters in new Holdrid ge life zones are relevant becaus e Zones 2 3 are on the Pacific slope, whereas Zone 5 is on the Caribbean slope , and Zone 4 includes both slopes and the continental divide. For example, Isthmohyla zeteki , a frog that lives in the upper canopy, is expected at low middle elevations on the Caribbean slope , within Zones 5 6. However, I acoustically recorded individuals of I. zeteki at middle high elevations on the Pacific slope, in Zones 3 4. Orientation of montane slope in Costa Rica, either Pacific or Caribbe an influences seasonal climate . Slope orientation is thus interc onnected with the Holdridge life zones , which are determined by mean annual temperature and precipitation values. To further address the relation between species distribution and slope orienta tion, expansive regions including more gradients and sites on both slopes should be used. A decrease in species richness with increasing elevation is common in herpetofaunal studies (Gifford and Kozak 2011), yet other studies display more complex trends (F u et al 2006, Khatiwada 2011). Within this study, species diversity showed no distinct pattern across the elevational gradient examined (Fig. 1). However, species richness in low (1134 1230m) elevations significantly varied from that in m iddle (1319 1396m) elevations. Comparatively, no significant difference between middle (1319 1396m) and high (1422 1700m) elevations was found (Table 2). Larger relative differences in species richness from low middle elevations is a pattern similarly foun d in other amphibi an studies. However, middle high elevational similarities in richness reject the concept that diversity declines with elevation. (Naniwadekar & Vasudevan, 2007, Wiafe and Agyei 2013). Within this study, I compa red my results to distribu tion ranges establ ished 20 30 years ago (Hayes and Pounds, 1989, Savage 2002) . Not only is it valuable to consider environmental changes, but also changes in research efforts and contributions. Interest in herpetofauna research may now be more common and i ntensive, while co ncepts may be more developed. For example, extensive guidebooks and previous studies have established a basis for distribution studies similar to this. Increased frequency of visual and auditory encounter surveys in all zones may be responsible for encount ers across new life zones and elevations . Additionally, it is worthy to note discrepancies due to altitude equipment, as I recorded the elevation of th e Estación Biológica de Monteverde as approximately 1422 meters. However, literature values and personal communication reveal an elevation of approximately 1570 meters.

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Anura Distributions McDonald 9 Despite this, the classification and use of Holdridge life zones within this study shou ld be accurate, as they were based on past studies and field guides. The overall aim of this study is to establish a current baseline of amphibian distribution in the Monteverde area that can be compared to past and future studies. Because factors such as habitat loss, infectious disease and climate change are contributing to major amphibian declines worldwid e, this study aims to understand and model richness and abundances of amphibians in local areas. Comparatively, baseline databases of amphibians and re ptiles specific to Monteverde exist, yet pressures from many factors have likely altered the distribution zones previously described (Hayes and Pounds, 1989). This study continues a trend of mapping and predicting changes to community ecology concepts. By monitoring and measuring how vulnerable species react to changing environmental conditions, we can better understand changes over time. A CKNOWLEDGEMENTS For the many muddy rubber boots and rainy nights, I would like to thank Yanelyn Perez and Felix Salazar amphibians, and enthusiasm on late night herping trips. I would like to acknowledge the Estación Santuario Ecológico, and LIFE Monteverde for providing the space to complete this study. To my advisor, F ederico, I want to recognize all the late nights spent making sure we made it home, and for the endless support throughout this process. Also, thanks f L ITERATURE C ITED Fu, C., Hua, X., Li, J., Chang, Z., Pu, Z. & Chen, J. 2006 . E levational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate e ffects. Echography , 29 (6): 919 927 Gifford, M.E, and Kozak, K.H. 2011 . Islands in the sky or squeezed at the t op? Ecological causes of elevational range limits in montane salamanders. Echography . DOI: 10.1111/j.1600 d0587.2011.06866. Published online, 07/14/2011 Haber, W.A, Nadkarni, N.M., Wheelwright, N.T. 2000 Monteverde e cology and c onservation of a t ropical c loud f orest. New York: Oxford University Press, pp.41 45 Hayes, M. P., Pounds, J. A., & Timmerman, W. W. 1989 . An a nnotated l ist and g uide to the a mphibians and r eptiles of Monteverde Costa Rica. Miami University, Oxford, Ohio: D. H. Taylor. Heyer W. Ronal d, et al. 1994 . Measuring and m onitoring b iological d iversity: s tandard m ethods f or a mphibians. Smithsonian Institution Press, pp.80 88. Holdridge, L.R., 1967. Life zone ecology. Revised edition. Tropical Science Center, San Jose, Costa Rica. pp. 206

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Anura Distributions McDonald 10 Khati wada, J. R. 2011 . A mphibian species richness and composition along an elevatio nal gradient in Chitwan, Nepal. Norwegian University of Life Sciences Leenders, T. 2016 . Amphibians of Costa Rica. Ithaca, New York: Zona Tropical Press, pp.6 8 : 144 146. Mattoon , A. 2000 . Amphibia Fading. [online] Available at: http://www.worldwatc h.org/system/files/EP134A.pdf [Accessed 29 Apr. 2019]. Naniwadekar, R. & Vasudevan, K. 2007 . Patterns in diversity of anurans along an elevational gradient in the Western Ghats, South I ndia. Journal of Biogeography, pp. 34 : 842 853 Pounds, A. and Puschendorf, R. 2004 . Clouded f utures. [online] Available at: http://www.geo.umass.edu/courses/geo458/ Readings/PoundsPuschendorf04_NAT.pdf [Acce ssed 29 Apr. 2019]. Pounds, J. A., Bustamante, M. R., Coloma, L. A., Consuegra, J. A., Fogden, M. P., Foster, P. N., La Marca, E., Masters, K. L., Merino Viteri, A., Puschendorf, R., Ron, S. R., Sanchez Azofeifa, G. A., still, C.J., & Young, B. E. 2006 . Widespread a mphibian e xtinctions from e pidemic d isease d riven by g lobal w arming. Nature, 439 : 161 167. Savage, J. M. 2002 . The a mphibians and r eptiles of Costa Rica. London: University of Chicago Press. Schlimm B. 2 007 . Herpetofauna d istribution a nd s pecies r ichness a long a n e levational g radient. D epartment of Biology, Villa Julie College Vredenburg, V. T., & Wake, D. B. 2007 . In Levin S. A. (Ed.), Global declines of amphibians. New York: Elsevier. doi://doi.org/10.1 016/B978 012226865 6/00578 X Wiafe, E.D. and Agyei, D. 2013 Species richness, diversity and distribution of amphibians along elevational gradient on mountain Afadjato, Ghana. Eurasian Journal of Forest Science: 68 76

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Anura Distributions McDonald 11 A PPENDIX : V ISUAL R EFERENCE OF A MPHIB IANS E NCOUNTERED Photography by Felix Salazar Rana (Lithobates) warszewitschii (10 July 2018) 4 Rhinella marina (8 April 2017) 2 D uellmanohyla rufiocu lis (10 May 2019, San Gerardo Biological Station) 3 Smilisca phaeota (7 May 2019) 5 Duellmanohyla uranochroa (11 May 2019) 1 Isthmohyla psuedopuma (14 May 2019)

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Anura Distributions McDonald 12 7 Espadarana prosoblepon (16 April 2017) 10 Hyalinobatrachium colymbiphyllum (11 May 2019) 11 Hyalinobatrachium talamancae (16 November 2019) 6 Isthmohyla zeteki (Photo by Marcos Guerra) 9 Pristim antis cruentus (16 April 2017) 8 Pristimantis ridens (14 Novemeber 2019)

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Anura Distributions McDonald 13 12 Craugastor crassidigitus (4 January 2018) 14 Diasporus diastema (12 Novemeber 2018) 13 Cruagastor underwoodi (12 November 2018)

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Anura Distributions McDonald 14 Current Amphibian Distributions in San Gerardo and Monteverde Areas The raw data is included of anuran encountered between 6 23 May 2019. General st udy sites, da tes, elevations, substrate, time and other notes about the encounter were included in observations. Individuals were identified and checked for identification with Amphibians of Costa Rica (Leenders, 2016). Species Site Dat e Elevation( m) Su bstrat e Tim e Tem p (C) Notes San Gerardo 6 May 2019; 7:30 9:00pm; upper and lower stream trails C. underwoodi San Gerardo 6 May 1191 leaf litter 7:4 3 20.9 R. warszewitschii San Gerardo 6 May 1185 leaf litter 7:5 8 C. fitzingeri San Gerardo 6 May 11 70 leaf litter 8:1 6 H. talamancae San Gerardo 6 May heard only C. podiciferus San Gerardo 6 May 1150 leaf litter 8:5 7 23.5 E. prosoblepon San Gerardo 6 May 1160 leaf of plant 21.3 blue humerus C.fitzingeri San Gerardo 7 May 1186 1m fern 7:3 0 2 6.4 San Gerardo 7 May 2019; 7:30 9:15pm; upper and lower stream trails R. taylori San Gerardo 7 May 1186 7:3 0 26.4 P.ridens San Gerardo 7 May 1179 1m leaf 7:3 4 24.4 female P.ridens San Gerardo 7 May 1176 leaf 7:3 6 24 male P.ridens San Gerardo 7 May 1167 0.1m leaf 7:4 3 25 male P.ridens San Gerardo 7 May 1161 1.5m fern 7:4 6 24.7 male E. prosoblepon San Gerardo 7 May 1159 1m leaf over stream 7:5 0 23.8 E. prosoblepon San Gerardo 7 May 1154 0.3m leaf over stream 7:5 3 23.5

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Anura Distributions McDonald 15 E. prosoblepon San Gerardo 7 May 1154 0.3m leaf over stream 7:5 3 23.5 D. rufioculis San Gerardo 7 May 1154 0.1m leaf over stream 7:5 8 23.8 D. diastema San Gerardo 7 May 1160 1m leaf of plant 8:1 1 21.9 pink/brown w/ tubercules P.ridens San Gerardo 7 May 1161 1m leaf o f plant 8:1 5 23.5 P.ridens San Gerardo 7 May 1161 1m leaf of plant 8:1 5 23.5 C. underwoodi San Gerardo 7 May 1158 leaf 8:2 0 23.3 usually in leaf litter, found on leaf P.ridens San Gerardo 7 May 1151 0.3m leaf of plant 8:2 8 23.2 D. rufioculis San Gerardo 7 May 1134 0.2m leaf over stream 8:4 6 23.6 D. rufioculis San Gerardo 7 May 1134 0.2m leaf over stream 8:4 6 23.6 S. phaeota San Gerardo 7 May 1165 1m branch 9:0 9 24 forest edge/pasture. Yellow I. psuedopuma San Gerardo 9 May 1215 bank manmad pond 7:3 8 19.7 Meadow tree frog San Gerardo 9 May 2019; 7:30 9:00pm; upper and lower stream trails P. ridens San Gerardo 9 May 1203 7:4 3 19.6 C. fitzingeri San Gerardo 9 May 1200 1m leaf of plant 8:0 6 20.2 masked P. ridens San Gerardo 9 May 1200 0 .3m leaf of plant 8:0 8 20.8 P. ridens San Gerardo 9 May 1200 1m leaf of plant 8:0 8 20.8 P. ridens San Gerardo 9 May 1196 0.1m leaf of plant 8:1 4 20.4 P. ridens San Gerardo 9 May 1184 0.1m leaf of plant 8:2 0 20.2 C. fitzingeri San Gerardo 9 May 1181 0.3m leaf of plant 8:2 2 20.2 P. ridens San Gerardo 9 May 1172 1m leaf of plant 8:4 3 19.7

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Anura Distributions McDonald 16 I. psuedopuma San Gerardo 10 May 1215 7:3 0 20 San Gerardo 10 May 2019; 7:30 9:30pm; upper and lower stream trails P.ridens San Gerardo 10 May 1209 7:4 8 19.6 D. diastema San Gerardo 10 May 1202 7:5 4 19.7 D. diastema San Gerardo 10 May 1190 8:0 6 19.5 C. fitzingeri San Gerardo 10 May 1176 8:1 3 19.2 C. fitzingeri San Gerardo 10 May 1176 8:1 3 19.2 E. prosoblepon San Gerardo 10 May 1173 8: 2 3 19.5 D. diastema San Gerardo 10 May 1170 8:2 4 19.7 E. prosoblepon San Gerardo 10 May 1170 8:2 4 20 P.ridens San Gerardo 10 May 1179 8:3 6 19.7 P.ridens San Gerardo 10 May 1175 8:4 2 20 C. fitzingeri San Gerardo 10 May 1175 8:5 2 20 D. rufioculis San Gerardo 10 May 1148 9:2 0 19.8 P. cruentus San Gerardo 10 May 1153 9:2 4 19.8 H. colymbiphyllu m San Gerardo 11 May 1700 over stream 7 San Gerardo 11 May 2019; unknown time; Continental divide, Felix's observations D. uranochroa Sa n Gerardo 11 May 1700 leaf valley near stream 1 E. prosoblepon San Gerardo 11 May 1600 ? 8 R. Taylori Estacion Biologica 13 May 1399 edge/ba nk of pond 8:3 0 19.2 more than 10 individua ls Estacion Biologica 13 May 2019; 8:00 9:15pm;

PAGE 17

Anura Distributions McDonald 17 man made pond wit h introduced species (Marvin's) R. marina Estacion Biologica 13 May 1395 edge of pond 8:3 4 19.7 13 May 1350 stream near waterfall No auditory or visual observations 13 May 1420 stream near upper trail No auditory or visual observations I. psuedop uma Estacion Biologica 14 May 1429 0.2m leaf over pond 8:2 7 23.8 in amplexu s Estacion Biologica 14 May 2019; 8:00 9:15pm; small man made pond in garden with introduced native species, Principal trail and stream I. psuedopuma Estacion Biologica 14 May 14 29 0.2m bromelia d 8:3 4 25 near pond I. psuedopuma Estacion Biologica 14 May 1429 0.3m leaf 8:3 5 25 near pond I. psuedopuma Estacion Biologica 14 May 1429 0.2m leaf over pond 8:3 7 25 near pond C. fitzingeri Estacion Biologica 14 May 1429 0.5m leaf 8:4 0 21.9 near pond P.ridens Estacion Biologica 14 May 1433 1m leaf 9:3 0 24 P.ridens Estacion Biologica 14 May 1433 9:3 6 24.3 C. fitzingeri Estacion Biologica 14 May 1428 9:4 3 23.2 I. psuedopuma Estacion Biologica 14 May 1429 road 9:5 3 22.4 P.rid ens Instituto de Montever de 15 May 1332 1m leaf 8:0 9 20.2 Instituto de Monteverde 15 May 2019; 8:00 9:00pm;

PAGE 18

Anura Distributions McDonald 18 P.ridens Instituto de Montever de 15 May 1332 0.2m branch 8:2 1 20.5 P.ridens Instituto de Montever de 15 May 1332 flower of heliconia 8:4 8 20. 3 P.ridens Instituto de Montever de 16 May 1396 0.2m branch 7:4 4 22 Instituto de Monteverde 16 May 2019; 7:20 9:00pm; Stream on trail P.ridens Instituto de Montever de 16 May 1396 1m leaf 7:4 6 22 E. prosoblepon Instituto de Montever de 16 May 1354 1.89m leaf fallen tree over stream 8:1 5 21.5 ~7 heard C. fitzingeri Instituto de Montever de 16 May 1382 0.2m leaf 8:3 5 21.7 H. fleischmanni Instituto de Montever de 16 May heard, single chirp near stream R. warszewitschii Café Montever de 17 May 11 70 leaf litter 7:0 3 21.7 Café Monteverde 17 May 2019: 7:00 8:30pm; trail along stream P. ridens Café Montever de 17 May 1169 0.2 m leaf near stream 7:5 6 22.4 D. diastema Café Montever de 17 May 1172 0.4m leaf 8:1 2 21.9 C. underwoodi Café Montever d e 17 May 1186 leaf litter 8:2 9 21.9

PAGE 19

Anura Distributions McDonald 19 C. podiciferus Café Montever de 17 May NOT SEEN or HEARD...last noted in 2013, usually higher elevation R. taylori Santuario Ecologia 19 May 1215 in/on bank of pond 8:1 5 24 more than 10 individua ls Santuario Ecolo gia 19 May 2019; 8:00 9:30pm; man made ponds R. warszewitschii Santuario Ecologia 19 May 1215 on water lettuce in pond 8:3 0 24 more than 5 individuals C. fitzingeri San Gerardo 20 May 1172 leaf over man made pond 7:1 6 San Gerardo 20 May 2019; 7:15 8:30pm; upper and lower stream trails I. psuedopuma San Gerardo 20 May 1172 leaf over man made pond 7:1 6 S. phaeota San Gerardo 20 May 1172 leaf over man made pond 7:1 6 C. underwoodi San Gerardo 20 May 1172 leaf litter 7:2 2 P. diastema San Ge rardo 20 May 1164 1m leaf 7:2 9 C. crasidigitus San Gerardo 20 May 1144 0.3m leaf 7:4 2 C. crasidigitus San Gerardo 20 May 1141 0.7m leaf over stream 7:4 5 E. prosoblepon San Gerardo 20 May 1138 0.5m leaf over stream 7:4 9 E. prosoblepon San Ge rardo 20 May 1138 1m leaf over stream 7:5 4 E. prosoblepon San Gerardo 20 May 1138 1m leaf over stream 7:5 5

PAGE 20

Anura Distributions McDonald 20 D.rufioculis San Gerardo 20 May 1138 1m leaf over stream 7:4 5 D. rufioculis San Gerardo 20 May 1135 0.1m leaf near stream 7:5 4 metamorph P. ridens San Gerardo 20 May 1145 1m leaf 8:1 8 P. ridens San Gerardo 20 May 1142 1m leaf 8:2 3 P. ridens San Gerardo 20 May 1137 0.2m leaf 8:3 3 amplexus P. ridens San Gerardo 20 May 1137 0.2m leaf 8:3 3 amplexus E. miliaria San Gerardo 20 May heard, not detected for 6 years prior H. talamancae San Gerardo 20 May 1141 heard P.ridens Santuario Ecologia 22 May 1235 0.1m leaf of plant 7:4 8 22.4 Santuario Ecologia 22 May 2019; 7:30 9:15pm; Emergencia trail, man made pond R. taylori Sa ntuario Ecologia 22 May 1235 in/near pond 8:0 0 22.4 atleast 10 individuals R. warschewittsc hii Santuario Ecologia 22 May 1235 on water lettuce in pond 8:1 5 22.4 5 individuals E. prosoblepon Instituto de Montever de 24 May 1319 0.5m leaf of plant 7:0 5 21.8 far from stream Instituto de Monteverde 24 May 2019; 7:00 8:30pm; Senderos, did not go to stream C. fitzingeri Instituto de Montever de 24 May 1380 1m leaf of plant 8:0 0 21.1 C. fitzingeri Instituto de Montever de 24 May 1382 0.4m on tree 8:1 5 22.8

PAGE 21

Anura Distributions McDonald 21 D. diastema Estacion Biologica 26 May 1542 heard only 7:5 8 20.8 atleast 10 individua ls Estacion Biologica 26 May 2019; 7:30 9:15pm; main trail I. zeteki Estacion Biologica 26 May 1617 heard only 8:2 2 19.4 I. zeteki Estacion Biologica 26 May 1500 he ard only 8:4 5 19.4 I. zeteki Estacion Biologica 26 May 1474 heard only 8:5 7 19.5 P. ridens Estacion Biologica 26 May 1422 0.1m fern frond 9:0 3 20 p. ridens Estacion Biologica 26 May 1422 0.1m fern frond 9:0 5 20


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