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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
and unpubl. known as quadrat sampling (plot design). Given the lack of time for the development of effective conservation programs.RÖDEL & ERNST essential for academic and practical purposes alike. Floristically the TNP belongs to the Guinean-Congo-Region (Guillaumet 1967. Parren & de Graaf 1995. ratures vary between 20 and 33° C. We thereby evaluated several methods that are already commonly used and present a set of techniques that. situated in western Côte d’Ivoire (5°08’6°07’N. Poynton 1999). The first. Transect paths were kept open so that walking at a constant speed was possible at all times. The minimum distance between adjacent transects was 200 m. It can be characterized as humid tropic seasonal (Riezebos et al. 6°47’-7°25’W). The maximum distance between transects was 6. known as transect sampling. The major rainy season stretches from September to October. uses linear transects instead of squares. four in secondary forest). The starting coordinate for each transect always marked the southeast corner to ensure identical geographic orientation between sampling units. 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. The mean annual temperature is about 25° C (Rompaey 1993). standardization of methods that yield broad-scale comparative data is now needed more than ever. We aim to present a methodology that i) provides quantitative and qualitative data useful for rapid surveys as well as long-term monitoring programs. 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. Transects were arranged in pairs. 1994. as well as their relative abundances and densities. thus ensuring that all habitat types of a certain area within the very inhomogeneous forest were covered. More detailed descriptions of TNP are provided by Rompaey (1993).3 km. also for para-ecologists. Mean annual precipitation in the study area was 1806 mm during 1988–2002 (± 297mm. followed by a short dry season in July/August. PACPNT 2000). (1994) and PACPNT (2000). data: Taï Monkey Project). 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. TNP is part of the Upper Guinea forest block. Our study sites are either pristine forest areas or have not been logged or cultivated since 1978. 7°20’W). Daily tempe2 . The core dry season lasts from November until February/March. Doan 2003) or total removal plots (Barbault & Trefaut Rodrigues 1978. b. Lawson 1986. transect sampling can provide similar data across habitat or dis- MATERIAL AND METHODS Study site The study was conducted in Taï National Park (TNP). Each had a north-south extension of 200 m and an east-west extension of 100 m. map “Projet OMS Forêt de Taï”). The rectangular transect design is a combination of two widely used standard techniques. Rodda et al. such as standard plot sampling (Allmon 1991. comm. consists of a series of small squares (quadrats) that are laid out at randomly selected sites within a habitat then thoroughly searched for amphibians. on the basis of our experience in the field as well as with consecutive data analyses. proved to be most effective for forest amphibians. Riezebos et al. that ii) is easy to handle. 5°50’N. 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. 2001) were a priori excluded from testing. A minor rainy season lasts from March/April to July. that stretches west of the Dahomey Gap from Ghana into Sierra Leone and Guinea (Schiøtz 1967. 2001. Africa: Howell 2002) reflect this need. Côte d’Ivoire. Whereas quadrat sampling can be used to determine the species present in a homogenous area. We avoided extensive cutting and thus manipulation of important habitat features. Every subunit was marked with numbered colored flag-tape. and iv) has a low environmental impact. Vonesh 2001. Parts of the habitats on the park’s periphery were subject to more or less intense logging and cultivation until 1998 (P. A number of recently published regional field monitoring manuals for amphibian surveys (Latin America: Lips et al. Richards 1996). as they may result in severe disturbance to the system under investigation. All evaluations and recommendations are based on a study of tropical amphibian assemblages conducted over four years (1999–2002) in Taï National Park. Thus widely used techniques. Formenty pers. The second. Precipitation is distributed across two more or less distinct periods. TNP is the largest remaining protected area of rain forest in West Africa. 1979a. iii) is efficient concerning time and money. For data acquisition the complete transect length of 600 m was subdivided in 25 m subunits (24 subunits/transect).
thereby recording all amphibians within a distance of 100 cm from either side of the path. Sampling was performed independent of prevailing weather conditions. Repeated controls of identical transects on consecutive days were avoided to ensure independence of samples. and to investigate correlations of species assemblages with environmental variables. Sampling methods and sampling effort Habitat characterization.AMPHIBIAN MONITORING IN TROPICAL FORESTS turbance gradients (Jaeger 1994. thus providing data that are useful to test for sex. substrate moisture was determined in four categories during every transect walk. divided into seven categories corresponding to particular densities. to assign them to particular guilds).5 m..5 2 2. lower tree stratum: 3–10 m. Categories of habitat variables measured on transects. 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. In order to quantify the availability of potential aquatic breeding sites. In addition to species identity we recorded sex and snout-vent-length (SVL) of every individual. These parameters included vegetation density in four strata (canopy: > 20 m.. Pearman 1997). Definitions of habitat variables are summarized in Table 1. Jaeger & Inger 1994).5 m). To assess habitat preferences of particular species (e. vegetation density was measured in four strata (see text). we characterized all habitats using several variables that were recorded at two defined points within each 25 m subunit (beginning and midpoint). bush and shrub stratum: 0. As far as possible all individuals were captured. Standardized visual transect sampling (SVTS). 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. for comparison of local with regional species pools). Sampling was interrupted for the duration of the recording in order not to overestimate locations in which animals had previously been captured.g. To avoid duplicate recordings. secondary forests.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 .5 3 3. 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.35 m/s). captured frogs were marked by TABLE 1. rectangular ones provide the possibility of easily up.5–1. whereas primary for- ests show increasing numbers of plants of larger dbh (Chatelain et al. Usually four to six transects per day were sampled during daytime. Additionally.or age-specific differences in habitat use.g. dbh = diameter of plants at breast height.or downscaling data without neglecting local habitat diversity (e. understory: < 0. Compared to linear transects. In addition.30–0. We assumed that the number of plants with small dbh is greater in degraded. Transects were intensively patrolled at a constant speed (0. dbh). surface and depth. A maximum of four transects was patrolled at night. Specimens heard but not seen within this distance were not searched for and not included in the SVTS data sets (see below). definition category 1 1. For a discussion of the advantages of rectangular sample units over quadratic or circular ones see Krebs (1989) and McCune & Grace (2002). both in time and space. 1996. Edaphic parameters were registered as general substrate types according to Lieberoth (1982).
1994. and to estimate relative abundances of species within an assemblage (Crump & Scott 1994). Coding schemes used for individual recognition were not applied. For that reason a maximum recording distance of 12. Recaptures were excluded from the analyses. beginning with segment 1: segment 1. According to Corn & Bury (1990). species composition.5 m high and stapled vertically onto wooden stakes. In concordance with these authors the methods were only used as a qualitative or semi-quantitative tool within the scope of our studies. Bufo species). and additionally in only one of these transects from January 2001 to September 2002. Likewise microhabitat description was not always possible. following the characterization routine used for general habitat description. An area or habitat is searched systematically for individuals in a defined time period. this technique allows the detecting of cryptic species that. Audio strip transects represent a commonly used method for acoustic monitoring (Zimmerman 1994). The width of the acoustic transect depends on the ability to detect each species’ advertisement call. Standardized acoustic transect sampling (SATS). Since in the majority of frog species males use species-specific calls to advertise their position to potential mates and rivals (Wells 1977). At capturing sites a thorough description of the microhabitat was recorded. VES can only provide information on the presence or absence of a species in an area but are inadequate for determining abundances. Dodd 1991). 8. 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. which can be adjusted to the complexity of the habitats being sampled.. From February 1999 to December 2000 we compiled 382.RÖDEL & ERNST toe clipping (Donnelly et al. individual marking should be restricted to studies in which individual recognition is indispensable. Anurans that are strong jumpers (e. In the transect corners...e. VES and AES are frequently used for rapid assessments and the evaluation of larger areas. Visual (VES) and acoustic (AES) encounter surveys. Counts can be used to estimate relative abundance of calling males. Total VES.5 m to either side of the transect was defined. Furthermore. 0. Drift fences consisted of durable green plastic gauze.g. keeping the transect routine identical for both techniques. This technique has been formalized as the time-constrained technique of Campbell & Christman (1982). As opposed to visual sampling. This device is useful to determine species richness of epigaeic organisms (Corn 1994). Trapping with pitfall or funnel traps along drift fences. It can be used to determine the species richness of an area and the species composition of a local assemblage. Ptychadena species) are more difficult to trap than terrestrial species that lack these abilities (e.g.and AES-effort was kept comparable to the time spent on SVTS and SATS surveys throughout the entire study period. We used these methods continuously in all habitats of the study area including the transect areas. and as the time-constrained searches of Corn & Bury (1990). Thus. Individuals below nine mm SVL were not marked due to their small size. 12 and 20). this behavior can be exploited for acoustic monitoring. only calls coming from the right-hand side in the first segments were registered (i. 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).5 4 hours of transect sampling. data can be expressed in numbers of individuals per time and surface units. In those cases. during SATS it was often impossible to determine individual parameters. habitat analyses can be based on data obtained from the general habitat characterization for single transects or segments. This corresponds to 765 transect walks. The resulting data are expressed in numbers of individuals of a certain species found in an area per unit time. Calls from greater distances cannot be unambiguously identified and the chance of duplicate recordings in neighboring segments increases. as well as breeding habitat use and breeding phenology of species (Zimmerman 1994). other than sex and species. thus creating 25 m x 25 m acoustic sampling plots. despite their potential abundance. Throughout transect walks a combination of visual and acoustic techniques was applied at all times. Pitfall traps with drift fences were installed in two of the primary forest transects from March to April 1999. Henle et al. For practical reasons “man-hours” can be used. Capture success may vary greatly between species (Corn & Bury 1990. Due to the simplicity of the method. An array of fences and traps consisted of a central trap (buckets: . may be underestimated when exclusively using visual techniques. 1997). For both SVTS and SATS. VES and AES thus could be used to evaluate if the randomly chosen transects represented the regional species pool.
Additionally. which are situated south and west of TNP respectively (Rödel & Branch 2002. one on either side. 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). Sampling success is referred to as number of species within families recorded. 285 mm top diameter. Trap = pitfall and funnel traps. Traps were checked at least on a daily basis. as sampling success and efficiency may vary with regard to the particular biology of a species. TC = species recorded during transect walks (SVTS and SATS combined). SATS = species recorded during acoustic transect walks. Duct tape was used to reduce the diameter of the buckets and construct funnel-like openings in order to impede escaping from traps. fossorial and aquatic species independent of families. Numbers above bars represent total number of species recorded with the respective method. Funnel traps summed to a total of 384 trap-days. In addition. using a particular sampling method. arboreal. results were analyzed for leaf litter. unpubl. Number of species per family recorded using different sampling methods. Branch & Rödel 2003). SVTS = species recorded during visual transect walks. Hillers et al. 1. data). In the course of this study. AES/VES = visual and acoustic encounter surveys (only species that have been recorded in forest habitats around SRET station). They were checked at least every morning. The ends of each segment were flanked with additional plastic buckets. funnel traps (see Branch & Rödel 2003) were installed in a primary forest transect from 18 to 27 September 2002. FIG. Pitfall trapping time summed to a total of approximately 4000 trap-days.AMPHIBIAN MONITORING IN TROPICAL FORESTS 275 mm deep.. 5 Ta ïF . we applied that method during a herpetological survey of the Haute Dodo and Cavally forests. 220 mm bottom diameter) and two triangular fence segments (total length 16 m). TaïF = all species recorded with all methods in forest habitats around SRET station. Each segment was tightened around a plastic bucket with an opening angle of 45°. and in a forest fragment (Paulé-Oula 2) outside TNP from 30 August to 12 September 2002 (A. The data from pitfall and funnel traps can be expressed as number of individuals per trap-day.
H.199. Sampling efficiency in families containing higher numbers of genera and/or species was only marginally lower. During transect sampling (SVTS.. Appendix 1). Hoplobatrachus occipitalis.) were recorded exclusively outside forest habitats (e.g. rather than actual sampling method insufficiency.0 % of TaïF) in eight families. AES/VES. Effectiveness in detecting species within given families differed significantly between methods (Friedman test. annulatus) were exclusively recorded during transect walks. TC. and P.g.3 % of TNP species). For abbreviations see Fig. 1. Sampling success and data quality varied between different methods. Perret 1988. bibroni. P. two species were each listed in two guilds (compare Appendix). SVTS. compare Appendix 1). mascareniensis. fusciventris. . Rödel et al. thus making overall sampling success very high (Tab. Hyperolius guttulatus). 2. SATS. Ten species (23. Bufo superciliaris) or simply absence in the area under investigation (e. χ2 = 19. 2.g. nienokouensis.. 2003. Rödel 2000. clearing around SRET) and thus ignored in the analysis. Number of species per amphibian guild recorded using different sampling methods. regularis.RÖDEL & ERNST FIG.2 %) were recorded with pitfall and funnel trapping. Trap). Taking all methods into account. Four arboreal frogs were exclusively recorded with these methods (Hyperolius picturatus. 56 amphibian species are known to occur in TNP (Schiøtz 1967. tested for TaïF. B. 2002. 2). N = 10. Using all methods we recorded 50 species in the region of the SRET (89. Families containing only 6 a few species in the TNP area showed the highest recording rates. p = 0. 1. see Appendix 1).002. H. Phrynobatrachus calcaratus. sp. df = 5. Numbers above bars represent total number of species recorded with the respective method. Ptychadena pumilio. Seven species (Bufo maculatus. VES/AES revealed 39 forest species (90. P. Two fossorial species were exclusively detected either with VES and pitfall traps (Hemisus sp. 1. 43 species were recorded in forest habitats within the study region (Fig. Three leaf litter species (Amnirana occidentalis. P. Failure to detect particular species in this region was most probably due to their general scarcity (e. Phrynobatrachus fraterculus. RESULTS In total. SATS) we recorded a total of 15007 individual amphibians belonging to 37 species (86..7 %) of nine families.) or pitfalls only (Geotrypetes seraphini). Phlyctimantis boulengeri ). Rödel & Ernst 2000. and between different amphibian families or guilds (Figs.
0 100.0 50. df = 4.0 100. quantitative data for all arboreal species were best assembled with SATS.7 0. As in TNP. SATS. In contrast. when being a part of the transect line itself.0 0. Quantification of these data. 7 .0 100. We only captured a few leaf litter frogs.0 86.0 100. Percentage of species within an amphibian family or guild that were recorded using a particular method. all were recorded using other methods as well. A distance of 1 m (visual) and 12..5 100. in a primary forest transect with none in the forest fragment (Hillers et al. For other abbreviations see text and Fig.0 33.4 100. nocturnal leaf litter frogs (arthloeptids.0 77.7 100.0 100.8 100.05.0 66.0 Trap 100.8 0. transect walks made it possible to search for amphibians in a very comparative way.0 100.0 100.0 68.0 81. the fossorial Geotrypetes seraphini was recorded with pitfall traps only.0 50.5 100. 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 0.0 36. 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.0 100. 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).0 72. However. unpubl.0 100. Branch & Rödel 2003).0 0.0 100.0 80.0 100.0 76. this was most probably due to the small overall sample size and high heterogeneity of species numbers in families and guilds (Bortz et al.294. 1990). Most specimens were captured with funnel traps (0.AMPHIBIAN MONITORING IN TROPICAL FORESTS TABLE 2.7 100. χ2 = 10.0 0. With the exception of Acanthixalus sonjae. Most arboreal frogs could be recorded with SVTS. presence and abundance of purely aquatic species and tadpoles of all species were underestimated with all methods used here.0 40. Schaich & Hamerle post hoc multiple comparisons revealed no significant differences between methods at α = 0.0 100. Reference is the number of species that were recorded using all methods in forested habitats around the SRET station (TaïF).0 77.0 31.7 0. All other species were recorded with VES/AES (Rödel & Branch 2002. Trap). Likewise. asytlosternids) were best recorded with SATS.0 100.0 100.8 0. data).067. Trapping success was highest in leaf litter frogs (56. but that concerned mostly single specimens.) vegetation density.0 76.0 100.0 100. No additional species were detected with funnel traps.47 specimens per trap-day in contrast to 0. N = 4. Diurnal ground-dwelling frogs were best recorded quantitatively using SVTS.0 100. p = 0. VES and AES revealed highest species numbers. TC. SVTS. other than based on time units.0 100.0 100.0 100. tested for TaïF. but very low in arboreal species (6.03 specimens per trap-day in pitfall traps).8 0.0 0.5 m (acoustic) proved to work equally well in all habitats investigated.0 100.0 100. was difficult as detectability of species varied considerably with (e.0 100.7 % of recorded arboreal species). which is mute.0 0.g.0 64.0 SATS 0.0 0.0 AES/VES 0.0 100. As aquatic sites were only taken into account.0 0. neither between given family or guild.0 100. VES/AES.2 66.0 SVTS 0. In TNP funnel traps were ineffective. 1.0 100.0 0.3 We could not detect a statistical difference in sampling efficiency between different amphibian guilds using the methods employed (Friedman test.5 % of recorded leaf litter frogs).4 0.0 100.
We found no additional species and only very few specimens by funnel trapping. 2003). The methods tested within this study proved to perform with varying degrees of efficiency. In this study this has been done by determining relative species richness with reference to families or guilds. and failed to detect aquatic and fossorial species. Bury & Corn 1987. such as the very abundant Arthroleptis sp. VES and AES are useful tools for the compilation of species inventory lists. Schiøtz 1999. VES and AES yielded highest species numbers. Methods of collecting quantitative data for aquatic amphibians and for tadpole assemblages have been described by Shaffer et al. these methods do not seem to sample a given fauna adequately well (Allmon 1991. such as species of the genus Acanthixalus. species not encountered during transect sampling. The advantage of transect sampling was especially obvious when focusing on leaf litter species. as an adequate measure. which are mute (Drewes 1984. We found it more realistic to apply a (clearly subjective) measure for habitat complexity on which to base collection effort during VES/AES. pitfall traps and drift fences .g. The differences in efficiency of visual versus acoustic sampling varied between taxa. Doan 2003). in order to sample particularly secretive leaf litter species. and iv: environmental impact. such as the Hylidae. Rödel 2003. acoustic sampling appeared to be indispensable since these species were more readily detected by their calls than 8 by sight. We found it impossible to quantify VES and AES other than in relation to time. genera or families. SVTS was generally the best method for detecting diurnal leaf litter frogs. Simple presence/absence data of aquatic species and stages can best be gathered by direct observation (surfacing individuals) and with dip-netting. during rapid assessment surveys (for recent West African examples see Rödel & Branch 2002. However. Rödel et al. The methods VES and AES in combination with pitfall traps with drift fences were useful in sampling additional. When including relative abundance measurements these methods were insufficient. According to Pearman et al. determined by using a particular sampling method. In addition. (1994). Nonetheless. whereas visual sampling appeared to be superior to acoustic sampling. could be detected exclusively using visual sampling methods. Thus even time-based quantification of these data is questionable. as they impose a severe disturbance on the system under investigation and thus were not suitable to our goals. e.g. especially arboreal and fossorial. Whereas transect walks provide the possibility of monitoring amphibians in a very comparative way.1. as well as SVTS alone. iii: efficiency. In general SATS was very efficient in detecting nocturnal leaf litter frogs and treefrogs but less successful in detecting diurnal leaf litter frogs. data acquisition varies considerably depending on habitat type in VES and even AES. Cardioglossa leucomystax. Rödel & Ernst 2003). Migrating pipid frogs can also be easily recorded with pitfall and funnel traps. spending more time in complex. a combination of SVTS and SATS. Vonesh 2001. However. The combined transect sampling methods provided a close approximation to the real presence and abundance of species. and will likewise hold true for other arboreal anurans throughout the tropics. Although widely used and recommended. (1997) and Rödel (1998a). this only holds true if species numbers alone are considered. Semlitsch et al. can then be compared to the real conditions in an area or habitat. 1996).2. Some taxa. Olson et al. but is certainly just as well suited for the investigation of tropical anuran assemblages in general.RÖDEL & ERNST DISCUSSION Choice of methods should reflect the best compromise between i: practicability. (1995) a simple comparison of species numbers may not be sufficient to evaluate the efficiency of different sampling methods. were nearly as successful in detecting species. 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.. or Phrynobatrachus alticola. such as species groups. but appear not to be appropriate when standardized quantitative sampling is required. Therefore transect sampling can be considered to be the method of choice when sampling leaf litter anurans. This was also true for most of the arboreal species belonging to the families Hyperoliidae and Rhacophoridae. especially in long-term field studies within temperate regions (e. inaccessible areas than in areas that are easy to monitor.. The percentage of species within supraspecific taxonomic units. Standard plot sampling and total removal plots were not tested. Pitfall trapping was only useful in detecting fossorial species that were not encountered during transect walks. Although the combination of AES and VES provided an adequate approximation to real conditions (see also Doan 2003). They suggest using relative species richness within taxonomic entities. ii: specific aims. Arthroleptis sp.
With the exceptions of aquatic and fossorial species they pro- vided data that cover the whole community very well. Doan 2003). They collected only six species (out of a total of 132 species of amphibians and reptiles being recorded). Data from both trapping methods can. in theory. However trapping success depends very much on choice of site. (2001) report on similar experiences using this method in a herpetological survey in Guyana.g.) that leopard (Panthera pardus) tracks TABLE 3. QS/TR = quadrat sampling/total removal sampling (data from Allmon 1991. e. A comparison of trapping data with data other than those of the observer in question seems to be difficult. Transects will have an environmental impact. Rodda et al.6 % of all individuals captured with this method.. Transect walks are time intensive. 2001. Costs with regard to time spent for data acquisition was highest in trapping and in transect walks. If the investigation is interrupted for some time. In our experience. accounted for 72. SVTS and SATS have been proven to be the only methods that provided quantitative data on forest amphibians with regard to space and time. be quantified in relation to time. as trapping success per trap per time unit. SVTS and SATS provided data on species and habitats und thus may reveal changes in composition and species abundance. Burger et al. Theoretically it might well be that trapping will result in dislocating at least some species from the area where traps have been established. as well as environmental changes.g. Thus data should be gathered throughout a whole season to obtain a thorough knowledge of the local fauna. 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 . for other abbreviations see text and Fig. one should test the efficiency at a particular site in advance before installing these traps on a broad scale. When considering the relatively high costs and time investment due to the high level of maintenance required (Parris 1999). Data can be used for comparisons between habitats. Parris 1999. VES and AES had probably the lowest environmental impact. Effectiveness of methodology in regard to type of data required. The impact of traps and drift fences was difficult to judge.AMPHIBIAN MONITORING IN TROPICAL FORESTS proved to be the least effective method. Vonesh 2001. the fossorial Hemisus perreti and the aquatic Silurana epitropicalis. However. We observed (e. However only two species. VES and AES are less time consuming. none of them uniquely found by this method. Donnelly et al. experience of investigator and prevailing weather conditions (see Branch & Rödel 2003). It is advisable to perform them randomly independent of prevailing weather conditions. the frequency of sampling can be reduced when general phenological traits of the species recorded have been clarified. however. generalization of data with regard to phenology or abundances is less possible. seasons and years. this method cannot be recommended for standardized sampling of tropical anurans. if not on amphibians then on other organisms. amphibian guild and environmental impact. in consecutive field seasons. traps have to be uninstalled or closed and reopened when starting again. (2004) had high trapping success with pitfalls in Gabon when considering numbers of individuals. Traps have to be checked at least daily. 1.
L.. noire. Department of the Interior.A. Ecol. Transect cutting and the use of transects should therefore be done with the necessary precautions. Sér. Evaluation of pitfall trapping in northwestern forests: trap arrays with drift fences. Hayek. Canopy fogging of an overstory tree – recommendations for standardization. 1979a.. 5: 37–53. Olson et al. Fish and Wildlife Service 13. Biodiv. Donnelly. & M. Trefaut Rodrigues..R... Linsenmair.-O. For simple short-term surveys we recommend VES and AES. Branch. R. J.J. possibly accompanied by opportunistic trapping. Standard methods for amphibians.. J. Grundzüge der Vegetationskunde.. Branch. U. R. Springer Verlag. C. Rev.. & M. 10 . 1982. Smithsonian Institution Press. western Ivory Coast. This might affect densities of other mammals including possible seed dispersers with. All this support is gratefully acknowledged.P. Bull.Y. a well visible path becomes established that at least in primary forests remains so for years. Halliday.S. Furthermore transects provide an effective method of investigating leaf litter frogs in particular. Trefaut Rodrigues.R. Alford. 1978. Gabon: Species turnover along an elevational gradient. Corn. Pp. Bury. 20: 64–77. R.G..E.J. Gbamlin. However. Lienert. Ptychadena maccarthyensis et Ptychadena oxyrhynchus. consequent long-term effects on forest structure.. Global amphibian declines: a problem in applied ecology. Pflanzensoziologie. Christman. 1990. 2003.S. and the “Taï Monkey Project” provided logistic support. Ecol. T. thus generating a much more complete picture of an amphibian community than is possible with other methods. 2004. Terre Vie 32: 441–452. Richards. Foster. Allmon. & S. Conserv. 1999. 01 LC0017).D. McDiarmid. Project W08 BIOTA-West..o.” G. Wildlife Research Report 13. Boehnke. A 41: 417–428. Herpetological survey of the Haute Dodo and Cavally forests.Côte d’Ivoire helped with transportation. 1996. Glos. & S. M. Amphibians and reptiles of Monts Doudou. Basset. Channing. G. Brazil. & M. W. Pp. Ecol. After using transects intensively for some time.C. W. & P..S. 1994.W. Jr. fond. A. Arthroleptis poecilonotus. Verteilungsfreie Methoden der Biostatistik. 1964. and B. Field techniques for herpetofaunal community analysis. (ed. Acad. 1991. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux IV. 109–117 in Heyer. M.. in the worst case. Phrynobatrachus accraensis. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux I. 2 ed. 1997. and C. & R.. Syst. Sci.. J. Memoir 28: 145–186. Gautier. J. 1987. P. Rödel 1998a). Salamandra 39: 91–110. & K. Floren.. Lips et al. P. The access permit to TNP was issued by the “Ministère de la Construction et de l’Environnement. Hammond. TROPENBOS . Part II: Trapping results and reptiles.. Campbell. Afr. Bortz.. J. Research permission was given by the “Ministère de l’Enseignement Supérieur et de la Recherche Scientifique. R. W.RÖDEL & ERNST were more often seen on the transect path than in undisturbed forest. Advantages and disadvantages of different methods are summarized in Tab. Ann. Barbault.1964. H. Ouoro provided invaluable help during fieldwork.. Trop. Central Amazon. 30: 133–165. Berlin a. Hillcoat provided valuable comments on the manuscript. (eds. & M.. R.B. U. Y.” of the Republic of Ivory Coast. Corn. J. ACKNOWLEDGMENTS This study is part of the BIOLOG-program of the German Ministry of Education and Science (BMBF. 2003) transects should be supplemented by specifically investigating aquatic breeding sites with the appropriate set of standardized techniques (Heyer et al..S.-A. L. Hofer. for specific questions (cf.): Herpetological communities. Trop.R. The “Projet Autonome pour la Conservation du Parc National de Taï”. Wildlife Management 51: 112–119. & M. A plot study of forest floor litter frogs.M.): Measuring and monitoring biological diversity. 939 pp.W. Burger. Calif... N. Braun-Blanquet. Wien. Springer Verlag.A. Various administrative services and lodging facilities in TNP were provided by the “Centre de Recherche en Ecologie” and the “Projet Autonome pour la Conservation du Parc National de Taï. Chatelain. Rödel. Barbault. 1998. 193–200 in Scott. & A. Spichiger.” Working in TNP was kindly permitted by the Commandant K. SVTS and SATS are especially useful for long-term studies.A. Straight-line drift fences and pitfall traps. R. 865 pp. 1979b. 364 pp. Barbault. Ecotropica 4: 93–97. 3. not only on their breeding sites but throughout the whole range of habitats used by them. Inst. N’Dri. 7: 503–522. W. Trefaut Rodrigues. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux III. A recent history of forest fragmentation in southwestern Ivory Coast. & K. The presented transect design proved to be adequate for representative sampling and appeared to be appropriate for most studies involving multivariate structure data. Washington & London. REFERENCES Adis. 1994.
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AMPHIBIAN MONITORING IN TROPICAL FORESTS APPENDIX. a = arboreal species. 5 = treated as full species and not as subspecies of Hyperolius fusciventris due to sympatric occurrence in TNP (see Rödel & Branch 2002). G = species recorded in southern TNP (Guiroutou. Perret 1988) but not recorded by us. a l l l l l 13 . Trap = species recorded with pitfall and funnel traps. TS = all species recorded on transects in secondary forest. f = fossorial species. TC = all species recorded on transects. TaïF = species recorded in forest habitats around the SRET station. 7°10’ W). 5°25’ N. SRET = species recorded with VES/AES on the clearing around the SRET station. 2 = determination not possible at present time. other than transects. 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. l = leaf litter species. 4 = Acanthixalus sonjae lives aquatically in large water filled tree holes but forages in surrounding vegetation (Rödel et al. 3 = Phrynobatrachus guineensis is a leaf litter frog but reproduces in small water-filled tree holes (Rödel 1998b). TP = all species recorded on transects in primary forest. VES/AES = species recorded via visual and acoustic encounter surveys in forest habitats near SRET. aq = aquatic species. List of amphibian species recorded in Taï National Park (TNP). 1 = known from TNP (Schiøtz 1967. 2 Ranidae Hoplobatrachus occipitalis Amnirana albolabris Amnirana occidentalis Aubria occidentalis Ptychadena pumilio Ptychadena bibroni Ptychadena mascareniensis Ptychadena superciliaris Ptychadena aequiplicata Ptychadena longirostris Ptychadena sp. species Gymnophiona Caecilidae Geotrypetes seraphini Anura Pipidae Silurana tropicalis Bufonidae Bufo regularis Bufo maculatus Bufo togoensis Bufo taiensis Bufo superciliaris Hemisotidae Hemisus sp. 6 = recorded north of area under investigation. 2003).
RÖDEL & ERNST Appendix continued species Phrynobatrachus calcaratus Phrynobatrachus taiensis 1 Phrynobatrachus villiersi Phrynobatrachus annulatus Astylosternidae Astylosternus occidentalis Arthropleptidae Cardioglossa leucomystax Arthroleptis sp. 12 Arthroleptis sp. 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. aq a 14 .