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Volume 10 2004 No. 1
ECOTROPICA 10: 1–14, 2004 © Society for Tropical Ecology
MEASURING AND MONITORING AMPHIBIAN DIVERSITY IN TROPICAL FORESTS. I. AN EVALUATION OF METHODS WITH RECOMMENDATIONS FOR STANDARDIZATION
Mark-Oliver Rödel 1,2 & Raffael Ernst 2
University of Mainz, Institute of Zoology, Department of Ecology, Saarstrasse 21, D-55099 Mainz, Germany 2 Theodor-Boveri-Institute, Biocenter of the University, Department of Animal Ecology and Tropical Biology (Zoology III), Am Hubland, D-97074 Würzburg, Germany
Abstract : The need for standardization of field methods within and across studies has been recognized by the majority of ecologists throughout the world. However, comparable studies based on a standardized protocol are still scarce. We provide a guideline for effectively sampling and monitoring tropical forest amphibians and give recommendations for standardization. Based on a four-year study on amphibians in Taï National Park, Côte d’Ivoire, we evaluate commonly used techniques, and offer a catalogue of efficient techniques, along with suggestions for improvement of particular methods, with regard to study objectives and specific amphibian guilds. For simple short-term surveys we recommend visual and acoustic encounter surveys, accompanied by opportunistic trapping. The transect design introduced here proved to be most adequate for representative sampling and appeared to be appropriate for most studies that involve multivariate data. It is especially useful for long-term studies. Transects furthermore provide an effective method of investigating at least leaf litter frogs, not only at their breeding sites but also throughout the whole range of habitats used by them, thus generating a much more complete picture of an amphibian community than is possible with other methods. Accepted 4 February 2004. Key words : Tropical amphibian monitoring, standardization, method evaluation, Taï National Park, West Africa.
In theory there is no disagreement about the necessity to standardize ecological field methods and data acquisition in order to guarantee comparability between different studies, as well as to enhance the power of predictions resulting therefrom. The need for standardization of techniques across and within studies has continuously been emphasized by several authors (e.g., Heyer et al. 1994, Adis et al. 1998). Those who are involved in community ecological field research know well that this is a crucial factor when it comes to generalizing results in order to make them
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accessible to those who urgently need scientific guidelines to back up their practical efforts. This becomes especially important when looking at phenomena such as the well known world-wide amphibian decline, which seems to have affected even populations in pristine habitats and continues to affect or even wipe out whole populations with terrifying rapidity (Houlahan et al. 2000, Kiesecker et al. 2001, Pounds 2001). In these cases synergistic efforts are urgently needed, calling for unified methods and data output (Alford & Richards 1999, Parris 1999, DAPTF 2002). However, in reality there still is a lack of studies that can be compared without reservations, especially studies that were conducted in different parts of the world, although precisely this kind of comparison is 1
b. It can be characterized as humid tropic seasonal (Riezebos et al. Transect paths were kept open so that walking at a constant speed was possible at all times. Côte d’Ivoire. Selection of sites and design of sampling units We chose sites within existing macrohabitats (primary/ secondary forest) to establish a total of ten rectangular transects (six in primary forest. The rectangular transect design is a combination of two widely used standard techniques. uses linear transects instead of squares. The intention of this paper is thus to provide a guideline to choosing effective and comparable methods for the monitoring of tropical forest amphibian assemblages in general. Doan 2003) or total removal plots (Barbault & Trefaut Rodrigues 1978. standardization of methods that yield broad-scale comparative data is now needed more than ever. comm. ratures vary between 20 and 33° C. also for para-ecologists. as well as their relative abundances and densities. We aim to present a methodology that i) provides quantitative and qualitative data useful for rapid surveys as well as long-term monitoring programs. 2001. Poynton 1999). 2001) were a priori excluded from testing. 6°47’-7°25’W). Riezebos et al. on the basis of our experience in the field as well as with consecutive data analyses. More detailed descriptions of TNP are provided by Rompaey (1993). Transects were arranged in pairs. Each had a north-south extension of 200 m and an east-west extension of 100 m. The second. followed by a short dry season in July/August.RÖDEL & ERNST essential for academic and practical purposes alike. Formenty pers. Precipitation is distributed across two more or less distinct periods. Daily tempe2 . known as quadrat sampling (plot design). The maximum distance between transects was 6. that stretches west of the Dahomey Gap from Ghana into Sierra Leone and Guinea (Schiøtz 1967. 1979a. situated in western Côte d’Ivoire (5°08’6°07’N. Richards 1996). Thus widely used techniques. Every subunit was marked with numbered colored flag-tape. TNP is part of the Upper Guinea forest block. TNP is the largest remaining protected area of rain forest in West Africa. Mean annual precipitation in the study area was 1806 mm during 1988–2002 (± 297mm. as they may result in severe disturbance to the system under investigation. The core dry season lasts from November until February/March. Lawson 1986. PACPNT 2000). four in secondary forest). proved to be most effective for forest amphibians. 5°50’N.3 km. We avoided extensive cutting and thus manipulation of important habitat features. The major rainy season stretches from September to October. Parts of the habitats on the park’s periphery were subject to more or less intense logging and cultivation until 1998 (P. For data acquisition the complete transect length of 600 m was subdivided in 25 m subunits (24 subunits/transect). We thereby evaluated several methods that are already commonly used and present a set of techniques that. All evaluations and recommendations are based on a study of tropical amphibian assemblages conducted over four years (1999–2002) in Taï National Park. such as standard plot sampling (Allmon 1991. that ii) is easy to handle. data: Taï Monkey Project). The starting coordinate for each transect always marked the southeast corner to ensure identical geographic orientation between sampling units. and iv) has a low environmental impact. thus ensuring that all habitat types of a certain area within the very inhomogeneous forest were covered. Whereas quadrat sampling can be used to determine the species present in a homogenous area. Vonesh 2001. Rodda et al. and unpubl. Floristically the TNP belongs to the Guinean-Congo-Region (Guillaumet 1967. A minor rainy season lasts from March/April to July. (1994) and PACPNT (2000). The mean annual temperature is about 25° C (Rompaey 1993). Our study site was located 23 km southeast of the small town of Taï and comprised about 30 km2 of primary and secondary rain forest around the Station de Recherche en Ecologie de Taï (SRET. 7°20’W). Given the lack of time for the development of effective conservation programs. 1994. Parren & de Graaf 1995. We report on our experiences with already existing methods and with methods that we thought needed modifications to enhance their effectiveness and to enable the use of new statistical analysis methods. Africa: Howell 2002) reflect this need. The minimum distance between adjacent transects was 200 m. consists of a series of small squares (quadrats) that are laid out at randomly selected sites within a habitat then thoroughly searched for amphibians. A number of recently published regional field monitoring manuals for amphibian surveys (Latin America: Lips et al. known as transect sampling. The first. iii) is efficient concerning time and money. Our study sites are either pristine forest areas or have not been logged or cultivated since 1978. map “Projet OMS Forêt de Taï”). transect sampling can provide similar data across habitat or dis- MATERIAL AND METHODS Study site The study was conducted in Taï National Park (TNP).
and to investigate correlations of species assemblages with environmental variables. Compared to linear transects.5 m.5 2 2. whereas primary for- ests show increasing numbers of plants of larger dbh (Chatelain et al. we characterized all habitats using several variables that were recorded at two defined points within each 25 m subunit (beginning and midpoint).. Categories of habitat variables measured on transects. Pearman 1997). A maximum of four transects was patrolled at night. To assess habitat preferences of particular species (e. Standardized visual transect sampling (SVTS). substrate moisture was determined in four categories during every transect walk. 1996. understory: < 0. To avoid duplicate recordings. lower tree stratum: 3–10 m.5–1. In order to quantify the availability of potential aquatic breeding sites. These parameters included vegetation density in four strata (canopy: > 20 m. We assumed that the number of plants with small dbh is greater in degraded. Jaeger & Inger 1994). Specimens heard but not seen within this distance were not searched for and not included in the SVTS data sets (see below). rectangular ones provide the possibility of easily up. dbh = diameter of plants at breast height.5 4 5 6 7 vegetation density absent transition gaps predominating transition closed areas predominating transition closed substrate types forest soil arenaceous forest soil loamy soil arenaceous soil sabulose soil muddy soil swampy soil dbh [cm] 0–5 6–10 11–20 21–50 > 50 leaf cover (%) 0–20 21–40 41–60 61–80 81–100 3 .or downscaling data without neglecting local habitat diversity (e. vegetation density was measured in four strata (see text). divided into seven categories corresponding to particular densities. thereby recording all amphibians within a distance of 100 cm from either side of the path.AMPHIBIAN MONITORING IN TROPICAL FORESTS turbance gradients (Jaeger 1994. captured frogs were marked by TABLE 1. Definitions of habitat variables are summarized in Table 1.35 m/s). A simplified method originally developed by Braun-Blanquet (1964) for determination of vegetation coverage in vegetation analyses was used to estimate the percentage of leaf litter coverage..g. As far as possible all individuals were captured. every aquatic habitat (lentic and lotic) located at a maximum distance of 25 m from either side of the transect was recorded with respect to type. bush and shrub stratum: 0. Sampling was interrupted for the duration of the recording in order not to overestimate locations in which animals had previously been captured. For a discussion of the advantages of rectangular sample units over quadratic or circular ones see Krebs (1989) and McCune & Grace (2002).or age-specific differences in habitat use. Usually four to six transects per day were sampled during daytime. Repeated controls of identical transects on consecutive days were avoided to ensure independence of samples. the vegetation of all 25 m segments within a distance of about 100 cm left and right of the transect was recorded by counting the number of plants belonging to a certain category (diameter at breast height. Sampling was performed independent of prevailing weather conditions. to assign them to particular guilds). Edaphic parameters were registered as general substrate types according to Lieberoth (1982). Transects were intensively patrolled at a constant speed (0. surface and depth.30–0.5 m). for comparison of local with regional species pools).5 3 3. In addition. both in time and space. thus providing data that are useful to test for sex. dbh). secondary forests. Sampling methods and sampling effort Habitat characterization.g. In addition to species identity we recorded sex and snout-vent-length (SVL) of every individual. definition category 1 1. Additionally.
Capture success may vary greatly between species (Corn & Bury 1990. This technique has been formalized as the time-constrained technique of Campbell & Christman (1982). Henle et al.and AES-effort was kept comparable to the time spent on SVTS and SATS surveys throughout the entire study period. VES can only provide information on the presence or absence of a species in an area but are inadequate for determining abundances. other than sex and species. Coding schemes used for individual recognition were not applied. For both SVTS and SATS. Trapping with pitfall or funnel traps along drift fences. Furthermore. Individuals below nine mm SVL were not marked due to their small size.. Recaptures were excluded from the analyses. species composition. 1997). Since in the majority of frog species males use species-specific calls to advertise their position to potential mates and rivals (Wells 1977). According to Corn & Bury (1990). It can be used to determine the species richness of an area and the species composition of a local assemblage. We used these methods continuously in all habitats of the study area including the transect areas. Drift fences consisted of durable green plastic gauze. despite their potential abundance. keeping the transect routine identical for both techniques. individual marking should be restricted to studies in which individual recognition is indispensable.5 4 hours of transect sampling. this behavior can be exploited for acoustic monitoring.e. following the characterization routine used for general habitat description.g. 8. At capturing sites a thorough description of the microhabitat was recorded. The width of the acoustic transect depends on the ability to detect each species’ advertisement call. Visual (VES) and acoustic (AES) encounter surveys. For practical reasons “man-hours” can be used. beginning with segment 1: segment 1. and also for evaluating if species that are rarely found on a transect belong to a particular local species pool or represent single migrating specimens that have habitat preferences not covered by the transects. VES and AES are frequently used for rapid assessments and the evaluation of larger areas. This device is useful to determine species richness of epigaeic organisms (Corn 1994). and additionally in only one of these transects from January 2001 to September 2002. Bufo species). VES and AES thus could be used to evaluate if the randomly chosen transects represented the regional species pool. Throughout transect walks a combination of visual and acoustic techniques was applied at all times. this technique allows the detecting of cryptic species that. Removal of additional toes may decrease the recapture rate of marked individuals by more than 6 –18 % for each additional toe removed after the first (Parris & McCarthy 2001). 0. habitat analyses can be based on data obtained from the general habitat characterization for single transects or segments.5 m to either side of the transect was defined. The resulting data are expressed in numbers of individuals of a certain species found in an area per unit time. Ptychadena species) are more difficult to trap than terrestrial species that lack these abilities (e. From February 1999 to December 2000 we compiled 382.RÖDEL & ERNST toe clipping (Donnelly et al. Thus.g. and to estimate relative abundances of species within an assemblage (Crump & Scott 1994). Anurans that are strong jumpers (e. An area or habitat is searched systematically for individuals in a defined time period. during SATS it was often impossible to determine individual parameters. Counts can be used to estimate relative abundance of calling males. may be underestimated when exclusively using visual techniques. In concordance with these authors the methods were only used as a qualitative or semi-quantitative tool within the scope of our studies. data can be expressed in numbers of individuals per time and surface units... Pitfall traps with drift fences were installed in two of the primary forest transects from March to April 1999. as well as breeding habitat use and breeding phenology of species (Zimmerman 1994). Audio strip transects represent a commonly used method for acoustic monitoring (Zimmerman 1994). 12 and 20). thus creating 25 m x 25 m acoustic sampling plots. Total VES. 1994. only calls coming from the right-hand side in the first segments were registered (i. In the transect corners. Standardized acoustic transect sampling (SATS). An array of fences and traps consisted of a central trap (buckets: .5 m high and stapled vertically onto wooden stakes. and as the time-constrained searches of Corn & Bury (1990). As opposed to visual sampling. Calls from greater distances cannot be unambiguously identified and the chance of duplicate recordings in neighboring segments increases. Due to the simplicity of the method. For that reason a maximum recording distance of 12. which can be adjusted to the complexity of the habitats being sampled. Likewise microhabitat description was not always possible. In those cases. This corresponds to 765 transect walks. Dodd 1991).
In the course of this study. and in a forest fragment (Paulé-Oula 2) outside TNP from 30 August to 12 September 2002 (A. data).. 1. FIG. using a particular sampling method. as sampling success and efficiency may vary with regard to the particular biology of a species. fossorial and aquatic species independent of families. TC = species recorded during transect walks (SVTS and SATS combined).AMPHIBIAN MONITORING IN TROPICAL FORESTS 275 mm deep. Trap = pitfall and funnel traps. 220 mm bottom diameter) and two triangular fence segments (total length 16 m). Numbers above bars represent total number of species recorded with the respective method. The data from pitfall and funnel traps can be expressed as number of individuals per trap-day. Sampling success is referred to as number of species within families recorded. 5 Ta ïF . Data from each sampling method were compared with each other and to a list of species records that we gathered through January 1999 to September 2002 for the forest parts of the SRET region (Appendix 1). Number of species per family recorded using different sampling methods. They were checked at least every morning. Branch & Rödel 2003). funnel traps (see Branch & Rödel 2003) were installed in a primary forest transect from 18 to 27 September 2002. which are situated south and west of TNP respectively (Rödel & Branch 2002. arboreal. unpubl. Funnel traps summed to a total of 384 trap-days. Duct tape was used to reduce the diameter of the buckets and construct funnel-like openings in order to impede escaping from traps. Pitfall trapping time summed to a total of approximately 4000 trap-days. SVTS = species recorded during visual transect walks. we applied that method during a herpetological survey of the Haute Dodo and Cavally forests. results were analyzed for leaf litter. TaïF = all species recorded with all methods in forest habitats around SRET station. The ends of each segment were flanked with additional plastic buckets. In addition. Traps were checked at least on a daily basis. Each segment was tightened around a plastic bucket with an opening angle of 45°. Additionally. one on either side. SATS = species recorded during acoustic transect walks. AES/VES = visual and acoustic encounter surveys (only species that have been recorded in forest habitats around SRET station). Hillers et al. 285 mm top diameter.
43 species were recorded in forest habitats within the study region (Fig. and P. 2. Sampling success and data quality varied between different methods. 2003. RESULTS In total. 2002..199.. Four arboreal frogs were exclusively recorded with these methods (Hyperolius picturatus. Using all methods we recorded 50 species in the region of the SRET (89. Phrynobatrachus calcaratus. Ptychadena pumilio. Three leaf litter species (Amnirana occidentalis. Ten species (23. 56 amphibian species are known to occur in TNP (Schiøtz 1967. Failure to detect particular species in this region was most probably due to their general scarcity (e. P. AES/VES. and between different amphibian families or guilds (Figs. During transect sampling (SVTS. SATS. compare Appendix 1). Two fossorial species were exclusively detected either with VES and pitfall traps (Hemisus sp.g. annulatus) were exclusively recorded during transect walks. sp. 2.RÖDEL & ERNST FIG. bibroni. see Appendix 1). Perret 1988. Taking all methods into account.g. mascareniensis. 1. Number of species per amphibian guild recorded using different sampling methods. 1.) were recorded exclusively outside forest habitats (e. P.2 %) were recorded with pitfall and funnel trapping. Hoplobatrachus occipitalis.002. Families containing only 6 a few species in the TNP area showed the highest recording rates. fusciventris. χ2 = 19. thus making overall sampling success very high (Tab. Seven species (Bufo maculatus. VES/AES revealed 39 forest species (90.3 % of TNP species). For abbreviations see Fig. H. Phrynobatrachus fraterculus. rather than actual sampling method insufficiency.0 % of TaïF) in eight families. P. regularis. Numbers above bars represent total number of species recorded with the respective method. Sampling efficiency in families containing higher numbers of genera and/or species was only marginally lower. SVTS. Trap).7 %) of nine families. Rödel 2000. 2). 1. p = 0. df = 5. tested for TaïF. Bufo superciliaris) or simply absence in the area under investigation (e. two species were each listed in two guilds (compare Appendix). clearing around SRET) and thus ignored in the analysis. . B.. N = 10.g. H. Rödel et al. Effectiveness in detecting species within given families differed significantly between methods (Friedman test. Hyperolius guttulatus). SATS) we recorded a total of 15007 individual amphibians belonging to 37 species (86. Phlyctimantis boulengeri ). Rödel & Ernst 2000. TC. nienokouensis. Appendix 1).) or pitfalls only (Geotrypetes seraphini).
N = 4. quantitative data for all arboreal species were best assembled with SATS. Diurnal ground-dwelling frogs were best recorded quantitatively using SVTS.0 72. df = 4.AMPHIBIAN MONITORING IN TROPICAL FORESTS TABLE 2..4 100. However. VES and AES revealed highest species numbers.0 100.0 0. Trapping success was highest in leaf litter frogs (56. The traps proved to be inadequate in qualitative and quantitative sampling since even small species such as Phrynobatrachus villiersi (SVL: 10–15 mm) managed to escape regularly (several direct observations).7 0. presence and abundance of purely aquatic species and tadpoles of all species were underestimated with all methods used here. Quantification of these data.0 100.0 Trap 100. the fossorial Geotrypetes seraphini was recorded with pitfall traps only. TC.5 100. data). Trap). tested for TaïF.0 81.8 100.0 100.0 36. Likewise.0 100. SATS.4 0. With the exception of Acanthixalus sonjae.0 100.0 100.0 100. No additional species were detected with funnel traps.5 % of recorded leaf litter frogs). Family / guild Caecilidae Pipidae Bufonidae Hemisotidae Ranidae Petropedetidae Astylosternidae Arthroleptidae Hyperoliidae Rhacophoridae leaf litter arboreal fossorial aquatic TaïF 1 1 2 1 5 11 1 3 17 1 22 18 2 3 TC 0.067. In contrast. In TNP funnel traps were ineffective.7 100. Branch & Rödel 2003). nocturnal leaf litter frogs (arthloeptids.0 76.0 100. 1.0 0.0 100. Most arboreal frogs could be recorded with SVTS.2 66.03 specimens per trap-day in pitfall traps).0 100. in a primary forest transect with none in the forest fragment (Hillers et al.3 We could not detect a statistical difference in sampling efficiency between different amphibian guilds using the methods employed (Friedman test.0 100. but very low in arboreal species (6.0 SVTS 0.0 SATS 0. For other abbreviations see text and Fig. all were recorded using other methods as well. which is mute.0 64. asytlosternids) were best recorded with SATS.0 77.g. but that concerned mostly single specimens.) vegetation density. was difficult as detectability of species varied considerably with (e. Schaich & Hamerle post hoc multiple comparisons revealed no significant differences between methods at α = 0. In Haute Dodo and Cavally forests we captured 38 % of the recorded amphibian species (16 of 42) with a combination of pitfall and funnel traps. All other species were recorded with VES/AES (Rödel & Branch 2002. neither between given family or guild.0 100.0 100.0 100.0 100.0 100.0 0.0 100. Percentage of species within an amphibian family or guild that were recorded using a particular method. when being a part of the transect line itself. As aquatic sites were only taken into account. Reference is the number of species that were recorded using all methods in forested habitats around the SRET station (TaïF).0 80.7 % of recorded arboreal species).8 0. VES/AES.0 100.8 0.7 0.0 50.0 0.0 66. p = 0.0 0.0 31. this was most probably due to the small overall sample size and high heterogeneity of species numbers in families and guilds (Bortz et al.0 0. We only captured a few leaf litter frogs.0 100. transect walks made it possible to search for amphibians in a very comparative way. other than based on time units. A distance of 1 m (visual) and 12.0 100.05.294.0 50.0 0.0 76.0 40. As in TNP. χ2 = 10.7 100.0 100.0 100.0 AES/VES 0.0 100. SVTS.0 77.5 100. unpubl.0 100. Most specimens were captured with funnel traps (0.8 0.47 specimens per trap-day in contrast to 0.5 m (acoustic) proved to work equally well in all habitats investigated.0 33.0 100.0 100.0 0. 1990).0 68. 7 .0 86.0 100.
could be detected exclusively using visual sampling methods. such as the very abundant Arthroleptis sp. Thus even time-based quantification of these data is questionable. Rödel et al. as well as SVTS alone. (1995) a simple comparison of species numbers may not be sufficient to evaluate the efficiency of different sampling methods. this only holds true if species numbers alone are considered. especially arboreal and fossorial. or Phrynobatrachus alticola. When including relative abundance measurements these methods were insufficient. Although widely used and recommended. SVTS was generally the best method for detecting diurnal leaf litter frogs. during rapid assessment surveys (for recent West African examples see Rödel & Branch 2002. 1996). We found it impossible to quantify VES and AES other than in relation to time. determined by using a particular sampling method.RÖDEL & ERNST DISCUSSION Choice of methods should reflect the best compromise between i: practicability. The methods VES and AES in combination with pitfall traps with drift fences were useful in sampling additional. Rödel 2003. Cardioglossa leucomystax. Pitfall trapping was only useful in detecting fossorial species that were not encountered during transect walks. In general SATS was very efficient in detecting nocturnal leaf litter frogs and treefrogs but less successful in detecting diurnal leaf litter frogs. In addition. The methods tested within this study proved to perform with varying degrees of efficiency. but is certainly just as well suited for the investigation of tropical anuran assemblages in general. (1997) and Rödel (1998a). Doan 2003). Some taxa. We found it more realistic to apply a (clearly subjective) measure for habitat complexity on which to base collection effort during VES/AES. Rödel & Ernst 2003). The percentage of species within supraspecific taxonomic units. VES and AES are useful tools for the compilation of species inventory lists. Standard plot sampling and total removal plots were not tested. pitfall traps and drift fences . such as the Hylidae. Olson et al. Schiøtz 1999. as they impose a severe disturbance on the system under investigation and thus were not suitable to our goals. especially in long-term field studies within temperate regions (e. Arthroleptis sp. but appear not to be appropriate when standardized quantitative sampling is required. ii: specific aims. such as species of the genus Acanthixalus. Nonetheless. Bury & Corn 1987. Compiling data on presence and abundance of aquatic species and tadpoles was outside the scope of this study and these species and stages were inadequately sampled with all methods used here.g. inaccessible areas than in areas that are easy to monitor. Although the combination of AES and VES provided an adequate approximation to real conditions (see also Doan 2003). data acquisition varies considerably depending on habitat type in VES and even AES.1. such as species groups. these methods do not seem to sample a given fauna adequately well (Allmon 1991. and iv: environmental impact. were nearly as successful in detecting species. However. a combination of SVTS and SATS. However. Vonesh 2001. The differences in efficiency of visual versus acoustic sampling varied between taxa. This was also true for most of the arboreal species belonging to the families Hyperoliidae and Rhacophoridae. species not encountered during transect sampling. VES and AES yielded highest species numbers. can then be compared to the real conditions in an area or habitat. in order to sample particularly secretive leaf litter species. They suggest using relative species richness within taxonomic entities. According to Pearman et al. as an adequate measure. Whereas transect walks provide the possibility of monitoring amphibians in a very comparative way. e. Simple presence/absence data of aquatic species and stages can best be gathered by direct observation (surfacing individuals) and with dip-netting.. iii: efficiency. The combined transect sampling methods provided a close approximation to the real presence and abundance of species. which are mute (Drewes 1984. Methods of collecting quantitative data for aquatic amphibians and for tadpole assemblages have been described by Shaffer et al. acoustic sampling appeared to be indispensable since these species were more readily detected by their calls than 8 by sight. whereas visual sampling appeared to be superior to acoustic sampling.2. and failed to detect aquatic and fossorial species. Semlitsch et al.g. (1994). Migrating pipid frogs can also be easily recorded with pitfall and funnel traps. genera or families. spending more time in complex. Therefore transect sampling can be considered to be the method of choice when sampling leaf litter anurans. and will likewise hold true for other arboreal anurans throughout the tropics. 2003). In this study this has been done by determining relative species richness with reference to families or guilds. We found no additional species and only very few specimens by funnel trapping. The advantage of transect sampling was especially obvious when focusing on leaf litter species..
Costs with regard to time spent for data acquisition was highest in trapping and in transect walks. be quantified in relation to time. however. experience of investigator and prevailing weather conditions (see Branch & Rödel 2003). SVTS and SATS provided data on species and habitats und thus may reveal changes in composition and species abundance. as well as environmental changes. if not on amphibians then on other organisms.AMPHIBIAN MONITORING IN TROPICAL FORESTS proved to be the least effective method. amphibian guild and environmental impact. However. one should test the efficiency at a particular site in advance before installing these traps on a broad scale.. 2001. seasons and years. We observed (e. for other abbreviations see text and Fig. VES and AES are less time consuming. in theory. (2001) report on similar experiences using this method in a herpetological survey in Guyana. A comparison of trapping data with data other than those of the observer in question seems to be difficult. When considering the relatively high costs and time investment due to the high level of maintenance required (Parris 1999). as trapping success per trap per time unit. Rodda et al. e. accounted for 72. QS/TR = quadrat sampling/total removal sampling (data from Allmon 1991. Donnelly et al. the fossorial Hemisus perreti and the aquatic Silurana epitropicalis. However trapping success depends very much on choice of site. traps have to be uninstalled or closed and reopened when starting again. 1. VES and AES had probably the lowest environmental impact. this method cannot be recommended for standardized sampling of tropical anurans. TC leaf litter frogs arboreal frogs fossorial amphibians aquatic frogs qualitative data quantitative data environmental impact costs (time) costs (material) utility (ecology) utility (conservation) +++ ++ – +++ +++ +++ low high low very high very high SVTS +++ ++ – +++ +++ ++ low high low high high SATS ++ ++ – – +++ ++ low high low high high AES/VES +++ +++ + +++ + + very low low very low low high Trap + – +++ + + – medium very high low very low low QS/TR +++ – – – +++ +++ very high high low very low low 9 . Burger et al. Thus data should be gathered throughout a whole season to obtain a thorough knowledge of the local fauna. Traps have to be checked at least daily. If the investigation is interrupted for some time. Transects will have an environmental impact. Theoretically it might well be that trapping will result in dislocating at least some species from the area where traps have been established. Effectiveness of methodology in regard to type of data required. They collected only six species (out of a total of 132 species of amphibians and reptiles being recorded). However only two species.g. With the exceptions of aquatic and fossorial species they pro- vided data that cover the whole community very well. the frequency of sampling can be reduced when general phenological traits of the species recorded have been clarified. in consecutive field seasons. Doan 2003). (2004) had high trapping success with pitfalls in Gabon when considering numbers of individuals. Data can be used for comparisons between habitats.6 % of all individuals captured with this method. SVTS and SATS have been proven to be the only methods that provided quantitative data on forest amphibians with regard to space and time. The impact of traps and drift fences was difficult to judge. In our experience. none of them uniquely found by this method. Parris 1999.g.) that leopard (Panthera pardus) tracks TABLE 3. generalization of data with regard to phenology or abundances is less possible. It is advisable to perform them randomly independent of prevailing weather conditions. Transect walks are time intensive. Vonesh 2001. Data from both trapping methods can.
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1 = known from TNP (Schiøtz 1967. f = fossorial species. 4 = Acanthixalus sonjae lives aquatically in large water filled tree holes but forages in surrounding vegetation (Rödel et al. Trap = species recorded with pitfall and funnel traps. species Gymnophiona Caecilidae Geotrypetes seraphini Anura Pipidae Silurana tropicalis Bufonidae Bufo regularis Bufo maculatus Bufo togoensis Bufo taiensis Bufo superciliaris Hemisotidae Hemisus sp. 2 = determination not possible at present time. 3 = Phrynobatrachus guineensis is a leaf litter frog but reproduces in small water-filled tree holes (Rödel 1998b). TC = all species recorded on transects. G = species recorded in southern TNP (Guiroutou. 2 Petropedetidae Phrynobatrachus accraensis Phrynobatrachus gutturosus Phrynobatrachus fraterculus Phrynobatrachus guineensis 3 Phrynobatrachus phyllophilus Phrynobatrachus liberiensis Phrynobatrachus alticola Phrynobatrachus alleni Phrynobatrachus plicatus TaïF G SRET TC TP TS VES/AES Trap guild 1 0 0 0 0 0 0 1 f 1 0 0 1 1 0 1 0 1 1 1 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 1 1 1 0 1 0 0 1 1 1 1 1 1 0 1 1 1 1 1 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 0 0 1 1 1 0 0 0 0 0 0 0 1 1 0 0 1 1 0 0 0 1 1 1 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 0 0 1 1 1 0 0 0 0 1 0 0 0 1 1 1 1 1 1 1 1 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 1 1 0 1 0 1 1 1 1 1 1 1 1 0 0 1 1 0 1 0 1 0 1 0 0 0 0 1 1 0 1 1 0 1 1 1 1 1 1 1 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 aq l l l l l f aq l l aq l l l l l l l l l l l. 7°10’ W). 6 = recorded north of area under investigation. 2003). 5 = treated as full species and not as subspecies of Hyperolius fusciventris due to sympatric occurrence in TNP (see Rödel & Branch 2002). VES/AES = species recorded via visual and acoustic encounter surveys in forest habitats near SRET. TaïF = species recorded in forest habitats around the SRET station. a = arboreal species. TP = all species recorded on transects in primary forest. TS = all species recorded on transects in secondary forest. aq = aquatic species. 2 Ranidae Hoplobatrachus occipitalis Amnirana albolabris Amnirana occidentalis Aubria occidentalis Ptychadena pumilio Ptychadena bibroni Ptychadena mascareniensis Ptychadena superciliaris Ptychadena aequiplicata Ptychadena longirostris Ptychadena sp. 5°25’ N. List of amphibian species recorded in Taï National Park (TNP). other than transects.AMPHIBIAN MONITORING IN TROPICAL FORESTS APPENDIX. a l l l l l 13 . SRET = species recorded with VES/AES on the clearing around the SRET station. l = leaf litter species. Perret 1988) but not recorded by us.
aq a 14 . 2 2 Hyperoliidae Leptopelis hyloides Leptopelis occidentalis Leptopelis macrotis Hyperolius concolor Hyperolius guttulatus 6 Hyperolius picturatus Hyperolius sylvaticus Hyperolius zonatus Hyperolius fusciventris 5 Hyperolius lamtoensis 5 Hyperolius nienokouensis Hyperolius wermuthi 1 Hyperolius chlorosteus Afrixalus dorsalis Afrixalus nigeriensis Afrixalus vibekae Kassina lamottei Phlyctimantis boulengeri Acanthixalus sonjae 4 Rhacophoridae Chiromantis rufescens total 56 species in TNP TaïF 0 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 43 G 1 0 1 0 1 1 1 1 1 1 1 1 0 1 1 1 0 1 1 0 1 1 1 0 1 0 0 1 41 SRET 0 0 0 0 0 0 1 0 1 0 0 1 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 16 TC 0 0 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 1 0 0 1 1 1 1 1 0 1 1 37 TP 0 0 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 1 0 0 1 0 1 1 1 0 0 1 33 TS VES/AES Trap 0 0 1 0 1 1 1 1 1 1 1 0 0 0 1 1 0 0 0 0 1 1 1 0 0 0 1 1 28 0 0 1 0 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 39 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 10 guild l l l l l l l l a a a a a a a a a a a a a a a a l a a. 12 Arthroleptis sp.RÖDEL & ERNST Appendix continued species Phrynobatrachus calcaratus Phrynobatrachus taiensis 1 Phrynobatrachus villiersi Phrynobatrachus annulatus Astylosternidae Astylosternus occidentalis Arthropleptidae Cardioglossa leucomystax Arthroleptis sp.
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